Vol. 63

January 1987

THE

No. 1

Pan-Pacific Entomologist

GRACE, J. K. and D. L. WOOD—Delusory Cleptoparasitosis: Delusions of arthropod

infestation in the home . 1

COHER, E. I.—Asian biting fly studies VI: Records and new species of Oriental Haematopotini

(Diptera: Tabanidae) from Nepal, Thailand, Laos and Cambodia. 5

GORDH, G.—A taxonomic study of Nearctic Meromyzobia Ashmead, 1900 (Hymenoptera:

Encyrtidae) . 16

STARR, C. K., P. J. SCHMIDT, and J. O. SCHMIDT—Nest-site preferences of the giant honey

bee, Apis (Hymenoptera: Apidae), in Borneo. 37

GIESBERT, E. F.—The genus Pachymerola Bates (Coleoptera: Cerambycidae) . 43

LUCKOW, M. and C. D. JOHNSON—New host records of Bruchidae (Coleoptera) from

Desmanthus (Leguminosae) from Texas and Mexico . 48

HOWELL, CHARLES D.—Sex-influenced protibial spines and synonymy in Dasytidae

(Coleoptera), study number three. 50

ADAMES, F. N. S.—A new species oiActaletes from Mexico (Collembola: Actaletidae) . 52

KIMSEY, L. S.—New species of Cleptes Latreille from Asia and North America (Chrysididae,

Hymenoptera) . 56

BUGG, R. L.—Observations on insects associated with a nectar-bearing Chilean tree, Quillaja

saponaria Mohna (Rosaceae). 60

SZCZYTKO, S. W. and R. L. BOTTORF— Cosumnoperla hypocrena, a new genus and species

of western Nearctic Isoperhnae (Plecoptera: Perlodidae) . 65

McELRAVY, E. P. and V. H. RESH—Diversity, seasonahty, and annual variabihty of caddisfly

(Trichoptera) adults from two streams in the Cahfornia Coast Range . 75

HYNES, C. D.—New species of the genus Styringomyia from the South Pacific and Southeast

Asia (Diptera: Tipulidae) . 92

SAN FRANCISCO, CALIFORNIA • 1987

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

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PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 1-4

Delusory Cleptoparasitosis:

Delusions of Arthropod Infestation in the Home

J. Kenneth Grace and David L. Wood

Department of Entomological Sciences, 201 Wellman Hall, University of

California, Berkeley, California 94720.

Abstract. —We describe two cases where individuals believed that their homes

were infected by nonexistent arthropods. These delusions of cleptoparasitosis (nest

parasitism) share many of the more dramatic symptoms associated with imagined

bodily infestation (delusory parasitosis). These symptoms may precede delusory

parasitosis and may be encountered by entomologists and pest control professionals

who are not usually involved with public health problems.

The term “delusory parasitosis” refers to a mental disorder in which an individual

has an unwarranted belief that insects or mites are infesting his or her body

(Waldron, 1962; Ebeling, 1975), i.e., literally a “delusion of parasitosis.” This

condition is distinct from “entomophobia” or an exaggerated fear of real insects and

from “illusions of parasitosis” (Waldron, 1972) where environmental factors such as

dust or static electricity are the true sources of physical discomfort attributed

erroneously to arthropod infestation.

Persons suffering from delusory parasitosis generally believe that their

environment is infested. Items cited by these individuals as sources of the arthropods

thought responsible for bodily infestation include automobiles, pieces of furniture,

and articles of clothing (B. Keh, pers. comm, and 1983).

In a recent review, Keh (1983) presents a thorough discussion of delusory

parasitosis from the point of view of a public health entomologist. However, neither

his review nor other references on the subject describe cases where individuals

imagine arthropod infestations of their living quarters without the accompanying

more dramatic delusion of bodily infestation. Public health entomologists probably

encounter such complainants, but would refer them to a pest control professional

without investigation unless health problems were mentioned (Keh, pers. comm.).

The delusory nature of the complaint would therefore probably not be documented

by pest control personnel or entomologists interested in structural pests or public

health entomologists. In an effort to correct this situation, we describe two cases

which illustrate our extension of the concept of delusory parasitosis to include

phenomena that we believe are best described in new terminology as “delusions of

cleptoparasitosis” (nest parasitism).

Case Histories

The first instance was encountered by JKG in California in 1976 as an employee of

a pest control company. A middle-aged female resident of a condominium complex

receiving monthly pest control service began calling daily, complaining of insect

infestation in her rugs and throughout her unit, and requesting immediate service.

PAN-PACIFIC ENTOMOLOGIST

Technicians responded to her request several times and chemically treated her

condominium unit. Each time, the woman expressed her anger at the supposed

inadequacy of the previous treatments and tearfully revealed details of personal

problems to the technicians. After several weeks, the pest control company refused

to respond to her repeated requests, despite complaints from the concerned manager

of the complex that the company was contractually obligated to respond to all

requests for service, whatever their basis.

The second case, which prompted this report, was encountered by us in California

in 1985. Despite assurances from several pest control companies and private

consultants, the male complainant was absolutely convinced that his home, which

was to go to his wife as part of a separation agreement, was infested by

wood-destroying insects. Old insect tunnels (varnished over) in a picture frame and

in a large cabinet were attributed by him to recent insect tunneling activity, and knots

falling out of a new redwood fence were thought to be eaten from within. A small

decorative box made from an undetermined hardwood also exhibited small tunnels.

This box had been stored in the freezer in an attempt to kill any resident insects.

Because these tunnels did not contain boring materials and had a dark stain

associated with them, we determined that they had been excavated by ambrosia

beetles (probably family Scolytidae or Platypodidae) when the tree was cut.

The tenant also provided us with samples of damaged wood and of insects,

determined to be the common dermestid Anthrenus verbasci (L.). This wood

was a 1 X 4 inch Douglas fir board which had been broken in half. Several of the

fractures (break lines) in the board had been marked for special attention as

suspected insect tunnels. One of the tenant’s friends had taken one of the pieces to

his home to observe further the imagined insect activity in the fracture zone. Wood

technologists determined that the wood had not been subject to decay but exhibited

an unusually large-sized early wood with large diameter tracheids.

Although dermestids and other pests of stored food products can damage wood

when an extensive infestation is present (Grace, 1985), only three specimens were

collected by the homeowner, two of them outside the structure. A. verbasci is a

common detrivore and has not been reported to damage wood.

Discussion

These two cases have several elements in common besides the conviction of the

complainants that their homes were infested by apparently nonexistent insects. Both

individuals were living alone at the time of the complaint, and volunteered details of

recent emotionally disturbing personal problems. The husband of the woman in the

first case had recently separated from her and she complained of having little contact

with her grown son. In the second instance, the male complainant volunteered that

he was involved in upsetting separation proceedings. Emotional trauma, particularly

from marital problems, is reported to be characteristic of delusory parasitosis (cf.

Ebeling, 1975; Keh, 1983).

If there is an emotional basis for the delusion of home infestation, it may be very

difficult for the entomological investigator to play a role in directing afflicted

individuals to sources of psychological counseling. In instances of imagined bodily

invasion, entomologists and pest control professionals are sometimes able to

communicate with the physician(s) that the victim has contacted. Physicians are

certainly in a better position to help these individuals obtain treatment for

VOLUME 63, NUMBER 1

psychological problems. When there has been no medical complaint, the

entomological investigator appears to have little choice but to determine whether or

not arthropods are present. If this conclusion is unacceptable to the affected

individual, he or she may be advised to consult another professional. However, the

complainant likely will resist recommendations for psychological counseling.

In the 1985 case we have described, our protocol was to inspect the premises and

“damaged” wood, and identify the insect samples provided to us. During our

telephone conversations and the inspection by DLW of the complainant’s home, no

mention was made of insects biting or parasitizing him. Several weeks after our initial

contacts with the complainant, he then began to claim that insects were burrowing

into his skin, a typical expression of delusory parasitosis. In an unusual development

(B. Keh and Professor R. S. Lane, pers. comm.), a male friend of the complainant,

who did not live in the house, expressed the same symptoms. We then referred both

men to Professor Lane, a medical entomologist. After being shown samples which

appeared to be skin scrapings, he interviewed both men and determined that no

parasitic arthropods were present. Apparently, these two persons from separate

homes were reinforcing each other’s mental condition and anxiety over the perceived

problem. Finally, we encouraged both complainants to seek medical (not necessarily

psychological) advice for their problems.

Our conclusions in this case were not acceptable to the complainants. They

continued to call and visit us without appointments and their manner towards us

became increasingly demanding and hostile. At the request of campus police, we

refused to engage in further conversation with the complainants and notified them

that any further contact would constitute harassment.

We do not know how often delusions of cleptoparasitosis are encountered by

entomologists and pest control professionals, since published reports involve only

imagined bodily infestations (cf. Keh, 1983). This may reflect the fact that medical

and public health investigators, who are familiar with the syndrome of delusory

parasitosis, are not usually contacted by individuals suffering from the delusion of

infested premises alone. This delusion could represent a different manifestation of

emotional problems similar to those associated with delusions of bodily infestation.

However, our experience in the 1985 case described here suggests that this complaint

may simply precede expression of the more dramatic symptoms of delusory

parasitosis. Contact with the complainant in the 1976 case was interrupted before

further symptoms could be observed.

Acknowledgments

We are grateful to R. P. Akers for establishing contact with the complainant in the

1985 case and providing technical assistance; R. S. Lane for analyzing the complaints

of bodily infestation; R. S. Beal, Jr. (Emeritus, Northern Arizona University,

Flagstaff, Arizona) for dermestid species determinations; and W. A. Dost (Head,

Wood Building Research Center, Richmond, California) for wood examination. We

thank B. A. Barr, L. W. Barclay, M. A. Hoy, W. C. Schaupp, and A. R. Weinhold

for describing their interactions with the complainants. G. W. Frankie, B. Keh and

R. S. Lane reviewed drafts of the manuscript and provided pertinent information.

R. P. Akers, J. W. Fox and L. D. Merrill also read early drafts of the manuscript and

offered helpful suggestions.

PAN-PACIFIC ENTOMOLOGIST

Literature Cited

Ebeling, W. 1975. Urban Entomology. Div. Agr. Sci., Univ. Calif., Los Angeles.

Grace, J. K. 1985. Three beetle families can wreak havoc in wood. Pest Control, 53(9):52,55,57.

Keh, B. 1983. Cryptic arthropod infestations and illusions and delusions of parasitoses. Pp. 165-185 in

G, W. Frankie and C. S. Koehler (eds). Urban entomology: interdisciplinary perspectives.

Praeger, New York.

Waldron, W. G. 1962. The role of the entomologist in delusory parasitosis (entomophobia). Bull.

Entomol. Soc. Am., 8:81-83.

-. 1972. The entomologist and illusions of parasitosis. Calif. Med., 117:76-78.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 5-15

Asian Biting Fly Studies VI:

Records and New Species of Oriental Haematopotini (Diptera:

Tabanidae)

from Nepal, Thailand, Laos and Cambodia

Edward I. Coher

Division of Natural Sciences, Southampton Center of L.LU., Southampton, New

York 11968.

The adult Oriental Haematopotini have been documented by Stone and Philip

(1974) to a point where it would seem little could be added to our knowledge of the

tribe without extensive specialized collecting. Nevertheless, the Oriental Region

continues to yield new records and new species in this unusual complex which has

radiated throughout the Indian and Malayan subregions into an impressive list of

closely related forms, several of which have been implicated as disease vectors. In

these two subregions only two genera are involved, Hippocentrodes Philip, 1961 and

Haematopota Meigen, 1803; the former apparently includes two species, one of

which is amply represented in this study. Based on the study by Stone and Philip,

Haematopota includes 160 species, 5 nomina dubia and two nomina nuda. Studies by

Thompson (1977) and Burger (1981) have added two species to this total.

From observations made in the present study, but requiring further examination,

the generic characterization of Haematopota should include statements on 1: the

form of the spermathecae which are not inflated apically and 2; the structure of the

spermathecal ducts which are much more membranous, longer (may reach into the

third abdominal segment) and more inflated than those of tabanine genera.

The following study of the distribution of 15 species of Oriental Haematopota

includes descriptions of five new species, one each from Cambodia, Laos and

Thailand and two from Nepal.

1. Hippocentrodes desmotes Philip, 1961

1961. Philip, Indian!. Ent.21(2):82-88,f.

1974. Stone and Philip, USDA Tech. Bull. 1489;28,f, figs. l,la,122.

1976. Matsumura and Takahasi, Japanese J. Sanit. Zool.27(3):297,f.

In 1912, Brunetti described a unique male Haematopota from Dehra Dun, India.

The dark wing with six very narrow incomplete hyaline transverse bands was distinct

compared with all other Asian species of that genus.

Philip (1961:82) described a female tabanid with six complete transverse pale wing

bands from Bengal, India. He placed it in a new monotypic genus Hippocentrodes

with the genotype H. desmotes. In 1974, Stone and Philip reviewed the

Haematopotini and transferred the Brunetti species striatipennis to Hippocentrodes.

Matsumura and Takahasi (1976:297) reported a single female of desmotes from

Nepal. Although stating that it is a “small blackish” species, the description given by

them refers to its dark brown appearance and it is particularly interesting to note that

they report no abdominal pattern at all.

During 1956-57, in dense lowland jungle of southern Nepal, I took a series of

PAN-PACIFIC ENTOMOLOGIST

tabanids with the distinctive wing pattern of transverse bands. Specimens were taken

both north and south of the Siwalik Hills. Comparison of this material with the type

of desmotes shows differences in color and setal pattern. The integument of the

Nepalese specimens is darker than the reddish-brown of Philip’s specimen. The

distal margin of the first four abdominal segments of the specimens from Nepal is

fringed with silvery setae. These setae are underlain on the first two tergites by silvery

pollinosity and laterally on the third and fourth segments. By contrast, desmotes has

yellowish setae, reduced pollinose and setal markings on the second tergite and no

markings on the third or fourth abdominal tergites. Philip described other

differences in the head and wing patterns in addition to which I note a size range of

6.5 to 8.5mm.

The question arises as to the relationship of the population from southern Nepal to

striatipennis and desmotes. Based on adult anatomical and pattern characteristics, it

is unlikely that these forms represent three distinct species. Owing to lack of material

from western India, it is not possible at this time to determine whether the type of

desmotes, or my Nepalese material, or both, represent the female of the Brunetti

species. My personal interpretation is that all these specimens could represent a

single population bearing the specific name striatipennis. The fine condition of my

material may be responsible for the apparent variation of color and pattern from the

two described forms. Indeed, Stone and Philip were convinced that one of my

specimens from Nepal was conspecific with desmotes. For the present, I will follow

their conclusion that the Nepal sample represents the westernmost population of

desmotes.

Records. —NEPAL, Amlekhganj, 19 July 1956,f; 8 July 1956,9f(Shannon trap,

500m.); 14 July 1957,f; Chisapani, 4 July 1957,2f (EIC and Ghan Sham).

2. Haematopota abacis Philip, 1960

1960b. Chrysozona abacis Philip, Fed. Malaya Inst. med. Res. St.29:29,f.

1963. Haematopota obscurata Philip, Pacific Insects 5:529,f,m.

1974. Stone and Philip, ibid:30,f,m,synonymy, figs. 50,173,174.

Record. —THAILAND, Chiengmai Province, Doi Sutep, 1000m., 8 August

1959.

This specimen was taken at the wat on Doi Sutep. Its external characteristics agree

well with the description of abacis by Stone and Philip; their notation in regard to the

dorsal margin of the subcallus, “upper margin nearly straight,” is more accurate than

their figure; also, the interanntenal spot is wider than shown in that figure.

3. Haematopota albofasciatipennis Brunetti, 1912

1912. Brunetti, Indian Mus. Rec. 7(5):458,m.

1974. Stone and Philip, ibid:41 ,f,figs.68,194.

1976. Matsumura and Takahasi, ibid:299,f, as H. albofasciatipennis.

A series of twenty-nine female specimens of this species shows the following

differences from the descriptions by Stone and Philip (1974) and Matsumura and

Takahasi (1976): frons of the specimens from north of the Siwalik Hills (Hetaura and

Chisapani) with gray to brownish pollinosity with vertexal area browner, the median

spot present or absent and either set off by light pollinosity or not; with a ring, or

partial ring, or virtually no silvery pollinosity around the lateral frontal spots which

VOLUME 63, NUMBER 1

are subquadrate; basal callus shiny dark brown; face immediately below antennae

slightly darker than remainder and gena. Stone and Philip describe the w/h of the

frons as subequal; my material differs with a w/h of 2 / 2 . Scutellar stripes indistinct.

Foretibia with basal third white; midtibia with two white bands more or less distinctly

divided, the basitarsus white with a narrow dark distal band; hindtibia with at least

one distinct white band, a second poorly defined band also present, basitarsus as for

midleg. Abdominal sternites dark with narrow light posterior margins.

Specimens from Mile 4, north of Amlekhganj and on the southern slope of the

Siwalik Hills, appear even lighter brown than other specimens. Scutal stripes are not

evident or are hardly developed. Basal callus brown to dark red-brown with a dark

median line from the point of the basal callus onto the blackish-gray pollinose frons.

The hindtibial white mark may be reduced to a single distinct basal band or with the

median white band reduced and best developed on the median aspect; the basitarsus

of both the mid and hindlegs is dark apically. Specimens from Amlekhganj show

variation similar to that of the material from Mile 4. The wing of the material from

north of the Siwaliks is proportionally broader and with a more broadly rounded anal

area than that from the southern slope. Limited dissections show variances in the

form of the spermathecae, those from Hetaura appearing much less sclerotized and

with reduced pigmentation. Both forms have medianly enlarged spermathecal ducts.

Records. —NEPAL, Rapti Valley, Hetaura, 520m., 14 April 1956,12f; Chisapani,

4 July 1957,2f. Amlekhganj, 520m., 6 June,f; 14 June,f; 19 June 1956,2f; 5 July

1957(500m.),7f. Mile 4,560m., 5 July 1957,3f. Some specimens were taken by hand

net, but those from Hetaura and two from Amlekhganj were taken in a Shannon trap

and one was taken at a light in Amlekhganj.

4. Haematopota assamensis Ricardo, 1911

1911. Ricardo, Indian Mus. Rec.4(9):343,f.

1974. Stone and Philip, ibid:50,f, figs. 17,138.

Recori/5.—NEPAL, Amlekhganj, 15 April 1956,f; 18 May 1957,f.

The fly taken in April was on a cow in an open pasture area. These records extend

the range of this species 600 miles westward from Assam.

5. Haematopota bealesi Coher, New Species

A unique specimen from Laos not clearly related to any Oriental species known to

me with the exception perhaps of H. tenasserimi Szilady, 1926.

Female. —8mm. Head: Frons (PI. 1) width/height subequal, with a large black

pollinose subtrapezoidal area, emarginate at dorsal margin where apex of vertex

intrudes, less so at ventral margin, vertexal area lighter with short black setae and

some shorter pilosity which produces silvery reflections; callus dark red-brown with a

dome-shaped margin intruding into the frontal pattern; subcallus wide, somewhat

lighter color than the callus, cleft and with narrow lateral projections which are

widened apically; face whitish pollinose with brownish subtriangular areas below

each antenna and with a median brownish longitudinal stripe that reaches their

confluence; parafacials whitish pollinose; beard white; antenna (PI. 1) with scape

narrow and slightly shorter than the flagellomere; apex of style narrowly dark; palpus

light brown with dark setae. Eye: (PI. 1) globose, particularly as compared to other

species. Thorax: scutum dark with black setae dominant on central disc and towards

PAN-PACIFIC ENTOMOLOGIST

PI. 1. Head, showing Irons and antenna of Haematopota bealesi, H. cynthiae, H. excipula, H. gobindai

and H. vimola.

the median anterior margin, golden setae dominant on lateral margins, posteriorly

and onto anterior scutellum which is otherwise dark with black setae. Wing: (PI. 2)

with a broad apical hyaline band in which there is an isolated spot intersecting the

upper branch of the third vein. Halter: stem and knob whitish. Legs: dark brown with

white of lighter markings as follows: basal five-sixth of foretibia whitish; basal half of

mid femur lighter brown-yellow; midtibia with two narrow yellowish rings, one of

which is about one-third the distance from the base, the second about one-third the

distance from the apex; hind femur somewhat lighter; hind tibia with a narrow

sub-basal whitish ring and a similar incomplete ring about two-thirds the distance

from the base. Abdomen: tergites dark brown with very narrow light pollinose

lae

bealesi

indai

VOLUME 63, NUMBER 1

PI.2

bealesi

cynthiae

gahindai

PI. 2. Wings oi Haematopota bealesi, H. cynthiae, H. excipula, H. gobindai andH. vimola.

posterior margins on TII-TVI, TI-TII with grayish lateral pollinosity; SI-SIII and

anterior SIV more reddish brown, SII-SVI with a hint of a lighter posterior margin.

Holotype female. —LAOS, Vientiane-Pak San Rd., 80 miles E., 21 June 1959

(P. F. Beales). In the collection of the California Academy of Sciences.

H. bealesi is most closely related to H. tenasserimi but is easily separable from that

species on the basis of the extent of the pollinose area of its frons and the subequal

w/h of the frons, the proportions of the antennal segments, and the pattern of the

apical hyaline area of the wing.

It gives me great pleasure to name this species for Dr. Peter F. Beales, W.H.O.,

who helped to collect many of the specimens in this study and who obliged me by

collecting material during trips he made to Laos and Cambodia.

PAN-PACIFIC ENTOMOLOGIST

6. Haematopota bicolor Stone and Philip, 1974

1974. Stone and Philip, ibid: 57,f,figs.41,165.

Records. —NEPAL, Amlekhganj, 1956,520ni., 6 June,2f; 14 June,f; 16 June,f; 29

July,2f; 30 July,f; 14 Aug.,5f; 28 Sept.,f. 500m., 8 July,2f; 13 July,3f; 29 Aug.

1957,2f; Baridamar, 6 Aug. 1956,2f.

These lowland forms of bicolor do not appear different from the boreal

populations reported from Assam and northcentral Nepal. However, one specimen

taken 8 August at Amlekhganj is longer by 2mm. and with a wider abdomen than the

remainder of the series; its wing markings in aggregate are the most different of the

entire series, but, its variations appear in other combinations in other specimens.

7. Haematopota cilipes Bigot, 1890

1890. Bigot. Paris Mus. d’Hist. Nat. Nouvelle Arch.(3)2:205,f.

1974. Stone and Philip, ibid:74,f,figs.7,128.

The five specimens taken in southern Thailand not only differ in some details from

the description given by Stone and Philip but also differ from each other. The

principal variations occur as follows: frontal callus with dorsal median point varying

in shape to as much as a broad rectangular area; frontal spots of four specimens more

subquadrate than the figure of Stone and Philip and with at least some light brown

pollinosity marginally; median frontal spot small and light-colored on four

specimens; scutal stripes developed only on the anterior margin of the scutum; scutal

and scutellar setae golden yellow; posterior scutellar crescents fused medially on

some specimens; mid and hind tibial bands not always well-defined and may appear

as a single long fight band; two of the specimens with a narrow median diagonal dark

stripe through the subapical hyaline band of the wing.

Records. —THAILAND, Trang Province(no data),f; Lamor Vill.#3, 8 June

1960, f; Chong, 15 June 1960,3f.

The place of capture for all the specimens with full data, places this as a jungle

inhabiting species.

8. Haematopota cynthiae Coher, New Species

Female. —7mm. Head: Frons (PI. 1) width/height 6/5, clothed with gray

pollinosity, subcircular lateral spots black, separated by half their diameter from the

callus and surrounded by whitish pollinosity with a sparse but noticeable tuft of white

setae below each; vertex gray pollinose, apical vertexal spot with white pollinosity

below and a narrow median white pollinose line above; callus red-brown with a

strongly bowed dorsal margin; subcallus cleft, dark brown; face white pollinose;

parafacials with a small dark comma-shaped lateral mark; beardless; antenna (PI. 1)

with scape/flagellomere 5/6, scape yellowish and slightly thicker than the width of the

widest part of the flagellomere which is brownish and narrow, style dark brown;

palpus dark cream-colored with brownish setae. Thorax: median gray scutellar stripe

short and narrow, sublateral gray stripes longer and broader, disc with short golden

setae intermixed with brown setae; scutellar setae brown; pleura whitish pollinose.

Wing: (PI. 2) much like albofasciatipennis but with apical hyaline band intruding into

the apex of the marginal cell; hind margin broadly hyaline including the anal region,

joined or not to subapical hyaline transverse band. Halter: cream-colored stem and

brown knob. Legs: brown with white markings as follows: foreleg dark brown.

VOLUME 63, NUMBER 1

foretibia with front surface two-thirds white, reduced obliquely to one-third of

lateral surface; midleg light brown with basal four-fifths of tibia white and basitarsus

white except for narrow distal brown ring, tarsal segments showing basal white; hind

leg light brown with basal third of tibia diffuse whitish, basitarsus white with a narrow

distal dark ring, tarsal segments with basal white. Abdomen: TII-TVI with

submedian rows of white pollinose spots within a more diffuse lateral white pollinose

area and with light-colored setae; TIII-TVI with a narrowly lighter posterior margin;

sternites darkening posteriorly, SII-SVI with narrow lighter posterior margins, setae

light-colored.

Holotype female. —NEPAL, 520m., Rapti Valley, Hetaura, 14 April 1956. In the

collection of the California Academy of Sciences.

Paratopotype. —female with the same data.

Both specimens were taken flying with albofasciatipennis. To the eye, they are

more robust and larger and the wings are not as deeply infuscated. The lateral frontal

spots are smaller and rounder and surrounded by silvery pollinosity. The pattern of

the vertexal area is reduced and silvery pollinosity lies below the median spot and

continues as a faint median vertexal line. The scutal pattern of stripes is strongly

developed. TIII-TVI show well-developed submedian light spots. The subapical spot

of the wing is much longer and somewhat broader and the hind margin of the wing is

at least twice as broadly hyaline.

This sibling of albofasciatipennis, flying with that species, is easily separated from

its ‘twin’ by the form of the frontal black spots, the length of the subapical hyaline

band and the wide posterior hyaline band on the posterior margin of the wing. I take

great pleasure in naming this species for my wife, also a twin.

9. Haematopota excipula Coher, New Species

Female. —9.5mm. Head: Frons (PI. 1) width/height 2/1, with brownish to silvery

pollinosity, subcircular lateral black pollinose spots touch or nearly touch callus and

eyes, spots in some light show small light-colored setae around each as well as around

the smaller median dark spot from which there is a median vertexal pollinose line;

vertexal area not clearly defined by pattern; callus dark brown, dorsal margin

horizontal with a small median triangular projection; subcallus dark brown and cleft;

face silvery pollinose with a small black spot below each antennal base; parafacials

with an extensive black pollinose area which touches eyes; beardless; antenna (PI. 1)

with scape about four-fifths as long as flagellomere and narrower than the greatest

width of the flagellomere, light brown; flagellomere brown, narrow, style slightly

darker; palpus cream-colored with brown setae. Thorax: with a narrow median

longitudinal silvery pollinose stripe to center of disc; submedian stripes shorter and

followed by a silvery spot, setae brown with smaller scattered golden setae; posterior

margin of disc silvery pollinose; scutellum silvery pollinose along anterior margin

and median area, fine brown scutellar setae; pleura silvery pollinose. Wing: (PI. 2)

pattern slightly variable in marginal cell, paratype with a broader rosette; in anal

region of paratype, hyaline stripes connected by a longitudinal hyaline stripe. Halter:

with stem and apical third of knob cream-colored, basal two-thirds of knob

brownish. Legs: dark brown, forecoxa with basal half silvery pollinose; forefemur

laterally silvery pollinose; foretibia with basal fourth lighter; midfemur with basal

two-thirds lighter; midtibia with a narrow light ring and two other indistinct rings;

midbasitarsus lighter except at apex; basal two-fifths of hindtibia lighter and a hint of

PAN-PACIFIC ENTOMOLOGIST

subapical light ring; basal portion of hind tarsus lighter. Abdomen: dark brown with

scattered golden setae, TII-TV with posterior margins silvery pollinose, TII with a

median T-shaped pollinose pattern; TIII-TVII with submedian elongated silvery

pollinose spots; sternites dark brown.

Holotype female. —CAMBODIA, Kbal Trach, 12 May 1958 (P. F. Beales). In the

collection of the California Academy of Sciences.

Paratopotype. —slightly damaged female with the same data.

This species is most closely related H. biroi Szilady, 1926 from which it may be

distinguished by its larger size, its well-defined parafacial spots, differences in the

apical hyaline spots of the wing and its lack of median light markings on any but the

second abdominal tergite.

10. Haematopota gobindai Coher, New Species

Although much of this specimen is obscured by fungus, it clearly represents a new

taxon which I take pleasure in naming for a close companion and co-worker for over

two years in the Nepal terai, Gobinda Prasad Joshi.

Female. —8.5mm. Head: Frons (Pl.l) with width/height 2/1 and with large oval

lateral black pollinose spots which do not touch margins of eyes or callus; callus

brownish yellow with a broad dorsal median triangle; subcallus reddish brown;

antenna with scape about four-fifths as long as flagellomere and as wide as widest part

of flagellomere and more yellowish than light brown flagellomere; style dark brown.

Thorax, legs and abdomen cannot be characterized. Wing: (PI. 2).

Holotype female. —NEPAL, Neghauli, 9 April 1957 (G. P. Joshi), at light. In the

collection of the California Academy of Sciences.

This species appears, based on wing characteristics, to be most closely related to

H. punctifera Bigot, 1891. It may be told from that species by its larger size, the w/h

proportion of the frons and the shape and pattern of the wing, particularly the

included dark spot within the wide apical hyaline band and the hyaline areas of all

posterior margin cells.

11. Haematopota howarthi Stone and Philip, 1974

1974. Stone and Philip, ibid: lll,f,m, figs.4,125.

This species has been recorded from Laos.

Records. —LAOS, Vientiane-Pak San Rd., 40 miles E., 21 June 1959,f; same

data, 80milesE.,f, (P. F. Beales).

12. Haematopota pachycera Bigot, 1890

1890. Bigot, Paris Mus. d’Hist. Nat. Nouv. Arch.(3), 2:206,f.

1974. Stone and Phihp, ibid: 155,f, figs.23,147.

Variation in the shape of the flagellomere of the antenna, the vertexal pattern and

its setation, the tomentum and setae of the scutellum as well as the wing pattern may

indicate that there is a complex of species involved rather than the wide variation I

presently attribute to this species.

The flagellomere is variously shaped and may either taper distally or be widened

following a taper. The vertex is somewhat variable in the extent of its pattern and

setation and the scutellar pattern is variable to the extent reported by Stone and

Philip. Although basically similar, the wing pattern is highly variable.

VOLUME 63, NUMBER 1

Records. —THAILAND, Chiengmai Province, 1959, Sarapee, 6 July,f; 30

July,2f; Chompu,27 July,f;NongQuai, 23Sept.,f;TrangProvince,Lamor, Vill. #4,

18 April 1960,f; 10 May 1960,f; Bang Mark, 29 April 1959,f. LAOS, 1959,

Vientiane-Pak San Rd., 21 June, 40 miles E.,f; 21 June, 80 miles E.,f; Pak San-Pak

Dang Rd., 21 June,f.

13. Haematopota singularis Ricardo, 1911

1911. Ricardo, Rec. Indian Mus.4:339,f, fig.

1963. Philip, Pacific Ins.5(3):530,f, fig.; as H. s. vietnamensis.

1974. Stone and Philip, ibid: 176,f,synonymy, figs.9,131.

My specimens show the following variations from those described by Ricardo and

by Stone and Philip. Paired frontal spots large and subquadrate, surrounded by a ring

of lighter pollinosity; median spot with a longitudinal light line pf pollinosity running

onto the vertexal area and also anteriorly on one specimen, plus a broad m-like

marking of light pollinosity reaching to the paired frontal spots; eye margins with

light pollinosity; callus dark brown; apex of scape plus pedicel dark. Thorax with the

light areas and stripes silvery; pleura with two dark bands, one extending from the

wing base to the spiracle, the second extending from the wing base around the

sternites and including the base of the forecoxa. Wing with extensive brown

markings in the oblique stripe of one specimen; halter light at apex. Legs with

midbasitarsus definitely lighter-colored except at tip; hindfemur densely clothed

with dark hairs along the entire length of dorsal and ventral surface; hind basitarsus

quite swollen. Abdomen with TVI and TVII with a pair of submedian light spots,

those on TVII much larger. Spermathecae narrow, pigmented and sclerotized, each

with the terminal portion differently shaped, one strongly and broadly narrowed, a

second narrowed less close to the tip and a third rounded.

Record.—CAMBODIA, KbalTrach, 12 May 1958,3f.

These three specimens, one of which is lacking head and abdomen, are noted as

being taken on carabao. The laterally split abdomen of one is due to engorgement

with blood, a condition not uncommon in other field caught flies. This is the first

record of this species in Cambodia.

14. Haematopota splendens Schuurmans Stekhoven, 1929

1929. Schuurmans Stekhoven, Treubia 6(Suppl.):95,f.

1974. Stone and Philip, ibid:179,f,m, figs.48,171.

Record/.—THAILAND, Pattalung Province, Khouw Pup Pah, 26 June 1960.

Taken flying with vimoli. Dr. Philip has kindly supplied me with a copy of the data

on the Thai collection of this species. Unfortunately, I have no better luck in

deciphering the cryptic data than he did. However, it is interesting to note that the

specimen was taken at 5500' on 7 April, 1939.

15. Haematopota vimoli Coher, New Species

Female. —8.5-9.5mm. Head: Frons (PI. 1) width/height about 2/1, pollinosity

variable, silvery to silvery-black to brownish-black and with brown setae; vertexal

area usually darker; lateral dark spots subcircular and variably surrounded by

lighter-colored pollinosity which forms a line running posteriorly through the median

line of the vertex; callus red-brown, with dorsal margin variable in shape; subcallar

PAN-PACIFIC ENTOMOLOGIST

area black medially and cleft; face medianly black pollinose and large lateral black

pollinose triangles on the parafacials; beard white; antenna (PI. 1) with scape slightly

longer than flagellomere, style brownish; palpus brown with whitish pollinosity.

Thorax: scutum with anterior narrowly silvery, disc and scutellum brown with golden

setae; pleura whitish pollinose. Wing: (PI. 2) much like H. helviventer. Halter: hght

stem and dark knob. Legs: dark brown with white markings as follows, basal third of

foretibia, basal four-fifths of midtibia and barely so on base of basitarsus, basal

two-thirds of hindtibia. Abdomen: TI-TVI dark brown with posterior margin of TII

entirely or partly light-colored and a narrow pollinose triangle projecting two-thirds

of way to anterior margin, clothed with golden setation particularly at posterior

margin of each segment; SI-SVI whitish with a darker median patch on SIII-SVI.

Holotype female. —THAILAND, Trang Province, Chong, 15 June 1960,

Shannon trap. In the collection of the California Academy of Sciences.

Paratopotypes. —All taken in Shannon trap. 1960: 10 May,f; 15 June,7f; 29

June,4f; Paratype: Pattalung Province, Khouw Pup Pah, 26 June 1960,f.

T. vimoli is clearly related to T. spenceri Stone and Philip, 1974 and T. helviv enter

Stone and Philip, 1974 but is clearly separable based on the w/h of the frons which is

wider than high, the presence of large black parafacial spots and the form of the long,

slim flagellomere of the very narrow antenna.

This jungle species is named for Dr. Vimol Notananda of Chiengmai, Thailand,

who so graciously supported my activities in that country.

It is of interest to note that certain species were taken flying at the same time and

place with other tabanids. The following listing is noted:

H. albofasciatipennis with Tabanus adhabarensis and Chrysops dispar on 4 July 1957

north of the Siwalik Hills and with T. albosetosus and T. nepalensis on 5 July 1957

south of the Siwalik Hills, Nepal.

H. bicolor With T. albosetosus on 6 June 1956, with T. jucundus on 16 June 1956, with

T. subcallosus, T. teraiensis, T. jacobarius and T. nepalensis on 30 July 1956, with T.

jacobarius on 8 July 1956 and 29 August 1957, all in Amlekhganj, Nepal.

H. cilipes with T. aurilineatus on 8 June 1960 in Trang Province, Thailand.

H. pachycera with T. konis on 27 July 1959 in Chiengmai Province and with T.

brunipennis on 29 April 1959 in Trang Province, Thailand.

H. vimoli with T. hybridus, T. subhybridus, T. brunnicolor, T. caduceus, T.

griseipalpis, T. macdonaldi, C. dispar and C. fixissimus on 15 June 1960; with H.

splendens on 26 June 1960; with T. hybridus and T. subhybridus on 29 June 1960. All

records from Trang Province, Thailand.

Literature Cited

Bigot, J. M. 1890. Dipteres. IN Pavie. Paris Mus. d’Hist. Nat. Nouv. Arch. (3)2:203-208.

Brunetti, E. 1912. New Oriental Diptera, I. Rec. Indian Mus. 7:445-513, figs.

-. 1924. Diptera of the Siju Cave, Garo Hills, Assam. Indian Mus. Rec. 26(1):99-106.

Burger,!. F. 1981. A review of the horse flies of Sri Lanka (Ceylon). Ent. scandinavica Suppl. 11:81-123,

figs. 1-10.

Chvala, M. 1969. Einige neue oder wenig bekannt Bremsen von Nepal. Acta Ent. Bohemoslavaca

66:39-54,1 pi.

Matsumura T. and H. Takahasi. 1976. Notes on some Nepalese Tabanidae with descriptions of two new

species. Japanese!. Sank. Zool.27:289-300.

VOLUME 63, NUMBER 1

de Meijere, J. Ch. 1911. Studien iiber sudostasiatische Dipteren. VI. Tijd. v. Ent.54;258-432, figs.

Moucha, J. 1969. Tribus Haematopotini. ActaEntomol. Mus. Nat. Pragae 38:213-235.

-. 1970. The Tribe Haematopotini in the Oriental Region. H. D. Srivastava Commem.

Vol.:391-394.

Philip, C. B. 1960a. Three New Tabanine Flies from the Orient. Indian J. Ent. 21:82-88, figs.

-. 1960b; Malaysian Parasites XXXV. Descriptions of some Tabanidae from the Far East. Stud.

Inst. med. Res. Malaya, No.29:1-32.

-. 1960c. Malaysian Parasites XXXVI. A Summary Review and Records of Tabanidae from

Malaya, Borneo and Thailand. Stud. Inst. med. Res. Malaya, No.29:33-78.

-. 1961. Three New Tabanine Flies from the Orient. Indian J. Entomol.21(2):82-88, figs.

-. 1963. Further Notes on Far Eastern Tabanidae III. Records and New Species oiHaematopota and

a New Chrysops from Malaysia. Pacific Insects 5(3);519-534, figs. 1-9.

Ricardo, G. 1906. Notes on the Genus Haematopota of the Family Tabanidae in the British Museum

Collection. Ann. Mag. Nat. Hist. Ser.7,18:94-127.

-. 1908. Descriptions of some New Species of Tabanidae with Notes on some Haematopota. Ann.

Mag. Nat. Hist. Ser.8,1:54-60.

-. 1911. A Revision of the Oriental Species of the Genera of the Family Tabanidae other than

Tabanus. Indian Mus. Rec., 4(9):321-397,401.

-. 1913. New Species of Tabanidae from the Oriental Region. Ann. Mag. Nat. Hist. Ser.8,11:546.

-. 1916. Two New Species of Haematopota from the Federated Malay States. Bull. Ent.

Res.6(4):403-404.

-. 1917. New Species of Haematopota from India. Ann. Mag. Nat. Hist. Ser.8,19:225-226.

Schuurmans Stekhoven, J. H. 1926. The bloodsucking arthropods of the Dutch East Indian Archipelago.

Treubia 6 (Suppl): 1-551, Pis. 1-18.

Senior-White R. 1927. Catalogue of Indian Insects. Pt. 12-Tabanidae. Pp. 1-70. Gov’t. India Publ.,

Calcutta.

Stone, A. and C. B. Philip. 1974. The Oriental Species of the Tribe Haematopotini. U.S.D.A. Tech. Bull.

No. 1489:1-240, figs. 1-249.

Stone, A. 1975. Family Tabanidae. IN Delfinado, M.D. and D.E. Hardy (editors). A Catalog of the

Diptera of the Oriental Region, Vol.2:i-ix, 1-459. Univ. Press Hawaii, Honolulu.

Szilady, Z. 1926. New and Old World Horseflies. Biol. Hungarica l(7):l-30.

Thompson, F. C. 1977. A New Haematopota from Nepal. Proc. Entomol. Soc. Washington 79:19-24,

figs.

Wiedemann, E. R. W. 1828. Aussereuropaische Zweiflugelige Insekten 1:1-608, figs.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 16-36

A Taxonomic Study of Nearctic Meromyzobia Ashmead, 1900

(Hymenoptera: Encyrtidae)

Gordon Gordh

Division of Biological Control, Department of Entomology, University of

California, Riverside, California 92521.

Abstract. —The Nearctic species of Meromyzobia Ashmead, 1900, are reviewed.

A generic diagnosis is provided and some important morphological characters are

discussed. Desantisella Subba Rao is synonymized with Meromyzobia (New

Synonymy). Meromyzobia americana Ashmead, based on a male, and M. flavicincta

Ashmead, based on a female, are synonymized (New Synonymy), flavicincta

recognized as the valid name. A Lectotype is designated for Ericydnus maculipennis

Ashmead, the type-species of Meromyzobia. Descriptive notes are given for M.

flavicincta, M. bifasciata, and M. maculipennis. Four New Species are described (M.

deserticola, M. melanosoma, M. pedicelata, andM. texana).

Introduction

The following taxonomic notes are part of a review of the Nearctic Encyrtidae and

are published here in response to requests for identifications of material by several

workers interested in ecological aspects of insects associated with salt marsh grasses

{Spartina spp.) in Florida. I do not consider this a revision because fewer than 150

specimens are involved. Detailed phylogenetical notes on the relationship of

Meromyzobia to other encyrtids will be published elsewhere (Gordh, in prep.).

Meromyzobia was characterized by Ashmead (1900) for a loosely described

assemblage of species, most of which had been placed in other genera in earlier

publications. Ashmead (1900) recognized six species in Meromyzobia from North

America and these have been carried in subsequent catalogs without taxonomic

study (Muesebeck et al., 1951; Peck, 1963; Gordh, 1979). Elsewhere, species

referable to Meromyzobia have been taken only in South America. DeSantis (1968,

1972) has described M. gripha from a male taken near Buenos Aires, and M. flavipes

from a female taken near Loreto (Missiones). DeSantis’ (1979) Catalog of

Neotropical Chalcidoidea (except Brazil) only includes these two species. Subba

Rao (1971) described Desantisella brasiliensis from two females taken at Nova

Teutonia, Brazil, and D. plaumanni from a male taken at the same locality.

Desantisella is synonymized with Meromyzobia (New Synonymy). Explanation of

this synonymy is given below. Otherwise, treatment of the South American species

awaits collection of more material.

Meromyzobia Diagnosis

Female. —Body moderately large (usually 1.50-3.25 mm. long), sometimes

elongate, sometimes robust. Coloration predominantly pale with weak to moderate

metallic reflections over various parts of the body. Head hypognathous to

VOLUME 63, NUMBER 1

subopisthognathous; in dorsal aspect with anterior margin broadly rounded and

continuous with compound eye margins laterad; vertexal margin broadly rounded,

medial margins of compound eyes parallel, posterior margins not contiguous with

posterior margin of head. Ocelli not close-set but centrally located on vertex; angle

formed by anterior ocellus greater than 90 degrees; lateral ocellus about one

diameter from medial margin of compound eye. Frontovertex moderately wide,

about 0.4-0.5 times as wide as head, weakly reticulate, sometimes shallowly and very

sparsely punctate. Head in frontal aspect transversely oval with toruli closest, near

ventral margin of compound eyes; scrobal impressions very shallow, short, and

poorly developed; malar sulcus complete and well formed. Antennae with scape

cylindrical in cross section or weakly compressed but not expanded ventrad; first two

funicular segments anelliform, distal four funicular segments not modified; club not

large or well differentiated, septa sometimes difficult to distinguish in point mounted

specimens. Mandible with one tooth and a broad truncation or with three equal-sized

teeth. Maxillary palpus four-segmented; labial palpus three-segmented.

Mesosoma with incomplete, short, straight parapsidal sutures; axillae meeting

mesad; propodeum smooth, sloping posteriad and with two medial, longitudinal,

subexocuticular carinae or lines of propodeal reinforcement. Wings macropterous,

frequently conspicuously fuscous, females sometimes brachypterous. Middle tibial

spur large, robust, and frequently enlarged distad.

Metasoma as large or larger than mesosoma; pygostyli near midline to apical

one-third of metasoma, ovipositor and gonostyli usually not strongly exserted,

sometimes exserted up to one-fifth length of metasoma. Seventh sternum from basal

to apical one-third of metasoma.

Male. —Typcally smaller than the female with antennal flagellar segments

enlarged. Body coloration resembles female. Body parts similar in shape to female

(including frontovertex). Males macropterous with hyaline wings. Ocelli not larger

than female ocelli.

Discussion

The correct systematic placement of Meromyzobia remains vague. Ashmead

(1900) summarized early attempts at higher classification of the Encyrtidae and

provided the modern concept upon which encyrtids were classified for more than 50

years. In his schema, Meromyzobia was assigned to the Ectromini, a Tribe

characterized by species with bidentate mandibles, a large (frequently

plowshare-shaped) seventh sternum, and a rather long marginal vein. Meromyzobia

clearly does not possess the essential features of the Ectromini. Mercet (1921)

abandoned the Tribal classification of Ashmead in favor of Subfamilies (the

Encyrtinae and Arrhenophaginae with 12 “groups” in the former), but did not place

Meromyzobia because it did not occur on the Iberian Peninsula. The genus was not

treated by Erdos and Novicky (1955) for similar reasons. Hoffer (1955) provided a

formal attempt at reorganizing encyrtid genera into Tribes. He did not consider

Meromyzobia, but his reappraisal of the Mirini Ashmead has been shown to have

merit and it is here that Meromyzobia belongs. Trjapitzin (1973A, B) reorganized

the Encyrtidae and modified the concept of Subtribes within the group (corrected to

Miraini). Meromyzobia was placed in the Subtribe Mayridiina by Trjapitzin and

Gordh (1978), but was listed Incertae Sedis by Gordh (1979). Placement by

Trjapitzin and Gordh is taken as correct pending further study.

PAN-PACIFIC ENTOMOLOGIST

I have not had the opportunity to study the holotype of Desantisella brasiliensis

Subba Rao, which should be in the British Museum. However, the extensive

description of that species and diagnosis by Subba Rao (1971) leave no doubt that

Desantisella is synonymous with Meromyzobia. The shape of the middle tibial spur

clearly fits within the concept of Meromyzobia as understood here. Other characters

regarded as diagnostic for Desantisella are within the range of variation expressed by

Meromyzobia.

Biologically, Meromyzobia appears diverse and does not fit the pattern of most

Miraini which, according to Trjapitzin (1973B), are primary and secondary parasites

of coccoids. Tachikawa (1978) reports Meromyzobia as parasitic on undetermined

Orthoptera eggs. Meromyzobia pedicelata New Species was undoubtedly reared

from Orthoptera eggs. The host plant was Tripsacum laxum Nash (Gramineae).

Meromyzobia melanosoma New Species has been repeatedly taken in association

with Spartina spp. (Gramineae) along the eastern seaboard of the United States.

Meromyzobia maculipennis has been taken from the puparia of Anthracophaga

ingrata (Williston), a chloropid. This fly was described from material taken by F. M.

Webster, who also collected the parasite. The host plant for the chloropid was

Muhlenbergia mexicana (L.) Trin. (Williston 1983). Meromyzobia flavicincta was

taken from unidentified galls on Aristida gyrans Chapman (Gramineae).

Meromyzobia deserticola New Species has been taken from Hilaria rigida (Thurb.)

(Gramineae). Meromyzobia texana New Species has been reared in association with

Antonina graminis (Maskell), an introduced pest of pasture, turf,

and lawn grasses. Subba Rao (1971) does not mention the biology of Desantisella,

but Noyes (1980) indicates its species are hyperparasites of Chamaemyiidae

attacking Aclerididae. Thus, overwhelming circumstantial evidence suggests that

Meromyzobia should be considered a New World genus of Miraini with an ecological

preference for grasses.

Morphologically, Meromyzobia has several interesting features which may not be

appreciated from the following taxonomic treatment, but which have importance in

higher classification of this large and difficult group. Thus, sexual dimorphism should

be studied carefully because it will prove helpful in understanding relationships.

Most groups of encyrtids express dimorphism in antennal conformation, ocellar size

and triangle shape, frontovertex size and shape, body size and coloration, and wing

size and coloration. Species of Meromyzobia express very weak sexual dimorphism

in head and antennal characters. The curious reduction of FI and FII of the antenna is

a synapomorphy shared by males and females. Additionally, the vestigal ring

segments are evident in several species (Figs. 4, 22, 35). Also of note is the lack of

sexual dimorphism in ocellar size or arrangement.

Meromyzobia may be added to the number of genera in which mandible dentition

is not constant. Most species are tridentate but the type-species and several others

have one tooth and a broad truncation. The functional significance of this character

and its character states must be considered before the character is considered of more

than specific importance. I suspect that tooth shape and number is correlated with

the context in which development occurs. The mandible shape of one tooth and a

broad truncation is functionally adapted to facilitate the parasite emerging from

galls. Gall wall architecture differs among plants but basically consists of a matrix of

fibers. I hypothesize that the anterior, short, mandibular tooth grasps the gall fibers

and loosens them from their position in the matrix. Subsequently, the conspicuous

VOLUME 63, NUMBER 1

Figures 1-6. 1. Meromyzobia maculipennis, female head, frontal aspect. 2. M. maculipennis,

female antenna, lateral aspect. 3. M. maculipennis, female middle tibial spur and basitarsus. 4. M.

maculipennis, female antennal pedicel, anellus (arrow), Funicular I-III. 5. M. maculipennis, female

middle tibial spur, medial aspect. 6. M. maculipennis, female middle tibial spur setae on medial surface.

truncations (which form a broad line of contact when mandibles are opposed) engage

the loose fibers which are then pulled free of the gall wall. A line of contact formed by

the truncations is functionally more efficient for engaging the loose fibers than

mandibular teeth in the form of conical projections. The line or truncated surface of

contact is nearest to the liberated gall fibers and engages long and short fibers with

equal facility. In comparison, conical mandibular teeth are less efficient because they

contact gall material only near the apex of the teeth and therefore can engage short

fibers over a limited portion of the potential surface area available for fiber

engagement. A partial explanation for tridentate and truncate mandibles in

PAN-PACIFIC ENTOMOLOGIST

Meromyzobia may lie in host associations and the matrix being processed by the

mandibles in emerging from pupal containment.

Parapsidal sutures are present in all species but constitute a plesiomorphous

character. Female brachyptery is seen in more than one species but males of all

species for which they are known are macropterous. The macropterous female wing

is sometimes infumated, but the male wing is invariably hyaline. This dimorphism

appears to be a plesiomorphy. In at least one species we see associated with the wing

development a concomittant reorganization of thoracic sclerites consistent with a

non-ecologically proximate wing reduction condition. (That is, wing reduction in

encyrtids can be in response to at least two types of conditions. Environmentally

induced brachyptery is a response to immediate conditions and seen in genera such as

Cheiloneurus where individuals of a species may be macropterous or brachypterous

depending on the time of the year, host, or both. Non-environmentally mandated

brachyptery is manifested in species which undergo radical reorganization of

thoracic sclerites to include elongation and enlargement of the pronotum at the

expense of the mesoscutum and scutellum. The different conditions imply different

levels of genetic control and complexity, and therefore should be evaluated as

different characters, not character states.)

The middle tibial spur is perhaps the most important structural character used in

defining Meromyzobia. Within the genus the character ranges from typically

encyrtid to balloon-like and inflated distally. The extreme condition seems best

expressed in South American forms. In North American Meromyzobia, the spur

holds different forms of cuticular ornamentation. In all species examined, the outer

surface appears pubescent. Light microscopy shows this pubescence as seta-like.

However, SEM shows the pubescence is composed of trichode-shaped acanthae, not

the characteristic trichogen-tormogen formation (Figs. 12, 13). In contrast, the

medial surface of all species studied have the classic seta-in-a-socket (Fig. 37). The

seta is modified in all species studied, but owing to the nature of the material

examined (old) and its availability (borrowed and in limited numbers), a thorough

study could not be made. Differences indicated under each species are taken as

species specific. The setae range from short and rather straight (Fig. 6) to long and

curved (Figs. 15,31). Most species display a seta which is longitudinally depressed or

spoon-shaped along one face (Figs. 6,15, 31) with the apical surface tined (Figs. 6,

15, 31). At least one species displays setae which are hyphae-like and not

spoon-shaped or apically tined (Fig. 37). In other respects, the morphology of

Meromyzobia is not remarkable.

A Key to the Nearctic Species of Meromyzobia Ashmead

1 (A). Males . 2

(B). Females .. 8

2 (A). Mandible with one tooth and a broad truncation; body robust; middle

tibial spur distally lobate; scutellum elongate, nearly three times as

long as median length of propodeum; subexocuticular prdpodeal

carinae widely separated and parallel . .Meromyzobia maculipennis

(Ashmead)

(B). Mandible with three teeth or other characters variable or not in the

combination above . 3

VOLUME 63, NUMBER 1

3 (A). Body not strongly dorsoventrally flattened and/or body predominantly

pale colored or reddish brown . 4

(B). Body rather conspicuously dorsoventrally flattened; body

predominantly dark colored with some weak metallic reflections ... 5

4 (A). Parapsidal sutures transverse (Fig. 33), not oblique to primary axis of

body; head viewed in dorsal aspect with posterior margins of

compound eyes nearly contiguous with posterior margin of head at

one point; propodeum pale ... Meromyzohia texana (New Species)

(B). Parapsidal sutures not transverse (Fig. 8) oblique to primary axis of

body; posterior margin of compound eyes separated from posterior

margin of head by at least two ocellar diameters; propodeum reddish

brown. Meromyzohia flava (Ashmead)

5 (A). Body large (ca. 3.5 mm); femora and tibiae all dary ... Meromyzohia

melanosoma (New Species)

(B). Body relatively small (ca. 2.5 mm); femora and tibiae not all dark .... 6

6(A). Pronotum about as long as median length of mesoscutum; head

subopisthognathous . Meromyzohia deserticola (New Species)

(B). Pronotum shorter than median length of mesoscutum; head

hypognathous. 7

7(A). Frontovertex < 0.50 times as wide as head; with shallow setigerous

punctures; body reddish brown . Meromyzohia flavicincta

(Ashmead)

(B). Frontovertex >0.50 times as wide as head; without shallow setigerous

punctures; body predominantly pale colored .. Meromyzohia flava

(Ashmead)

8 (A). Metasoma in lateral aspect with gonostyli and ovipositor projecting well

beyond apex of metasoma, at least 0.5 times length of middle tibial

spur. 9

(B). Metasoma in lateral aspect with gonostyli and ovipositor considerably

shorter, not projecting conspicuously beyond apex of metasoma

(Note: some distortion may give the impression of weak exsertion,

but genostylus never as long as 0.5 times middle tibial spur length) . 10

9 (A). Metasoma slightly shorter than mesosoma; forewing with one fuscous

sport posterior of marginal and stigmal veins; axilla yellow or pale

colored; propodeum brownish with subexocuticular longitudinal

carinae evident and parallel; mesopleuron pale. Meromyzohia

unifasciata (Ashmead)

(B). Metasoma clearly longer than mesosoma (Figs. 17, 18); forewing with

two fuscous clouds separated by a hyaline stripe; propodeum nearly

black or if somewhat more pale and subexocuticular carinae visible,

then they diverge posteriad; mesopleuron dark brown or black ....

Meromyzohia melanosoma (New Species)

10(A). Mandible with one tooth and broad truncation; macropterous;

subexocuticular propodeal carinae parallel or nearly so . 11

(B). Mandible tridentate; macropterous or brachypterous; subcuticular

propodeal carinae diverging posteriad . 13

11 (A). Pedicel elongate; propodeum with elevated median cuticular carina;

pronotum less than 0.35 times medial length of mesoscutum.

Meromyzohia pedicellata (New Species)

PAN-PACIFIC ENTOMOLOGIST

(B). Pedicel normal; propodeum medially smooth, without carinae;

pronotum at least 0.40 times as long as median length of mesoscutum 12

12 (A). Pronotum 0.5 times as long as medial length of mesoscutum; scutellum

about 1.10 times longer than wide; frontovertex 0.45 times as wide as

head . Meromyzobia bifasciata (Ashmead)

(B). Pronotum about 0.45 times medial length of mesoscutum; scutellum

about 1.25 times longer than wide; fronto vertex 0.42 times as wide as

head . Meromyzobia maculipennis

13 (A). Macropterous; forewing weakly infuscated beneath marginal vein;

parapsidal sutures transverse (Fig. 33), not oblique to primary axis of

body; propodeum with incomplete, weak, median longitudinal

Carina . Meromyzobia texana (New Species)

(B). Brachypterous; forewing infuscation obscure; parapsidal sutures

oblique to primary axis of body; propodeum medially polished and

without longitudinal carina. 14

14 (A). Body predominantly pale colored; pronotal median length considerably

longer than mesoscutum, but posterior margin transverse or nearly

so, clearly not forming a broad, inverted “V”; mesosoma flattened;

gonostyli concealed or not exserted.

. Meromyzobia deserticola (New Species)

(B). Body predominantly dark reddish-brown; pronotal median length

shorter than mesoscutum with posterior margin forming a very

broad, inverted “V”; mesosoma not conspicuously flattened;

genostyli slightly exserted .... Meromyzobia flavicincta (Ashmead)

The following comments are included under names previously recognized as valid.

Meromyzobia americana (Ashmead). 1888. Entomol. Amer. 41:16. Male.

This species was based on a male specimen taken in Florida. It has not been

recovered or reported (except catalog entries) since its description. The original

description is misleading because it states that the scutellum of the holotype is “large,

highly convex and finely grooved. ” In fact, the scutellum is of normal size for a male,

not robust, and narrowly and longitudinally reticulate along postero-medial 0.60

with the pattern larger elsewhere; apex polished. Similarly, the pedicel is not

unusually small, but rather the flagellar segments are disproportionately large.

Otherwise, the description is accurate. Additional characters include: Frontovertex

0.48 times as wide as head; surface reticulate with scattered, shallow, setigerous

punctations. Middle tibial spur not enlarged distad. Propodeum with two distantly

separated, longitudinal carinae.

Comparison of the holotype with the type of M. flava suggests that they are similar,

although colored very differently. The specimens resemble one another in the shape

of the head, conflguration of the antenna, scrobal cavity, size and shape of the

mesopleuron. Both bear a non-dilated middle tibial spur. The frontovertex:

headwidth ratio is 0.53 for M. flava, and scuteller sculpture patterns are different.

Synonymy is not implemented here because too few specimens are available for

study, and the holotypes are not in perfect condition for comparison.

I feel confident that the male described as americana is conspecific with the female

described sls flavicincta (New Synonymy). Both were taken from the same locality

VOLUME 63, NUMBER 1

Figures 7-15. 7. M. flavicincta, female head, frontal aspect. 8. M. flavicincta, female mesosoma

(arrow to parapsidal suture). 9. M. flavicincta, female mesosoma and metasoma. 10. M. flavicincta,

female antenna, lateral aspect. 11. M. flavicincta, female antennal pedicel, anellus, Funicular I-IV.

12. M. flavicincta, female middle tibial spur and basitarsus, lateral aspect. 13. M. flavicincta, female

middle tibial spur acanthae. 14. M. flavicincta, female middle tibial spur, medial aspect. 15. M.

flavicincta, female middle tibial spur setae on medial surface.

PAN-PACIFIC ENTOMOLOGIST

and described in the same publication. The dimorphism expressed in this species is

identical to that found in M. deserticola New Species, including female aptery and

male macroptery. As first revisor, I select Meromyzobiaflavicincta as the valid name

because the type is based on the female sex and I regard this sex as more important in

encyrtid classification.

Meromyzobia bifasciata (Ashmead). 1890. Bui. Colo. Biol. Assoc. 1:28,46. Female.

This species was based on one female taken at West Mountain Valley, Colorado,

by T. D. A. Cockerell and originally placed in Homalotylus. The point-mounted

holotype stands in the USNM collection (Type-number 4720), and lacks the club of

one antenna and all flagellar segments of the other antenna. The description was

based primarily on coloration, the precise nature of which cannot now be confirmed.

Two points in error are corrected here; the body length is 2.32 mm, not 2.20 as

indicated by Ashmead, and the frontovertex is not closely punctate with a few larger

punctures but rather minutely reticulate with scattered, shallow, setigerous

punctures. Characters here considered important, but not included in the original

description include: frontovertex 0.45 times as wide as head, mesosoma robust,

pronotum 0.5 times as long as medial length of mesoscutum, posterior margin

forming broad, inverted “V”; scutellum about 1.10 times wider than long, posterior

0.30 polished and flattened. Forewing macropterous with extensive infuscation.

Middle tibial spur lobate distad. Metasoma with gonostyli and ovipositor very

slightly exserted, pygostyli and posterior margin of seventh sternum near midline of

metasoma.

I am not convinced that bifasciata is distinct from maculipennis as I can find no

reliable structural characters to differentiate them. The former is represented in the

USNM collection by the holotype only; the latter is represented by the “type” and

fewer than ten specimens identified by earlier workers. This material is in poor

condition and many characters are difficult to observe or measure. They are probably

synonyms, but synonymizing them seems more appropriate when more and better

curated material is available for study. Characters used to separate females in the key

may be an artifact of small sample size and poor preservation.

Meromyzobia deserticola, New Species

Female. —1.86 mm long. Body predominantly pale yellow with following parts

darker: posterior 2/3 of mesopleuron, lateral 2/4 propodeum, metasomal terga IV-V

reddish brown. Antennal scape concolorous with head, remaining segments

uniformly darker. Basal 2/3 tegula white, remainder dusky, prepectus nearly

transparent. Fore and hind coxae yellow, hind coxa with vestiture of conspicuous

white setae. Middle coxa dusky; pretarsi dusky. Hind femur reddish brown, basal 1/5

tibia white, remainder reddish brown; pretarsi dusky, remaining tarsomeres nearly

white. Wings hyaline.

Head subopisthognathous, in dorsal aspect with frontovertex 0.46 times as wide as

head, 0.75 times as long as head. Ocelli small, forming a very broad, obtuse triangle

with lateral ocellus about one diameter from medial margin of compound eye.

Posterior margin of head very broadly rounded; head in frontal aspect 1.27 times

wider than long; vertex weakly arched above imaginary line continuous between

lateral margins of compound eyes. Ventral margin of head broadly arched. Head in

lateral aspect 1.7 times longer than wide. Compound eye with minute setae.

VOLUME 63, NUMBER 1

Figures 16-24.16. M. melanosoma New Species. Female head, frontal aspect. 17. M. melanosoma,

female mesosoma. 18. M. melanosoma, female metasoma. 19. M. melanosoma, male antenna, medial

aspect. 20. M. melanosoma, female antenna, medial aspect. 21. M. melanosoma, male antennal

pedicel. Funicular I-III. 22. M. melanosoma, female antennal pedicel, anellus (arrow). Funicular I-III.

23. M. melanosoma, female middle tibial spur and basitarsus, medial aspect. 24. M. melanosoma,

middle tibial spur setae on medial surface.

PAN-PACIFIC ENTOMOLOGIST

posterior margin of eye diverging from posterior margin of head ventrad. Malar

sulcus complete and conspicuous. Entire surface of head minutely reticulate with

moderate vestiture of pale, short setae. Antenna as illustrated (Fig. 39). Mandible

tridentate.

Thoracic notum flattened. Pronotum campanulate, as wide as long; mesoscutum

0.5 times as long as pronotum, 0.83 times as long as scutellum. Parapsidal sutures

difficult to discern, but short, straight, oblique to primary axis of body and

subparallel to lateral margin of mesoscutum. Propodeum 0.65 times as long as

scutellum, median portion with two subparallel, subsurface carinae which diverge

posteriad. Mesopleuron smooth, 2.82 times longer than wide. Middle tibial spur

enlarged but apically tapered, not pointed. Wings micropterous.

Metasoma 1.15 times as long as mesosoma; pygostyli at apical 0.64 of metasoma.

Ovipositor and gonostyli not exserted; gonostyli broad and apically rounded.

Seventh sternum at basal 0.33 of metasoma; posterior margin medially incised.

Tergum I as long as following three terga combined, reticulate; sculptural pattern

less pronounced on terga II-VII; all terga with moderate vestiture of conspicuous

pale setae.

Male. —1.44 mm long. Habitus as female, differing in that posterior aspect of head

reddish brown, anterior aspect more pale; axillae black, remainder of mesosoma and

metasoma reddish brown. Antenna concolorous with forecoxa; hind femur dusky;

fore and middle femora and tibiae yellow; basal half of hind tibia nearly white; apical

half nearly yellow. Tarsomeres 1-3 white, tarsomere 4, pretarsi dusky. Wings

hyaline, macropterous. Ocelli larger than female. Antenna as illustrated (Fig. 38).

Wings macropterous (Fig. 40), projecting beyond apex of metasoma. Thoracic notal

sculpture not as bold as female.

Described from six females and three males taken at Seeley, CA, 25 March 1965

on Hilaria rigida (Thurb.) (Gramineae) by R. A. Flock and J. Pineda. Holotype

female, allotype male, and paratypes deposited in USNM collection.

Meromyzobia deserticola most nearly resembles M. flavicincta and may be

distinguished from that and other species based on characters given in the key.

Meromyzobia flava Ashmead, 1900. Proc. U.S. Natl. Mus. 22:350. Male.

According to the original description and USNM type-catalog, this species was

based on a male taken in the District of Columbia. It has not been recovered

elsewhere or at a later time. The holotype is point mounted and stands in the USNM

collection (Type-number 4723) and bears a label reading “Arlington, VA.” The

original description is not particularly accurate or informative. Study of the specimen

shows that it is morphologically similar to M. americana. Characters incorrect or not

included in the original description include body length 1.46 mm, head yellow with

three darker spots each anterior of an ocellus, forewing slightly infuscated beneath

marginal vein. Frontovertex 0.53 times as wide as head, without scattered setigerous

punctures.

Meromyzobia flavicincta (Ashmead), 1888. Entomol. Amer. 4:17. Female.

This species was described from one female taken at Jacksonville, Florida. The

intact holotype is mounted on a card and deposited in the USNM collection. The

brief description is not particularly informative or accurate and descriptive notes are

provided here. The specimen is 1.90 mm long (0.75 in, not 0.80 as stated in the

VOLUME 63, NUMBER 1

original description). The coloration is not as the description but the life-like color

cannot be accurately given owing to the age of the specimen. Based on a specimen I

identify as M. flavicincta, the head, pronotum, and mesoscutum appear tan, antenna

somewhat darker; axilla, scutellum, propodeum, mesopleuron dark reddish brown;

metasoma predominantly dark with pale yellow with transverse band near base.

Coxae, tibia, femora dusky; tarsomeres white, pretarsus dusky.

Head as shown (Fig. 7); antenna (Figs. 10, 11) not particularly short or slender.

Frontovertex 0.48 times as wide as head, reticulate with several shallow setigerous

punctures. Pronotum 0.75 times as long as medial length of mesoscutum; posterior

margin forming broad, inverted “V.” Parapsidal sutures distinct, but incomplete

(Fig. 8). Wings brachypterous but projecting slightly beyond posterior margin of

propodeum; distal margin of wing weakly infuscated. Propodeum with subcuticular

carinae diverging posteriad. Middle spur distally lobate. Ovipositor and gonostyli

very slightly exserted beyond apex of metasoma (Fig. 9). Mesopleuron 1.66 times

longer than wide; surface very weakly reticulate. Middle tibial spur (Figs. 12-15)

enlarged distad; pubescence on outer surface composed of elongate acanthae (Figs.

12,13); medial-surface setae long, curved, apically tined (Figs. 14,15).

Although the female is brachypterous, I consider it conspecific with the male

described by Ashmead as Prionomastix americana as noted above. The shape of the

head in male and female is similar and the relative length of the pronotum to the

other components of the thorax are identical. The propodeum is the same size,

shape, and has two subcuticular, longitudinal carinae which diverge posteriad. The

middle tibial spur in both sexes is large, robust, but apically pointed in the male and

enlarged in the female. The parapsidal sutures are similarly developed in the male

and female. Both specimens were taken at Jacksonville, Florida, probably about the

same time as judged from the identically printed locality labels and curiously

constructed microscopic card mounts. Neither specimen pin carries supplemental

collection information. The USNM type-catalog number 4721 indicates that

specimen came from the Ashmead collection and was taken from Florida.

Type-catalog number 4719 carries the entry for americana. It also reports that

specimen is from the Ashmead collection, taken from Florida without a specified

locality, and erroneously logs the sex of the specimen as a male. Nevertheless, the

synonymy proposed here is taken as correct and the male is nearly identical with the

male described asflava, but synonymy is not implemented. Five female specimens

taken at Miami, Florida, from gall on Aristida gyrans also stood in the USNM

collection appear conspecific with the holotype.

Meromyzobia maculipennis (Ashmead), 1893. Bui. Ohio Agr. Exp. Sta., Tech. Ser.

1:162-163.

Understanding the concept of this species is important for several reasons.

Originally described by Ashmead as Ericydnus maculipennis, it was based on several

specimens which Ashmead considered as males. The type-series was of

indeterminant size and reared from Chlorops ingratus Williston (= Anthracophaga

ingrata) by F. M. Webster in Ohio. A holotype was not designated. In the USNM

collection is one specimen in the type-collection with labels which read: “Columbus

O,” “F. M. Webster,” “Type 4722 USNM” and “Ericydnus maculipennis Ashm.”

The last label carries the word “Ashm.” twice and a male symbol which has been

replaced with a female symbol. This label is in Ashmead’s handwriting whereas the

PAN-PACIFIC ENTOMOLOGIST

other three labels are typeset. The specimen on the pin is a male. Four other

specimens in the collection carry handwritten labels “5272° Par: on Chlorops.” At

least two of these specimens are females. The number on the latter three pins is a

Webster Number, but that card entry is missing from the Webster Number Catalog in

the USNM. The USNM type catalog entry indicates that three specimens were

received from Webster, which with published information and information on the

collection labels suggests that the card mounts with the handwritten labels are part of

the type-series used by Ashmead to describe the species. Study of all four specimens

shows them to be conspecific. To comphcate matters a microscope slide with

fragments is labeled “Meromyzobia maculipennis Ashmead S ? types” in the

handwriting of A. A. Girault. Under the circumstances the specimen in the

type-collection has been remounted on a larger card (owing to its precarious

position) with a leg and forewing remaining on the original point. It has been labeled

LECTOTYPE. The remaining specimens have been labeled as Meromyzobia

maculipennis, but not designated paralectotypes.

The original description of maculipennis is misleading in many respects.

Specimens range in size from 2.30-2.65 mm in length, not 3.0-3.1 as reported by

Ashmead. Antennal segments (Figs. 2,4) are not subfiliform as noted in the original

description, but rather females bear the characteristic two anelliform funicular

segments and the male apparently does as well, although this cannot be confirmed

from the specimens available for study. Other characters not mentioned by Ashmead

in the original description, but important in recognizing this species include:

mandible with one tooth and a broad truncation; head (Fig. 1) with frontovertex 0.42

times as wide as head, reticulate with a few scattered, shallow, setigerous punctures.

Pronotum 0.43 times as long as medial length of scutellum; posterior margin forming

a broad, inverted “V” of an angle more than 90 degrees. Scutellum 1.25 times longer

than wide, predominantly reticulate with apical 0.30 polished and flattened. Middle

tibial spur (Figs. 3, 5, 6) large with setae on medial surface short, straight, apically

tined. Ovipositor and gonostyli not exserted or very slightly exserted (due to

distortion of metasoma after death). Stylus less than half as long as middle tibial spur.

Pygostyli and posterior margin of seventh sternum near an imaginary transverse line

bisecting metasoma.

Subba Rao (1971) apparently had not seen Meromyzobia maculipennis when he

characterized Desantisella. This genus was proposed for two South American

species, brasilensis and plaumanni. Desantisella is coincident with Meromyzobia in

all salient features, including head shape, antennal configuration, wing coloration,

shape, and venation, and size of the middle tibial spur. Other characters, including

the three toothed mandible, fall within the scope of variation expressed for other

species included in the genus. Desantisella here is recognized as a junior synonym of

Meromyzobia (New Synonymy).

Meromyzobia melanosoma, New Species

Female. —3.72 mm long. Body elongate (Figs. 17, 18) macropterous with wing

projecting near apex of metasoma but not beyond distal portion of gonostyli. Body

predominantly dark brown to black. Head reddish brown, pronotum dusky;

anteromedial portion of mesoscutum dark brown, remainder yellow; anterior half of

tegula yellow, posterior half dusky; axillae, scutellum, propodeum black;

mesopleuron dark brown; metasomal tergum I anterolaterally dusky, remainder

VOLUME 63, NUMBER 1

weakly yellow; remainder of metasoma brown; gonostyli contrastingly pale brown.

Antenna somewhat darker than head. Fore wing predominantly fuscous with clouds

over wing blade. Coxae, femora, tibiae concolorous with mesopleuron; fore

tarsomeres brown; middle tibial spur, tarsomeres 1-3 of middle and hind leg white;

tarsomere 4 and pretarsus of middle and hind leg dusky.

Head hypoganthous, in dorsal aspect with frontovertex 0.42 times as wide as head

and 0.85 times medial head length; frontovertex minutely reticulate, setigerous

punctures very shallow and sparse, nearly absent. Ocelli forming a large triangle

whose anterior angle exceeds 90 degrees; lateral ocellus less than one diameter from

medial margin of compound eye, about four diameters from occipital margin.

Occipital margin broadly rounded. Head in lateral aspect about 0.51 times taller than

wide; malar sulcus conspicuous and complete; compound eye with scattered minute

setae, posterior margin diverging from posterior margin of head ventrad. Head in

frontal aspect (Fig. 16) 1.06 wider than long, with toruli close-set, separated by less

than torular width; interantennal prominence weak and broadly rounded; scrobal

impression weakly developed, nearly absent (Fig. 16). Mandible tridentate.

Maxillary palpus four-segmented; labial palpus three segmented. Antenna as

illustrated (Figs. 20, 22).

Mesosoma rather elongate (Fig. 17) but shorter than metasoma (Fig. 18).

Pronotum weakly, minutely reticulate; with moderate vestiture of darkened setae;

medial length 0.31 times as long as medial length of mesoscutum; posterior margin

forming an angle of about 110 degrees. Mesoscutum with reticulate polygonal

sculptural pattern somewhat larger than pronotal pattern but similar vestiture of

setae; parapsidal sutures incomplete but converging posteriad. Axillae reticulate,

broadly joined mesad; mesoscutellum 1.22 times longer than wide, predominantly

reticulate with apex polished; mesopleuron 2.13 times longer than wide;

predominantly reticulate but pattern minutely and longitudinal reticulate anteriad

and expanding in size and diminishing in boldness posteriad; posterodorsal margin

polished. Coxae with conspicuous vestiture of long pale setae on the ventral-facing

surface. Middle tibial spur shorter than basitarsus (Fig. 23) with rather long, curved,

apically tined and compressed setae along medial surface (Fig. 24).

Metasoma lanceolate (Fig. 18), 1.60 times longer than mesosoma. Terga weakly

reticulate. Syntergum apically pointed and projecting over base of gonostyli.

Gonostyli slightly shorter than middle tibial spur (0.92), apically broadly rounded.

Pygostyli near midline of metasoma. Apical sternum terminating near basal third of

metasoma.

Male. —2.32 mm long. Similar to female habitus, coloration, and sculpture;

differing in the following features: frontovertex 0.28 times as wide as head; 0.92

times as head medial width. Antenna as illustrated (Figs. 19, 21).

Material Examined. —Holotype, Female. FLORIDA, Levy Co., 1 Female

16. hi.75; 1 FE 30.hi.75; 1 male, 1 female 25.iv.1975; 1 male 13.iv.1975; 1 female

23.xi.1975; 2 females, 2 males 9.xi.l975; 8 females, 1 male 7.xii.l975; 4 females, 1

male 11.i.1976; 1 female, 1 male 8.ii.l976; 2 males 16.ii.l976, all specimens taken on

Spartina alterniflora Lois. (E. E. Grissell). GEORGIA, Sapela Island: 1 female

17. V.1963 (E. P. Odum), 1 female, 3 males on Spartina (H. Kale), 3 males 9.x.1963

on Spartina (E.P. Odum), on Spartina. NORTH CAROLINA, Cartaret Co.,

Newport R., 1 female 14.xi.l959 (L. V. Davis) on Spartina alterniflora; 3 females

15.i.1960 on Spartina alterniflora (L. V. Davis) NEW JERSEY, Atlantic Co.,

PAN-PACIFIC ENTOMOLOGIST

Oceanville, 6 females, 5 males, 9.viii.l950 (no collector specified); 4 females, 4

males, 31.viii.l959 (no collector specified).

FLORIDA, Wakulla Co., 12.vi.l980 on Spartina alterniflora, 6 females, (P. D.

Stiling); 8.vi.l980 on Spartina alterniflora, 6 females (P. D. Stiling); 8.vi.l980 on

Spartina alterniflora, 3 females, 1 male (P. D. Stiling). All material deposited in U.S.

National Museum collection.

Meromyzobia melanosoma most nearly resembles M. unifasciata based on the

three-toothed mandible, exserted ovipositor and gonostyli; apically lobate middle

tibial spur, and seventh sternum which terminates near the basal one-third of the

metasoma. The new species differs from unifasciata most conspicuously in the

elongate metasoma which is clearly longer than the mesosoma, the body coloration,

and the forewing with two large fuscous areas separated by a hyaline stripe. The male

of unifasciata remains unknown, and the species remains known only from the

holotype. The species here called melanosoma appears widespread in Atlantic

Coastal situations and taken frequently with the salt marsh grass Spartina

alterniflora. The holotype and other specimens taken by P. D. Stiling were reared

from fly puparia (Stiling numbers 628C, 628D).

Meromyzobia pedicelata, New Species

Female. —2.30 mm long. Head predominantly pale brown; intertorular

projection, clypeus tan. Anterior face of pronotum reddish brown, posterior

one-third tan; mesoscutum predominantly brown; region posteriad of parapsidal

sutures tan. Axilla, scutellum, propodeum dark reddish brown; anterior half of

tegula nearly white, posterior half dark reddish brown; mesopleuron pale brown.

Anterior half of first metasomal tergum yellow, posterior half of first and second

through sixth terga reddish brown; posterior margin of fifth and sixth terga nearly

black, each forming a conspicuous “V.” Basal terga pale yellow; distal portion of

seventh sternum brown. Antenna tan. Coxae and trochanters yellow or nearly so;

fore and middle femora tan; hind femur dark brown or nearly black; fore and middle

tibiae dusky; hind tibia with basal one-third tan, distal two-thirds concolorous with

hind femur; middle tibial spur yellow; tarsomeres dusky with pretarsi somewhat

darker in certain plays of light. Forewing hyaline with a large fuscous cloud posteriad

of marginal and stigmal veins which expands toward remigium and which is

interrupted by a pale, transverse line corresponding roughly to a cubital vein; hind

wing hyaline. Gonostyli white or nearly so.

Head in dorsal aspect with frontovertex 0.50 times as wide as head; head in frontal

aspect (Fig. 25) minute and weakly reticulate with several scattered, shallow,

setigerous punctures; lateral ocellus less than one diameter from medial margin of

compound eye; posterior-most margin of compound eye less than one ocellar

diameter from vertexal margin. Antenna as illustrated (Figs. 28,29). Mandible with

one tooth and a broad truncation.

Pronotum about 0.33 times as long as medial length of mesoscutum. Mesoscutum

weakly and uniformly reticulate (Fig. 26), nearly polished; sculptural pattern nearly

identical on axillae and scutellum. Scutellum as long as wide, rather robust.

Mesopleuron polished with very weak reticulate sculpture along anterior margin, 1.7

times longer than wide. Propodeum with a complete, well developed, longitudinal

median carina; subcuticular carinae parallel. Middle tibial spur slightly longer than

VOLUME 63, NUMBER 1

Figures 25-31. 25. M. pedicelata New Species. Female head, frontal aspect. 26. M. pedicelata,

female mesosoma. 27. M. pedicelata, female mesosoma and metasoma. 28. M. pedicelata, female

antenna, medial aspect. 29. M. pedicelata, female pedicel. Funicular I-III. 30. M. pedicelata, female

middle tibial spur and basitarsus, medial aspect. 31. M. pedicelata, female middle tibial spur setae on

medial surface.

basitarsus; distally lobate (Figs. 30, 31); setae on medial surface moderately long,

curved, with apical tines absent or very weakly developed.

Metasoma (Fig. 27) about as long as mesosoma; ovipositor and gonostyli very

weakly exserted.

Metasoma 1.33 times as long as mesosoma. Terga weakly and uniformly

reticulate; pygostyli just posteriad of imaginary line bisecting metasoma. Seventh

PAN-PACIFIC ENTOMOLOGIST

Figures 32-37. 32. M. texana New Species. Female head, frontal aspect. 33. M. texana, female

mesosoma (arrow to parapsidal suture). 34. M. texana, female antenna, lateral aspect. 35. M. texana,

female antenna, annelus (arrow), Funicular I-III. 36. M. texana, female middle tibial spur, medial

aspect. 37. M. texana, female middle tibial spur setae on medial surface.

Sternum near imaginary line bisecting metasoma; posterior margin transverse.

Ovipositor and gonostyli visible when specimen viewed in lateral aspect, but not

exserted beyond apex of metasoma.

Male: —Unknown.

Described from six females reared from Orthoptera eggs attached to Tripsacum

laxum Nash taken at Kicco, Florida, on 3 February 1932 by R. D. Kennedy. This

VOLUME 63, NUMBER 1

Figures 38-40. 38. M. deserticola New Species. Male antenna, medial aspect. 39. M. deserticola,

female antenna, medial aspect. 40. M. deserticola, male forewing.

plant is native to Central America and not common in the United States. Holotype

and paratypes card-point mounted and deposited in the USNM collection. One

paratype dissected and mounted on a microscope slide.

This species appears related to the type-species, M. maculipennis, and M.

bifasciata, based on the large, robust body, forewing coloration and venation,

mandible with one tooth and a broad truncation, and large, distally enlarged, middle

tibial spur. It may be distinguished from those species based on the elongate pedicel,

distinctive medial propodeal carina, and the host association. According to Dr. D.

Nickle (pers. comm.), the host was probably a tettigoniid. The host association

appears solid. Preserved with the type-series are the eggs from which the parasites

emerged that contain the pupal exuviae of the parasites. Other species of

Meromyzobia for which the biology is known develop within fly puparia.

Meromyzohia texana, New Species

Female. —2.05 mm long. Body predominantly lemon yellow with following areas

darker: anterior face and postero-lateral margin of pronotum, posterior half of

tegula; metanotum, propodeum reddish brown; fourth metasomal tergum forming a

broad, transverse dark brown dusky stripe; fifth tergum dusky, metasoma with a

dusky spot on either side of apex. Antenna dusky. Forewing predominantly hyaline

with infuscation distad of speculum and beneath marginal vein and expanding

toward remigium, interrupted by a hyaline, transverse stripe at region of imaginary

cubital vein; distal margin with an occasional darkened area. Coxae, fore and middle

PAN-PACIFIC ENTOMOLOGIST

femora, all tibiae yellow; basal half of hind femur and tibia pale, distal half dusky to

dark brown; middle tibial spur white, tarsomeres off-white; apices of pretarsi dusky.

Head in dorsal aspect with frontovertex 0.41 times as wide as head; surface

minutely reticulate with several shallow setigerous punctures; lateral ocellus

separated from medial margin of compound eye by about one ocellar diameter;

posterior margin of compound eye nearly confluent with posterior margin of head,

separated by less than ocellar diameter. Head in frontal aspect (Fig. 32) as wide as

tall; toruli just beneath imaginary transverse line extending between ventral margins

of compound eyes. Toruli short and shallow. Clypeal margin transverse. Antenna as

illustrated (Figs. 34, 35). Mandible tridentate.

Mesosoma (Fig. 33) with pronotum 0.30 times as long as mesoscutum; posterior

margin broadly indented; surface weakly reticulate with moderate vestiture of dark

setae. Mesoscutum with similar sculpture and vestiture of setae; parapsidal sutures

transverse but not meeting mesad. Scutellum as long as wide, moderately robust.

Middle tibial spur (Figs. 36, 37) as long as middle basitarsus; lobate distad. Setae on

medial surface long and apically clubbed. Propodeum with subcuticular longitudinal

carinae divergent posteriad, medially bearing a very weak, incomplete superficial

Carina.

Metasoma 1.25 times as long as mesosoma. Gonostyli very slightly exserted.

Seventh sternum apically transverse, along with pygostyli just anteriad of an

imaginary line bisecting metasoma.

Male. —1.76 mm long. Apical half of metasoma dusky, otherwise identical to

female in habitus, chaetotaxy and coloration.

Described from seven females and one male taken at Weslaco, Texas, on

15/11/1949 by P. T. Riherd from rhodesgrass infested with A. graminis, and two

females taken by the same collector on Chloris gay ana Kunth infested with A.

graminis at the same locality on 14/1/1947. Material deposited in the USNM

collection. All specimens card-point mounted except one female which is slide

Discussion

Placement of this species is difficult. I have run it out among the smaller species in

the genus because I believe that is where its affinities lie. This being despite the fact

that M. texana is macropterous while M. deserticola and M. flavicincta are

micropterous as females. Perhaps more meaningful in distinguishing texana from

these two species is the curious directly convergent nature of the parapsidal sutures, a

feature not seen in other Meromyzobia. An incipient medial propodeal carina

suggests it may be related to M. pedicelata, but that species has mandibles with one

tooth and a broad truncation. I no longer doubt that rhodesgrass scale could be a host

of this parasite, but I have not seen it from other collections of this parasite taken

from rhodesgrass scale in Texas. If texana is a parasite of graminis, the parasite

shifted from a native host or was originally from the Orient, the natural range of

graminis. The suggested host range of Meromyzobia is too broad to rule out any

possibility for the origin of texana at this time. Curiously, A. graminis has been well

studied in Texas and has other imported encyrtid parasites (Clausen, 1978),

including other Miraini and mealybug parasites. Meromyzobia has not been

reported from the Orient (Noyes and Hayat, 1984). Given very limited data, I

suspect texana is probably attacking Diptera puparia in grasses, and that the species

is native to North America.

VOLUME 63, NUMBER 1

Meromyzobia unifasciata (Ashmead), 1900. Proc. U.S. Natl. Mus. 22:350. Female.

This species was based on one female taken at Utica, Mississippi, and has not been

recovered subsequently. The holotype is point mounted and stands in the USNM

collection (#4724). The original description is generally accurate, but relies

exclusively on coloration. More important diagnostic characters useful in identifying

this species include the distally lobate middle tibial spur, conspicuously exserted

ovipositor and gonostyli, parallel, subexocuticular, longitudinal propodeal carinae,

frontovertex 0.72 times as wide as head is long and 0.39 times as wide as head.

Mandible tridentate. Macropterous, forewing weakly infumated beneath marginal

vein. Metasoma, excluding exserted gonostyli, as long as mesosoma. Coloration

faded, but taken as generally accurate in the original description.

Scutellum with median length 1.18 times maximal width. Mesopleuron weakly

reticulate with pattern fading posteriad and absent along posterodorsal margin.

Metasoma, excluding exserted portion of gonostyli, 0.89 times as long as

mesosoma. Fuscous cloud beneath marginal and stigmal veins of forewing not

projecting to posterior margin of wing.

The male remains unknown.

Literature Cited

Ashmead, W. H. 1900. On the genera of the chalcid-flies belonging to the subfamily Encyrtinae. Proc.

U.S. Natl. Mus. 22:323-412.

Clausen, C. P. (Ed.). 1978. Introduced parasites and predators of arthropod pests and weeds: A world

review. USD A Handbk. 480. 545 pp.

DeSantis, L. 1968. Adiciones a la fauna Argentina de Encirtidos. II. (Hymenoptera). Rev. Mus. La Plata

(Zool.) 10:149-154.

-. 1972. Adiciones a la fauna Argentina de Encirtidos. III. (Hymenoptera: Chalcidoidea). Rev. Per.

Entomol. 15:44-60.

-. 1979. Catalogo de los himenopteros calcidoideos de America al sur de los Estados Unidos. Prov.

Bs. Aires Com. Inv. Cient. Esp. Pub., 488 pp.

Erdos, J., and S. Novicky. 1955. Genera Encyrtidarum regionis palaearcticae. Beitr. Entomol.

5:165-202.

Gordh, G. 1979. Family Encyrtidae, in K. V. Krombein et al. (eds.), Catalog of Hymenoptera in America

north of Mexico, GOP, Washington, 1:890-967.

Hoffer, A. 1955. The phylogeny and taxonomy of the family Encyrtidae (Hym., Chalcidoidea). Acta

Mus. Nat. Prague (B) 11:3-21.

Mercet, R. G. 1921. Himenopteros Fam. Encirtidos. Mus. Nac. Cien. Nat. Madrid. 732 pp.

Muesebeck, C. W. F., et al. 1951. Hymenoptera of America north of Mexico. Synoptic Catalog, USDA

Agr. Monogr. 2,1420 pp.

Noyes, J. S. 1980. A review of the genera of Neotropical Encyrtidae (Hymenoptera: Chalcidoidea). Bull.

Br. Mus. (Nat. Hist.) Entomol. Ser. 41:107-253.

-, and M. Hayat. 1984. A review of the genera of Indo-Pacific Encyrtidae (Hymenoptera:

Chalcidoidea). Bull. Br. Mus. (Nat. Hist.) Entomol. Ser. 48:131-395.

Peck, O. 1963. A catalog of the Nearctic Chalcidoidea (Insecta: Hymenoptera). Can. Entomol.

30:1-1091.

Subba Rao, B. R. 1971. New genera and species of encyrtids (Hymenoptera: Encyrtidae). Jour. Nat.

Hist. 5:209-224.

Tachikawa, T. 1978. Host of the Encyrtidae in the world (Hymenoptera: Chalcidoidea). Trans. Shikoku

Entomol. Soc. 14:43-63.

Trjapitzin, V. A. 1973A. The classification of parasitic Hymenoptera of the family Encyrtidae

(Hymenoptera, Chalcidoidea). Part I. Survey of the systems of classification. The subfamily

Tetracneminae Howard, 1892. Entomol. Rev. 52:118-125.

-. 1973B. Classification of the parasitic Hymenoptera of the family Encyrtidae (Chalcidoidea). Part

11. Subfamily Encrytinae Walker 1837. Entomol. Rev. 52:287-295.

PAN-PACIFIC ENTOMOLOGIST

-, and G. Gordh. 1978. Review of the genera of Nearctic Encyrtidae (Hymenoptera, Chalcidoidea).

II. Entomol. Rev. 57:437-448.

Williston, S. W. 1893. Description of a species of Chlorops reared from galls on Muhlenbergia mexicana,

by F. M. Webster. Ohio Agr. Exp. Sta. Tech. Bull. 1:157-158.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 37-42

Nest-site Preferences of the Giant Honey Bee, Apis dorsata

(Hymenoptera: Apidae), in Borneo

Christopher K. Starr% Patricia J. Schmidt^, and Justin O. Schmidt3

(CKS) Biology Department, De La Salle University, P.O. Box 3819, Manila,

Philippines; (PIS) Department of Entomology, University of Arizona, Tucson,

Arizona 85721; (JOS) Carl Hayden Bee Research Center, U.S. Dept, of

Agriculture, 2000 E. Allen Road, Tucson, Arizona 85719.

Abstract. —The characteristics oiApis dorsata (Hymenoptera: Apidae) nest-sites

in Sabah, Borneo, are described on the basis of 15 nest-bearing trees. Nests were

consistently high in very tall trees, exposed, and commonly aggregated. These

features are consistent with what has been reported from mainland populations of

A. dorsata, but not with giant honey bees from the Philippines. This supports the

hypothesis that these latter are a distinct species.

The giant honey bees form a distinct species-group within Apw and are sometimes

treated as a subgenus, Megapis Ashmead. At the most conservative modern

estimate, the group contains two species, the Himalayan A. laboriosa Smith and

Indomalaysian A. dorsata Fabricius. On the other hand, Maa (1953) divided the

latter species into three: A. dorsata, A. binghami Cockerell (= A. zonata Smith) and

A. breviligula Maa. This assessment is tentatively accepted by Sakagami et al.

(1980), and Roubik et al. (1985), though not generally by other authors.

The name A. binghami applies to the little-known populations of Megapis on

Celebes and associated smaller islands; A. breviligula applies to all populations in the

Philippine islands aside from the Palawan group; and A. dorsata applies to all other

non-laboriosa populations, including those of Borneo and neighboring Palawan. The

distinctness of A. binghami and A. breviligula is currently under biometrical

investigation (S.F. Sakagami, pers. comm.). For convenience, we refer to all

Megapis from the Philippines proper as A. breviligula, without implying that the

taxonomic question is settled, and to all non-laboriosa Megapis as the A. dorsata

group.

Wallace (1869) was among the early naturalists who commented on these

conspicuous bees and his remarks will serve as a summary for what many later

travelers noted. A. dorsata in Borneo, he said, “build huge honeycombs, suspended

in the open air from the underside of the lofty branches of the highest trees.” He

remarked especially on an aggregation of three nests that he watched being robbed

by a team of men. The man who climbed the tree and cut down the combs protected

himself with a heavy cloth wrapping and a smoke-torch, but was nonetheless

repeatedly stung.

Morse and Laigo (1969) provided most of what is known of the biology of

A. breviligula and reviewed the literature on both A. breviligula and A. dorsata.

Since then, Deodikar et al. (1977), and Seeley et al. (1982; summarized by Seeley

PAN-PACIFIC ENTOMOLOGIST

1983) have contributed important new information on the biology of dorsata on

the Asian mainland.

In this paper we describe nest-sites and colony aggregations of A. dorsata from

Sabah and compare these taxonomically with data for A. breviligula.

Materials and Methods

All observations from Sabah were made during May, 1985. Trees with active

colonies of A. dorsata or which had had colonies in them were identified along the

highways between Tamparuli-Marak Parak via Kota Maruda and between Ranau

and Sandakan (lat. 6°N, long. 117°E). In most cases initial discoveries were made

from the road. The nests were then often observed through binoculars and by

approach on foot to the tree. A. dorsata trees were initially recognized by the

presence of active colonies or parts of a honey hunting ladder, and in one case by

vestiges of a nest (comb-scar).

Results

Table 1 lists characteristics of the 15 nesting trees we found. Although we did not

specifically search low vegetation, we saw no indication of A. dorsata nesting

anywhere except on tree branches, an observation supported by the local people we

contacted. All the trees were very tall, and we estimate that none had branches lower

than 15m. Moreover, unlike many other trees of similar size and shape, the trees with

bees were all clean and free of epiphytes or lianas on the trunk and main branches

(Figs. 1, 3, 4). Almost all the nests were in open, unencumbered zones, without

vegetation close to them. As seen in Table 1, a majority of the trees were

smooth-barked. These had light-medium gray bark and appeared to be a single

species. The usual tree for A. dorsata in that area is reported to be Koompassia

[Fabaceae], (Orolfo 1965; Anthea Phillips pers. comm.). The description of

K. excelsa (Becc.), one of the world’s tallest known angiosperm tree species, agrees

well with the trees, we observed. In addition. Corner (1952) remarked of K. excelsa

that, “the branches often bear large combs of wild bees” in Malaya and cites a local

name for it, tualang, meaning “tree of swarming bees.” Roepke (1948) also noted a

tendency for A. dorsata to nest in Koompassia. The bark of the three putative

Koompassia trees that we closely inspected was smooth, hard, and compact, with no

sign of flaking (Figs. 1, 2).

Many areas in which the bee trees were found had great numbers of tall, standing

dead trees, as a result of habitat destruction through logging. Nonetheless, all eight

trees with active bee colonies were living, as were six of the seven trees with signs of

previous colonies (Table 1).

At least 10 of the 15 trees contained a ladder or vestiges of one going up the trunk

to the bottom branches (Table 1, Fig. 2). Some of the other trees also might have had

ladders, on a side hidden from us. The main part of the ladder consists of a series of

sharp stakes with fire-hardened tips, from an exceptionally dense, hard dipterocarp

tree, driven directly into the bee tree (Charles Jackson, pers. comm.). These are then

lashed with rattan (Ca/flmw5sp. [Palmae]) to a bamboo upright (Fig. 3). Inasmuch as

at least three of the trees with active colonies in them had vestiges of ladders—an

indication that they had probably been hunted successfully at least once before—^it

seems likely that some trees are hunted repeatedly.

On eight trees we observed 22 active nests of bees in groups of 1-7 nests per tree

VOLUME 63, NUMBER 1

Table 1. Characteristics of nesting trees oiApis dorsata in Sabah.

Tree

Bark

Ladder

Number of

Active Colonies

Remarks

rough

—

possibly one or two

other hidden nests

semi-rough

—

smooth

dead tree

smooth

on two adjoining trees

smooth

—

definite comb-scar

smooth

—

smooth

—

smooth

—

% of total trees

60% ladders

53% active

(Table 1). In those trees with multiple nests the nests appeared to be widely dispersed

amongst the lower branches (Fig. 4), and certainly, no strong clumping within a tree

was evident.

In the Philippines, one of us (CKS) has seen five swarms of A. breviligula on the

island of Leyte and two of A. dorsata on the island of Palawan. These were all on the

undersides of gently sloping branches. They resembled colonies with nests, but were

more compact, i.e. forming a shallow, distinctly broader mass (Fig. 5). In a

residential area of Kota Kinabalu, Sabah, we collected a small swarm of A. dorsata

on a tree branch about 4m from the ground (Fig. 5). It appeared to have

approximately one-tenth the volume of the five more uniform swarms mentioned

above. Our collection, which appeared to encompass virtually all the bees in the

swarm, comprised 1114 workers and 171 males. No queen was among them.

Discussion

Deodikar et al. (1977) reported observations on 1860^. dorsata nests in India. Of

these, 55% were in trees, the rest on human-made structures. The majority of nests

were between 6 and 12 m above ground, with only 6% lower down. Seeley et al.

(1982) described 15 trees in Thailand with A. dorsata nests. Fourteen of these were

straight, smooth-barked, and limbless for at least 13m. Although the lowest nest was

only 3.5m above the ground, most nests were located at heights between 13 and 27m,

and all were in open vegetation. These observations agree with ours from Sabah, as

well as with various other reports on the nesting of A. dorsata (Morse and Laigo,

1969).

Seeley et al. (1982) also reported a significant tendency for colonies to aggregate,

with up to 24 nesting in a single tree. Morse and Laigo (1969) reported that most

colonies seen by Morse in India were also aggregated, the largest aggregation being

PAN-PACIFIC ENTOMOLOGIST

Figures 1-3. 1. Typical bee tree, showing height, smooth bark, and lack of epiphytes. A. dorsata

colony is visible (arrow). 2. Vestiges of honey-hunting ladder on bee tree trunk. This ladder allowed

access to the position of the lowest colony (arrow), and also went higher into the tree. 3. Detail of one

step of honey-hunting ladder, just beginning to disintegrate.

34 colonies, and Deodikar et al. (1977) likewise reported nests generally aggregated.

Even larger aggregations, the greatest number being 156 in a single tree (see Morse

and Laigo, 1969, for citations), have been noted. These observations led Morse and

Laigo to conclude that trees with 20-30 colonies of A. dorsata are not rare

throughout most of its range.

The nesting characteristics of A. breviligula as reported by Morse and Laigo (1969)

from 30 colonies in Luzon, Philippines, and corroborated by one of us (CKS) from

five colonies in Leyte, are in strong contrast to the above. A number of consistent

differences are apparent:

a. A. breviligula nests are lower; they are rarely found in high trees, and often the

bottom of the nest is within Im of the ground.

b. It shows no preference for smooth-barked trees.

c. It tends to nest less in the open, often in the midst of fairly dense vegetation, so

that the nests are much less conspicuous from a distance. Some nests even had

small branches projecting through them.

d. Colonies of A. breviligula are single, not aggregated.

Morse and Laigo (1969) explicitly noted the first and last of these contrasts

between A. breviligula and A. dorsata and remarked on these “subtle differences.”

In our view, they are important and lend support to the hypothesis that A. breviligula

is a distinct species. The significance of our observations is strengthened by the fact

that they were made in Sabah, very near the Philippines proper. If the two forms

were the same species, we would expect bees from Sabah to be intermediate between

those of Thailand and Luzon. With regard to nesting characteristics, this prediction is

not corroborated. The distinct-species hypothesis further predicts that bees in

Palawan will not differ significantly from those of Sabah, in the direction of

resembling those of Luzon. This has yet to be tested.

Seeley et al. (1982) account for the nesting habits of A. dorsata as part of their

defensive strategy against vertebrate enemies. The keys to this strategy are

VOLUME 63, NUMBER 1

Figure 4. Crown of bee tree with four active colonies and parts of one old nest in view (arrows).

Figure 5. A. dorsata colony at Aborlan, Palawan, Philippines, a. As a normal colony, with bees

covering a nest. b. As a swarm, resettled on the same branch after they had been driven away with smoke

and the nest removed.

inaccessibility and a readiness to launch a massive attack. They reason that the

depredations of humans, in particular, over the millenia must have constituted

strong natural selection. The use of honey-hunting ladders today in Sabah indicates

that this selection continues.

A. breviligula can launch comparable attacks against intruders (Morse and Laigo,

1969) but its nests are commonly quite accessible. It is not easy to explain this

difference, unless A. breviligula had few natural vertebrate predators before the

PAN-PACIFIC ENTOMOLOGIST

arrival of humans. At present, A. breviligula colonies located in inhabited areas are

rarely left undisturbed; instead they are usually destroyed or driven out with smoke

and fire within a few days of discovery (Morse and Laigo, 1969; pers. obs.). It may be

that they gain some cryptic protection by nesting lower and in denser vegetation, but

the more likely hypothesis is that selection pressure from humans has until recently

not been very strong in the Philippines proper.

The reason for colony aggregation in A. dorsata is obscure. In light of the scant

available evidence, Seeley et al. (1982) tentatively concluded that the scarcity of

suitable nest substrates best accounts for aggregation. Our own observations do not

support this hypothesis; suitable unoccupied trees appeared plentiful in the vicinity

of those with two or more active colonies.

Acknowledgments

We thank the governments of Malaysia and Sabah for permission to do research in

Sabah, granted on exceptionally short notice. This study also benefited from advice

and other assistance from George Lo, Vincent Au, Charles Jackson, Anthea Phillips

and the Eugene Fuller family. We thank Steve Buchmann, Jim Cane, Roger Morse

and Shoichi Sakagami for critical comments and Lucille Valente and Cristina

Bramley for manuscript preparation.

Literature Cited

Corner, E. J. H. 1952. Wayside Trees of Malaya. 2nd Edition. Gov’t. Printing Office, Singapore.

Deodikar, G. B., A. 1. Ghatge, R. P. Phadke, D. B. Mahindre, K. K. Kshirsagar, K. S. Muvel and C. V.

Thakar. 1977. Nesting behavior of Indian honeybees. III. Nesting behavior oiApis dorsata Fab.

Indian Bee J. 39:1-12.

Maa, T. C. 1953. An inquiry into the systematics of the tribus Apidini or honeybees (Hym.). Treubia

21:525-640.

Morse, R. A., and F. M. Laigo. 1969. Apis dorsata in the Philippines (including an annotated

bibliography). Phil. Assoc. Ent., Monogr. No. 1:96pp.

Orolfo, P. 1965. Gathering jungle honey. Sabah Soc. J. 2:165-168.

Roepke, W. 1948. Observations sur les abeilles des Indes. Rev. Franc. Apic. Abeille Miel 28:567-569,

29:596-599, 30:618-620.

Roubik, D. W., S. F. Sakagami and 1. Kudo. 1985. A note on the distribution and nesting of the

Himalayan honey bee Apis laboriosa Smith (Hymenoptera: Aposidea). J. Kansas Ent. Soc.

(58):746-749.

Sakagami, S. F., T. Matsumura and K. Ito. 1980. Apis laboriosa in Himalaya, the little known world’s

largest honeybee (Hymenoptera, Apidae). Ins. Matsumurana (N. S.) 19:47-77.

-, R. H. Seeley and P. Akratanakul. 1982. Colony defense strategies of the honeybees in Thailand.

Ecol. Monogr. 52:43-63.

Wallace, A. R. 1869. The Malay Archipelago. MacMillan, London (Reprinted 1962 by Dover, NY).

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 43-47

The Genus Pachymerola Bates

(Coleoptera: Cerambycidae)

Edmund F. Giesbert

9780 Drake Lane, Beverly Hills, California 90210.

Abstract .—The monobasic genus Pachymerola Bates is examined and

redescribed; P. vitticollis is discussed and briefly redescribed. A new species and

subspecies are proposed: P. ruficollis ruficollis from western Mexico to Honduras,

and P. ruficollis humeralis from Costa Rica. The latter is figured. A change in tribal

placement is proposed, assigning Pachymerola to the tribe Hyboderini.

Bates (1892) described Pachymerola vitticollis from a single male specimen taken

at Chilpancingo (alt. 4600 ft.), Guerrero, Mexico, and assigned this new genus to the

tribe Compsocerini, based on an observed similarity of the male to that of Coremia

Serville. The modified abdomen of the female, herein described, necessitates

reassignment of Pachymerola to the Hyboderini.

S. A. Fragoso (1978) has attempted to clarify the tribal classification of the North

American Cerambycinae, primarily based on studies of the terminalia of both sexes.

This work may result in a future change of status for the tribe Hyboderini.

Specimens of Pachymerola are fairly commonly taken on blossoms of Croton, and

other flowering woody plants, although no larval host associations are recorded. The

genus is thus far known from Sinaloa in western Mexico to the Cordillera de Tilaran

of Costa Rica.

Examination of the large quantity of material at hand has failed to reveal any

specimen agreeing with the unique type of P. vitticollis.

Genus Pachymerola Bates

Pachymerola Bates 1892:161; Blackwelder 1946:580 (list).

Form moderately small, elongate. Pubescence fine, short, sericeus, denser on

ventral surfaces. Head coarsely punctate, with front canaliculate, clypeal suture

deeply impressed; eyes finely facetted, deeply emarginate; antennae slender, not

longer than body, with scape clavate, second segment short, fourth segment longest,

outer segments subserrate. Pronotum pyriform, about as long as broad, sides

obtusely angulate, widest behind middle; disk uneven, asperate-punctate;

prosternum with intercoxal process narrow, procoxal cavities closed behind;

mesosternal process wider; metasternum longitudinally impressed; metacoxae

enlarged, tumid. Abdomen with first sternite elongate in both sexes; female with first

abdominal sternite as long as remaining sternites together, second sternite deeply

emarginate, with transverse brush composed of scooplike setae arising from distal

border, lateral setae longer, incurved, pointed, remaining sternites shorter, concave,

hairy. Elytra somewhat flattened, nearly parallel sided, asperate-punctate,

alutaceous, apices obtusely acuminate. Legs with femora moderately clavate;

PAN-PACIFIC ENTOMOLOGIST

metafemora enlarged, thickened, exceeding elytral apices, alutaceous, asperate;

metatibiae asperate.

Type Species. —Pachymerola vitticollis Bates (monobasic).

Key to Separate the Species and Subspecies of Pachymerola Bates

1. Pronotum black with a yellowish-gray pubescent vitta on each side of disk.

Antennae of male about as long as body, with 8th segment equal to 3rd.

Guerrero, Mexico. P. vitticollis

- Pronotum orange or reddish, rarely infuscated, lacking pubescent vittae.

Antennae of male reaching at most to apical Vs of elytra, with 8th

segment distinctly shorter than 3rd .2.

2. Elytra black, sometimes indistinctly suffused with orange or reddish at

base, without distinct triangular orange humeral maculae. Femora, at

least anterior pair, usually orange at base. Mexico to Honduras .

. P. ruficollis ruficollis

— Elytra black, with distinct orange triangular humeral maculae. Legs black.

Costa Rica . P. ruficollis humeralis

Pachymerola vitticollis Bates

Pachymerola vitticollis Bates 1892:161; Blackwelder 1946:580 (list).

Male. —Form moderately small, nearly parallel sided. Integument black.

Antennae about as long as body, 8th segment subequal to 3rd. Pronotum

asperate-punctate, with a yellowish gray pubescent vitta on each side of disk. Elytra

somewhat flattened, asperate-punctate, alutaceous, with very fine, short, sericeous

pubescence. Body beneath with somewhat more dense silvery-white pubescence;

abdomen simple. Legs with metafemora enlarged, thickened, asperate, alutaceous,

denticulate-asperate distally on underside, metatibiae sulcate, asperate. Length

8.5mm.

Female. —U nkno wn.

Type Locality. —Chilpancingo, Guerrero, Mexico.

Remarks. —Known from a unique male specimen, this species appears to differ

from all but the most melanic individuals of the following new species by the black

color, and from all individuals of that species by the vittate pronotum, and longer

antennae.

Pachymerola ruficollis ruficollis Giesbert, New Species

Male. —Form small to moderately small, nearly parallel sided, with sides feebly

incurved behind middle. Integument piceous to black, with pronotum usually orange

or reddish, rarely infuscated, and often with occipital area of head, bases of elytra

indistinctly, femoral bases, coxae, and parts of sternum suffused with orange.

Antennae reaching apical 1/5 of elytra, with outer segments shortened, 8th segment

shorter than 3rd. Pronotum opaque, usually densely asperate-punctate, without

pubescent vittae. Elytra somewhat flattened distally, distinctly asperate-punctate,

alutaceous, with fine, short, sericeous pubescence. Body beneath with somewhat

denser sericeous pubescence; abdomen simple, with first sternite comprising 1/3 to 1/2

VOLUME 63, NUMBER 1

of abdominal length. Legs with metafemora enlarged, thickened, asperate,

alutaceous, often somewhat nitid and denticulate distally beneath. Length 6-llmm.

Female. —Similar to male, but with antennae shorter, reaching just past middle of

elytra. Abdomen modified with scopate second sternite. Length 7-13mm.

Types.—Holotype male, allotype (California Academy of Sciences), and 19

paratypes (7 male, 12 female) from 5 mi. N Mazatlan, Sinaloa, Mexico, July

28-August 3, 1973 (E. Giesbert). 167 additional paratypes as follows: MEXICO:

Sinaloa; 26 males, 30 females, 5 mi. N Mazatlan, July 21-August 1,1972, on flowers

of Buddleia wrightii and Jatropha curcas (J. & M. A. Chemsak, A. & M.

Michelbacher); 1 male, 5 mi. N Mazatlan, August 15, 1966 (J. A. Chemsak & J.

Doyen); 1 male, 2.5 mi. N Mazatlan, August 12, 1970 (M. Wasbauer); 3 males, 1

female, 2.5mi. NMazatlan, August 10-11,1970 (J. A. Chemsak);2males, 1 female,

5 mi. N Mazatlan, August 9-15,1970 (J. A. Chemsak); 11 males, 14females, 5 mi. N

Mazatlan, July 25-August 1,1973 (J. Chemsak, E. G. Linsleys & Michelbachers); 24

males, 4 females, 5 mi. NMazatlan, July 30-August 8,1983 (F. T. Hovore); 1 male, 5

mi. N Mazatlan, August 9, 1983 (E. Giesbert); Nayarit: 6 males, 2 females,

Ahuacatlan, July 18-22,1951, on flowers of Donnellsmithia hintoni M. & C. (P. D.

Hurd, H. E. Evans); 3 males, Arroyo Santiago nr. Jesus Maria, July 5, 1955

(B. Malkin). Jalisco: 9 males, 3 females, Est. Biol. Chamela, July 10-20, 1985, on

flowers of Croton. (E. Giesbert); 2 males, Est. Biol. Chamela, July 8-16, 1985

(J. Chemsak, H. Katsura, A. & M. Michelbacher); 1 male, Chamela, June 19,1983

(S. H. Bullock); 1 male, Chamela, July 21, 1984, on flowers of Croton (J. A.

Chemsak). Oaxaca: 2 males, 4 females, 30 mi. NE Tehuantepec, July 8, 1955

(D. Giuliani); 1 male, 6 mi. N Juchitan, July 3 1955 (Univ. Kans. Mex. Expedition);

1 male, 8 mi. N La Ventosa, July 20, 1963 (W. A. Foster); 1 female, Isth. of

Tehuantepec, (F. Sumichrast); 2 males, 56 mi. NW Tehuantepec, July 27, 1963

(J. Doyen). Chiapas: 1 female, 8 mi. E Rizo de Oro, June 22, 1985 (D. Heffern).

Quintana Roo: 3 males, 2 females, 18 km N Felipe Carillo Puerto, May 27-June 1,

1984 (J. E. Wappes); 1 male, 10 km N Puerto Morelos, June 15-16, 1983

(E. Giesbert). EL SALVADOR: 1 male, 1 female. Dept. Ahuachapan, Bosque El

Imposible (745m), June 18, 1979 (R. D. Cave). HONDURAS: 1 female,

Tegucigalpa, June 12,1918 (Dyer).

Remarks. —Integumental color is somewhat unstable within populations in this

species. The Sinaloa population exhibits the greatest tendency toward reduced

melanism throughout, with pronota and femoral bases consistently orange. The

Jalisco population, also with somewhat reduced melanism, exhibits pronota of a rich

red color, sometimes partially infuscated. Specimens from further south in Mexico

exhibit an orange pronotum, which is sometimes infuscated to varying degrees, legs

with only the procoxae and bases of profemora usually suffused with orange, and

consistently black elytra. The small amount of material available from Central

America is also quite melanic: specimens from El Salvador are entirely black with

orange pronota, the Honduran example similar, but with orange profemoral bases.

A single entirely black specimen was seen from Tehuantepec, but generally, it would

appear that melanism tends to increase clinally north to south.

In addition, the Sinaloan population exhibits a more nitid, less alutaceous surface

on the underside of the enlarged metafemora, and northern populations exhibit a

tendency toward a more densely asperate pronotal surface.

Figures 1. Pachymerola ruficollis humeralis Giesbert; male.

VOLUME 63, NUMBER 1

Pachymerola ruficollis humeralis Giesbert, New Subspecies

(Fig-1)

Male. —Form moderately small, with sides slightly incurved behind middle.

Integument black, with pronotum and large triangular maculae on humeri orange.

Antennae reaching apical 1/5 of elytra, with 8th segment shorter than 3rd. Pronotum

opaque, somewhat indistinctly asperate-punctate, without pubescent vittae.

Abdomen simple, with first sternite comprising 1/2 of abdominal length. Legs with

metafemora elongate, thickened, asperate-punctate, alutaceous, with underside

denticulate distally; metatibiae asperate. Length 9-12mm.

Female. —Similar to male, but with antennae shorter, just surpassing middle of

elytra. Abdomen modified with scopate second sternite. Length 9.5-13.5mm.

Types.—Holotype male, allotype (California Academy of Sciences), and 23

paratypes (15 males, 8 females), from 6 km S Santa Elena, 1100m, Puntarenas prov.,

Costa Rica, June 5-7, 1980, on flowers of Croton (E. Giesbert). 36 additional

paratypes, all from the same locality, as follows: 9 males, June 2, 1979 (H. & A.

Howden); 8males, 7females, June 4—7,1980 (J. E. Wappes); 7 males, 1 female, June

6-7, 1983 (E. Giesbert); 2 males, June 6-7, 1983 (J. E. Wappes); 2 males. May 18,

1984 (F. T. Hovore).

Remarks. —This apparently isolated and genetically stable population is

characterized by the large orange triangular humeral patches of the elytra, and by a

slightly larger average size than Pachymerola r. ruficollis.

Acknowledgments

I wish to thank John A. Chemsak of The University of California, Berkeley, for

the loan of material and review of the manuscript, Frank T. Hovore and James E.

Wappes for specimen data, and R. D. Pope of the British Museum (Nat. Hist.) for

allowing me to examine and photograph the type. I would also like to thank Dr.

Henry Howden for generous collecting information on the Costa Rican locality.

Literature Cited

Bates, H. W. 1892. Additions to the Longicornia of Mexico and Central America, with remarks on some

of the previously recorded species. The Transactions of the Entomological Society of London,

1892:143-183, Ulus.

Blackwelder, R. E. 1946. Checklist of the Coleopterous Insects of Mexico, Central America, The West

Indies, and South America. Part 4. Bulletin of the United States National Museum. 185:551-763.

Fragoso, S. A. 1978. Male and female terminalia as a basis for tribal classification of the subfamily

Cerambycinae of America north of Mexico. Unpublished PhD dissertation. University of Florida

1978.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 48-49

New Host Records of Bruchidae (Coleoptera)

From Desmanthus (Leguminosae) From Texas and Mexico

Melissa Luckow and Clarence Dan Johnson

(ML) The Department of Botany, The University of Texas at Austin, Austin,

Texas 78713-7640; (CDJ) Department of Biological Sciences, Northern Arizona

University, Flagstaff, Arizona 86011.

Abstract .—Nine species and two varieties of Desmanthus are reported as hosts

from new localities in Texas and Mexico for four species of Bruchidae. New records

reported include Acanthoscelides compressicornis feeding in the seeds of

Desmanthus painteri, D. pringlei, and D. virgatus var. glandulosus; A. pectoralis in

the seeds of D. reticulatus; and A. desmanthi in the seeds of D. subulatus. Other

species of Desmanthus reported as hosts for bruchids are D. covillei, D. obtusus, D.

velutinus, D. virgatus ^ 20 :. depressus, D. virgatus var. virgatus, and D. fruticosus.

The bruchids Stator pruininus and several species in the genus Acanthoscelides

have been reported to feed in the seeds of species of Desmanthus from the United

States to Central America (Bottimer, 1969; Center and Johnson, 1974, 1976;

Johnson, 1970, 1977, 1983, 1984; Johnson and Kingsolver, 1976). It also has been

observed by CDJ that bruchids feed in the seeds of Desmanthus in northern South

America.

Recently a number of new host records from new localities in Texas and Mexico

were obtained by ML in the course of her work on the systematics of Desmanthus.

Because of ongoing research on bruchids and their hosts, these records are published

here to make them available for studies on bruchid-host interactions. New records

found and reported upon here include Acanthoscelides compressicornis feeding in

the seeds of Desmanthus painteri, D. pringlei, and D. virgatus var. glandulosus; A.

pectoralis in the seeds of D. reticulatus; and A. desmanthi in the seeds of D.

subulatus.

Desmanthus virgatus var. glandulosus Turner is believed to be a distinct and

separate species more closely related to D. velutinus and D. cooleyi than D. virgatus

by ML so we are listing it here under its currently accepted name pending further

study.

Acknowledgments

Partial support for this paper was provided by an Institute of Latin American

Studies-Tinker Foundation Grant and NSF Predoctoral Grant BSR85-14314 to ML;

and NSF Grant BSR82-11763 to CDJ.

New Host Records

Acanthoscelides compressicornis (Schaeffer)

1. Desmanthus covillei (Britton & Rose) Wiggins ex Turner: Mexico. Sonora:

Mpio Cd. Obregon: 3 mi NW Santini on Hwy 15, 26 August 1985 (ML 2806).

VOLUME 63, NUMBER 1

2. Desmanthus obtusus S. Wats.: Texas. Upton Co.: 2 mi W Rankin on Hwy 67, 5

August 1985 (ML 2737). Reagan Co.: 9 mi W Big Lake on Hwy 67, 5 August 1985

(ML 2738).

3. Desmanthus painteri (Britton & Rose) Standley: Mexico. Coahuila: Mpio

Arteaga: 8.1 mi SE turnoff to San Antonio de Las Alazana on Rte 57, 2 July 1985

(ML 2641).

4. Desmanthuspringlei (Britton & Rose) Hermann: Mexico. Nuevo Leon: Mpio

Allende: 1.7 mi N of Allende on Hwy 85,2 July 1985 (ML 2640).

5. Desmanthus velutinus Scheele: Texas. Reeves Co.: 10.6 mi W jctn Hwy 290 on

Hwy 10, 21 August 1985 (ML 2750). Travis Co.: Big Sandy Creek Park outside of

Austin, 31 July 1985 (ML 2709). Val Verde Co.: Pecos River near intersection with

Hwy 10, 2 August 1985 (ML 2721).

6. Desmanthus virgatus var. depressus (H. & B.) Turner: Texas. Uvalde Co.:

Sabinal Cemetery, 1 mi E Sabinal on Hwy 90, 2 August 1985 (ML 2713).

7. Desmanthus virgatus var. glandulosus Turner: Texas: Brewster Co.: Iron Mt.

Ranch, plateau just below Gilliland Peak near the main ranch house, 3 August 1985

(ML 2729).

Acanthoscelides pectoralis (Horn)

1. Desmanthus reticulatus Benth.: Texas. Uvalde Co.: Sabinal Cemetery, 1 mi E

Sabinal on Hwy 90, 2 August 1985 (ML 2712).

Acanthoscelides desmanthi Johnson

1. Desmanthus subulatus (Britton & Rose) Wiggins ex Turner: Mexico. Sinaloa:

Mpio Mazatlan: Hwy 15,3.8 mi NW of Rio San Lorenzo, 8 January 1986 (ML 2980).

2. Desmanthus virgatus (L.) Willd. var. virgatus: Mexico. Guerrero: Mpio

Tecpan: 12 mi W of bridge at San Luis on Hwy 200,13 March 1985 (ML 2615).

Statorpruininus (Horn)

1. Desmanthusfruticosus Rose: Mexico. Baja Sur: Mpio Todos Santos: 2.4 mi S of

Todos Santos on Hwy 19, 30 August 1985 (ML 2823).

Literature Cited

Bottimer, L. J. 1969. Two Acanthoscelides (Coleoptera: Bmchidae) from southern United States

with notes on related species. Can. Entomol., 101:975-983.

Center, T. D. and C. D. Johnson. 1974. Coevolution of some seed beetles (Coleoptera: Bruchidae) and

their hosts. Ecology, 55:1096-1103.

-, and-. 1976. Host plants andparasitesof some Arizona seed-feeding insects. Ann. Entomol.

Soc. Amer., 69(2): 195-201.

Johnson, C. D. 1970. Biosystematics of the Arizona, California, and Oregon Species of the Seed Beetle

Genus Acanthoscelides Schilsky (Coleoptera: Bruchidae). Univ. Calif. Publ. Entomol., 59:1-116.

-. 1977. Two new species oi Acanthoscelides (Coleoptera: Bruchidae) from North America and new

bruchid host records from Desmanthus and Hoffmanseggia (Leguminosae). Pan-Pac. Entomol.,

53:60-73.

-. 1983. Ecosystematics of Acanthoscelides (Coleoptera: Bruchidae) of Southern Mexico and

Central America. Misc. Publ. Entomol. Soc. Amer., 56:1-370.

-. 1984. New host records and notes on the biology of Stator (Coleoptera: Bruchidae). Coleopterists

Bull.,38(l):85-90.

-, and J. M. Kingsolver. 1976. Systematics of Stator of North and Central America (Coleoptera:

Bruchidae). U.S. Dept. Agric. Tech. Bull., 1537.101 pp.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 50-51

Sex-influenced Protibial Spines

And Synonymy in Dasytidae (Coleoptera),

Study Number Three

Charles D. Howell

San Bernardino County Museum, 2024 Orange Tree Lane, Redlands, California,

and The University of Redlands 92373.

Abstract. —Protibial spines are sex-influenced in many populations of

Emmenotarsus and Trichochrous. In the extreme case, they are found to be

sex-limited to females in Listropsis, an emmenotarsus-like genus. This led to the

discovery that Listropsis and Trichochroides are synonymous, and Trichochroides

must be suppressed on basis of priority.

Considerable confusion exists in the systematics of Dasytidae. Synonymy of some

species was noted by Fall (1901) and more can be expected since many species were

based by Casey (1895) on only one specimen. What effect the synonomy may have on

the total number of species is uncertain, for new collections continue to reveal

populations so different from those described as to require the addition of new

specific names (e.g. Howell, 1979).

Adding to the confusion, some populations contain a tremendous variety of

morphs, species, and even genera. Howell (1985) reports finding seven genera of

Dasytidae in the flowers of one bush of Ceanothus; and nine morphs of one genus,

Eschatocrepis, in a small local area. The present study describes a situation in which

males and females of the same species can be keyed to different genera.

This came about by using the characteristic “protibiae beset with spines” in

current keys (Casey, 1895; Blaisdell, 1938) to separate two fairly large divisions of

Dasytidae. Among the genera with spines is Trichochrous, the largest dasytid genus

according to Casey. Among those lacking them is Amecocerus {Listrus, in Blaisdell,

1921), which is the second largest.

Blaisdell (1938) subdivided Trichochrous into several genera, among which

Emmenotarsus was set apart for its bristling fringes and shaggy appearance. At the

same time he selected Trichochrous sexualis Casey, also emmenotarsus-like, as the

type of the genus Trichochroides which is defined by a male-limited feature, a

striking impression on the fifth abdominal sternite. Earlier, Blaisdell (1924a, 1924b)

described species of Listropsis, based on males lacking protibial spines. This latter

feature led him to underestimate the obvious emmenotarsus-like appearance of

Listropsis based on males lacking protibial spines. This latter feature led him to

underestimate the obvious emmenotarsus-like appearance of Listropsis and to place

it among listrus-like genera.

A study of the number of spines on protibiae led me to discover that in many

species of Trichochrous and Emmenotarsus the number of spines on the protibiae is

sex-influenced, females having significantly more than males. The number may vary

VOLUME 63, NUMBER 1

from zero to over a dozen, so that specimens at the extremes, at least, would end up

in widely different genera if keyed individually.

This challenged me to devise a key avoiding a major reliance on protibial spines.

Such a key is in use in our laboratory and appears to be as reliable as Blaisdell’s

(1938) key. It brings Trichochrous and Amecocerus close to each other, which is

concordant with their gross similarity. It also brings Listropsis close to

Emmenotarsus, which resemble each other.

The significance of this latter relation was long undetected, for my early collections

of these two genera consisted of small collections composed wholly or largely of one

sex. Later, obtaining a larger bisexual collection, I sexed it and discovered that all the

males keyed into Listropsis and all the females into Emmenotarsus. This collection

was made above the 5000 foot elevation on Mt. Pisgah, near Yucaipa, San

Bernardino Countv, CA. On reviewing all my collections from this same area, I

found a total of 109 Emmenotarsus-like specimens in seventeen separate collections

made between 1969 and 1981.

A disproportionate sex ratio was found in these 109 specimens with 80 males and

only 29 females. The feature, “protibiae beset with spines” was sex-limited to

females, which had an average of 6.1 spines per protibia, as compared with 0.3 in

males. All the males had the fifth sternite impressed. Therefore, I concluded that the

specimens represented a population of Trichochroides according to Blaisdell, and

that Listropsis and Trichochroides are synonyms.

This conclusion was further substantiated on examination of specimens available

to me, which Blaisdell had labelled Trichochroides. In all examples in which males

were present, they keyed to Listropsis, and the females to Emmenotarsus. On

reviewing Blaisdell’s descriptions of Trichochroides (1941) I found that he never

once made a reference to protibial spines, and thus missed the relationship of these

two genera.

These observations all support the conclusion that Listropsis and Trichochroides

are synonyms. By rules of priority Trichochroides must be suppressed and be

replaced by Listropsis.

The Mt. Pisgah collection is believed to be Listropsis virilis (Blaisdell, 1941),

formerly Trichochroides.

Literature Cited

Blaisdell, E. F. 1921. New species of Melyridae and Chrysomelidae and Tenebrionidae (Coleoptera)

from the Pacific Coast with notes on other species. Stanford U. Publ. Biol. Series I, No. 3,

137-231.

-. 1924a. Two new species of Melyridae from California and one from British Columbia, including

two new genera. Can. Ent., 56:1-5.

-. 1924b. Studies in the Melyridae (Coleoptera) Number four. Trans. Amer. Ent. Soc., 64:313-318.

-. 1938. A generic synopsis and generic revision of the Tribe Dasytini of North America, north of

Panama. Trans. Amer. Ent. Soc., 64:1-32 (two plates).

-. 1941. A monographic study of the species belonging to the Melyrid genus Trichochroides

(Coleoptera). Trans. Amer. Ent. Soc., 66:238-306 (one plate).

Casey, Thomas E. 1895. Coleopterological Notices VI. N.Y. Acad. Sci., 8:435-838.

Fall, H. C. 1901. List of the Coleoptera of Southern California, with notes on habits and distribution and

description of new species. Calif. Acad. Sci. Occasional Papers, 8:1-282.

Howell, Charles D. 1979. Studies of Dasytidae No. 1: New species of Pristocelis (Coleoptera). Pan. Pac.

Ent. 55:41-45.

-. 1985. Diverse populations and subspecies in Eschatocrepis (Coleoptera: Dasytidae). Ent. News,

96:129-141.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 52-55

A New species of Actaletes from Mexico

(Collembola: Actaletidae)

Felipe N. Soto Adames

Department of Biology, University of Puerto Rico, Mayagiiez, Puerto Rico 00708.

Abstract.—Actaletes nemyops new species is described from Sonora, Mexico.

Males possess differentiated setae on the third and fourth antennal segments but lack

the metatibiotarsal spur present in other New World species.

Thanks to the kindness of Drs. Kenneth Christiansen and Peter F. Bellinger I was

able to study several specimens of Actaletes from Mexico which represent the new

species described below. Only four species of Actaletes have been described so far,

one from Europe (France) and three from America (Jamaica, Venezuela and

Mexico). All species are strictly littoral and rarely seen or collected.

The species name, nemyops, refers to the absence of a metatibiotarsal spur in the

male.

Actaletes nemyops New Species

Habitus typical of genus. Length to 1.24mm (x for females 1.03mm; only male

0.83). Head, body and appendages pale brown. Proportions of antennal segments

1-4 as 10:27:22:20. Apex of fourth antennal segment (Ant. 4) with a pin seta on a

papilla (Fig. 3), 3 apically curved setae, 5-6 stout setae and several blunt

sensillae—one being much larger than the others (Figs. 13-14). Ant. 4 of male with a

large seta expanded as a lamella (Fig. 12). Ant. 3 sense organ of 2 mushroomlike

setae on a shallow depression (Fig. 1); a sensillar triangle is opposed to this organ

(Fig. 2). Medial region of Ant. 3 with a prominent spinelike seta (Fig. 5). Apex of

Ant. 2 with a short blunt spine similar to that on A. venezuelensis. Eyes 8 + 8; 6-8 hairs

within eye patch (Fig. 4). Postantennal organ 1.1 x wider than the lower innermost

eye (Fig. 9). Right mandible with 6 teeth (Fig. 18), left mandible with 4 teeth (Fig.

17). Terminal seta of outer maxillary lobe somewhat smaller than basal seta. Labial

triangle with 4 setae; 10-11 setae in an irregular row almost perpendicular to the

cephalic groove (Fig. 16). Male without profemoral or protibiotarsal spinelike setae;

metatibiotarsal spur absent (Fig. 10). Tenent hair lamellar. Ungues with a small

tunica (Fig. 6). One small tooth on unguis II, without teeth on ungues I and III. Inner

margin of unguiculi I and II concave (Figs. 6-7), inner margin of unguiculus III

convex (Fig. 8). Fourth abdominal segment dorsally 2x longer than segments 1-3

combined; with 4 pairs of bothriotricha (Fig. 15). Tergal sutures almost reaching

bothriotrix IT Colophore with 3 + 3 setae. Tenaculum with 5 setae, upper pair smaller

than the others. Ratio length dens: mucro 70:10 in female and 54:10 in male.

Distribution of dental spines as in Figure 19. Inner margin of dens with 4-5 long

setae. Mucro tridentate (Fig. 20), basal and subapical teeth close together but not

facing each other. Mucronal seta present. Male genital plate with 2 differentiated

setae (Fig. 11). Female genital plate without such setae.

VOLUME 63, NUMBER 1 53

Figs. 1-11. Actaletes nemyops new species. 1. Ant. 3 sense organ. 2. Ant. 3 apical triangle of blunt

setae. 3. Apex of Ant. 4. 4. Left eye patch. 5. Ant. 3, ventral view. 6. Mesothoracic claw. 7.

Pro thoracic unguiculus. 8. Metathoracic unguiculus. 9. Lower innermost eye and postantennal organ.

10. Dorsal view of metatibiotarsus. 11. Differentiated seta of male genital plate.

PAN-PACIFIC ENTOMOLOGIST

Figs. 12-20. Actaletes nemyops new species. 12. Modified setae on mid-ventral region of Ant. 4. 13.

Normal blunt seta of Ant. 4. 14. Large blunt seta of Ant. 4. 15. Tergum of fourth antennal segment

showing bothriotrichal pattern and tergal suture. 16. Cephahc groove and seta behind labial triangle.

17. Apex of left mandible. 18. Apex of right mandible. 19. Arrangement of dental spines, thickest

circles represent spines seen directly from above, other circles represent smooth setae. 20. Mucro and

insertion of mucronal seta.

VOLUME 63, NUMBER 1

Diagnosis.—Actaletes nemyops new species can be distinguished from A.

venezuelensis Najt y Rapoport 1972 (Venezuela) by the color of the body, antennal

chaetotaxy, absence of teeth on unguis I, form of third unguiculus and by the distance

between the basal and subapical mucronal teeth. Actaletes nemyops is easily

separated from A. calcarius Bellinger 1962 (Jamaica) and A. boneti Parisi 1972

(Mexico) because males of the latter species possess a conspicuous metatibiotarsal

spur. Actaletes neptuni Giard 1889 (France) can be distinguished from A. nemyops by

the labial chaetotaxy, form of the third unguiculus, relative position of basal and

subapical mucronal teeth and by the presence of sexual dimorphism in^. nemyops.

Material examined. —Mexico, Sonora, on surface of tide pool, 20.11.1974. V.

Roth and W. Brown. Holotype male and 4 paratypes on slides, one paratype in

alcohol. The specimen in alcohol remains in my collection, the holotype and

paratypes are deposited in the Museum of Comparative Zoology, Cambridge,

Massachusetts.

Acknowledgments

I wish to thank Dr. Jose A. Mari Mutt for his guidance and critical review of the

manuscript.

Literature Cited

Bellinger, P. F. 1962. A dimorphic species oi Actaletes (Collembola). J. N. Y. Entomol. Soc., 70:88-91.

Giard, A. 1889. Sur un nuveau genre de Collembole marin et sur I’espece type de ce genre Actaletes

neptuni Gd. Le Naturaliste, 11(53):123.

Najt, J. y H. E. Rapoport. 1972. Una nueva especie de Actaletidae de Venezuela (Insecta, Collembola).

Physis,31(82):219-221.

Parisi, V. 1972. Actaletes boneti —a new species of Actaletidae (Collembola) of the Mexican coast of the

Pacific. Accad. Naz. Lincei, Rome, 171:43-46.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 56-59

New Species of Cleptes Latreille from Asia and North America

(Chrysididae, Hymenoptera)

Lynn Siri Kimsey

Department of Entomology, University of California, Davis, California 95616.

Abstract. —Three new species of Cleptes are described, asiana and townesi from

Taiwan, and canadensis from western Canada.

While examining miscellaneous unidentified Chrysididae from the American

Entomological Institute (GAINESVILLE) and Canadian National Collection

(OTTAWA), 3 new species of Cleptes Latreille became apparent. Two of these are

from Taiwan and the third is from western Canada.

The following abbreviations are used: F-I etc. = flagellomere I and so on,

MOD = midocellus diameter, PD = puncture diameter.

Cleptes asianus Kimsey, New Species

Holotype female. —Body length 5 mm. Face (Fig. 4) punctures small and 1-3 PD

apart; least interocular distance as long as head length from midocellus to antennal

socket; malar space 1.5 MOD long, clypeal truncation 2 MOD wide at apex;

ocellocular distance about three-fourths ocelloccipital distance; F-I length 2.1 times

breadth; F-II 1.1 times as long as broad; pronotum (Fig. 3) somewhat flattened in

profile with row of deep pits across posterior margin, with medial pair largest,

punctures small and 2-3 PD apart; scutal punctures sparser than on pronotum;

mesopleuron (Fig. 7) with deeply impressed somewhat foveate scrobal sulcus

forming a loop with obhque mesopleural carina, mesopleural punctures tiny and 3-5

or more PD apart; propodeum coarsely punctate, lateral angles obtuse; forewing

radial cell about 2.5 times as long as wide. Head, thorax, abdomen, scape, pedicel,

femora and tibiae purple; flagellum and tarsi dark brown to blackish; wings evenly

brown-tinted.

Male. —Unknown.

Holotype female. —TAIWAN: Wushe, 1150 m, 15 May 1983 (H. Townes,

GAINESVILLE).

Discussion. —Based on Tsuneki (1959) asiana most closely resembles seoulensis

Tsuneki a.nd fudzi Tsuneki, based on the mesopleuron having a long foveate scrobal

sulcus connected to a foveate oblique mesopleural carina and the entirely purple

body. However, asiana differs from these species in the following characteristics:

genae only slightly converging, mandibles reddish, clypeal truncation not trilobate,

anterior two-thirds of pronotum parallel-sided and the entire body including femora

and tibiae purple without other tints or highlights.

Cleptes canadensis Kimsey, New Species

Holotype male. —Body length 6 mm. Face (Fig. 1) with deep medial sulcus

extending from midocellus to clypeal margin; punctures 1 PD apart; least interocular

VOLUME 63, NUMBER 1

Figures 1,2,4. Front view of face. 3,5,6. Dorsal view of pronotum. 7. Mesopleuron and midcoxa

showing scrobal sulcus (ss).

distance 1.3 times head length from midocellus to antennal socket; malar space and

clypeal truncation at apex 1.3 MOD long and wide; ocellocular distance equal to

ocellocciptal distance; F-12.8 times as long as broad, F-II length 1.7 times breadth;

pronotum (Fig. 5) evenly rounded with medial groove or posterior groove or pits,

punctures 1-3 PD apart; scutal punctures 4-6 PD apart; mesopleuron smooth with

PAN-PACIFIC ENTOMOLOGIST

deep scrobe, punctures slightly striatiform, about 1 PD apart; propodeum coarsely

and irregularly reticulate, lateral tooth acute but apically blunt. Head, thorax, scape

and femora coppery with strong green highlights becoming darker and bluish on

propodeum, with erect blonde setae; wings evenly and lightly brown stained; rest of

antennae dark brown; tibiae, tarsi and abdominal segments I-II red; abdominal

segment III red basally and black apically; segments IV and V black.

Female. —Same as male, except legs entirely red, with forefemur coppery; F-11.8

times as long as broad; F-II 0.8 times as long as broad, and scape, pedicel and

F-I-III red, F-IV to apex dark brown.

Holotype male. —CANADA: Saskatchewan, 28 June 1950, reared from the

tenthredinid Pikonema alaskensis (Rohwer) (Dalton, OTTAWA). Paratypes:

CANADA: British Columbia, Ft. Nelson, 14 June 1948, W. R. M. Mason (1 male);

Northwest Territory, Norman Wells, 12 July 1949, W. R. M. Mason (1 male and 1

female).

Discussion. —This species is closest to the palearctic species nitidula, based on the

lack of a posterior or medial groove on the pronotum, the reddish basal abdominal

segments and the smooth mesopleuron. Aside from these characteristics canadensis

can be distinguished by the coppery green color of the head and thorax in both sexes,

the male F-I nearly 3 times as long as broad, female F-I twice as long as wide; and

propodeal dorsal surface finely reticulate.

Cleptes townesi Kimsey, New Species

Holotype male. —Body length 5.5 mm. Face (Fig. 2) punctures shallow 0.3-0.5

PD apart; least interocular distance 1.1 times head length from midocellus to

antennal socket; malar space 0.8 MOD, clypeal truncation 2 MOD wide at apex;

ocellocular distance subequal to ocelloccipital distance; F-I length 2.4 times

breadth; F-II 1.8 times as long as broad; pronotum (Fig. 6) flattened in profile, with

posterior subapical transverse groove, punctures 1-2 PD apart; scutal punctures 3-5

PD apart; mesopleuron smooth with small scrobal pit and punctures 0.5-1.0 PD

apart; propodeum sparsely reticulate with large blunt lateral angles; propodeum

laterally and metapleuron smooth and impunctate; T-V medially emarginate. Face

with blue tints; rest of head, thorax, abdomen, antenna and femora dark brown;

body covered with erect blonde setae; wings lightly and evenly brown-tinted.

Female. —Unknown.

Holotype male. —TAIWAN: Wushe, 1150 m, 15 May 1985, H. Townes

(GAINESVILLE). Paratypes: 1 male, same data as type, 10 May 1983; 3 males

TAIWAN: Wu-feng, 10 April 1983, C. Townes.

Discussion. —The most distinctive feature of townesi is the non-metallic black

body. This species most closely resembles crassiceps Tsuneki and thaiensis Tsuneki,

based on the posterior pronotal groove and dark coloration. C. townesi can be

distinguished from these and other species of Cleptes by the simple mesopleuron,

metallic coloration restricted to the face, the pronotum without a medial groove and

the ocellocular distance subequal to the ocelloccipital distance.

Acknowledgments

This study was made possible by Henry Townes at the American Entomological

Institute and Lubomir Masner at the Canadian National Collection, and was funded

by NSF Research Grant No. BSR-8407392.

VOLUME 63, NUMBER 1

Literature Cited

Tsuneki, K. 1959. Contributions to the knowledge of the Cleptinae and Pseninae faunae of Japan and

Korea. Mem. Fac. Lib. Arts, Fukui Univ. (ser. 2, Nat. Sci.) (9):l-78.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 60-64

Observations on Insects Associated with a

Nectar-bearing Chilean Tree,

Quillaja saponaria Molina (Rosaceae)

Robert L. Bugg

Department of Entomology, University of California, Davis, California 95616.

Abstract. —Several species of entomophagous insects were observed feeding on

floral nectar of a specimen of soapbark tree, Quillaja saponaria Molina (Rosaceae), a

landscape plant introduced into northern California from Chile. Entomophaga

observed in relatively large numbers included a green lacewing {Chrysoperla carnea

(Stephens)), convergent ladybeetle (Hippodamia convergens Guerin-Meneville),

and a brown lacewing (Hemerobius sp. (prob. ovalis Carpenter)), as well as various

unidentified parasitic Hymenoptera. Contingency table analyses of weekly vacuum

samples indicated that members of each taxon were significantly more abundant

during, as opposed to after, flowering. Samples taken at different times of day

indicated that the brown lacewing was mainly a nocturnal visitor, whereas the green

lacewing was present at similar densities at all hours tested. These findings suggest

that the soapbark tree should be included in experimental schemes for enhancing

biological control of agricultural pests.

The use of nectar-bearing trees and shrubs in windbreaks and hedgerows has been

suggested as a means of enhancing biological control (Solomon, 1980; Altieri and

Letourneau, 1983). The present study concerns attendance by various

entomophagous insects at the flowers of soapbark tree, Quillaja saponaria Molina

(Rosaceae), a landscape tree introduced into northern California from Chile. These

initial observations serve as a first step in assessing the possible value of this tree in

enhancing biological control of agricultural pests.

In its native Chile, the soapbark tree is said to be responsible for the production of

abundant and exquisite honey (Munoz Pizarro, 1973). In Davis, CA, the plant

flowered from early June through early July, and it appeared to be

andromonoecious, featuring hermaphroditic flowers early in the blossoming period,

and exclusively male flowers thereafter. The flowers exuded a shiny, sticky nectar

that was fed upon by a diverse assemblage of insects. The vast majority of flower

visitors observed were feeding upon nectar rather than pollen. A sampling program

was set up to investigate insect attendance at the tree, with special emphasis on

potential biological control agents.

Materials and Methods

All data were recorded from one specimen of Q. saponaria, approximately 12 m in

height and 4 m in basal diameter, growing at the Environmental Horticulture

Department grounds. University of California, Davis. Specimens were also present

VOLUME 63, NUMBER 1

Table 1. Counts for weekly vacuum samples from Quillaja saponaria.

No. Collected/Subsample/Date

Taxon

Quadrant

6/4

In Flower

6/11

6/18

Out of Flower

6/25 7/2

West

Hippodamia

South

convergens

East

West

Chrysoperla

South

carnea

East

West

Hemerobius sp.

South

(prob. ovalis)

East

West

Parasitic Hymenoptera

South

East

at the Berkeley and Santa Cruz campuses of University of California, but none of

these trees, to my knowledge, flowered appreciably during the year of this study.

In the immediate vicinity of the experimental site were various trees and shrubs

suitable for landscaping in Mediterranean climates; within ca. 2 km were fields of hay

alfalfa and winter wheat. The agricultural fields were likely sources of ladybeetles

and lace wings observed in the present study.

Using the U.C. Vac suction device (Summers et al., 1984), insects were sampled

on 4, 11, 18, and 25 June, and 2 July, 1982. The first three dates occurred during

flowering, whereas the latter two occurred after blossom fall. Samples were taken

during the early afternoon hours, 1300-1530 PDT. The insects were vacuumed from

the flowers and foliage during three one-minute intervals, each corresponding to one

of the three accessible quadrants of the tree (east, south, and west—the north

quadrant was obstructed by an adjoining shrub). During each one-minute episode,

the tree was sampled from ground level to a height of about 2 m. Vacuum samples

were retained in organdy net bags, placed on ice, taken to the laboratory, and frozen.

Samples were later sorted, and the arthropods in the different taxa counted.

In addition to the regular weekly samples mentioned above, supplementary

samples were taken on 5 June at 1200,7 June at 1800,5 June at 2215,12 June at 1800,

and 12 June at 2300 (PDT). Together with samples from one of the regular weekly

visits (11 June), these latter data were used to compare the diel patterns of

attendance by a green lacewing, Chrysoperla carnea (Stephens) (Neuroptera;

Chrysopidae), and a brown lacewing, Hemerobius sp. (prob. ovalis Carpenter)

(Neuroptera: Hemerobiidae).

Visitation by nectarivorous insects was assessed both qualitatively and

quantitatively. A species list was compiled based on the vacuum samples and

observation or collection of flower visitors by insect net. The regular weekly vacuum

data were analyzed using separate chi-square analyses for 1 x 2 contingency tables

constructed for each of the three taxa, C. carnea, convergent ladybeetle

{Hippodamia convergens Guerin-Meneville; Coleoptera: Coccinellidae), and

PAN-PACIFIC ENTOMOLOGIST

parasitic Hymenoptera (all species pooled). As Hemerobius were seldom

encountered in the regular weekly samples, which were all taken during early

afternoon hours, that predator was excluded from these analyses. Comparisons were

made for numbers obtained during flowering vs. those obtained after blossom fall.

Expected values were generated based on the 3 to 2 ratio of sample dates during vs.

after flowering, and the assumption that under the null hypothesis the observed

numbers of insects should follow the same ratio. A significant deviation in favor of

higher attendance during flowering would be taken to indicate that these

entomophaga were attracted to flowers, and not to some unrelated feature of the

tree.

In order to determine whether the green and brown lacewings exhibited

significantly different diel patterns of attendance, the relevant data were subjected to

contingency table analyses via the BMPD-4F program (Dixon, 1983). These data

were arrayed in a three-dimensional contingency table employing species, date, and

time of day (early afternoon, late afternoon, and late evening) as the variables, and

all possible loglinear models were reviewed (see Fienberg, 1977). In the event that a

time X species interaction term were required to explain the data, the green and

brown lacewings could be said to differ significantly in their diel patterns of

attendance.

Results

Several species of insects were found foraging for nectar on the soapbark tree.

These included convergent ladybeetle and the green and brown lacewings

mentioned. Among the parasitic Hymenoptera, unidentified Diapriidae and

Chalcidoidea were most frequently observed. The former were subject to vacuum

sampling, whereas the latter tended to pass through the mesh of the net bags.

Honeybees, Apis mellifera L. (Hymenoptera: Apidae), were commonly collected

from soapbark tree flowers. I also observed various aphidophagous hover flies

(Diptera: Syrphidae) taking nectar; these included Scaeva pyrastri (L.), Eupeodes

volucris Osten Sacken, and Metasyrphus sp. Also, two chloropid flies, probably the

aphidophagous Thaumatomyia glabra (Meig.) and T. rubida (Coquillett), were

observed at the flowers, as was Argentine ant, Iridomyrmex humilis Mayr

(Hymenoptera: Formicidae), and European earwig, Forficula auricularia L.

(Dermaptera: Forficulidae) (earwigs and ants were encountered principally at

night). Minute pirate bug, Orius tristicolor (White) (Hemiptera: Anthocoridae) was

also commonly found. Neither syrphids nor anthocorids were subject to reliable

collection by the vacuum method: the former were too quick to fly, while the latter

were so small that many escaped through the organdy mesh of the net bags.

The results for weekly afternoon suction samples are presented in Table 1. The

mean counts (± S.E.) obtained during flowering were 20.22 ± 3.8, 9.11 ± 1.39, and

10.44 ± 2.44 for/f. convergens, C. carnea, and parasitic Hymenoptera, respectively

(n = 9 for each estimate). The corresponding figures obtained after flowering were

0.67 ± 0.42, 0.50 ± 0.22, and 5.00 ±1.61 (n = 6 for each estimate). Hemerobius

were scant in these diurnal samples that they were not included in these assessments.

The contingency table analyses (d.f. = 1 for each test) revealed highly significant

differences (p < 0.01) for all three taxa assessed {H. convergens,

chi-square = 111.03; C. carnea, chi-square = 47.1; and pooled parasitic

VOLUME 63, NUMBER 1

Table 2. Diel attendance patterns for Chrysoperla carnea and Hemerobius sp. (prob. ovalis) on flowering soapbark

tree.

Total no. per vacuum sample/time/date^

First week in June

Second week in June

Species

LA LE

Chrysoperla

carnea

45 20

Hemerobius sp.

(prob. ovalis)

3 26

®EA = early afternoon, LA = late afternoon, LE = late evening.

Hymenoptera, chi-square = 12.908). These results indicate significantly higher

numbers during flowering for all three taxa.

The data for lacewing attendance, presented in Table 2, indicate a strong

nocturnal tendency for the brown lacewing, whereas the green lace wing was

abundant on the plant at all hours assessed. Totals of 64,71, and 64 were observed for

green lacewings for early afternoon, late afternoon, and late evening, respectively.

The corresponding totals for brown lace wing were 8, 4, and 56. The chi-square

statistics of all possible loglinear models were reviewed, and the most parsimonious

model acceptable under the conventional criterion of p > 0.05 involved all three

main effects plus time x date and time x species interaction terms

(Likelihood-Ratio Chi-Square = 3.84; d.f. == 3; p = 0.2790). The need for the

time X species interaction term indicates that the green and brown lacewings

exhibited significantly different temporal patterns of attendance on the flowering

tree.

The results do not necessarily indicate that the green lacewing adults were

nectar-feeding throughout the day. In fact, the green lacewings were most commonly

observed feeding at flowers during the evening hours, so most individuals were

probably resting in the tree’s foliage during the afternoon and early evening hours.

Perhaps the brown lacewing merely rested diurnally in a different stratum than did

the green, and so was seldom collected except late at night, when feeding. These

questions warrant further investigation.

In summary, the flowering soapbark tree investigated here attracted large

numbers of nectar-feeding predators, several of which are known to be important in

reducing agricultural pests, and which are known to feed on nectar or honeydew (see

Hagen, 1962; New, 1975; Sundby, 1967; Neuenschwander and Hagen, 1980). These

results are suggestive, but because of the limited scope of this study, they should be

viewed with caution. Given its reputation as a “honey plant,” the soapbark tree

might be of some value in urban apiculture, quite apart from any role it might play in

pest management. The nutritional value of the nectar should also be explored.

Finally, the tree should be included in experimental trials of windbreak, hedgerow,

and urban landscape vegetation to determine whether it can enhance biological

control by the predators that feed so avidly at its blossoms.

Acknowledgments

I wish to thank the following people for their assistance: P. Adams, D. A. Andow,

M. A. Altieri, S. Bartholow, L. E. Ehler, R. Fissell, P. Foley, K. S. Hagen

PAN-PACIFIC ENTOMOLOGIST

(identification of coccinellids, neuropterans, and syrphids), M. Hertlein, A. W.

Johnson, J. Johnson (identification of neuropterans), J. Ketcham, W. H. Lange, Jr.

(identification of chloropids), D.K. Letourneau, M. Miller, A. Moldenke, D.

Simser, H. Sauter, W. Settle, W. Roberts (identification of the tree), C. A. and M. J.

Tauber, K. Thorarinsson, L. T. Wilson, S. Whitworth, andF. Zalom. This work was

funded in part by a Graduate Research Award and by the University of California

Appropriate Technology Program.

Literature Cited

Altieri, M. A. and D. K. Letourneau. 1983. Vegetational management and biological control in

agroecosystems. Crop Prot., 1:405-430.

Dixon, W. J. (Chief Editor). 1983. BMPD Statistical Software, 1983Printing With Additions. University

of Cahfornia Press. Berkeley, CA. 734 pp.

Fienberg, S. E. 1981. The Analysis of Cross-Classified Categorical Data. M.I.T. Press. Cambridge, MA.

198 pp.

Hagen, K. S. 1962. Biology and ecology of predaceous Coccinellidae. Ann. Rev. Entomol., 7:289-326.

Munoz Pizarro, C. 1973. Chile: Plantas En Extincion. Editorial Universitaria. Santiago, Chile. 247 pp.

Neuenschwander, P. and K. S. Hagen. 1980. Role of the predator Hemerobius pacificus in a

non-insecticidal treated artichoke field. Environ. Entomol., 9:492-495.

New, T. R. 1975. The biology of Chrysopidae and Hemerobiidae (Neuroptera), with reference to their

usage as biocontrol agents: a review. Trans. Entomol. Soc. Lond., 127:115-140.

Solomon, M. G. 1980. Windbreaks as a source of orchard pests and predators. In: J. M. Thresh (ed.).

Pests, Pathogens and Vegetation: The Role of Weeds and Wild Plants in the Ecology of Crop Pests

and Diseases. Pitman Advanced Pubhshing Program. Boston, MA. 517 pp.

Summers, C. G., R. E. Garrett, and F. G. Zalom. 1984. A new suction device for sampling arthropod

populations. J. Econ. Entomol., 77:817-823.

Sundby, R. A. 1967. Influence of food on the fecundity of Chrysopa carnea Stephens (Neuroptera:

Chrysopidae). Entomophaga, 12:475-479.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 65-74

Cosumnoperla hypocrena, a new genus and species of western

Nearctic Isoperlinae (Plecoptera: Perlodidae)*

Stanley W. Szczytko and Richard L. Bottorff

*Study supported in part by the Univ. of Wisconsin Faculty Research Fund #5413.

(SWS) College of Natural Resources, Univ. of Wisconsin, Stevens Point,

Wisconsin 54481; (RLB) Dept, of Land, Air and Water Resources, Univ. of

California, Davis, California 95616.

Abstract.—Cosumnoperla hypocrena, a new genus and species of Isoperlinae is

described from specimens of male, female, nymph and ova. The major diagnostic

characters of this new genus include: 1. male 10th tergum with a broad, elevated,

notched triangular process; 2. small bulbous male supra-anal process; 3. male vesicle

absent; 4. male paraprocts reduced, flat and lightly sclerotized; 5. female subgenital

plate large with a deep posteromedian notch; 6. ova oblong, collar and eclosion line

absent; 7. nymphal mandibles with 6 teeth and lacinia bidentate; 8. adult and

nymphal mesosternal Y-ridge attached to posterior ends of furcal pits, transverse

ridge connecting anterior ends of furcal pits, and 9. gills absent. Based on these

characters this genus is thought to be most closely related to Calliperla Banks.

Isoperlinae now includes the endemic Nearctic genera Calliperla, Cascasoperla

Szczytko and Stewart, Clioperla Needham and Claassen, and Cosumnoperla, the

Holarctic genus Isoperla Banks, and the Palearctic genera Kaszabia Rauser and

Mesoperlina Klapalek.

Isoperlinae is a small group of 6, mostly monotypic genera including

Palearctic-Oriental endemics {Mesoperlina Klapalek and Kaszabia Rauser),

Nearctic endemics {Calliperla Banks, Cascadoperla Szczytko and Stewart, and

Clioperla Needham and Claassen), and the large Holarctic genus Isoperla Banks.

Ricker a Jewett and Bulgaroperla Rauser have recently been placed in Perlodinae

(Szczytko and Stewart, 1984; Stark and Szczytko, 1984).

In the Nearctic and Palearctic Regions Isoperla contains the greatest diversity of

Isoperlinae species with ca. 95% of the Nearctic and 91% of the Palearctic species.

The number of Isoperla species in North America is divided into western and eastern

faunal segments generally delineated by the eastern edge of the Rocky Mountains

and the Great Plains. The eastern faunal segment is the most diverse with ca. 67% of

the North American Isoperlinae species. The western genera Calliperla and

Cascadoperla have fairly restricted distributions along the coastal range while the

eastern Clioperla is widely distributed (Szczytko and Stewart, 1984).

Recently while R. L. Bottorff was studying the ecology of stoneflies in the

Cosumnes River, El Dorado County California, a series of Isoperlinae were

discovered which did not key to any known genus. Detailed study of the adults

indicated that these specimens represented a new genus and species. This discovery

parallels other recent finds of rare and unique stoneflies in similar habitats (eg.

Viehoperla Stark and Stewart, 1982a; Oconoperla Stark and Stewart, 1982b).

PAN-PACIFIC ENTOMOLOGIST

Materials and Methods

Illustrations of adult and nymphal structures were drawn using a Wild M8 stereo

dissecting microscope equipped with lightfield-darkfield base and camera lucida.

Nymphal mouthparts were drawn from scanning electron micrographs made using

an ISI Super III SEM.

Male and female terminalia were treated for study according to the methods

described by Szczytko and Stewart (1981). Aedeagal armature was studied using the

method of Szczytko and Stewart (1984).

Ova dissected from preserved gravid females were prepared for SEM study as

described by Szczytko and Stewart (1979). Scanning electron micrographs of ova

were made with an ISI Super III SEM.

Terminology used to describe antennal and cereal sensory structures follows

Kapoor (1985); egg terminology follows Stark and Szczytko (unpub.)

Results and Discussion

Cosumnoperla, New Genus

Type species. — Cosumnoperla hypocrena Szczytko and Bottorff Generic

Characters: Adult and nymphal mesosternal Y-ridge arms meet posterior corners of

furcal pits; transverse band connects anterior corners of furcal pits (Fig. 7). Gills

absent.

Male. —Tenth tergum entire with broad, elevated notched triangular process

(Fig. 3). Supra-anal process small, bulbous and lightly sclerotized (Fig. 3).

Paraprocts reduced and lightly sclerotized (Figs. 3, 5). Vesicle absent. Aedeagus

membranous (Fig. 2).

Female. —Subgenital plate large with deep posteromedian notch (Fig. 4).

Ova. —Outline oblong, cross-section circular. Collar absent (Fig. 20).

Nymph. —Lacinia bidentate (Figs. 13, 14). Mandibles with 6 teeth, not deeply

incised (Figs. 8, 9, 16, 18). Abdominal terga with 3 longitudinal dark stripes and 2

median and 6 lateral longitudinal rows of dots (Fig. 10).

Distribution. —This monotypic genus is known only from the Cosumnes River in

the Sierra Nevada Mountains of northern California.

Cosumnoperla hypocrena, New Species

Male. —Macropterous. Body length ll-13mm; forewing length 10-llmm.

General body color light creamy yellow with dark brown markings on the head and

thorax. Dorsum of head with wide dark brown patch connecting lateral and anterior

ocelli, light, small inverted U-shaped spot extending midlength of interocellar area;

lighter brown patches separated medially, extending from lateral ocelli to occiput,

few scattered small spots in light area between light brown patches; light brown

patches extending from dark brown ocellar patch to bases of antennae, thin

U-shaped, light patch above anterior ocellus with dark brown median patch

extending to frons (Fig. 1). Pronotum with median light stripe, disks medium brown,

rugosities dark brown (Fig. 1). Meso- and metanota medium brown. Antennal

flagella medium brown, pedicle light; surface of flagellar segments covered with long

and short sensilla trichodea (Fig. 30) and sensilla companiformia (Fig. 24). Cerci

light basally, progressively darker apically, cereal segments covered with long

sensilla trichodea and with posterior whorl of short and 4 long posteroventral sensilla

trichodea (Fig. 22). Maxillary and labial palpi dark brown. Wings medium smoky

VOLUME 63, NUMBER 1

v wv^'^

V/ w w

Figures 1-5. Cosumnoperla hypocrena. 1. Adult head and pronotum; line = 0.9mm. 2. Male

aedeagus, lateral aspect; line = 0.5mm. 3. Male terminalia, dorsal aspect (A.-tenth tergal process,

B.-supra-anal process, C.-paraprocts); line = 0.5mm. 4. Female terminalia, ventral aspect;

line = 0.8mm. 5. Male paraprocts, dorsal aspect; line = 0.5mm.

PAN-PACIFIC ENTOMOLOGIST

brown with dark brown veins. Dorsal surface of femora dark brown, most of tibia

and tarsi dark brown. Abdomen creamy yellow. Eighth tergum with posteromedian

patch of short stout setae (Fig. 3). Ninth tergum with median patch of stout reddish

brown spinulae (Fig. 3). Elevated triangular process of 10th tergum heavily

sclerotized, darker than rest of tergum, highest at anterior margin with 2 sharply

pointed anterior lobes, margins fringed with long fine setae, base depressed (Fig. 3).

Supra-anal process with dark stout apical spinulae, basal area with scattered small,

fine, light spinulae (Fig. 3). Paraprocts nearly flat, broadly triangular, positioned

below supra-anal process (Figs. 3, 5). Aedeagus with large anteromedian and

posteroventral lobes, small finger-like lobe above posteroventral lobe, apical section

expanded into large balloon-like tip; posterior patch of small stout, golden brown

spinulae above small finger-like lobe, patch of longer stout spinulae below large

posteroventral lobe, patch of medium spinulae on large anteromedian lobe which

grade into short stout spinulae laterally, patch of lateromedian small stout spinulae

posterior to large anteromedian lobe (Fig. 2).

Female. —Macropterous. Body length ll-15mm; forewing length ll-14mm.

Body coloration and external morphology similar to male. Subgenital plate broadly

truncate extending to near posterior margin of 9th sternum, base extending to

midlength of sternum 8 (Fig. 4).

Nymph .—Body length of mature nymph 10-13mm. General body coloration

medium brown. Dorsum of head with distinct broad, light M-shaped band anterior to

median ocellus; broad median brown patch connecting ocelli extending across head

anteriorly to frons with light spots anterior to lateral ocelli and also anterior to light

M-shaped band; interocellar area with irregular shaped light spot; irregular

reticulate light patches posterior to occiput; frontoclypeus light (Fig. 6); occiput with

irregular sinuous row of short spinulae. Dorsum of head and thorax with scattered

fine black clothing hairs. Lacinia triangular with 4-5 long axillary setae between

apical teeth; shoulder with 7-8 long stout spine-like setae below subapical tooth; row

of 5-6 long finer marginal setae below shelf; 5-6 small fine marginal setae scattered

to lacinal base; subapical tooth ca. length of apical tooth (Figs. 13,14); teeth with

fine striations (Fig. 17). Mandibles with median ventral row of long setae from base

of outer tooth extending to near mandibular base; inner mandibular surface concave

with row of long stout marginal setae; dorsal surface with median row of medium

length setae from base of inner tooth to base of marginal setal row; brush of short

stout, thick setae from base of inner teeth to marginal setal row (left mandible with

thicker, longer setal brush) (Figs. 8,9,16,18); inner tooth and subapical tooth with 2

rows of small, shallow crenulations (Fig. 14). Antennae light, distal margin of

flagellar segments with thin and thick wall sensilla trichodea and coniform sensilla

complexes consisting of 4-6 cuticular spines (Figs. 23, 25); conical poreless

coeloconic pegs scattered on flagellar surface (Figs. 27, 29); pedicle surface covered

with scattered thick-walled and thin-walled sensilla basiconica and distal margin with

complete row of long, stout sensilla basiconica (Fig. 23); first flagellar segment with

sparse median row of long sensilla basiconica (Fig. 23). Pronotum with light median

stripe and lateral margins; disks light brown; rugosities light; margin completely

fringed with short stout setae; occasional longer setae at posterior margins.

Meso-metanota medium brown with irregular reticulate light areas. Thoracic and

abdominal sterna with numerous chloride cells on integumental membranes (Fig.

28). Femora light brown with continuous row of long, fine dorsal setae and scattered

VOLUME 63, NUMBER 1

Figures 6-12. Cosumnoperla hypocrena. 6. Nymph head and pronotum; line = 1.0mm. 7.

Nymph mesosternum; line = 0.7mm. 8. Nymph right mandible, ventral aspect; line = 0.3mm. 9.

Nymph left mandible, ventral aspect; line = 0.3mm. 10. Nymph abdomen, dorsal aspect;

line = 1.4mm. 11. Nymph right, hind femora, dorsal aspect; line = 0.7mm. 12. Nymph right, hind

tibia, dorsal aspect; line = 0.7mm.

PAN-PACIFIC ENTOMOLOGIST

Stout setae; dorsal 1/3 of outer surface with numerous medium length, stout setae and

numerous fine black setae (clothing hairs); thin light stripe void of setae below dorsal

1/3; ventral 2/3 with sparser medium length, stout setae and numerous fine black

clothing hairs; no long ventral fringe; inner surface with few scattered medium

length, stout setae (Figs. 11). Tibia with sparse dorsal fringe of long setae; ventral

margin with fringe of medium length stout setae; outer surface covered with long

black clothing hairs (Fig. 12). Abdominal terga with 2 noncontinuous light median,

and 2 lateral longitudinal bands between dark median bands, usually enclosed by

thin dark irregular anterior and posterior bands (may vary to some degree between

individuals) (Fig. 10). Terga with posterior fringe of medium length stout setae and

numerous scattered intercalary spinules (Fig. 19). Cerci light, segments with

posterior whorl of short and several scattered long sensilla basiconica; longitudinal

grooves at posterior margins (Fig. 26).

Ovum. —Length 0.8-0.9mm; width 0.4-0.5mm (fresh ova with living embryos

are 0.6-0.7mm wide). Color light green. Chorion covered with irregular hexagonal

follicle cell impressions (FCFs); FCI walls thick, raised; floor flat, irregularly shaped

(quadrangular to hexagonal). Micropylar row subequatorial; orifices small without

lips, set in floor of FCI’s, some associated with FCI rosettes. Eclosion line absent

(Figs. 20,21).

Distribution. —This species is known only from the type locality.

Material Examined. —USA. California. El Dorado Co., 5.8 km E. of Somerset,

Unnamed trib., 500m upstream from Sweeneys Crossing Bridge on North Fork

Cosumnes River, 28-V-1983, R. L. Bottorff (3 females), 4-VI-1983 (2 females),

24-VI-1983 (1 female), ll-V-1984 (6 nymphs, 3 males, 1 female),

27-V-18-VI-1984 (3 males, 2 females), 22-VI-1984 (1 female), 24-VI-1984 (1

female), 29-V-1985 (1 male, lab-reared), 22-VI-1985 (2 females, lab-reared),

24-VI-1985 (1 male, 7 females lab-reared).

Types. —Holotype male, allotype female and 1 paratype nymph from the above

locality deposited in the U.S. National Museum, one male, one female and one

nymph paratypes deposited in the Brigham Young University collection. Six male,

18 female and 4 nymph paratypes deposited in the collections of S. W. Szczytko and

R. L. Bottorff.

Etymology. —The genus was named in honor of the Cosumnes River and a tribe of

Miwok Indians in central California. The species name is derived from the

“hypocrenon” (head water tributary stream) of lilies and Botosaneanu’s (1963) river

classification scheme, as a habitat descriptor of this species.

Biological Notes. —The type locality is shallow spring water flowing over moss

covered rocks and is heavily shaded. The stream only flows about 7 months

(November-June) each year, then is dry in summer and autumn. Intensive

collecting along the main Cosumnes River, its North Fork, and a few smaller

tributaries, involving slit traps (Kuusela and Pulkkinen, 1978) and searching

methods, as part of a larger ecological study of all Cosumnes River stoneflies, failed

to produce any additional specimens of this species. The restricted habitat

preference of C. hypocrena has, most likely, precluded previous collection by other

workers.

Emergence begins the first week in May and extends through late June, or until the

stream dries up. Males emerge before females and continue into early June. Females

begin emerging in late May.

VOLUME 63, NUMBER 1

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Figures 13-21. Cosumnoperla hypocrena. 13. Nymph right, lacina, ventral aspect; line = 263

microns. 14. Detail of inner left mandibular tooth, ventral aspect; line = 26 microns. 15. Nymph left

lacina, ventral aspect; line = 263 microns. 16. Detail of nymph left mandibular teeth, ventral aspect;

line = 132 microns. 17. Detail of right, apical lacinial tooth, ventral aspect; line = 4 microns. 18.

Detail of nymph right mandibular teeth, ventral aspect; line = 132 microns. 19. Nymph second and

third abdominal terga, dorsal aspect; line = 208 microns. 20. Ova; line = 208 microns. 21. Detail of

ova chorion and micropyle; line = 25 microns.

PAN-PACIFIC ENTOMOLOGIST

This species has the largest egg of any described Isoperlinae species. Most Isoperla

species have eggs which range from 0.21-0.45 mm-length and 0.13-0.36 mm-width

(Jop and Szczytko, 1984; Nelson and Kondratieff, 1983; Szczytko and Stewart, 1976,

1978,1979 and 1984). The egg of Clioperla clio (Newman) is 0.42 mm-length and 0.31

mm-width (Szczytko and Stewart, 1981) and the egg of Calliperla luctuosa (Banks) is

0.36 mm-length and 0.28 mm-width (Szczytko and Stewart 1984). The egg is also

fairly atypical of most Isoperlinae species in that the collar and eclosion line are

absent, although C. luctuosa also has an egg with no collar or eclosion line (Szczytko

and Stewart, 1984).

Females of this species have the lowest fecundity of any reported Isoperlinae

species. Eggs from 9 gravid females with abdomens extended with well developed

eggs, were counted and mean fecundity was 39 with a range of 19-58 eggs. Harper

(1973) reported mean fecundities of 441 (range-113-788) for C. clio, 95

(range-23-190) for Isoperla transmarina (Newman), 177 (range-17-392) for I.

cotta Ricker and 146 (range-18-277) for /. frisoni Prison. These data were

generated from lab-reared and field-collected specimens and the determined means

are most likely low, due to field oviposition. The low fecundity of C. hypocrena is

probably related to the large size of the egg and the physical restriction of space

within the female abdomen.

Several nymphs were dissected and the gut contents examined for food items. Two

nymphs had several culicid larvae, as well as diatoms in the hind gut, indicating that

this species is probably omnivorous in late instars.

Diagnosis. — Cosumnoperla and Calliperla share the following synapomorphies;

mostly membranous male supra-anal process, reduced male paraprocts not recurved

to level of 10th tergum and large oblong ova without collar and ecolosion line.

Cosumnoperla males can be distinguished from Calliperla by aedeagus lacking

tubular, striated apical portion, vesicle absent from 8th sternum, supra-anal process

without finger-like apical section, 10th tergal process elevated and triangular and

head pattern with interocellar light spot. Females can be separated by the large,

truncated, deeply notched subgenital plate and light interocellar spot of the head

pattern. Nymphs can be differentiated by the longer subapical lacinal tooth, 4-5

axiallary setae and '^/s long stout spine-like setae on shelf, cereal segments without

long dorsal setae, transverse ridge connecting anterior corners of mesosternal furcal

pits, dark lateral abdominal bands without light spots and head pattern with light

interocellar spot. Ova can be separated by larger size, FCI floors not punctate,

micropyles positioned in FCI floors and some associated with FCI rosettes.

Conclusions

Cosumnoperla is apparently most closely related to Calliperla. This relationship is

supported by synapormorphies exhibited in males, females and ova discussed above.

Cosumnoperla also shares characters with Mesoperlina such as modified male 10th

tergum, vesicle absent, distinct spinule patches on male 9th tergum, presence of

supra-anal process and transverse ridge connecting anterior corners of mesosternal

furcal pits. The phylogenetic relationships of Cosumnoperla and other Nearctic

Isoperhnae genera with Mesoperlina and Kaszabia are suspect at this time due to the

paucity of material available for critical study of all life stages.

Cosumnoperla shares characters with other Isoperlinae genera such as absence of

gills and mesosternal Y-ridge attached to posterior corners of furcal pits, but also

VOLUME 63, NUMBER 1

Figures 22-30. Cosumnoperla hypocrena. 22. Adult cerci; line = 250 microns. 23. Nymph

antennae, basal segments; line = 132 microns. 24. Detailed adult antennal segment (A.-sensilla

companiformia); line = 8 microns. 25. Detailed adult antennal segment with coniform sensialla

complex and cuticular spines; line = 18 microns. 26. Nymph cerci, ventral aspect; line = 125 microns.

27. Nymph antennal segment (A.-conical poreless coeloconic peg, B.-thick-wailed sensilla basiconica);

line = 26 microns. 28. Nymph chloride cell; line = 4 microns. 29. Detailed nmph antennal segment

with conical, poreless coeloconic peg; line = 4 microns. 30. Adult antennal segment; line = 63

microns.

PAN-PACIFIC ENTOMOLOGIST

shares characters such as male supra-anal and 10th tergal processes, with some

Perlodinae.

Acknowledgments

We thank Katherine A. Clarke-Girolamo for the nymph and adult head pattern

and nymph abdomen drawings and T. Remnsen and the Great Lakes Research

Facility for use of their SEM and lab. We also thank Drs. B. P. Stark, A. W. Knight,

R. W. Baumann and N. N. Kapoor for helpful suggestions and review of the

manuscript.

Literature Cited

Harper, P. P. 1973. Emergence, reproduction and growth of setipalian Plecoptera in southern Ontario.

Okios, 24:94-107.

lilies, J. and L. Botosaneanu. 1963. Problemes et methodes de la classification et de la zonation

ecologique des eaux courantes, considerees surtout du point de vue faunistique. Mitt. Int. Verein.

Theor. Angew. Limnol., 12:1-57.

Jop, K. and S. W. Szczytko. 1984. Life cycle and production of Isoperla signata (Banks) in a central

Wisconsin trout stream. Aquatic Insects, 6:81-100.

Kapoor, N. N. 1985. External morphology and distribution of the antennal sensilla of the stonefly,

Paragnetina media (Walker) (Plecoptera: Perlidae). Int. J. Insect Morphol. Embryo!.,

14:273-280.

Kuusela, K., and H. Pulkkinen. 1978. A simple trap for collecting newly emerged stoneflies (Plecoptera).

Oikos, 31:323-325.

Nelson, C. H. and B. C. Kondratieff. 1983. Isoperla major, a new species of eastern Nearctic Isoperlinae

(Plecoptera: Perlodidae). Ann. Entomol. Soc. Amer., 76:270-273.

Stark, B. P. and K. W. Stewart. 1982a. The nymph of Viehoperla ada (Plecoptera: Peltoperlidae). J.

Kans. Entomol. Soc., 53:494-498.

-, and K. W. Stewart. 1982b. Oconoperla, a new genus of North American Perlodinae (Plecoptera:

Perlodidae). Proc. Entomol. Soc. Wash., 84:746-752.

-, and S. W. Szczytko. 1984. Egg morphology and classification of Perlodinae (Plecoptera:

Perlodidae). Annls. Limno., 20:99-104.

Szczytko, S. W., and K. W. Stewart. 1976. Three new species of Nearctic Isoperla (Plecoptera). Great

Basin Natur., 36:211-220.

-, and -. 1978. Isoperla bilineata: Designation of a neotype and allotype, and further

descriptions of egg and nymph. Ann. Entomol. Soc. Amer., 71:212-217.

-, and -. 1979. The genus Isoperla of western North America: holomorphology and

systematics, and a new stonefly genus Cascadoperla. Mem. Amer. Entomol. Soc., 32:1-120.

-, and-. 1981. Reevaluation of the genus Clioperla. Ann. Entomol. Soc. Amer., 77:563-569.

-, and-. 1984. Descriptions of Calliperla Banks, Ricker a Jewett, and two new western

Nearcticspecies (Plecoptera: Perlodidae) Ann. Entomol. Soc. Am., 77:251-263.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 75-91

Diversity, Seasonality, and Annual Variability of

Caddisfly (Trichoptera) Adults from Two Streams in

the California Coast Range

Eric P. McElravy and Vincent H. Resh

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

California 94720

Abstract .—Faunal composition and fluctuations in abundance of Trichoptera

adults were analyzed from three years of pan trap collections (1979-1981) made

during the dry season (April-October) at two streams (Big Sulphur Creek, Sonoma

County; Big Canyon Creek, Lake County) located 11 km apart in the California

Coast Range. At least 57 species in 15 families were identified among the 2003

individuals collected; 24 species were common to both sites. The length of the flight

period of these adults, on average, was significantly longer than that observed at four

temperate sites in eastern North America. Most probably, this is related to the

warmer temperature regime associated with the Mediterranean climate of the

California Coast Range. When years with varying amounts of precipitation were

compared, the annual variability of Coast Range Trichoptera (as measured by

year-to-year changes in population abundances) was found to be similar to annual

variability observed for other temperate caddisfly faunas. When precipitation

amounts were similar, year-to-year differences in caddisfly abundance were reduced.

Introduction

Environments that have stable climatic characteristics have been assumed to

exhibit greater faunal and floral diversity (Klopfer, 1959). As a result, tropical areas

have been assumed to contain faunas that are less variable temporally than their

temperate counterparts (MacArthur, 1972). Wolda (1977, 1978, 1980a, b) and

Wolda and Fisk (1981) examined the temporal variability of insect faunas from

(mostly terrestrial) habitats worldwide and concluded that tropical insects are often

less seasonal than temperate insects, but they also observed that tropical insect

populations are not generally more stable on a year-to-year basis (i.e., climatic

stability is not reflected in the annual variability of the insect populations).

Temperate/tropical comparisons of seasonality and annual variability for aquatic

insects are more difficult because most research has been done in areas with

continental temperate climates (hot summers, cold winters) and over relatively short

terms (^2 years). McElravy et al. (1981,1982) compared the diversity and temporal

variability of an adult caddisfly fauna from a climatically stable “non-seasonal”

tropical environment in Panama with several faunas from continental temperate

climates in North America and Europe. They found that tropical caddisflies

exhibited higher diversity and have longer flight periods than adults of most

temperate species, but that these caddisflies did not show an increase in the stability

of their populations on a year-to-year basis.

PAN-PACIFIC ENTOMOLOGIST

Within the temperate-zone areas of the world there are regions with climatic

differences that are analogous to those that occur between the temperate and

tropical zones. For example, the streams of the California Coast Range are not

tropical, but they are located in a region with a Mediterranean-type climate (i.e.,

hot, dry summers and mild, wet winters). Subfreezing temperatures are uncommon

as a result of maritime influences, and insect faunas in these areas are subject to less

seasonal temperature variation than is found in most other temperate-zone sites.

Temperature, along with photoperiod, is known to be an important factor in

determining several life history features of aquatic insects (e.g., Hynes, 1970; Butler,

1984; Sweeney, 1984).

Does the diversity and temporal variability of caddisflies from temperate-zone

streams in regions with Mediterranean climates differ from those in more typical

temperate systems (i.e., hot summers, cold winters)? This paper presents the results

of a three-year study of Trichoptera adults from two nearby streams in the California

Coast Range, and compares the diversity and temporal variability of this fauna with

that of other temperate-zone caddisfly communities.

Study Area

Big Sulphur Creek, Sonoma County, California, is a third-order stream that flows

northwesterly through The Geysers Known Geothermal Resources Area (KGRA).

The stream traverses a steep-sided valley characterized by an unstable terrain

(Brown and Jackson, 1974; Neilson, 1975). In this region, more than 95% of the

annual precipitation occurs during the rainy season (late September-early May).

Following cessation of rains in the spring, streamflows gradually recede, reaching a

minimum level by early September. Extensive blooms of the green alga Cladophora

glomerata (L.) Kutzing can occur within the stream over the summer. Collections of

caddisfly adults were made near the confluence of Big Sulphur Creek and a major

tributary. Little Geysers Creek (38° 46'N, 122° 45' W, elevation 680m, gradient

47m/km).

Big Canyon Creek is located in a separate drainage basin approximately 11 km

northeast of Big Sulphur Creek in Lake County, California. As at Big Sulphur

Creek, most precipitation and maximum discharge occurs from late September to

early May. Samples were collected near the headwaters of Big Canyon Creek (38° 51'

N, 122° 41' W, elevation 570m, gradient 61m/km). At this location, the watershed is

covered with a mixed hardwood-conifer forest, which provides dense shading for a

large portion of the stream.

Although Big Sulphur Creek and Big Canyon Creek are both within The Geysers

KGRA and are subject to similar climatic influences, some differences exist between

the two stream systems. First, the liquid-dominated, subsurface, geothermal

reservoir in the Big Canyon Creek watershed produces alkaline conditions in the

overlying streams; in contrast acidic conditions predominate at Big Sulphur Creek

where the geothermal reservoir consists of dry steam (McColl et al., 1978). Second,

the presence of numerous springs in the upper reaches of the Big Canyon Creek

watershed (and near the study site) reduce seasonal variation in discharge by

maintaining higher summer streamflows than in Big Sulphur Creek. Third, much of

the substrate at Big Canyon Creek consists of large particles that are “cemented” in

place. Compared to the loose gravel-boulder substrate at Big Sulphur Creek, the

compacted substrate of Big Canyon Creek is less likely to be displaced during the

VOLUME 63, NUMBER 1

spates that occur in both watersheds during the wet season; however, this substrate

provides less interstitial habitat. Finally, the riparian vegetation at the Big Canyon

Creek study site is largely deciduous and quite dense, forming a canopy that provides

shading during most of the day. In contrast, the canopy at Big Sulphur Creek is

intermittent and the stream is only partially shaded, particularly during the middle of

the day. The increased shading, lack of surface geothermal inputs, and presence of

numerous springs produce a temperature regime at Big Canyon Creek that is more

constant on a seasonal basis than the temperature regime of Big Sulphur Creek.

Further descriptions of these streams are provided by Lamberti (1983), Lamberti

and Resh (1983), McColl et al. (1978), and McMillan (1985).

Methods

Rainfall .—During most of the period covered by this study daily precipitation

data for The Geysers, California, were available from National Oceanic and

Atmospheric Administration records (NOAA, Hourly Precipitation Data,

California, 1979-1981). Estimates of rainfall for periods with missing data were

calculated by the normal-ratio method (Gilman, 1964) using records from

surrounding stations.

Collection Methods .—Caddisfly adults were collected using pan traps, which are

28 X 40 cm aluminum pans supported 20-30 cm above the stream surface by a metal

frame. The pans are filled with a 50-50 mixture of ethylene glycol and water, and

capture and preserve adult insects for periods up to one month (see Resh et al.

[1984a, b] for pictures of pan traps and a further description of their use). Although

fewer adult insects are collected with pan traps than with traditional attractive traps

(such as light traps), we have found that these traps are generally nonselective and

that samples obtained provide an accurate, relative estimate of adult insect

abundance. Jones and Resh (unpublished data) compared pan trap and Malaise trap

captures in a Montana stream; with one exception, pan traps collected all species that

had > 1 individual in the Malaise trap collections.

At Big Sulphur Creek, four pan traps were placed streamside and continuous

collections were made from the end of the rainy season (mid April-early May) until

the end of the summer dry season (September) in 1979 and 1981, and until the end of

October during 1980. At Big Canyon Creek, four pan traps were used from July until

the end of August in 1979, and two to four pan traps were operated from early May

until late October in 1980 and until late September in 1981. Pan traps were usually

emptied bi-weekly by draining through a 250 p.m sieve. Caddisflies were separated

from other insects, collected, transferred to alcohol, and identified.

Data Analysis .—Seasonal Range (SR) (Wolda, 1979) is a statistic that measures in

weeks the length of the adult flight season of a population, correcting for sample size

(SR can be calculated for samples N 5= 6 in a given year, Wolda, 1979). Higher values

of SR indicate less seasonality for a species (i.e., the adults have longer flight periods

during the year). Seasonal Range was calculated for those species in which six or

more individuals were collected in a given year (SR was not calculated for Big

Canyon Creek species during 1979 because of the short collecting season). In order

to compare the seasonality of the fauna at each site with other temperate sites, the SR

values for the populations at a given site are arranged in a frequency distribution and

differences between sites are analyzed with tests.

Year-to-year differences in Seasonal Range values were examined for species in

PAN-PACIFIC ENTOMOLOGIST

which SR values were available for two or more years. The Annual Variability

statistic (AV) of Wolda (1978) provides a measure of the stability of the community

in terms of the variability of its constituent populations. This parameter was

calculated for the caddisfiies at Big Sulphur Creek for the years 1979-1980,

1980-1981, and 1979-1981, and at Big Canyon Creek for 1980-1981; the AV

statistic was not calculated for other years due to low numbers of captures. Species

for which ^ 10 individuals were collected in the two years that were being compared

were included in the analysis. To make years comparable, data from October 1980

and April 1981 were omitted from Big Sulphur Creek samples, whereas the number

of captures at Big Canyon Creek in 1981 was doubled because only two pans were

operated (cf. four pans in 1980). The values obtained for both sites were compared

with previously reported values of this statistic for temperate and tropical caddisfly

communities.

Results and Discussion

Rainfall .—Precipitation data for the rainy season (September-May) at The

Geysers indicates a 25-yr. mean (±SD) of 145 ±38.3 cm yr“k Total precipitation

during the 1979-1980 rainy season (155.1 cm) was near this average. However, the

1978- 1979 and 1980-1981 totals were below average (98.4 cm and 90.5 cm,

respectively); hence the 1979 and 1981 collections were made during “dry” years.

The Fauna.—During the three-year sampling period 1979-1981 at Big Sulphur

Creek, 1039 Trichoptera adults were collected. Although females could not be

associated with males for 183 (18%) of these adults, the remaining 856 specimens

represent 39 species in 14 families (Table 1), At least 10 of the species (26%) appear

to be undescribed (D. Denning, pers. comm.). Since some of the unassociated

females are probably not represented by males in the 39-species count, the actual

number of species collected is most likely higher.

At Big Canyon Creek, a total of 964 adult Trichoptera was collected during

1979- 1981, including 317 unassociated females (33%). The remaining 647

individuals represented 42 species in 15 families (Table 2). At least seven of these

species (17%) appear to be undescribed (D. Denning, pers. comm.). A total of 24

species was common to both streams, and at least 57 species were collected from

these two sites.

The Trichoptera diversity of Big Sulphur Creek and Big Canyon Creek may be

compared with other sites by examining the relationship between the number of

species and the number of individuals collected (i.e., the species-abundance curve).

These two California sites are near the center of the diversity-abundance

distributions for temperate-zone sites in North America and Europe examined by

McElravy et al. (1981) and, in terms of diversity, they represent typical temperate

caddisfly communities. The proportion of undescribed species (17-26%), however,

is much higher than usual (cf. Resh et al. [1975] for a Kentucky stream, Morse et al.

[1980] for a South Carolina stream).

Seasonal Occurrence .—The seasonal occurrence of the caddisfly adults at Big

Sulphur Creek that composed ^ 1% of the total number collected from 1979 to 1981

is presented in Fig. 1. Although year-to-year variation is apparent for most of these

species, some general patterns can be observed. Most species have adults emerging

and active over a fairly long time period in at least one of the years. Only two species,

Rhyacophila angelita Banks diVid Amiocentrus aspilus (Ross) have short flight periods

VOLUME 63, NUMBER 1

BIG SULPHUR CREEK

APRIL MAY JUNE JULY AUGUST SEPTEMBER OCTOBER

I_I_I_I_I_I_I_I

Wormaldia undescr. sp. A

Tinodes consuetus

Hydropsyche philo

Hydropsyche occidentalis

Hydropsyche oslari

Rhyacophila arrgelita

Hydroptila arctia

Hydroptila sp. nr. rono

Amlocentrus aspllus

Gumaga nigricula

Helicopsyche borealis .

Figure 1. Seasonal occurrence of 11 numerically dominant 1% of total 1979-1981) species of

Trichoptera collected at Big Sulphur Creek 1979-1981. Values of Seasonal Range (SR) calculated

according to Wolda (1979). Dotted lines indicate periods when pan traps were not operated, and for which

flight information is not available.

in the spring. The patterns of seasonal occurrence of the 13 species of Trichoptera

adults that composed ^ 1% of the total collected from 1979-1981 at Big Canyon

Creek (Fig. 2) were similar to those observed for the Big Sulphur Creek fauna. Only

the Rhyacophilidae (except for Rhyacophila vao Milne) and Amiocentrus aspilus

have short flight periods in the spring.

The seasonal occurrence of the Trichoptera faunas at the two northern California

sites was compared with other temperate-zone sites by examining the frequency

distributions of the SR statistic between two sites using tests. When the

distribution of SR for the Big Sulphur Creek fauna (Fig. 3a) is compared with the

four temperate sites in eastern North America described in McElravy et al. (1982),

the fauna at Big Sulphur Creek is significantly less seasonal in all cases (p < 0.05,

Table 3); that is, caddisfly adults at Big Sulphur Creek have longer flight periods than

the other sites. When a similar comparison is made for the Big Canyon Creek fauna

(Fig. 3b), all sites except Mahoning River, Ohio, St. #3, were also significantly less

seasonal (p < 0.05, Table 3).

These California sites are in an area that is climatically different from the other

sites. Most obviously. Big Sulphur Creek and Big Canyon Creek have higher mean

air temperatures than the other sites during late fall to early spring (Table 3). Asa

result, these California streams are not subjected to long periods of near-freezing or

subfreezing conditions over the winter, which occur at the other sites that are, at least

in respect to temperature, located in a much more severe climate. We assume that

these higher temperatures permit longer flight periods for the adult caddisflies by

PAN-PACIFIC ENTOMOLOGIST

BIG CANYON CREEK

APRIL MAY JUNE JULY AUGUST SEPTEMBER OCTOBER

1_I_I_I-1-1-1-1

t97»|

Dolophilodes dorcus issol

1981-I

Hydropsyche occidentalis

Hydropsyche oslari

Rhyacophila pellisa

Rhyacophila vao

Agapetus taho

Hydroplila arctia

Hydroptila sp. nr. rono

Amlocentrus aspllus

MIcrasema undescr. sp. A

Lepidostoma cantha

Marilia Ilexuosa

Helicopsyche borealis .

Figure 2. Seasonal occurrence of 13 numerically dominant (> 1% of total 1979-1981) species of

Trichoptera collected at Big Canyon Creek 1979-1981. Values of Seasonal Range (SR) calculated

according to Wolda (1979). Dotted lines indicate periods when pan traps were not operated, and for which

flight information is not available.

providing longer periods of favorable weather conditions during early spring and late

autumn.

Annual Variability. —At Big Sulphur Creek, the number of taxa varied little over

the three years of this study (Table 1). However, the total numbers of individuals

varied greatly. For example, the relative abundance of Helicopsyche borealis

(Hagen), which as larvae are important grazers in this stream (Lamberti and Resh,

1983), and Hydropsyche spp. was fairly constant over 1979-1980, but was reduced in

1981 (Table 1). Gumaga nigricula (McLachlan), another numerically dominant

macroinvertebrate in Big Sulphur Creek (Resh et al., 1981), was less abundant

during 1980 than in 1979 or 1981. The 1980 decrease in G. nigricula may be related to

the higher rainfall totals and increased substrate disturbance the previous winter,

which has been shown to adversely affect G. nigricula densities (Resh et al., 1981).

Further evidence that substrate disturbance resulting from high winter discharge

events can reduce caddisfly densities was obtained when Big Sulphur Creek pan trap

collections were made in 1986. The 1985-1986 rainy season included two 50-year

floods, both occurring in February, that resulted in considerable rearrangement of

stream bed materials, including displacement of large boulders and logs. Only 10 G.

nigricula adults were collected during 1986, one-half of the 1980 total and less than

one-sixth of the numbers of G. nigricula collected in 1979 or 1981. A similar trend

was observed for total numbers of individuals (89 in 1986; cf. 501 in 1979,252 in 1980,

VOLUME 63, NUMBER 1

0 10 20 30 40 50

SEASONAL RANGE

Figure 3. Histograms showing distribution of Seasonal Range (SR) as percentages. Values along abcissa

from left to right indicate decreasing seasonality. Trichoptera from A. Big Sulphur Creek (N = 24); B. Big

Canyon Creek (N = 15).

and 286 in 1981) as well as number of taxa (14 in 1986; cf. 23 in 1979,25 in 1980, and

26 in 1981).

At Big Canyon Creek, the number of taxa collected was similar during the two

years for which sampling was complete (1980 and 1981) (Table 2). However, the total

number of individuals collected decreased from 1980 to 1981. Over this period, the

relative abundance of the Lepidostomatidae increased, while that of the

Brachycentridae declined. Little change in abundance of Rhyacophilidae was

observed (Table 2).

Year-to-year changes in the seasonality of the Trichoptera fauna at Big Sulphur

Creek and Big Canyon Creek can be examined with between-year plots of SR for

those species with N ^ 6 in two or more years. Since SR is corrected for sample size,

this procedure allows data, such as presented in Figs. 1 and 2, to be compared. In

both 1979-1980 and 1980-1981, the numerically dominant caddisfly species at Big

Sulphur Creek did not exhibit similar seasonality patterns over the two years (Fig.

4a,b). This may be related to variation in abiotic and/or biotic factors caused by

differences in hydrologic regimes between the two relatively dry years (1979, 1981)

82 PAN-PACIFIC ENTOMOLOGIST

Figure 4. Between-year comparison of seasonal occurrence of Trichoptera adults (measured by SR) at

two sites in the central California Coast Range. A. Big Sulphur Creek 1979-1980; B. Big Sulphur Creek

1980-1981; C. Big Sulphur Creek 1979-1981; D. Big Canyon Creek 1980-1981.

and a wetter year (1980). When the two dry years are compared (Fig. 4c), less

year-to-year variability in the length of the species’ active seasons is evident. Mean

wet season stream temperature was probably less important than precipitation in

producing this pattern; temperature did not vary between 1979-1980

(October-April mean temperature = 11.4° C) and 1980-1981 (October-April

mean temperature = 11.3°) and was lower during 1978-1979 (October-April mean

temperature = 9.6°). At Big Canyon Creek (Fig. 4d), a comparison of SR values for

1980-1981 also showed considerable variation for three of the six species examined;

these three species had considerably longer active seasons in 1980 than in the

relatively dry year of 1981.

Year-to-year variability of the entire fauna at a particular site can be assessed using

the Annual Variability statistic (AV) (Wolda, 1978), which takes into account

changes in total numbers of each species over the years of comparison. When this

statistic is calculated for Big Sulphur Creek (Table 4), the pattern of faunal variation

is similar to that seen above when the SR values were plotted, i.e., less variation

occurs when the two drier years (1979 and 1981) are compared than when a dry and a

VOLUME 63, NUMBER 1

Table 1. Species present, number of individuals collected, and abundance as percent of individuals collected in each

year, from Big Sulphur Creek, 1979-1981.

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Philopotamidae

Chimarra undescr. sp. A

0.3

0.0

0.7

Wormaldia undescr. sp. A

8.8

4.7

7.3

Psychomyiidae

Tinodes consuetus McL.

4.8

3.9

3.5

Tinodes siskiyou Denning

0.0

0.8

0.0

Polycentropodidae

Polycentropus halidus Milne

0.0

0.4

0.0

Unassociated Polycentropus

Females:

Polycentropus sp. #1

0.6

0.0

Polycentropus sp. #2

0.2

0.0

Hydropsychidae

Cheumatopsyche mickeli

Denning

0.6

2.0

0.0

Homoplectra oaklandensis

(Ling)

0.2

0.0

Hydropsyche californica Banks

0.0

0.8

0.0

Hydropsyche philo Ross

10.2

9.8

4.9

Hydropsyche occidentalis

Banks

1.0

2.7

0.7

Hydropsyche oslari Banks

0.8

2.4

Hydropsyche sp. A

0.2

0.0

0.3

Parapsyche almota Ross

0.2

0.0

1.0

Rhyacophilidae

Rhyacophila angelita Banks

0.2

1.2

4.2

Rhyacophila vedra Milne

0.0

0.3

Rhyacophila vocala Milne

0.2

0.8

1.4

Rhyacophila undescr. sp. A

0.2

0.0

0.7

Rhyacophila undescr. sp. B

0.0

0.4

0.3

Rhyacophila undescr. sp. C

0.0

0.4

0.0

Unassociated Rhyacophila

Females:

Rhyacophila sp. #1

0.2

0.8

1.4

Rhyacophila sp. #2

0.0

1.2

5.6

Rhyacophila sp. #3

0.0

0.3

Rhyacophila sp. #4

0.0

0.3

Glossosomatidae

Unassociated Agapetus

Female:

Agapetus sp. #1

0.0

0.3

PAN-PACIFIC ENTOMOLOGIST

Table 1. continued

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Glossosoma califica Denning

0.2

0.0

Glossosoma oregonense Ling

0.0

0.3

Unassociated Glossosoma

Females:

Glossosoma sp. #1

0.0

0.8

0.0

Glossosoma sp. #2

0.0

1.0

Hydroptilidae

Hydroptila arctia Ross

6.2

18.7

7.7

Hydroptila sp. A nr.

rono Ross

3.6

3.5

Unassociated Hydroptila

Females:

Hydroptila sp. #1

7.8

4.7

2.8

Hydroptila sp. #2

0.0

0.4

1.4

Ithytrichia sp. A

0.0

0.4

0.0

Neotrichia sp. A

0.2

0.0

Neotrichia undescr. sp. A

0.0

0.4

0.0

Unassociated Neotrichia

Female:

Neotrichia sp. #1

2.2

0.8

0.0

Ochrotrichia undescr. sp. B

0.0

0.3

Ochrotrichia undescr. sp. C

0.0

0.3

Ochrotrichia \md&sci. sp. D

0.0

0.7

Ochrotrichia undescr. sp. E

0.2

0.0

0.3

Unassociated Ochrotrichia

Female:

Ochrotrichia sp. #1

0.8

0.3

Oxyethira sp. A

0.0

2.0

0.0

Unassociated Hydroptilidae

Females:

Hydroptilidae sp. #1

0.0

1.2

0.0

Brachycentridae

Amiocentrus aspilus (Ross)

0.8

1.2

0.7

Unassociated Amiocentrus

Females:

Amiocentrus sp. #1

0.3

0.0

Amiocentrus sp. #2

1.0

1.2

4.9

VOLUME 63, NUMBER 1

Table 1. continued

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Lepidostomatidae

Lepidostoma strophis Ross

1.4

0.0

0.3

Unassociated Lepidostoma

Females:

Lepidostoma sp. #1

3.0

1.2

0.7

Lepidostoma sp. #2

0.0

0.4

0.0

Sericostomatidae

Gumaga nigricula (McL.)

15.6

7.9

21.7

Odontoceridae

Mariliaflexuosa Ulmer

0.0

0.4

0.0

Unassociated Marilia

Females;

Marilia sp. #1

0.0

0.4

0.0

Marilia sp. #2

0.8

0.0

Unassociated Odontoceridae

Female:

Odontoceridae sp. #1

0.0

0.3

Helicopsychidae

Helicopsyche borealis

(Hagen)

126

25.1

20.2

14.7

Calamoceratidae

Heteroplectron californicum

McL.

0.0

0.4

0.7

Leptoceridae

Mystacides alafimbriata

Hill-Griffin

0.0

0.8

0.3

Ylodes frontalis

(Banks)

1.2

0.8

0.0

Unassociated Ylodes

Females:

Ylodes sp. #1

0.0

0.3

Ylodes sp. #2

0.8

0.4

0.3

Total N

Total No. Taxa (Excluding

Unassociated Females)

No. Unassociated Females

501

252

286

PAN-PACIFIC ENTOMOLOGIST

Table 2. Species present, number of individuals collected, and abundance as percent of individuals collected in each

year, from Big Canyon Creek, 1979-1981.

No.

1979

Percent

of total

No,

1980

Percent

of total

No.

1981

Percent

of total

Philopotamidae

Dolophilodes dorcus (Ross)

16.7

0.0

0.6

Wormaldia cruzensis (Ling)

0.0

0.2

0.0

Wormaldia\xnd&%cx. sp. A

0.0

0.4

0.0

Psychomyiidae

Nyctiophylax sp. A

0.0

0.3

Tinodes siskiyou Denning

0.7

0.2

0.0

Unassociated Tinodes

Female:

Tinodes sp. #1

0.0

0.4

0.0

Polycentropodidae

Polycentropus halidus Milne

0.0

0.4

0.3

Unassociated Polycentropus

Female:

Polycentropus sp. #3

0.0

0.2

0.0

Hydropsychidae

Hydropsyche philo Ross

0.0

0.6

Hydropsyche occidentalis

Banks

7.6

0.6

0.3

Hydropsyche oslari Banks

8.3

4.4

2.9

Parapsyche almota Ross

0.0

0.4

0.9

Rhyacophilidae

Rhyacophila narvae Navas

0.0

0.6

Rhyacophila pellisa Ross

0.0

7.3

6.4

Rhyacophila sierra Denning

0.0

0.3

Rhyacophila vao Milne

1.4

2.7

0.6

Rhyacophila vedra Milne

0.0

1.0

0.0

Rhyacophila vocala Milne

0.0

0.2

0.3

Rhyacophila undescr. sp. A

0.0

0.2

0.3

Rhyacophila\xr\dQS>cx. sp. C

0.0

0.2

0.0

Unassociated Rhyacophila

Females:

Rhyacophila sp. #2

0.0

0.2

0.0

Rhyacophila sp. #5

0.0

0.6

2.9

Rhyacophila sp. #6

0.0

7.3

7.9

Rhyacophila sp. #7

0.0

0.8

1.7

Rhyacophila sp. #8

0.0

0.3

Rhyacophila sp. #9

0.0

1.3

0.0

Rhyacophila sp. #10

2.1

0.4

0.0

VOLUME 63, NUMBER 1

Table 2. continued

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Glossosomatidae

Agapetus taho Ross

16.0

0.0

2.0

Unassociated Agapetus

Female:

Agapetus sp. #1

0.0

0.6

0.3

Anagapetus sp. A

0.7

0.4

0.0

Glossosoma ore gone ns e Ling

0.0

0.2

0.0

Unassociated Glossosoma

Female

Glossosoma sp. #1

0.0

1.7

1.2

Hydroptilidae

Hydroptila arctia Ross

6.3

2.9

4.4

Hydroptila sp. A nr.

rono Ross

8.3

0.2

0.9

Unassociated Hydroptila

Females:

Hydroptila sp. #1

5.6

1.9

1.5

Hydroptila sp. #2

0.0

0.2

0.3

Neotrichia sp. A

3.5

0.2

0.0

Unassociated Neotrichia

Female:

Neotrichia sp. #1

3.5

0.0

Ochrotrichia undescr. sp. A

0.7

0.0

Ochrotrichia undescr. sp. E

0.7

0.0

Ochrotrichia undescr. sp. F

0.0

0.6

Unassociated Ochrotrichia

Female:

Ochrotrichia sp. #1

0.7

0.6

0.3

Oxyethira sp. A

0.0

0.3

Unassociated Hydroptilidae

Females:

Hydroptilidae sp. #1

0.7

0.0

0.3

Brachycentridae

Amiocentrus aspilus (Ross)

0.0

3.0

3.8

Unassociated Amiocentrus

Females:

Amiocentrus sp. #1

0.0

20.1

15.7

Amiocentrus sp. #2

0.0

0.3

Amiocentrus sp. #3

0.0

2.3

Micrasema undescr. sp. A

0.0

123

25.8

11.1

PAN-PACIFIC ENTOMOLOGIST

Table 2, continued

1979

1980

1981

No,

Percent

No.

Percent

No.

Percent

of total

Lepidostomatidae

Lepidostoma cantha Ross

0.0

7.5

19.5

Lepidostomafischeri Denning

0.0

0.6

Lepidostoma strophis Ross

0.0

0.3

Unassociated Lepidostoma

Females:

Lepidostoma sp. #1

0.0

1.5

Lepidostoma sp. #2

0.0

0.3

Limnephilidae

Limnephilidae sp. A

0.0

0.2

0.0

Sericostomatidae

Gumaga nigricula (McL.)

0.7

0.4

0.6

Odontoceridae

Marilia flexuosa Ulmer

Nerophilus californicus

1.4

3.4

4.1

(Hagen)

0.0

0.2

0.0

Parthina vierra Denning

0.7

0.0

Helicopsychidae

Helicopsyche borealis

(Hagen)

11.8

0.4

0.3

Calamoceratidae

Heteroplectron

californicum McL.

0.7

0.4

0.6

Leptoceridae

Mystacides alafimbriata

HiU-Griffin

0.0

0.2

0.0

Oecetis sp. A

1.4

0.2

0.0

Total N

144

477

343

Total No. Taxa (Excluding

Unassociated Females)

No. Unassociated Females

173

126

wet year are compared. This statistic also allows the Trichoptera fauna at Big

Sulphur Creek and Big Canyon Creek to be compared with other faunas. The

1979-1980 and 1980-1981 values of 0.104 and 0.098, respectively (each comparing a

dry with an average year), are within the range of AV values for 14 temperate and

tropical sites reported by McElravy et al. (1982) for Trichoptera; a comparison

between the two dry years (1979 and 1981 AV = 0.063), however, is lower than any

previously reported value.

VOLUME 63, NUMBER 1

Table 3. Comparison of Seasonal Range (SR) frequency distributions between Big Sulphur Creek (BSC) and Big

Canyon Creek (BCC), and with four sites in North America having a continental temperate climate. Number of species

for which SR was calculated is given below each site. Temperature data from the National Oceanic and Atmospheric

Administration.

Site

Location

Elevation

(m)

Latitude

Mean Temp.

(Oct.-April)

°C

with BSC

with BCC

Big Sulphur

Creek

(N = 24)

N. Calif.

680

38° 46'

12.2

p = 0.49

Big Canyon

Creek

(N = 15)

N. Calif.

570

38° 51'

9.3

p = 0.49

Mahoning

R. St. #2

(N = 33)

N.E. Ohio

347

41° 12'

2.9

p = 0.01

p = 0.02

Mahoning

R. St. #3

(N = 30)

N.E. Ohio

326

41° 14'

2.9

p = 0.05

p = 0.16

Linesville

Creek

(N = 50)

N.W. Pa.

317

41°40'

1.9

p = 0.01

Fourmile

Creek

(N = 59)

N.W. Pa.

259

42° 7'

2.2

p<0.01

p = 0.01

Table 4. Annual Variabihty (AV) of caddisfly faunas of Big Sulphur Creek and Big Canyon Creek as indicated by the

mean and variance of the net reproductive rate (sensu Wolda, 1978). Calculations based on species for which the total

number in both years ^ 10.

Site

Years

Compared

Mean

Variance

(=AV)

Big Sulphur Creek

1979-1980

-0.35

0.104

Big Sulphur Creek

1980-1981

0.09

0.098

Big Sulphur Creek

1979-1981

-0.29

0.063

Big Canyon Creek

1980-1981

0.36

0.373

The 1980-1981 AV for Big Canyon Creek, 0.373, is higher than that of Big

Sulphur Creek for the same year. This value is exceptionally high for Trichoptera. In

fact it is exceeded only by an AV of 0.925 that was reported for a Kentucky stream

(Haag et al., 1984), in a study that compared pre-impact 1973 and impact 1979 faunas

at a single site affected by siltation. Since we have only one set of comparative of data

(from a dry to average pair of years) we do not know if AVs are generally higher in

Big Canyon Creek than in Big Sulphur Creek. The benthic (immature) stages of the

Trichoptera at both Big Sulphur Creek and Big Canyon Creek respond to

year-to-year differences in precipitation (Resh et al., 1981; McElravy et al.,

PAN-PACIFIC ENTOMOLOGIST

unpublished data). In all probability, changes in numerical abundance of adult

Trichoptera also result from the same conditions that affect the benthic stages of

these species.

Faunal studies, in addition to providing descriptive accounts of insect populations

and communities, are often done to make spatial (e.g., temperate/tropid;

upstream/downstream) or temporal (e.g., changes in populations and communities

following perturbation) comparisons. Wolda (1979) has emphasized that quantifying

fluctuations in abundances is important in understanding the ecology of insect

populations and communities. Quantitative indices such as Seasonal Range and

Annual Variability may be used to summarize faunal information and permit

statistical tests of temporal parameters among different data sets. The separation of

seasonal or annual variability from variability produced by perturbation is a major

concern in aquatic biology. If sampling programs for faunal studies can be designed

and data reported in such a way that seasonality and/or annual variability measures

can be applied, the value of such studies to researchers interested in partitioning this

variability can be greatly enhanced.

Acknowledgments

We wish to thank John Wood for his assistance in the collection and sorting of

samples, Mrs. Carol Hills for use of the Boggs Intermountain Coast Range Reserve

property at Big Canyon Creek, and Dr. Donald Denning, Moraga, California, for

confirming identifications. The research leading to this report was supported in part

by the United States Department of the Interior, under the Matching Grant Program

of Public Law 95-467, Project No. B-200-CAL, and the University of California

Water Resources Center, Project UCAL-WRC-519.

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Butler, M. G. 1984. Life histories of aquatic insects. Pp. 24-55. In: V. H. Resh and D. M. Rosenberg

(eds.), The ecology of aquatic insects. Praeger Publishers, New York, 625 pp.

Gilman, C. S. 1964. Rainfall. Pp. 9-1—9-68. In: V. T. Chow (ed.). Handbook of applied hydrology.

McGraw-Hill, New York.

Haag, K. H., V. H. Resh, and S. E. Neff. 1984. Changes in the adult caddisfly (Trichoptera) community

of the Salt River, Kentucky. Trans. Ky. Acad. Sci., 45:101-108.

Hynes, H. B. N. 1970. The ecology of running waters. University of Toronto Press, Toronto, 555 pp.

Klopfer, P. H. 1959. Environmental determinants of faunal diversity. Amer. Nat., 93:337-342.

Lamberti, G. A. 1983. Interactions among benthic insects, algae, and bacteria in a geothermally

influenced stream. Ph.D. thesis. University of California, Berkeley, California, 291 pp.

Lamberti, G. A., and V. H. Resh. 1983. Geothermal effects on stream benthos: separate influences of

thermal and chemical components on periphyton and macroinvertebrates. Can. J. Fish. Aquat.

Sci., 40:1995-2009.

MacArthur, R. H. 1972. Geographical ecology. Harper and Row, New York, 269 pp.

McColl, J. G., L. M. Gallagher, and C. P. Martz. 1978. Geochemical origin of surface waters in a

geothermal area. Pp. 365-382. In: D. C. Adriano and 1. L. Brisbin, Jr. (eds.). Environmental

chemistry and cycling processes. U.S. Dept. Energy, Technical Information Center, Washington,

D.C.

McElravy, E. P., V. H. Resh, H. Wolda, and O. S. Flint, Jr. 1981. Diversity of adult Trichoptera in a

“non-seasonal” tropical environment. Proc. 3rd Int. Symp. Trichoptera, Ser. EntomoL,

20:149-156.

VOLUME 63, NUMBER 1

McElravy, E. P., H. Woida, and V. H. Resh. 1982. Seasonality and annual variability of caddisfly adults

(Trichoptera) in a “non-seasonal” tropical environment. Arch. Hydrobiol., 94:302-317.

McMillan, L. E. (ed.). 1985. Geysers-Calistoga KGRA-ARM program 1982-1983 annual report. 2 vols.

Morse, J. C., J. W. Chapin, D. D. Herlong, and R. S. Harvey. 1980. Aquatic insects of Upper Three

Runs Creek, Savannah River Plant, South Carolina. Part I: Orders other than Diptera. J. Georgia

Ent.Soc., 15:73-101.

Neilson, D. 1975. Final environmental impact report for geothermal leasehold of Union Oil Co. at The

Geysers, Sonoma County, California. Ecoview Environmental Consultants, Napa, CA.

Resh, V. H., K. H. Haag, and S. E. Neff. 1975. Community structure and diversity of caddisfly adults

from the Salt River, Kentucky. Environ. Entomol., 4:241-253.

Resh, V. H.,T. S. Flynn, G. A. Lamberti, E. P. McElravy, K. L. Sorg, and J. R. Wood. 1981. Responses

of the sericostomatid caddisfly Gumaga nigricula (McL.) to environmental disruption. Proc. 3rd

Int. Symp. Trichoptera, Ser. Entomol., 20:311-318.

Resh, V. H., G. A. Lamberti, E. P. McElravy, J. R. Wood, and J. W. Feminella. 1984a. Quantitative

methods for evaluating the effects of geothermal energy development on stream benthic

communities at The Geysers, California. California Water Resources Center Contr. No. 190, 57

pp.

Resh, V. H., G. A. Lamberti, and J. R. Wood. 1984b. Biology of the caddisfly Helicopsyche borealis

(Hagen): A comparison of North American populations. Freshwater Invert. Biol., 3:172-180.

Sweeney, B. W. 1984. Factors influencing life-history patterns of aquatic insects. Pp. 56-100. In: V. H.

Resh and D. M. Rosenberg (eds.). The ecology of aquatic insects. Praeger Publishers, New York,

625 pp.

Woida, H. 1977. Fluctuations in abundance of some Homoptera in a neotropical forest. Geo-Eco-Trop.,

3:229-257.

Woida, H. 1978. Fluctuations in abundance of tropical insects. Amer. Nat., 112:1017-1045.

Woida, H. 1979. Seasonality parameters for insect populations. Res. Popul. Ecol., 20:247-256.

Woida, H. 1980a. Seasonality of tropical insects. 1. Leafhoppers (Homoptera) in Las Cumbres, Panama.

J. Anim. Ecol., 49:277-290.

Woida, H. 1980b. Fluctuationes estacionales de insectos en el tropico: Sphingidae. Mem. VI Congr. Soc.

Colombiana Entomol. (1979, Cali, Colombia), pp. 11-58.

Woida, H., andF. W. Fisk. 1981. Seasonality of tropical insects. II. Blattaria in Panama. J. Anim. Ecol.,

50:827-838.

PAN-PACIFIC ENTOMOLOGIST

63(1), 1987, pp. 92-97

New Species of the Genus Styringomyia

from the South Pacific and Southeast Asia (Dip ter a, Tipulidae)

C. Dennis Hynes

Department of Biological Sciences, California Polytechnic State University, Sah

Luis Obispo, California 93407

This paper reports on new species of Styringomyia from specimens collected in

Fiji, New Caledonia, Ryuku Islands, Sarawak (British North Borneo), Solomon

Islands, and Vietnam. I wish to thank Dr. Neal Evenhuis for the privilege of studying

specimens from the Bernice P. Bishop Museum Collections, in Honolulu, Hawaii.

All types are being returned to the Bishop Museum. Thanks are also due to Dr. Van

L. Johnson, Professor Emeritus of Classics, Tufts University, for reviewing the Latin

forms used in the names of the species.

The genus Styringomyia is considered an abberant group within the tribe

Eriopterini, and its affinities to other groups in the tribe have not been well worked

out. Although the larvae have not as yet been described, the cervical sclerite on the

adult of all species appears as an L-shaped plate, which places the genus in, or very

close to, the genera which comprise the Eriopteraria.

Styringomyia bidentata New Species

Male. —Length, 5.8 mm; wing, 3.4 mm. Rostrum yellow-brown. Antennae with

scape light brown, pedicel and flagellar segments fulvous. Head yellow-gray. Thorax

with nota brown, with two narrow, dark brown lines on either side of center. Legs

missing. Wings with brown veins, lighter in cell areas; a dark brown stripe along Cu

from base to midwing. A yellow tinge is found along costal and subcostal regions.

Cell 2nd M 2 weakly petiolate; vein 2A curves gently to the margin of the wing.

Abdomen with terga and sterna brown, unpatterned; last segment and hypopygium

(Fig. 1) yellow. The apex of ninth tergite terminating with a setiferous lobe that is

subtended by small lobes. Ninth sternite truncated at tip, each outer apical angle with

a long, slender seta. Basistyle terminating with a small thick tubercle bearing two

long, modified setae, the bases of which are contiguous. Outer dististyle flat, broad,

bearing abundant retrorse setae with the upper and inner edge glabrous. Inner

dististyle with the outer lobe a flattened, quadrate blade, and with the inner surface

covered with setae. The inner lobe a U-shaped rod. The posterior arm obtuse with

three to four strong setae emerging at tip; the anterior arm longer, ending in two

strong teeth, the upper tooth thicker and with one or two setae emerging from

posterior surface.

Holotype. —(Male) New Caledonia, Mt. Koghi, 27.i. 1963 (C. Yoshimoto and N.

Krauss) BPBM Slide 2042.

Styringomyia bidens New Species

Male. Length 5.3 mm; wing 4.3 mm. Antennae with scape and pedicel brown,

remainder broken. Palps light brown, the tip yellow. Head, thorax, and abdomen

brown with light yellow stripe down midline, continuing to hypopygium. Halteres

VOLUME 63, NUMBER 1

light brown at bases, yellow at tips. Hind coxae yellow, other coxae brown. Front

femur yellow with light brown ring at the three-quarter mark, remainder of legs

broken. Wings with yellow tinge, veins light brown, with branches of Rs, r-m, m-cu

and 2A darker; a barely perceptible brownish cloud over cord and r-m. Venation

standard, with 2nd Cell M 2 sessile; vein 2A strongly curved to margin. Hypopygium

(Fig. 2) with basistyle having two modified setae at tip of the distal tubercle. Ninth

sternite parallel-sided, the apex with a deep U-shaped notch, the lateral angles

produced into slender lobes, not tipped with setae. The ninth tergite is broad at the

base, forming a triangle at the setiferous tip, which extends posteriorly to the level of

the tubercles of the basistyles. The outer dististyle an elongate blade, the distal end

with one short and one elongate seta directed cephalad. Inner dististyle with the

outer lobe a small, triangular plate, with about 14-15 sclerotized teeth on the inner

surface. The middle lobe with a row of sclerotized teeth at the base, six teeth near the

apex, the remainder of the teeth as in Fig. 2. The outer surface of the middle lobe

platelike and glabrous; the inner lobe a long heavily sclerotized bar which forks into

two thick points, the ventral point thicker than the dorsal.

Holotype. —(Male) Ryuku Islands, Iriomote I., Mt. Ushiku, 350 m, 2.xi.l963

(G. A. Samuelson) BPBM Slide 2106.

Styringomyia digitostylus New Species

Male.—Length 5.6 mm; wing 5.0 mm. Antennae with scape and pedicel yellow on

ventral half, brown on dorsal half. Head dark brown, pronotum slightly lighter.

Mesonotum with three brown stripes separated by yellow to wing base, remainder of

mesonotum and metanotum brown with a yellow stripe at midline. Coxae and

trochanters yellow, remainder of legs missing. Wings tinged with yellow, veins light

brown with brown clouds over fork of Rs, the basal portion of Cell 1st M 2 , m, and

r-m. Venation standard with Cell 2nd M 2 sessile. First abdominal segment brown,

remainder of segments yellow brown with a darker brown ring at posterior border of

each segment. Hypopygium (Fig. 3) with basistyle having one strong modified seta at

tip of tubercle; tubercle and seta subequal in length. Ninth sternite elongate, slender,

with two setae at tip. Ninth tergite short and truncate. Outer dististyle an elongate

blade, with long setae at tip. Inner dististyle with outer lobe an elongate blade having

short, blunt, black teeth bordering distal end. Mesal lobe also an elongate blade with

short, blunt, and black spines at distal margin; the inner margin directed dorsad and

mesad forming a small membranous tooth, darkened at point. Beneath the distal

spines is a bulbous area, from which extends a prominent, curved, and coiled

fingerlike projection.

Holotype. —(Male) Ryuku Islands, Ireomote I., Mt. Ushiku, 350 m, 7-10.xi. 1963

(G. A. Samuelson) BPBM Slide 2105.

Styringomyia rostrostylus New Species

Male .—Length 4.7 mm; wing 3.5 mm. Antennae with scape and pedicel dark

brown ventrally, light yellow dorsally, flagomeres yellow. Palpi ochreous, darker at

tips. Head yellow. Pronotum, mesonotum, metanotum, and first abdominal segment

light brown. Remainder of thorax, coxae, and trochanters yellow. Dorsum of

abdominal segments yellow, dark brown on latero-posterior margin; sterna yellow.

Legs yellow. Front femur with an incomplete dark brown band at midlength, another

smaller band at three-quarter length; hind femur with same coloration and markings;

PAN-PACIFIC ENTOMOLOGIST

front tibia with complete dark brown band at midlength, another at distal end; tarsi

yellow, darker at tips, last segment dark brown; remainder of legs broken. Wings

tinged with yellow, brighter along costal edge. Veins yellow with dark brown

markings at anterior arculus, fork of Rs, r-m, m-cu, fork of M 3 ^ base of 2nd Cell

M 2 , and distal section of 2A. Venation standard, with 2nd Cell M 2 sessile; 2A curved

sharply to margin. Hypopygium with one modified seta extending from the

basistyles; the ninth sternite and tergite as indicated in Fig. 4. The outer dististyle an

elongate blade, curved slightly mesad at midlength, with one short and one elongate

seta at the tip, the latter directed cephalad. Inner dististyle as in the drawing. The

basal extension of what appears to be the inner lobe elongate, expanded and rounded

at the tip, and a thick spine extending from the ventral edge, appearing much as a

beak or rostrum from the head of a bird.

Holotype. —(Male) British N. Borneo, Forest Camp, 19 km N. of Kalabaken,

25.x. 1962 (K. J. Kuncheria) BPBM Slide 2104.

Styringomyia vietnamensis New Species

Male. —Length 5.1 mm; wing 6.5 mm. Antennae with scape, pedicel, and first

flagellomere yellow, remainder of flagellomeres missing. Palps yellow. Head yellow,

vertex mottled. Prothorax reddish brown with lateral yellow markings on either side

and along posterior margin. Mesothorax reddish brown with gray pruinosity over

surface; inner area of scutellum marked with yellow; paratergites yellow.

Metanotum brown. Abdomen yellow, only slightly patterned with brown dorsally

and ventrally. Wings tinged with yellow, lighter in cells. Veins yellow. No markings

on wings except a slight darkened at r-m. Venation standard with Cell 2nd M 2

scarcely sessile. Foreleg yellow with no rings; last tarsal segment abruptly dark

brown. Remainder of the legs missing. Hypopygium (Fig. 5) slightly darker yellow

than the remainder of the body. Basistyle with two modified setae from tubercles,

one slightly behind the other. The ninth tergite and oval-shaped hirsute lobe abruptly

expanding on the lateral edges. The ninth sternite truncate with a large seta at either

corner of the porterior margin. Outer dististyle bladelike, clear, with an elongate

seta at the tip directed cephalad. Inner dististyle with the three lobes having a

continuous comb of peglike spines on outer edges; the inner surface of the lobes also

with peglike spines. At the outer tip of what appears to be the middle lobe is an

elongate, slightly sinuate spine.

Holotype. —(Male) Vietnam, Fyan, 900-1000 m, ll.vii-9.viii.l961 (N. R.

Spencer) BPBM Slide 2109.

Paratype. —(Male) same information as given for holotype.

Styringomyia ysabellae New Species

Male. —Length 5.7 mm; wing 4.5 mm. Antennae with scape and pedicel light

yellow dorsally, brown ventrally; flagellomeres missing. Palps fulvous at base

becoming pale yellow at tips. Head dark brown. Thorax ochreous; halt ere yellow at

base, lighter at tip. Coxae and remainder of legs yellow; femur with incomplete

brown bands at midlength, three-quarter length, and very tip; tibia with brown band

at one-third length and very tip. Tarsi light yellow, abruptly dark brown on last

segment. Middle legs missing. Abdomen yellow with posterior margin of each

vergum dark brown. One strong black seta above each inner edge of the antennal

scape, directed laterad and cephalad; one on each side of the vertex at about the level

VOLUME 63, NUMBER 1

of the posterior margin of the eye, erect and directed cephalad; two on each

antero-lateral edge of the antepronotum, subtended by two to three smaller setae,

both sets directed cephalad; one at each postero-lateral corner of the postpronotum,

directed cephalad. At the lateral end of the preascutal suture are smaller, but still

strong, pencils of two setae each; on either side of the midline behind the transverse

suture are two pairs of setae, the larger on the scutellum near the base of the wing,

the other on the postscutellum. Venation standard, cell 2nd M 2 sessile and vein 2A

strongly curved to the edge of the wing. Dark brown marks at anterior arculus, fork

of Rs, r-m, fork of M 3 ^ 4 , junction of m, and all tips of veins reaching lateral and

posterior margins of wing. Membrane surrounding r-m also with dark brown cloud.

Hypopygium (Fig. 7) yellow. Basistyle with two strong modified setae, the dorsal

thicker and curved medially. The ninth tergite elongate, triangular, covered with

numerous, very small setae. The ninth sternite truncate, with a small protrusion at

midline. Outer dististyle bladelike with two setae at tip, one small, the other elongate

and directed cephalad; at midlength a puffed cushion covered with setae; on the

outer edge, at the base of the style, is a row of four to six small, dark, peglike setae.

The inner dististyle is a flattened plate narrowing distally to a point. The distal third

has a row of about 20 peglike spines; at midlength on the inner edge occurs an

expanded cushion from which many setae project medially. The inner edge with a

dorsal, stout, sharp, triangular tooth and a ventral dark, thick, rectangular,

projection with the outer corners forming small teeth.

Holotype. —(Male) Solomon Islands, Santa Ysabel SE, Tatamba, 0-50 m,

7.ix.l964 (R. Straatman) BPBM Slide 2117.

Paratype. —(Male) Solomon Islands, Santa Ysabel SE, Tatamba, 0-50 m,

8.ix.l964 (R. Straatman).

Styringomyia dilinhi New Species

Male.—Wing 4.1 mm. Head missing. Thorax and abdomen dark brown with

yellow on the paratergite and on the katepisternum, giving the appearance of broken

stripes on the pleural area. Coxae and trochanters brown. All legs missing except for

femur of foreleg. The femur is light brown with two broad dark brown bands, one

midlength, the other at distal end; halteres light brown. Venation standard. Cell 2nd

Ml sessile and vein 2A strongly curved to edge of wing. Wing with brownish tinge,

veins brown, darker marking at r-m, extending on to surrounding membrane.

Hypopygium (Fig. 8 ) brown with one thick, modified seta coming off a tubercle, both

subequal in length and directed mesad. The ninth sternite is slender, elongate, with

two setae at the tip. The ninth tergite is broad at its base, which then narrows into a

short protrusion covered by many fine setae. The outer dististyle is a very slender,

curved, blade with two setae at the tip, one very short, the other noticeably elongate,

nearly three-fourths the length of the blade and directed cephalad. The inner

dististyle arises at the base of the tubercle of the basistyle, extending mesad at a right

angle. The inner lobe is a flattened plate with dark teeth at its apex. Two rows of nine

to ten peglike setae are beneath the teeth. The outer lobe is also flattened and

elongate. The tip is expanded into a triangular plate with four to six short, dark teeth

at the most posterior point. At midlength, along the inner edge, arise several strong,

elongate setae, directed basad.

Holotype. —(Male) Vietnam, Di Linh (Djfring), 1200 m, 22-28.iv.1960 (L. W.

Quate) BPBM Slide 2066 (genitalia).

PAN-PACIFIC ENTOMOLOGIST

Styringomyia labuanae New Species

Male .—Length 4.8 mm; wing 3.4 mm. Antennae with scape yellow, pedicel

brown, flagellomeres missing. Palpi and head light brown, darker on either side of

midvertex to posterior margin. Pronotum brown on sides, light brown medially.

Praescutum dark reddish brown, yellow along lateral margins and including

paratergites. Scutellum and metanotum dark reddish brown, yellow spots at base of

wing and edges of metanotum. Halteres light brown. Pleura, coxae, and trochanters

yellow. Hind legs yellow, darkened at tip of femur; remainder of legs broken.

Abdomen yellow, darker at hypopygium. Wings yellow with dark brown markings,

which include both veins and surrounding membranes in areas r-m, m-cu, base of cell

1st M 2 , and at bend of 2A, and tips of veins reaching outer margin of wing. Venation

standard, with cell 2nd M 2 short petiolate, 2A bent sharply at a right angle to the

margin of the wing, a spur at the angle. Hypopygium (Fig. 6) with one modified seta

from tubercle of basistyle (spines on holotype broken). The ninth sternite elongate,

coming to a blunt point from which two short, thickened setae extend. The ninth

tergite a small plate, tip obtuse, and covered with fine setae. Outer dististyle flat,

slender (probably with an elongate seta at the tip, although these are broken on the

type). Inner dististyle with three lobes; the outer with short setae fining the margins,

the middle lobe with 10-11 large teeth on margins, the inner lobe a short triangular

plate, with a brush of setae directed laterally on its inner surface.

Holotype. —(Male) British N. Borneo, Labuan Island, 28-29.xi.1958 (L. W.

Quate) BPBM Slide 2110 (genetalia).

Paratype. —(Male) same data as given for holotype.

Styringomyia idioformosa New Species

Male. —Length 6.0 mm; wing 5.3 mm. Antennae with scape large and fight brown,

pedicel and flagellomeres darker; palpi dark brown. Head dark brown with yellowish

tinge on vertex. Thorax dark brown; scutum with a fight yellowish brown stripe on

either side of the midfine; scutellum slightly fighter, with a yellowish tinge. Abdomen

dark brown. Legs dark brown with a yellowish ring about three-quarter length and at

base of tibia; remainder of leg fighter. Hind leg with the first tarsal segment abruptly

white, remainder of the tarsi brown. Venation standard, with m-cu gently sigmoid,

cell 2nd M 2 sessile; 2A curved gently to margin. Wing dark brown with white

transverse stripes at base, middle, and extreme tip. Hypopygium (Fig. 9) yellow

brown, dististyles darker. Basistyle with three strong modified setae, two at the end

of an elongate tubercle, one slightly posterior. The ninth tergite a dark brown, wide,

elongate plate from which emerges a small obtuse plate covered with numerous,

small, golden setae. The sternite brown, short, obtuse. Phallosome an elongate rod

with two black dots just before the blackened tip. Outer dististyle a long, brown

blade with one short seta and one long seta extending cephalad; the inner face with a

group of short, spinous setae about midlength; the base with six to seven black,

peglike setae along the inner edge. Inner dististyle with the outer lobe a flattened,

oval plate, glabrous on the outer surface; setae over entire inner surface, more

numerous at tip and along edges. Inner lobe a small triangular plate with several

setae along the basal margin.

Holotype. —(Male) Solomon Islands, Bougainville near Crown Prince Rs., 900 m.

11.vi. 1956 (J. L. Gressitt).

VOLUME 63, NUMBER 1

Paratype. —(Male) same data as given for the holotype. BPBM Slide 2122

(genitalia).

The species Styringomyia idioformosa (idio = strange, formosus = pretty) is very

interesting in having banded wings and three modified spines extending from the

basistyle, features found in no other species of this genus described to date.

EXPLANATION OF FIGURES

Figures 1-9. 1. Styringomyia bidentata n. sp. 2. S. bidens n. sp. 3. 5. digitostylus n. sp. 4. S.

rostrostylus n. sp. 5. S. vietnamensis n. sp. 6. S. labuanae n. sp. 7. S. ysabellae n. sp. 8. lS. dilinhi n.

sp. 9. 5. idioformosa n. sp. (b = basistyle, t = ninth tergite, s = ninth sternite, od = outer dististyle,

id = inner dististyle, il = inner lobe of inner dististyle, ol = outer lobe of inner dististyle, p = phallosome).

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Vol. 63

Spring 1987

THE

No. 2

Pan-Pacific Entomologist

DALY, H. V., C. D. MICHENER, J. S. MOURE, and S. F. SAKAGAMI—The relictual bee

genus Manuelia and its relation to other Xylocopinae (Hymenoptera: Apoidea). 102

PARKER, F. D.—Nests of Callanthidium from block traps (Hymenoptera; Megachilidae). 125

EVANS, H. E.—Observations on the prey and nests of Podalonia occidentalis Murray

(Hymenoptera: Sphecidae). 130

KROMBEIN K. V.—Biosystematic studies of Ceylonese wasps, XVIII: The species of

Trachepyris Kieffer (Hymenoptera: Bethylidae: Epyrinae). 135

CHEMSAK, J. A.—A new Mexican species of Linsleyella Chemsak (Coleoptera:

Cerambycidae) . 145

GIESBERT, E. F.—A new genus and species in the tribe Macrotomini (Coleoptera:

Cerambycidae) from Costa Rica . 147

HOVORE, F. T.—A new genus and species of Cerambycidae from Costa Rica (Coleoptera) ... 151

ALEXANDER, B. and J. G. ROZEN, JR.—Ovaries, ovarioles, and oocytes in parasitic bees

(Hymenoptera: Apoidea) . 155

SNELLING, R. R.—A revision of the bee genus Aztecanthidium (Hymenoptera: Megachilidae) 165

PARKER, F. D., J. H. CANE, G. W. FRANKIE, and S. B. VINSON—Host records and nest

entry by Dolichostelis, a kleptoparasitic anthidiine bee (Hymenoptera: Megachilidae) ... 172

BOHART, G. E. and T. L. GRISWOLD—A revision of the dufoureine genus Micralictoides

Timberlake (Hymenoptera: Halictidae) . 178

THORP, R. W.—A new species of Andrena {Onagrandrena) from Utah’s San Rafael Desert

(Hymenoptera: Andrenidae) . 194

Proceedings of the Pacific Coast Entomological Society, 1986 . 199

Corrigenda . 205

Publications received. 150

SAN FRANCISCO, CALIFORNIA • 1987

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

EDITORIAL BOARD

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R. S. Lane, Associate Editor

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THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is pubUshed quarterly (January, April, July,

and October) for $20.00 per year by the Pacific Coast Entomological Society, California Academy of Sciences,

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This issue mailed June 15,1987

The Pan-Pacific Entomologist (ISSN 0031-0603)

PRODUCED BY A-R EDITIONS, MADISON, WISCONSIN 53703, U.S.A.

Dedicated

E. Gorton Linsley

Professor Emeritus

University of California, Berkeley

I have a twofold purpose in this dedication to E. Gorton Linsley. The first is to

acknowledge on behalf of the Pacific Coast Entomological Society his dedication,

efforts, and contributions, not only to the Society, but to the field of entomology as

well. The second, more personal, is to express my gratitude, gratification, and

satisfaction for the years of friendship and collaboration. It has been an honor and a

privilege to have been closely associated with Gort for the past 30 years.

It is not my intended purpose to build a chronological list of Professor Tinsley’s

accomplishments in the field of entomology, since even a neophyte is aware of these

to some degree. The more than 400 published scientific papers, books and articles

attest to his academic productivity. Although a large portion of Gort’s career was

spent on the study of systematics and biologies of the Apoidea and wild bee

pollination studies, the systematics of the Cerambycidae are probably his first love. It

was due to his efforts that the “Monograph of the Cerambycidae of North America”

was initiated. This project, now nearing completion, will surely remain a monument

in the forthcoming years.

Gort has been an inspiration to those closely associated with him, both as students

and as collaborators. His associations with the author, R. F. Smith, John MacSwain,

P. D. Hurd, Jr., A. E. Michelbacher and R. L. Usinger are some examples of not

only working, but close personal relationships.

No tribute to Gort would be complete without mention of Juanita Linsley who has

been wife, companion and field assistant for over 50 years. Their relationship is an

excellent example of the harmonious blending of professional and personal life.

On this, the occasion of Gort’s 77th birthday, the entire community of the field of

entomology wishes him well and many happy returns. Also our thanks for setting an

example of personal integrity and professionalism of the highest caliber.

John A. Chemsak, Editor

Pan-Pacific Entomologist

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 102-124

The Relictual Bee Genus Manuelia and Its Relation to Other

Xylocopinae

(Hymenoptera: Apoidea)

Howell V. Daly, Charles D. Michener, J. S. Moure, S. F. Sakagami

(HVD) Division of Entomology and Parasitology, University of California,

Berkeley, California 94720; (CDM) Departments of Entomology and of Systematics

& Ecology and Snow Entomological Museum, University of Kansas, Lawrence,

Kansas 66045; (JSM) Sec^ao de Zoologia, Universidade Federal do Parana, 80.000

Curitiba, Parana, Brazil; (SFS) Institute of Low Temperature Science, University of

Hokkaido, Sapporo 060, Japan.

Abstract. —The bee genus Manuelia Vachal (subfamily Xylocopinae) is

redescribed, and its three species, all found in Chile and Argentina, are also

characterized. Although Manuelia looks superficially like Ceratina, it is intermediate

in its character combination between the tribes Xylocopini and Ceratinini, and has a

few synapomorphies of its own. Its nests are more similar to those of Xylocopa than

to those of Ceratina. It appears to be a relict genus, nearer than any other to the

ancestral group from which Xylocopini and Ceratinini diverged. The three species of

Manuelia are redescribed and emphasis is placed on striking morphological

characters that make them as different from one another as subgenera in other

Xylocopinae.

Ceratina herbsti Friese is shown to be a new synonym of Manuelia gayatina.

Because of nomenclatural confusion the type species of Manuelia and of the halictid

generic name Corynogaster Sichel are designated.

A genus of small xylocopine bees, Manuelia, consisting of three species found in

Chile and Argentina, is so similar superficially to the widespread genus Ceratina that

it has attracted relatively little attention. Our study, however, shows that it is a relict

type, almost as close to Xylocopa as to Ceratina.

This study was first made by two of us (HVD and JSM) over 20 years ago, then

independently repeated with additional material by SFS and CDM; the two

manuscripts were amalgamated by CDM since the coverage and conclusions were

too similar to justify independent publication. Each manuscript was stronger than

the other in some features, and each recorded some significant characters not noted

in the other. The illustrations except for the maps were made by SFS.

Nomenclatural Problems

The taxonomic history of the three species is confused and involves the generic

name Manuelia as well as the names Corynura and Corynogaster in the Halictidae.

Spinola (1851) described the three species of Manuelia as species of Halictus that

possessed certain distinctive characters not shared by other Halictinae. The bees

were Halictus gayi (p. 208, no. 10, female, male), H. posticus (p. 208, no. 11, female,

male) and H. gayatinus (p. 209, no. 12, female). In the same volume he described the

102

VOLUME 63, NUMBER 2

103

male of a genuine halictid bee, Corynura gayi (p. 301, no. 1), but placed it in the

Thynnidae. His associated female was a wasp, but the male was described first and

the name Corynura has been used for the bee; the problem was discussed by Alfken

(1926). Following Spinola, Smith (1854:424—425) listed all three species of Manuelia

as Halictus. Sichel (1867:146) described as new “Halictus {Augochlora Smith)

chrysurus”; it is apparently a synonym of H. gayi. Correctly recognizing the male of

Corynura gayi as a bee, he also erected a new genus, Rhopalictus, to include

Corynogaster gayi Spinola” (p. 301, no. 1) and others. Spinola did not mention

Corynogaster and we must assume that Sichel meant Spinola’s Corynura gayi of the

same page and number. This was also the interpretation of Herbst (1917) and is

supported by the fact that Sichel recognized the difference between the Halictus gayi

and the Corynura gayi of Spinola. Eickwort (1969), in listing Corynogaster, noted

''lapsus for CorynuraT' Rhopalictus is a synonym of Corynura although its type

species is C. flavofasciata Spinola, not C. gayi.

Until 1896 the use of the specific name gayi for two unrelated bees, one a

xylocopine incorrectly placed in Halictus, the other a true halictid, had not resulted in

confusion of the two. However, while correctly listing Halictus posticus and H.

gayatinus under Halictus, Dalla Torre (1896) placed H. gayi in Corynura and gave

the reference as Halictus (Corynura) gayi Spinola, “p. 208 n. 10 & p. 301 n. 1.”

Placement of the Corynura gayi and the Halictus gayi of Spinola as the same species

by Dalla Torre, without the recognition that they were different species (indeed they

are different families), began a long line of confused nomenclature. Dalla Torre

further listed Rhopalictus gayi Sichel (“p. 146, n. 1”) as a synonym (the reference

should have been to no. 3). As shown above, Rhopalictus is concerned only with

Corynura, not with Manuelia.

Alfken (1904:141), recognizing that they are long-tongued bees not related to

Halictus, placed Halictus gayi, H. posticus, and H. gayatinus in Ceratina. Friese

(1910) concurred by redescribing H. gayatinus as Ceratina herbsti. Vachal (1905)

erected a new genus, Manuelia, in honor of M. Manuel-J. Rivera, for the three

species. Vachal did not mention Alfken’s transfer, but arrived at the same general

conclusion, namely, that these species are near Ceratina. Cockerell (1905) listed

Halictus posticus and H. gayatinus with the comment that Alfken placed them in

Ceratina and Vachal erected a new genus for them. He incorrectly placed H. gayi as a

synonym of Rhopalictus gayi, i.e., Corynura gayi Spinola (p. 301, no. 1), but

Cockerell objected to Dalla Torre’s association of Corynura with this species. On the

other hand, Friese (1916:554) listed Halictus gayi (Spinola p. 208), but his

description indicates that he had Corynura gayi (Spinola p. 301) and he could not

distinguish his specimens from Halictus rubellus Haliday, a species of Corynura.

Sandhouse (1943), apparently following Dalla Torre (1896), designated Halictus

(Corynura) gayi Spinola as the type species of each of the following generic names:

Corynura, Corynogaster, and Manuelia. She listed the last two as synonyms of the

first. The combination "Halictus (Corynura) gayi Spinola,” however, involves two

different species (and families) of bees described by Spinola on different pages. To

avoid further confusion, we regard Halictus (Corynura) gayi Spinola used by Dalla

Torre (1896) and Sandhouse (1943) as a meaningless combination. It is not possible

to say whether Sandhouse designated Halictus gayi Spinola or Corynura gayi Spinola

as the type species of the three generic names concerned. Therefore she did not

designate valid type species for these genera. The type of Corynura was designated

104

PAN-PACIFIC ENTOMOLOGIST

by Alfken before Sandhouse’s work, but for the other genera we designate type

species here, using Sandhouse’s format:

Corynura Spinola (= Corynogaster Sichel)

Historia fisica y politica de Chile . . . por Claudio Gay, Zool., vol. 6, p. 296,

1851. Two species.

Type species.— Corynura gayi Spinola, 1851, Ibid. , p. 301. (By designation of

Alfken, 1926, Deutsche Entom. Zeitschr., 1926: 146.)

( Corynogaster Sichel) = Corynura Spinola

Novara-Expedition. Zoologischer Theil, vol. 2, Hymenoptera, Fossoria et

Mellifera. Suppl. p. 146,1867. Two species.

Type species.— Corynura gayi Spinola, 1851, Historia fisica y politica de Chile

. . . por Claudio Gay, Zool., vol. 6, p. 301. (Present designation.)

Manuelia Vachal

Bull. Soc. Ent. France, p. 25,1905. Three species.

Type species.— Halictus gayi Spinola, 1851, Historia fisica y politica de Chile

. . . por Claudio Gay, Zool., vol. 6, p. 208. (Present designation.)

Sandhouse also lists Halictus gayatinus Spinola as the type species of “Presbia

Spinola.” In his discussion of the possible generic relationships of H. gayatinus,

Spinola states (p. 209) that this species does not belong to ''Presbia Ilhg.” Like

Sandhouse, we have been unable to discover Illiger’s publication of Presbia and we

regard this name as a nomen nudum. Therefore H. gayatinus is not the type species of

a genus Presbia Spinola, as indicated by Sandhouse.

Genus Manuelia Vachal

Description. —Many specific characters are indicated in this description in order to

show the variability within the genus. Characters not relating to all three species are

marked with the abbreviated names of the species that possess them, as follows:

ga = M. gayi, po = M. postica, and gt = M. gayatina. Other characters are indicated in

Table 1 and in the illustrations. Some important features are italicized. Terga and

sterna are abbreviated T and S and numbered as metasomal structures. Features

resembling Ceratinini are annotated “(Q”5 those resembling Xylocopini, “(X),” and

special features of Manuelia, “(M).” An asterisk indicates that variation exists so

that the statement of similarity, while generally true, breaks down in certain cases.

Female. —Body slender, 4.9-8.5 mm long (C). Color dull metallic blue (ga, similar

to Pithitis unimaculata) or black with apical segments reddish (po) or not (gt); legs,

tegula, mandible tending to be brownish, tergal margins not depigmented. No pale

maculations (X*). Wings grayish hyaline, veins and stigma dark brown to black.

Pilosity sparse, not hiding surface except fringe of lateral lobe of pronotum and (ga)

area on each side of pygidial plate; predominantly whitish to pale yellow, ferruginous

on T6 (ga,po).

Head (Figs. 1-4,16) shghtly wider than long, more elongate than in most Ceratina

and Xylocopa but less so than in Braunsapis. Outer and inner orbits convergent

below, more so than in most Ceratina and Xylocopa. Antennae shghtly above middle

of eyes. Vertex seen frontally gently convex. Preoccipital and paraocular carinae

absent. Circumalveolar depression wide, from antennal base down nearly to level of

lower margin of supraclypeal area, up an equal distance, and laterally nearly to inner

orbit in its median one third. Antennocellar triangle much larger than ocellar

triangle. Supraclypeal area above with narrow frontal carina, extending upward as

VOLUME 63, NUMBER 2

105

Figures 1-15. 1. Face of Manuelia gayi, female. 2-4. Faces of males, M. gayi, postica, and gayatina;

shaded areas are yellow. 5. Labrum of female, M. postica. 6-11. Bases of left hind tibia, even numbers,

females, odd numbers, males; 6,7. M. gayi', 8 ,9. M. postica; 10,11. M, gayatina. 12-15. Hind tarsi; 12,13.

M. gayatina, female, male; 14, 15. M. gayi, female, male. In these and all other illustrations, the scale

line = 0.25 mm.

106

PAN-PACIFIC ENTOMOLOGIST

linear, shallow frontal sulcus nearly attaining anterior ocellus. Clypeusflat, its upper

half contrasting with gently convex, raised supraclypeal area. Lower margin of

clypeus straight, extending beyond lower end of eye; lower lateral part only

obliquely and briefly bent backward at side of labrum; lateroclypeal carina

represented by weak ridge. Lateral clypeal margin mildly concave, giving shape of

clypeus neither typically inverted T form as in Ceratina nor hourglass form as in

allodapines (X*). Tentorial pit at upper third of clypeus (X). Upper margin of

clypeus nearly straight (ga, gt) or gently convex (po); summit of clypeus about as

wide as paraocular area at same level. Mandibular axis shghtly behind ocular axis.

Malar area linear, much shorter than scape width. Gena (Fig. 16) narrower than eye,

moderately convex, seen laterally not declivous immediately behind summit of eye.

Labrum (Figs. 1,5) in repose at right angle to clypeal surface, about twice as broad as

long, semicircular, apex with wide tuft of long, dense simple orange-brown hairs (M

but resembling X). Labrum basally with triangular, slightly elevated, hairless smooth

“disc” as in some Xylocopa (X*). Mandible tapering from base, not abruptly

narrowed as in Ceratina (X). Mandible apically bidentate (Figs. 1, 5, 16), fringed

beneath with long, white or yellowish simple hairs. Maxillary palpus 6-segmented,

reaching nearly to apex of galea (X), segments approximately equal in length,

progressively more slender apically (Fig. 16). Galea six or more times as long as

broad (C). Stipes with comb of moderate strength (C). Mentum over five times as

long as wide (C). Flabellum with posterior surface smooth, setal row in middle to

near apex of flabellum (C). Proboscidial fossa with sclerotic roof (C). Antenna

relatively long, much longer than in Ceratina, surpassing middle of mesoscutum (M).

Scape relatively short (Figs. 1^), attaining lower margin of anterior ocellus (about as

in Ceratina), about 4 times as long as wide (C). Flagellum (Figs. 25-30) 2.5 (ga), 2.7

(gt) or 3.0 (po) times as long as scape (M). First flagellomere as long as or shorter

than pedicel (C).

Mesosoma generally smooth and polished, partly coriaceous (especially in ga),

punctures rather sparse and coarse. Pronotum not carinate. Prosternal apophyseal

arms with apices separate (C) (fused in Xylocopa). Mesoscutum anteriorly not

c3ima.tQ,notaulus strong, parapsidal sulcus shorter than in Ceratina, Braunsapis, and

Xylocopa, less than half length of tegula which is fairly large, only slightly shorter

than half scutal length (Fig. 20). Metasternum projecting considerably behind lower

condyle of hind coxa (X). Wings apically pubescent, not papillate (C). Stigma broad

(Fig. 17), shorter than costal margin of marginal cell (only slightly so in po) (C).

Marginal cell broad with apex rounded, apart from wing margin (C). Submarginal

cell 2 distinctly and 3 shghtly (or much in gt) shorter than 1 (C). Recurrent veins 1 and

2 respectively near to transverse cubitals 2 and 3. Basal vein slightly apical to cu-v.

Jugal lobe (Figs. 18,19) less than one fifth as long as vannal lobe (M). Hamuli 7-10;

hamular sinus as deep as wide (C). Fore coxa with apical hairy spine short (C*) (long

in gt). Strigilis as in Ceratina and Braunsapis (C). Arolia large; claws bifid. Fore and

mid tibial spines acute, hind tibia without spine (C). Fore and midtibial spurs normal

(as in Ceratina and Braunsapis) (C). Tibial scopa of sparsely plumose (often trifid)

hairs (C), moderately developed (ga, po. Fig. 31) or sparser (gt. Fig. 32). Hind tibial

spurs microserrate, apically gently (gt. Fig. 32) or rather strongly bent (ga, po. Fig.

31). Basitibialplate distinct (Figs. 6,8,10), simple (C), with short white hairs basally.

Hind basitarsus produced as bluntly pointed process beyond base of second tarsal

segment (X) (Figs. 12, 14). Coxae, trochanters, and femora with white plumose or

VOLUME 63, NUMBER 2

107

Figures 16-24.16. Lateral view of head and mouthparts, Manuelia gayi female. 17. Part of forewing of

same. 18,19. Posterior basal margins of hind wings, M. gayi and postica. 20. Pronotum and scutum, M.

gayi female. 21,22. T7, male, M. gayi, dorsolateral and ventral views. 23,24. S6, male, inner view, M. gayi

and M. gayatina.

simple hairs; fore and mid tibiae and tarsi with abundant, white to yellowish, mostly

simple hairs; apex of second trochanter and base of second femur each with a mesal

patch of dense, short, erect, white, simple hairs.

Metasoma widest at middle. Metasomal terga transversely microlineolate and

dully polished, punctures rather sparse, coarser on apical terga. T1 basally oblique,

basal part not sharply declivous or differentiated. Graduli 2-5 laterally not much

extending posteriorly beyond spiracles; apical terga not flattened as in allodapines.

Sterna without recognizable glandular areas (X); ventral hairs sparse, not forming

scopa. T6 not slanting down apically, with narrow, spine-like, apically upraised,

dorsally flattened pygidial plate (X) surrounded with dense plumose hairs (pygidial

fimbria) (X) (Fig. 57) which become longer and sparser laterad.

Male (differences from female only).—Face with yellow markings (Figs. 2-4).

Inner orbits below more convergent in po and gt (Figs. 3,4). Head (and other parts.

108

PAN-PACIFIC ENTOMOLOGIST

Figures 25-35.25-26. Pedicel and base of flagellum, Manuelia gayatina, female and male. 27,28. Same,

M. postica. 29, 30. Same, M. gayi. 31, 32. Outer side of hind leg of female, M. postica and gayi. 33-35.

Profiles of posterior part of thorax and propodeum of females, M. gayi, postica, and gayatina.

too) generally less coriaceous and more polished than in female. Labrum uniformly

gently convex and punctate. Mandible bidentate (ga) or simple (po, gt). Scape not

attaining anterior ocellus.

Legs slender, without special modifications, hind tibia and basitarsus with white

plumose hairs, in ga densely hairy, suggesting female scopa. Basitibial plate with

edge strong but less developed than in female (ga, Fig. 7) or absent (po. Fig. 9, gt Fig.

VOLUME 63, NUMBER 2

109

11), the plate represented by gentle swelling. Hind tibia with two apical spurs (C).

Hind basitarsus apically produced as in female in ga and po, not in gt (Figs. 13,15).

Metasomal T7 directed downward, apical margin broadly rounded, slightly raised

along margin so that disc is depressed (Figs. 21,22); apparent apex of T7 formed by a

short, wide, thin, shelf-like, sub apical extension with true apex recessed beneath. S6

apically rounded, without subapical modification as in Ceratina (X); gradulus

reduced, shown merely by different coloration in ga (Fig. 23) or present in gt and po

(Fig. 24). S7 short, transverse, with elongate lateral arms and no apical process or

lobes (Figs. 43, 47, 56) (M). S8 with sclerotized vessel-like main body, robust lateral

apodeme and hollow, apical, sparsely hairy process (Figs. 41,42,48,49, 54,55) (M).

Gonocoxite robust with wide, dorsal emargination, ventroapically not produced.

Volsella absent but “cuspis” (? sense of Marikovskaya, 1975) distinct, small, with

fine hairs in ga and po (Figs. 44,50) but not in gt (Fig. 39). Gonostylus unornamented,

semisclerotized, not fused with gonocoxite (C*). Penis valve stout, not rod-like,

sclerotized, basal bridge strongly sclerotized. Spatha absent (though ventral side of

penis weakly sclerotized in po. Fig. 44) or present in gt (Figs. 39, 40).

The “cuspis” is similar to that of Euglossini and may be homologous to some of the

anthophorine structures so labelled by Marikovskaya (1975). In addition to those

structures, which may not be homologous among themselves, there are other

structures in the same vicinity such as the ventroapical plate of the gonocoxite in

Allodapini (Michener, 1975) and even the squama between the gonocoxite and

gonostylus of Bombini. Further investigation of these structures is needed in order to

reliably determine the homologies.

Biology. —Claude-Joseph (1926) described the nesting biology of M. gayatina and

M. gayi. Unfortunately he provided no data by which one might judge the reliability

of his statements. For example one does not know how many nests he examined.

However, he obviously examined several and perhaps many for each species. As

with the morphological characters, the biological ones differ considerably between

the two species that have been studied.

The nests are branching burrows in dead stems or rotting wood, sometimes

utilizing abandoned beetle burrows (Jaffuel and Pirion, 1926). M. gayatina nests in

dry stems or twigs; in spring females may clean and reuse old nests or construct new

burrows in dead stems of brambles {Rubusl). A female does not enter at a broken

end of a stem as do females of Ceratina and Allodapini. Instead, she cuts into the side

of a stem, then makes one branch burrow going down, the other up. Occasionally

one bee uses the lower branch, another the upper, with common use of the entrance,

but usually there is only one bee per nest entrance. In about a month the first

generation emerges and is composed of both sexes. Claude-Joseph (1929) noted that

both sexes return to the old galleries for the night. The females of this generation may

select, in addition to bramble, dry twigs of wicker, willow, peach, or southern hazel.

Toward the end of fall, the second generation metamorphoses, but is said to be

composed only of females. The septa between cells are gradually destroyed and the

adults move about together in the burrow until spring.

M. gayi makes branching burrows in rotten logs of poplar and weeping willow,

starting in November. Each branch is said to be made by one female; obviously there

is a common entrance burrow used by several females. Claude-Joseph indicates that

up to eight or ten may use one entrance. The burrows enter across the grain of the

wood but then turn parallel to the grain. Gazulla and Ruiz (1928) record this species

PAN-PACIFIC ENTOMOLOGIST

also nesting in dry stems of zarzamora (Rubus ulmifolius) ; this is as in M. gayatina.

Apparently there is only one generation per year, for Claude-Joseph indicates that

the young grow through December and transform to adults near the end of summer

and that both sexes hibernate in their cells until spring. Both sexes may return to their

burrows to spend the night (Claude-Joseph, 1929).

The cells of both species are in series in the burrows, separated by partitions made

of particles of wood cemented together, evidently as in Xylocopa or Ceratina. Both

upper and lower surfaces of the partitions are illustrated as concave. At least in M.

gayi the cells are barrel-shaped, narrowed at each end as in Xylocopa, rather than

cylindrical as usual in Ceratina.

The females of M. gayatina bring pollen from diverse kinds of flowers to form the

elongate firm pollen loaf with a depression in the upper part (or lower part in the

upper branch burrow in a vertical stem). The egg is laid in this depression. The food

masses are rather similar in shape to those of Ceratina, but the egg is evidently on the

upper (i.e., toward the nest entrance) rather than the lower part of food. In M. gayi,

however, the food mass is described and illustrated as a ball occupying the lower end

of the cell, with the egg laid on top of it. This is as in many other bees but is unique for

the Xylocopinae. The observation needs to be verified.

M. gayatina overwinters as groups of adult females (only), crowded together in the

burrows in which cell partitions have been destroyed. This is as in Ceratina and

Xylocopa’, such groups have been called prereproductive assemblages by Michener

(1985 and in press). In M. gayi, however, each adult bee overwinters in its cell with

the cell partitions left intact. Overwintering in cells is otherwise unknown in the

Xylocopinae and may be an ancestral feature of M. gayi, for such behavior is well

known in various other bees.

In both species, after hibernation, old burrows are reused while other females

excavate new nests. Claude-Joseph believed that in M. gayatina, with only females

surviving the winter, the first brood is produced parthenogenetically. It is much more

-likely that the overwintering females mate in the preceding summer or autumn. In

view of the longevity now known for other Xylocopinae (Michener, 1985 and in

press), it is probable that females of M. gayatina live much longer than

Claude-Joseph supposed.

Collecting records indicate long seasons of flight. Specimens of all three species

have been taken every month from August to March. The rather numerous floral

records indicate broad polylecty. There are records of all species from various

introduced as well as native flowers. Families reported include the following:

Compositae, Cruciferae, Euphorbiaceae, Hydrophyllaceae, Labiatae, Malvaceae,

Rosaceae, Scrophulariaceae, Verbenaceae.

Geographical Distribution. —The three species are broadly sympatric in central

Chile and the lake district of Argentina. Thus Manuelia is characteristic of the

Araucanian faunal region (Ringuelet, 1961) which is faunistically so different from

the rest of the neotropics that it might be considered a separate faunal realm if it were

larger. Among bees such genera as Diphaglossa, Cadeguala, Corynura, and

Neofidelia are restricted to it. Such large genera as Alioscirtetica (Eucerinae) and

Chilicola (Xeromelissinae) are most abundant in the Araucanian region although

extending into other temperate or xeric parts of the neotropics. The Araucanian is

also the region inhabited by most of the archaic South American types that show

faunal or floral connections with Australia, e.g., Paracolletini among bees and

among trees, Nothofagus and Araucaria. The region also has some faunal

VOLUME 63, NUMBER 2

111

resemblances to Africa, exemplified among bees by the Fideliidae which are found

only in Africa and central Chile.

All three species range from rather xeric Coquimbo Province south to moist cool

temperate Osorno or Llanquihue Provinces (Figs. 36-38), and from sea level to over

Figures 36-38. Locality records for Manuelia gayatina, postica, and gayi, based on specimens seen by us

or by Haroldo Toro of Valparaiso. On the map for M. gayi the circle represents unknown localities in

Chubut Province, Argentina.

112

PAN-PACIFIC ENTOMOLOGIST

1000 m altitude. While they may attain higher altitudes in the north than in the south,

the meager altitude data probably do not show this, for M. gayi has been taken at

1400-1600 m altitude at its southernmost known locality in Llanquihue Province.

Both M. gayi and postica have been taken at 1700-2200 m in Santiago Province.

Unfortunately most collectors have not recorded altitudes.

There are two locality records that indicate that Manuelia also ranges across

Argentina. Specimens of all three species in the Snow Entomological Museum are

labelled Fundo Malcho, Parral, Cordoba, Argentina. They were taken on various

dates in 1956. In the same collection are specimens of M. gayi and gayatina labelled

San Isidro, Buenos Aires Province, Argentina (M. Senkute), also collected on

different dates. Could these labels be wrong? Elizabeth Chiappa T. of the

Universidad Catolica de Valparaiso writes that there is a Parral, Fundo Malcho, in

Chile (Valparaiso area), a finding that suggests bad labelling. Alternatively, might

they represent introductions, which could easily occur with nests in wood? There are

no old reports of Manuelia from eastern or central Argentina, so far as we are aware,

e.g., in the works of Holmberg (1903), Friese (1908), and Jorgensen (1912a, b).

Recent collections by bee collectors (A. Roig A., R. B. Roberts, and J. F. Neff) at

San Isidro and elsewhere in eastern and central Argentina do not include Manuelia.

We have therefore chosen to regard the records for eastern and central Argentina as

probable errors or possible introductions that did not persist, and have omitted these

localities from our maps. Future collectors, however, should watch for Manuelia in

these areas.

The Species of Manuelia

The three species of Manuelia seem about as different from one another as

subgenera in other xylocopine bees. Indeed two of us at one time prepared a

manuscript providing a subgeneric name for each species. Such multiplication of

genus-group names seems unnecessary, but the distinctiveness of the species should

be remembered. Table 1 summarizes the more striking characters including those of

the male terminalia which are illustrated, but not included in the specific

descriptions.

The characters of the three species are further summarized in the key below. We

have not worked out a meaningful cladistic pattern for them because polarity of the

specific variables is difficult to determine. For example, for bees in general, a

horizontal metanotum and propodeal base like those of most wasps is plesiomorphic

relative to the apomorphic slanting or vertical orientations of these surfaces or parts

of them, as in M. postica. These apomorphies are part of the development of a

relatively spherical thorax associated with the rapid flight of many bees. But slender

bodies are characteristic of various small bees that nest in narrow burrows in wood or

twigs, and such a body form results in reversion to a horizontal metanotum and

propodeal base. Examples are Chelostoma, Heriades, and Hoplitis in the

Megachilidae; Hylaeus and its relatives (especially Heterapoides) in the Colletidae;

and Ceratinini and Allodapini in the Anthophoridae. Hence the horizontal base of

the propodeum of M. gayatina, the smallest and most slender species of Manuelia,

could be primitive features retained from primitive bees or derived features adaptive

to life in narrow burrows. Similar problems exist in the interpretation of several other

variables.

VOLUME 63, NUMBER 2

113

Table 1. Major characters of Manuelia species. Asterisks (*) mark variables common to males and females.

Variable

gayi

postica

gayatina

Female

* Coloration

dark metallic blue

black with reddish

black

apical segments

Lateral margin of

straight

-strongly concave -

labral disc

* Lateral angle of

protuberant above

not above level of

protuberant above

pronotum

middle of collar

*Rear thoracic

beginning at rear edge

beginning in middle of

beginning before

declivity

of scutellum;

metanotum;

middle of metanotum

propodeum straight in

propodeum nearly

but interrupted by

profile

straight in profile

horizontal propodeal

base

Basitibial plate

marginal carina high,

marginal carina high.

marginal carina strong

erect apically

oblique apically

but low

Male

Mandibular apex

bidentate

.simple -

Vertex

posterior ocelli in front

-posterior ocelli on summit .

of summit

Flagellomere 3

-much longer than broad, like 4 -

much broader than

long, hke 2

Apex of hind

- produced -

not produced

basitarsus

Basitibial plate

present

absent, hairs sparse

absent, hairs dense

Sternum 6

gradulus evanescent

-gradulus complete -

Sternum 7

apical margin truncate

apical margin gently

rounded

pointed medially

Sternum 8

— basally rounded; apical process basally wide, —

basally truncate; apical

tapering apicad, with sparse hairs

process slender.

densely haired

Gonobase

wide

narrow

wide

Gonocoxite

.long, ventrally with

“cuspis” but without -

short, ventrally

preapical process

without “cuspis” but

with small preapical

process

Gonostylus

with sparse but

with sparse and minute

moderately long hairs;

hairs; base simple

base simple

base bifurcated

Penis valves

robust

-rather slender -

Penis

not sclerotized

-partly semichitinous -

Spatha

-absent--

present

114

PAN-PACIFIC ENTOMOLOGIST

Key to the Species of Manuelia

1. Entirely black with sparse white pubescence; body length about 5 mm; profile of

propodeum curved, with anterior portion horizontal and posterior portion

subvertical; metanotum horizontal; anterior coxa with a long median apical spine;

true apex of T7 of male without hairs; gonostyli long, equal in length to

gonocoxite. gayatina

— Blue or black, at least in female with some caudal integument or hairs

orange-brown; body length about 8 mm; profile of propodeum slightly curved or

straight, steeply sloping; metanotum sub vertical; anterior coxa with small median

apical spine; true apex of T7 of male with hairs; gonostyli short, about half length

of gonocoxite.2

2. Black with at least two caudal terga and sterna orange-brown; profile of

propodeum slightly curved; curvature of metanotum forming summit of posterior

thoracic declivity; labrum of female with basal triangle confluent with subapical

margin by a narrow median area; mandible of male without teeth; male without

basitibial plate. postica

— Metallic blue, last exposed tergum of female with orange-brown hairs; profile of

propodeum straight; posterior margin of scutellum at summit of posterior thoracic

declivity, the metanotum dechvous; labrum of female with basal triangle elevated

apically and separated from subapical margin by a complete transverse punctured

area; mandible of male bidentate; male with small bastibial plate. gayi

Manuelia gayatina (Spinola)

(Figs. 4,10-13,2^26, 32, 35, 36, 39-43)

Halictus gayatinus SpmoXdi, 1851:209; Cockerell, 1905:355.

Ceratina gayatina: Alfken, 1904:141.

Manuelia gayatina: Vachal, 1905:26; Claude-Joseph, 1926:216; Jaffuel and Pirion,

1926:370; Moldenke and Neff, 1974:9,29.

Ceratina herbsti Friese, 1910:703 (new synonym).

A female “typus” of Ceratina herbsti Friese in the American Museum of Natural

History appears to be a cotype; it is^from Concepcion, the type locality, collected by

Herbst in 1903. Although the synonymy of herbsti seems not to have been published,

it was recognized long ago for there is a note by Cockerell indicating the synonymy on

a specimen in the California Academy of Sciences.

Female. —Average and range of forewing lengths (5 specimens): 4.79 mm;

4.50-5.15 mm. Head. Integument shiny dark brown to black; antenna, labrum,

mandible black. Circumalveolar depression rather shallow. Subapical margin and

basal triangle of labrum shiny and impunctate, confluent by a narrow median area

and separated laterad by a transverse punctured area bearing long, yellowish, simple

hairs; lateral margin of basal triangle concave. Hypostomal carina low. Thorax.

Pronotum with lateral angle protuberant above. Thoracic capsule elongate;

posterior declivity beginning at scutellar-metanotal suture, scutal-scutellar tangent

not touching metanotum; metanotum and anterior dorsal portion of propodeum

sloping in same plane, posterior portion of propodeum with steeper slope and clearly

not in the same plane as the anterior portion. Integument shiny dark brown to black;

anterior dorsal portion of propodeum impunctate and roughened, lateral and

posterior portions punctured and with appressed, short, white, plumose hairs; lateral

VOLUME 63, NUMBER 2

115

Figures 39-43. Manuelia gayatina, male. 39, 40. Genitalia, ventral, dorsal, and lateral. 41, 42. S8, dor¬

sal, ventral, and lateral. 43. S7.

portions of pronotum and metepisternum with similar vestiture; remainder of thorax

punctured throughout, dorsum with erect, white, simple hairs, longer laterally;

mesepisternum with sparser, longer, simple to plumose hairs. Tegula brown;

number of hamuli (5 specimens): 7. Legs dark brown, paler distally; hairs yellowish

distally; scopa sparse. Basitibial plate broad, width about one-half distance from

base of tibia to apex of plate, apical edge evenly rounded, strong but low; hind tibial

spurs only gently curved apically. Metasoma. Elongate ovoid; shiny dark brown to

black; T1 and 2 sparsely punctured, remaining terga more densely punctured, with

short, ascending to erect, white, simple hairs. T6 with black spine (pygidial plate)

flanked by dense, yellowish white, plumose hairs which become longer and sparser

laterad. Sl-5 with ascending, white, simple or plumose hairs which are longest

subapically on each sternum; S6 with appressed to ascending, yellowish, simple to

plumose hairs which are dense at apex.

Male .—Forewing length (two specimens): 4.0 mm, 4.4 mm. Head. Coloration as

in female but the following pale yellow: labrum, paraocular area up to base of

antenna, entire clypeus and lower half of supraclypeal area. Mandible simple.

Posterior ocelli on summit of vertex. Third flagellomere much broader than long.

116

PAN-PACIFIC ENTOMOLOGIST

Thorax. As in female. Bastibial plate not perceptible. Hind basitarsus not produced

at apex. Metasoma. Dark brown to black, shiny above, paler and duller beneath;

sparsely punctured, especially anteriorly; dorsum and venter with short ascending to

erect, white, simple hairs which become longer and frequently plumose laterad. T7

with true apex without hairs. Terminalia as illustrated.

This is the least common of the three species in most collections but the large

number of specimens in the collection of Prof. Haroldo Toro in Valparaiso indicates

that it is abundant. Probably its small size results in less frequent capture as

compared to the larger species. The distribution is shown in Figure 36; the

northernmost locality is Vicuna, Coquimbo Prov., the southernmost in Chile is

Valdivia, Valdivia Prov. The only locality in western Argentina is El Bolson, Rio

Negro Prov. Altitudes range from sea level to 1100 m at Cabreria, Cordillera

Nahuelbuta, Malleco Prov.

Manuelia postica (Spinola)

(Figs. 3, 5, 8, 9,18,19, 27, 34, 37, 44-49)

Halictusposticus Spinola, 1851:208; Cockerell, 1905:355.

Ceratinapostica: Alfken, 1904:141.

Manuelia postica: Vachal, 1905:26; Jaffuel and Pirion, 1926:370; Gazulla and Ruiz,

1928:301.

Female. —Average and range of forewing lengths (64 specimens): 6.92 mm,

6.10-7.60 mm. Head. Integument shiny black; antenna, labrum, and mandible

black. Circumalveolar depression rather shallow. Subapical margin and basal

triangle of labrum shiny and impunctate, confluent by a narrow median area and

separated laterally by a transverse punctured area bearing long, yellowish, simple

hairs; lateral margin of basal triangle concave. Thorax. Pronotum with lateral angle

not elevated above level of middle of pronotal collar. Thoracic capsule ovoid;

propodeum slightly convex and steeply sloping; metanotum at summit of posterior

thoracic dechvity, touched by a scuto-scutellar tangent. Integument colored like

head; punctures close around margin of scutum, on scutellum and on metanotum,

less dense on mesepisternum, sparse on dorsum, absent in triangle at summit of

propodeum; surface shiny except propodeum which is finely roughened; surface with

short, erect, white, plumose hairs on dorsum, hairs longer laterally and longest

ventrally; propodeum and pronotum laterally with additional sparse, long, erect,

white, plumose hairs. Tegula black; average and range of hamuli (64 specimens):

8.2,7-10. Tarsi dark brown, remainder of legs shiny black with yellowish white hairs;

scopa moderately developed. Basitibial plate elongate, width about one-third of

distance from base of tibia to apex of plate; apex acute, marginal carina high and

oblique apically; hind tibial spurs strongly curved at apices. Metasoma. Elongate

ovoid; Tl-3 shiny black, punctured throughout, and with sparse, short, erect, white,

simple hairs; coloration of subsequent exposed terga shows variation apparently

uncorrelated with geography in the 69 specimens before us: 24 have basal

three-quarters of T4 with color and vestiture like preceding terga while the apical

quarter and subsequent exposed terga are translucent orange-brown; 43 have this

color restricted to fifth and sixth terga; and in the remaining two bees, T5 and 6 are

dark and only faintly orange-brown. Hairs on these terga colored like their

corresponding terga and longer and denser caudad. T6 with black spine (pygidial

VOLUME 63, NUMBER 2

117

Figures 44-49. Manueliapostica, male. 44. Genitalia, ventral, and dorsal. 45. Inner view of gonostylus

and adj acent structures (sketch). 46. Genitalia, lateral view. 47. S7,48,49. S8, lateral, dorsal, and ventral.

plate) flanked by abundant, ascending, orange brown, simple or plumose hairs which

become longer and sparser laterally. Sterna with vestiture like corresponding terga;

S6 with apical portion clothed with dense plumose hairs.

Male .—Average and range of forewing lengths (17 specimens): 6.85 mm, 5.85-7.05

mm. Head. Coloration as in female but the following areas pale yellow: base of

mandible, labrum, paraocular area up to half-way between summit of clypeus and

lower margin of antennal socket, and a variable area of clypeus. Sometimes dark

color of upper head extends down along lateral portions of epistomal suture to below

anterior tentorial pits and mesad from this suture to almost one-third of clypeal

width; others show reduction of the dark extensions and expansion of yellow above

clypeus. Mandible simple, dark brown apically. Posterior ocelli on summit of vertex.

Third flagellomere longer than broad. Thorax. Largely as in female. Mesepisternum

below scrobal suture with circular patch of appressed, short, white, plumose hairs;

118

PAN-PACIFIC ENTOMOLOGIST

average number and range of hamuli (17 specimens): 7.9, 7-9. Bastibial plate not

perceptible. Hind basitarsus with apex produced. Metasoma. Shaped as in female;

Tl-3 and Sl-3 shiny black with short, ascending, white, simple hairs; coloration of

subsequent exposed terga and sterna varies in 18 males before us: 1 has T4-7 and

S4-6 translucent orange-brown; 6 have the basal portions of T4 and S4 black, the

apical margins and subsequent segments orange-brown; 8 have T5-7 and S5, 6

orange-brown; 1 has the apical portions of T5 and S5 and subsequent terga and sterna

so colored; and two have the apical segments dark with orange-brown colors only

faintly expressed. Hairs white on black areas, orange on orange brown areas. T7 with

true apex bearing short, orange-brown, plumose hairs. Terminalia as illustrated.

The distribution of this species is shown in Figure 37. In Chile the northernmost

locality is Cuesta Cavilolen, Illapel, Coquimbo Prov., and the southernmost is

Osorno, Osorno Prov. An Argentine record, Isla Victoria, Neoquen Prov., is

slightly farther south than any Chilean locality known to us. Altitudes of collections

range from near sea level to 125 m in Cautin Prov., 1500 m in Linares Prov. and

1700-2200 m in Santiago Prov.

Manuelia gayi (Spinola)

(Figs. 1, 2, 6, 7,1^18, 20-23,29-31, 33, 38, 50-57)

Halictus gayi Spinola, 1851:208.

Corynura gayi: Dalla Torre, 1896:93 (part).

Ceratina gayi: Alfken, 1904:141.

Manuelia gayi: Vachal, 1905:26; Jaffuel and Pirion, 1926:370; Claude-Joseph,

1926:220, 1929:417; Gazulla and Ruiz, 1928:301; Moldenke and Neff, 1974:29;

Michener and Brooks, 1984:56, 59.

Female: Average and range of forewing lengths (66 specimens): 6.17 mm, 5.50-6.55

mm. Head. Integument shiny metallic blue; scape colored like head, flagellum ruddy

brown beneath, brown above; labrum and mandibles black. Circumalveolar

depression relatively deep. Subapical margin and elevated basal triangle of labrum

shiny and impunctate; lateral margin of basal triangle of labrum straight. Median

part of hypostomal carina expanded, almost lamella-like, sloping inward over

proboscidial fossa. Thorax. Pronotum with lateral angle protuberant above.

Thoracic capsule roughly spherical; metanotum and propodeum steeply sloping.

Integument colored as head; punctures close around margins of scutum, on

scutellum and on metanotum, becoming less dense on mesepisternum, sparse on

dorsum of scutum and absent in triangle at summit of propodeum; surface shiny

except for propodeum which is finely roughened; pubescence short, erect, white,

plumose on dorsum, hairs longer laterally and longest ventrally; propodeum with

long, erect, white, plumose hairs which become short and appressed laterally.

Tegula brown with bluish reflections; average and range of hamuli (66 specimens):

8.6, 7-10. Tarsi dark brown, remainder of legs brown with bluish reflections; scopa

moderately developed. Basitibial plate broad, its width about half distance from base

of tibia to apex of plate, apical edge bluntly pointed, marginal carina high, erect

apically; hind tibial spurs strongly curved at apices. Metasoma. Ovoid; Tl-4 colored

like head and thorax and bearing on their dorsa short, erect, white, simple hairs

which become longer laterally; base of T5 with coloration and vestiture like

preceding terga, but apical margin translucent and orange-brown; T6 dark brown

with the black spine (pygidial plate) flanked by orange-brown, mossy plumose hairs

VOLUME 63, NUMBER 2

119

Figures 50-57. Manuelia gayi. 50, 51. Male genitalia, ventral, dorsal, and lateral. 52. Sketch of dorsal

apical view of male gonocoxite, gonostylus, and penis valve. 53. Sketch of dorsal apical view of male

genitalia. 54. S8, male, lateral view. 55. Same, dorsal and ventral views. 56. S7, male. 57. Apex of T6,

female, dorsal view with lateral view at left.

which grade laterally into longer, white, plumose hairs. Sl-5 with less bluish

reflections than terga, each clothed subapically with ascending, white, simple hairs;

S6 like preceding in color, but with abundant, medio-apical, orange-brown, mossy

plumose hairs.

Male .—Average and range of forewing lengths (6 specimens): 5.76 mm, 5.40-6.55

mm. Head. Coloration as in female but labrum, most of clypeus, and lower

paraocular areas up to level of summit of clypeus light yellow; dark color of head

extends down along the epistomal suture to anterior tentorial pit. Mandible

bidentate. Posterior ocelli in front of summit of vertex. Flagellomere 3 longer than

broad. Thorax. Largely as in female. Average and range of hamuli (7 specimens):

8.4,7-10. Third leg with distinct basitibial plate in position and form similar to that of

120

PAN-PACIFIC ENTOMOLOGIST

female. Metasoma. Slightly more elongate than in female; Tl-6 with color and

vestiture as in Tl-4 of female; T7 with median impunctate area and lateral long,

white, simple hairs; true apex of T7 bearing short, white, plumose hairs. Sl-6 as in

female. Terminalia as illustrated.

The distribution of this common species is shown in Figure 38. In Chile the

northernmost locality in Vicuna, Coquimbo Prov.; the southernmost is Colegual,

Llanquihue Prov. In Argentina the southernmost located collection site is El Bolson,

Rio Negro Prov., but specimens are labelled El Hoyo and El Turbio, Chubut Prov.

(New York, Lawrence); the probable vicinity is indicated by a circle in Figure 38.

Altitudes of most collections are not given on the labels but range from near sea level

to 1400-1600 m (Llanquihue Prov.), 1100 m (Talca Prov.) and 1700-2200 m

(Santiago Prov.); in the lake district of Argentina, altitudes are indicated as 650 to

850 m.

Relationships of Manuelia to Other Xylocopinae

The anthophorid subfamily Xylocopinae as delimited by Michener (1944, p. 269),

Hurd and Moure (1963) and others includes both the large, robust bees of the tribe

Xylocopini and the smaller and usually slender forms of the tribes Ceratinini and

Allodapini. The last, recently segregated from Ceratinini (Michener, in press), has

many derived characters and is not particularly relevant to Manuelia. The

resemblances of Manuelia to Xylocopini (Xylocopa, Lestis, Proxylocopa) and to

Ceratinini (Ceratina, Pithitis, Megaceratina) have been indicated in the generic

description above, using for brevity the letters X (Xylocopini), C (Ceratinini), and M

(for special features of Manuelia). In that description 22 characters are marked C;

12, X; and 6, M. Thus Manuelia is most similar to Ceratinini, but also has numerous

features like Xylocopini in spite of its ceratinine appearance. A cladistic approach to

relationships among tribes of Xylocopinae is presented by Sakagami and Michener

(in press).

In some other features, not listed because they are variable and therefore not

generic characters of Manuelia, this genus is nonetheless intermediate between

Xylocopini and most Ceratinini. For example, the dechvity of the posterior part of

the thorax extends downward vertically from the posterior edge of the scutellum in

some Xylocopini while in most Ceratinini and in Manuelia gayatina (Fig. 35) the base

of the propodeum is more or less horizontal. The situation for the other species of

Manuelia is shown in Figures 33 and 34.

The nest characteristics of Manuelia are similar to those of Xylocopini. The

branching burrows and barrel-shaped cells are unlike those of any other small

Xylocopinae, but resemble those of Xylocopini.

Since most bees, and more specihcally most Anthophoridae, nest in the ground,

the Xylocopinae probably arose from ground-nesting forms. Malyshev (1913:55-56)

and Hurd (1958:368) attached much importance to the ground-nesting habits and

certain anatomical conditions (basitibial and pygidial plates) of Proxylocopa that

also occur in ground-nesting anthophorines but not in other Xylocopinae. Because

Proxylocopa is a member of the tribe Xylocopini, Malyshev regarded that tribe as

more closely related to the ground-nesting ancestor than is the Ceratinini. This view

is supported by the clustered cells and constructed cell walls of Proxylocopa,

described and illustrated by Gutbier (1915), and by the brood-cell linings secreted by

VOLUME 63, NUMBER 2

121

Dufour’s gland also found in Proxylocopa but not known in other Xylocopinae

(Kronenberg and Hefetz, 1984).

The nesting habits of Proxylocopa were assumed by Malyshev and Hurd to be

primary, although there are examples in both tribes of flexible behavior which could

have led to a return to the ground as a nesting site. In the large carpenter bees, Hurd

(1978) cites reports of nests in bricks and other soft substrates, Lucas (1868) noted a

nest in a copper tube, and Hardouin (1943) found that an individual of a species

presumably unaccustomed to bamboo accepted a bamboo tube offered

experimentally. Of the small carpenter bees, allodapines are reported by Brauns

(1926) to nest in the ground in the absence of suitable plants or in vacant beetle

galleries in wood. While constructed cell walls and linings derived from Dufour’s

gland are usual in soil-nesting anthophorids, and might seem unlikely to reappear in

Proxylocopa if it reverted to the soil, the partitions between cells in Xylocopa are in

reality constructed cell walls that do not extend to the sides of the cell in a wood

substrate. Reversion to soil could result in extension of partition-construction to

produce cell walls. Less likely, probably, would be reversion to production of the

hydrocarbon-rich hydrophobic cell lining. Thus while the nests of Proxylocopa offer

a basis for considering the genus as similar to the ancestral Xylocopinae, there is the

possibility that nesting in the ground is an adaptation to desertic environments

lacking plants for nesting. Independently, Hurd (in personal communication to

Daly) entertained the same explanation. Furthermore, Maa (1954) considered the

species of Proxylocopa to be closely related on the basis of morphology; such

similarity does not suggest the antiquity to be expected of ancestors of the other

Xylocopinae. Proxylocopa is noteworthy for its close resemblance to other genera of

the tribe Xylocopini.

Although Manuelia shares its nesting pattern with the Xylocopini, Ceratina and

Pithitis are quite different in their nesting activities and social organization

(Michener, in press), giving little indication of ground-nesting ancestry.

In the majority of anatomical features the Ceratinini rather than the Xylocopini

show the most plesiomorphic features, as judged by Michener’s (1944:228-229) list:

short first flagellar segment, horizontal metanotum, propodeum with horizontal

basal area, large pterostigma, long notaulus, relatively long second abscissa of vein

M + Cu in the rear wing, long jugal lobe, and hairy wings. The Xylocopini show more

specialized conditions of the same variables. Some ceratinine characters of the wings

and thorax may relate to the general body size and shape, associated with nesting in

small burrows. Nevertheless, other characters, presumably not related to size and

slenderness, also indicate the more primitive anatomy of the small carpenter bees.

For example, male Ceratinini have two hind tibial spurs and, in Manuelia and

Ceratina (Euceratina), have gonostyli, while the large carpenter bees have only one

tibial spur and no gonostyli. The basitibial plate is near the base of the tibia in females

of ground-nesting bees; Ceratinini have the plate usually in this position when it is

present, while only Proxylocopa, among the Xylocopini, has it in this location. Wille

(1958) found the dorsal circulatory vessel to be straight in the Ceratinini, a condition

considered by him to be primitive among the bees. Xylocopa possesses a specialized

condition with the thoracic portion arching between the longitudinal muscles and the

petiolar portion coiled.

In view of the primitive features occurring in members of both tribes (Ceratinini

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PAN-PACIFIC ENTOMOLOGIST

and Xylocopini), we cannot agree that the Xylocopini is closer to the stem of the

subfamily than Ceratinini. Moreover, we do not think that Proxylocopa is

particularly close to the ancestral Xylocopini; undoubted specialized features of

Proxylocopa include the reduced notauli. The ancestor for all modern members of

the subfamily Xylocopinae should combine the nesting pattern of the Xylocopini

with the primitive anatomical features of both Ceratinini and the Xylocopini.

It seems reasonable to suppose, as did Malyshev, that the ancestor nested in

relatively thick pieces of dead plant tissue (soft or rotten wood). The ancestor of the

large carpenter bees continued nesting in the thicker pieces of wood. Once the outer

layer has been broken, this medium places little restriction on the dimensions or

pattern of the nest. There was probably a trend of increasing body size and more

powerful jaws, together with the changes in the thorax and wings which impart the

specialized facies to the modern tribe Xylocopini. The nesting habits, however,

remained largely unaltered. The smaller forms of the ancestral stock could nest in

small beetle burrows and slender stems and consequently there was the trend to

smaller body size and slender form which led to the modern Ceratinini.

We believe that Manuelia is a surviving remnant of the early small carpenter bees.

Moreover, it is the most primitive of the subfamily Xylocopinae and nearer than any

other form to the phyletic dichotomy which separated the tribes Ceratinini and

Xylocopini. The restriction of the species to Central Chile and Argentina, the faet

that the genus is made up of three species fully as different as subgenera elsewhere

among bees, together with the anatomical and biological relations enumerated

above, support the conclusion that Manuelia is a relict genus. In view of its presumed

antiquity, Manuelia exhibits an interesting use of introduced plants both for

provisions and as nesting substrates. This is not surprising, however, since most

Xylocopinae are polylectic feeders and seem to select nesting sites according to their

physical characteristics without regard to the kind of plant concerned.

Acknowledgements

We welcome this opportunity to acknowledge our respect and admiration for Dr.

E. Gorton Linsley. We have shared a long association with him as colleagues and

friends. Dr. Linsley often encouraged fruitful collaboration. We trust our

contribution, the product of international cooperation, is in keeping with his

example.

Specimens of Manuelia were examined in various collections by Daly and Moure,

and were lent to Daly and Michener from certain collections. The following is a list of

the collections to which we are indebted either for the opportunity for study or for the

loan of material: American Museum of Natural History, New York, U.S.A.;

Academy of Natural Sciences of Philadephia, Philadelphia, Pennsylvania, U.S.A.;

British Museum (Natural History), London, U.K.; California Academy of Sciences,

San Francisco, California, U.S.A.; University of Kansas, Lawrence, Kansas,

U.S.A.; Museo Civico di Storia Naturale, Genova, Italy; Museum National

d’Histoire Naturelle, Paris, France; Museo National, Santiago, Chile; National

Museum of Natural History, Washington, D.C., U.S. A. We are grateful to the late

Dr. Paul D. Hurd, Jr., who arranged the loan of specimens of Proxylocopa from Dr.

E. S. Ross of the California Academy of Sciences. Mr. Rudolfo Wagenknecht

generously made his collecting notes available to us. Information on the nativity of

VOLUME 63, NUMBER 2

123

the various host plants was provided by the late Dr. L. H. Shinners of Southern

Methodist University. We especially appreciate the hard work of Elizabeth Chiappa

T. of the Universidad Catolica de Valparaiso who provided an extensive list of

material in the collection of Professor Haroldo Toro and that of the same University.

To both Ms. Chiappa and Prof. Toro we are indebted for the opportunity to consider

the extensive material in Valparaiso as well as for a map of localities and other

helpful information. Joetta Weaver provided help with editorial work and typing.

HVD’s research was supported in part by research grants from the National

Science Foundation (GB-7933, GB-34089).

CDM’s part in this study was possible thanks to National Science Foundation

(U.S.A.) grant DEB82-12223. JSM wishes to thank the Campanha Nacional de

Aperfeigoamento de Pessoal de Nivel Superior, Rio de Janeiro, for a travel grant

making possible his visit to North America and Europe; the Rockefeller Foundation,

New York, for a grant permitting his stay in the United States, and the National

Science Foundation (U.S.A.) for a grant to the University of Kansas which

permitted his study in European museums. SFS’s part in this paper was supported by

a grant-in-aid for Special Project Research on Biological Aspects of Optimal

Strategy and Social Structure from the Japan Ministry of Education, Science and

Culture.

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Deutsch. Entom. Zeitschr., 1926:145-163.

Brauns, H. 1926. A contribution to the knowledge of the genus Allodape, St. Farq. & Serv.; Order

Hymenoptera, Section Apidae (Anthophila). Ann. S. African Mus., 23:417-434.

Claude-Joseph, F. 1926. Recherches biologiques sur les Hymenopteres du Chili (Melliferes). Ann. Sci.

Nat., ZooL, (10)9:113-268. (Translated into Spanish but without some illustrations by J. Herrara

and M. Etcheverry, 1960, Investigaciones biologicas sobre himenopteros de Chile (Meliferos).

Publ. Centro Estudios Entom., Univ. de Chile, Santiago, no. 1:1-64.)

Claude-Joseph, F. 1929. Le repos nocturne chez quelques Hymenopteres du Chili. Zool. Anz.,

82:414-421.

Cockerell, T. D. A. 1905. Notes on some bees in the British Museum. Trans. Amer. Entom. Soc.,

31:309-364.

Dalla Torre, C. G. de. 1896. Catalogus Hymenopterorum, 10:i-vii+1-643, Leipzig.

Eickwort, G. C. 1969. A comparative morphological study and generic revision of the augochlorine bees

(Hymenoptera: Halictidae). Univ. Kansas Sci. Bull., 48:325-524.

Friese, H. 1908. Die Apidae (Blumenwespen) von Argentina nach den Reisenergebnissen der Herren

A. C. Jensen-Haarup und P. Jorgensen in den Jahren 1904-1907. Flora og Fauna, 10:1-94.

Friese, H. 1910. Neue Biennenarten aus Siid-Amerika. Deutsche Entom. Zeitschr., 1910:693-711.

Friese, H. 1916. Die Halictus-Arten von Chile (Hym.). Deutsche Entom. Zeitschr., 1916:547-564.

Gazulla, P. and F. F. Ruiz Perez. 1928. Los insectos de la Hacienda de “Las Mercedes.” Rev. Chilena

Hist. Nat., 32:288-305.

Gutbier, A. 1915. Essai sur la classification et sur le developpement des nids des guepes et des abeilles.

Horae Soc. Entom. Rossicae, 41(7): 1-57, pis. 1 and 2 [in Russian].

Hardouin, R. 1943. Ethologie variable de la Xylocope. Bull. Soc. Entom. France, 48:156.

Herbst, P. 1917. Nachtrag zur Synonymie Chilenischer Apidae (Hymenoptera). Deutsche Entom.

Zeitschr., 1917:293-294.

Holmberg, E. L. 1903. Delectus hymenopterologicus argentinus. An. Mus. Nac. Hist. Nat. Buenos

Aires, (3)2:377-512.

Hurd, P. D., Jr. 1958. Observations on the nesting habits of some New World carpenter bees with

remarks on their importance in the problem of species formation (Hymenoptera: Apoidea). Ann.

Entom. Soc. Amer., 51:365-375.

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Hurd, P. D., Jr. 1978. An annotated catalog of the carpenter bees (genus Xylocopa Latreille) of the

western hemisphere, pp. 1-106. Smithsonian Inst. Press, Washington, D.C.

Hurd, P. D., Jr. and J. S. Moure. 1963. A classification of the large carpenter bees (Xylocopini). Univ.

CaliforniaPubl. Entom., 29;i-vi-l-1-365.

Jaffuel, P. P. F. and A. Pirion. 1926. Himenopteros del Valle de Marga-Marga. Rev. Chilena Hist. Nat.,

30:362-383.

Jorgensen, P. 1912a. Aculeatos de la Provincia de Mendoza. An. Mus. Nac. Hist. Nat. Buenos Aires,

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Jorgensen, P. 1912b. Revision der Apiden der Provinz Mendoza, Repubhca Argentina. Zool. Jahrb.

(Syst), 32:89-162 and 33:643-644.

Kronenberg, S. and A. Hefetz. 1984. Comparative analysis of Dufour’s gland secretions of two carpenter

bees (Xylocopinae: Anthophoridae) with different nesting habits. Comp. Biochem. Physiol.,

79B:421-425.

Lucas, M. H. 1868. Vie evolutive de la.Xylocopa violacea. Ann. Soc. Entom. France, 37:727-736.

Maa, T. 1954. The 3rd Danish Expedition to Central Asia. Zoological results 14. The xylocopine bees

(Insecta) of Afghanistan. Vidensk. Meddel. Dansk Naturhist. For. Kpbenhavn, 116:189-231.

Malyshev, S. J. 1913. Life and instincts of some Ceratina-h&es (Hymenoptera, Apidae). Russkoe Entom.

Obshchestvo (Horae Soc. Entom. Ross.)., 40(8): 1-58 [in Russian; English summary].

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Kazak SSR (Alma-Alta), 3:16-24 [in Russian].

Michener, C. D. 1944. Comparative external morphology, phylogeny, and a classification of the bees.

Bull. Amer. Mus. Nat. Hist., 82:151-326.

Michener, C. D. 1975. A taxonomic study of African allodapine bees. Bull. Amer. Mus. Nat. Hist.,

155:67-240.

Michener, C. D. 1985. From solitary to eusocial: need there be a series of intervening species, pp.

293-305, in B. Hdlldobler and M. Lindauer, eds.. Experimental behavioral ecology and

sociobiology, Fortschritte der Zool., vol. 31.

Michener, C. D. in press. Caste in xylocopine bees, in W. Engels, Caste and Reproduction in Social Insect

Evolution, Springer Verlag.

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Entom. Inst., 22(1): 1-73.

Moldenke, A. R. and J. L. Neff. 1974. Studies on pollination ecology and species diversity of natural

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1-134.

Ringuelet, R. A. 1961. Rasgos fundamentales de la zoogeografia de la Argentina. Physis [Buenos Aires],

22:161-170.

Sakagami, S. F. and C. D. Michener. In press. Relationships of the tribes of xylocopinae to the Apidae

(Hymenoptera: Apoidea).

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92:519-619.

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Vespiden, Sphegiden, Pompihden, Crabroniden und Heterogynen, Reise der Osterreichischen

Fregatte Novara. . ., Zool. Theil, vol. 2.

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6, Paris.

Vachal, J. 1905. Manuelia, un noveau genre d’Hymenopteres melliferes. Bull. Soc. Entom. France, pp.

25-26.

Wille, A. 1958. A comparative study of the dorsal vessel of bees. Ann. Entom. Soc. Amer., 51:538-546.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 125-129

Nests of Callanthidium from Block Traps

(Hymenoptera: Megachilidae)

Frank D. Parker

USD A, Agricultural Research Service, Bee Biology and Systematics Laboratory,

Utah State University, Logan, Utah 84322-5310.

Abstract. —The nesting habits of Callanthidium formosum (Cresson) are

described for the first time. Nests were obtained from block traps set at 2000-3000 m

in northern Utah. Information on nest construction, cocoon formation, sex ratio,

adult weights, and mortality is presented. Additional information is presented on the

adult weights, cell lengths, and nest associates of Callanthidium illustre (Cresson).

Species of Callanthidium are among the largest members of the tribe Anthidiini,

and our two species are marked with bright yellow integumental bands that contrast

with their brownish-black body color. Many adults have been collected from a

variety of flowers (Hurd et al., 1979), but nests are less commonly found and only

those of C. illustre (Cresson) have been reported (Johnson, 1904; Hicks, 1929;

Parker and Bohart, 1966). Recent biological studies using block traps (Parker,

1985a) have added information on nests of C. illustre and C. formosum (Cresson),

and in this paper the nesting habits of C. formosum are described for the first time.

Additional data are presented on sex ratios and adult weights of both species.

Methods

One-meter stakes were driven partially into the soil and the block traps attached

near the top of the stake with the holes facing southeast. The trap blocks (see Parker

1985a for details of trap design) were set out in May and recovered in November of

the same year. The blocks were then taken apart and the layers split with a knife to

expose the nest contents. Each nest was measured, described, and photographed;

samples of pollen were removed to identify floral resources. A radiograph (Stephen

and Undurraga 1976) was made of intact nests that had been removed from the block

traps. Individual cell contents were placed and stored in 000 gelatin capsules, labeled

and held at 3° C to break overwintering diapause. During March of the next year, the

capsules were placed in a 27° C incubator, and when the adults emerged they were

weighed alive, killed, mounted and identified.

C. formosum

Nesting Sites. —All nests were recovered from block traps set at higher elevations

(2000-3000 m) in Logan Canyon, Farmington Canyon and along the southwestern

shore of Bear Lake in northern Utah. The predominant shrubs and trees at these

locations included junipers, mahogany, scrub maple, boxelder, sagebrush and

ceanothus; perennial and annual forbs were abundant and diverse.

Nest Construction. —Twelve nests containing 36 cells were recovered from 10 mm

diameter borings and a 2-cell nest from an 8 mm boring. The number of cells/nest

125

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PAN-PACIFIC ENTOMOLOGIST

ranged from 1 to 4 and averaged 3.2 (SD 1.2). The average length of male cells was

15.4 mm (SD 1.0 mm, n = 7) and this figure for female cells averaged 13.6 mm (SD

1.0 mm, n = 7). These differences between sexes and average lengths of cells were

significantly different (P < 0.004).

The plant source of the cotton-like fibrillose material used to line the cells was

undetermined. Cells were constructed from fibers that were formed into pouch-like

chambers that held the provisions. These chambers were 7-8 mm wide and 11-13 mm

long. The cells were separated initially by 4-5 mm thick partitions of fibers, but the

partitions were compacted during cocoon formation to 1-2 mm. Above the last cell

the nest was plugged with fibrillose material that averaged 34.0 mm (SD 14.3 mm

long, n = 4). Most nests were capped with an entrance plug of small white pebbles

stuck together with masticated leaf pulp (Fig. 1). Plugs averaged 6.0 mm (SD 1.4 mm

thick, n = 4), were disc-shaped and were placed at the entrance to the nest.

Provisions. —The cup-shaped mixture of pollen and nectar (Fig. 1) was tacky when

probed. The composition was about 60% mint and 40% legume pollens. The host

egg was laid across the top of the provision.

Cocoon. —The mature larva initiated formation of the cocoon by flattening the

fecal material and remaining provisions against the cell walls (except at the upper

rim) and then lining the walls with a shellac-like layer of silk; this layer had a

cone-shaped and hollow nipple at the top. Inside this layer, the second layer was

barrel-shaped (see radiograph) and was formed from strands of whitish silk

deposited in a cross-hatched pattern; this internal layer was brown. Beneath the

nipple, the second layer was formed into a mat-like pad from coarse, brownish silk

strands. A third layer of coarse, whitish silk strands that resembled cellophane

covered the entire inner surface of the cocoon, including the underside of the nipple.

Average length of cocoons from which males emerged was 12.3 mm (SD 0.8 mm)

long, and 8.1 mm (SD 0.3 mm) wide and those from which females emerged

averaged 11.3 mm (SD 0.5 mm) long and 7.1 mm (SD 0.2 mm) wide. These size

differences were significantly different (P < 0.001).

Overwintering. —All cells contained prepupae in diapause when examined in

November and all surviving overwintering prepupae pupated and emerged when

incubated in March.

Adult Weights and Sex Ratio. —Average weight of males was 142.9 mg (SD 10.4

mg, range 130.3-160.5 mg, n = 7) and females averaged less, 113.8 mg (SD 18.5 mg,

range 91.3-139.3 mg, n = 7). The observed and expected sex ratios (see Torchio and

Tepedino 1980 for methods of calculation) were identical: 1.26 females: 1 male.

There was a significant relationship between cell length and individual weight among

females (r = 0.78, n = 7, P<0.04) but not in males (r = 0.51, n = 7, P<0.24).

Placement of the sexes within cell series differed from most wood-nesting bees

because males were in the bottom cells and females were above. Some nests,

however, had all male or all female cells (Fig. 1).

Mortality. —Mortality of immature stages from unknown causes averaged 14.3%

of the total cells. No nest associates were found nor were any of the nests superseded

by other aculeates.

C. illustre

Nesting in this species has been described by several authors. Johnson (1904)

reported C. illustre nesting in clay banks in Denver, Colorado. Hicks (1929) reported

VOLUME 63, NUMBER 2

127

Fig. 1. Radiograph of nests of Callanthidium formosum and C. illustre illustrating placement of cells,

form of the cocoon and nest entrance plugs. XL = dead larva, XE^dead egg. TO = Trichodes ornatus

predation, ASC-Ascosphaera parasitism, NP = nest entrance plug.

that this bee nested in old, dead yucca flower stalks near Pasadena, California; he

described their nests, adult activity and possible nest associates. Parker and Bohart

(1966) reported nests in holes in wood (block traps). Grigarick and Stange (1968)

summarized the biology and included photographs of the nest and cocoon. Four

additional nests were obtained recently from trap blocks placed near Santa Clara in

southern Utah.

All nests were made in 10 mm diameter holes. The 4 nests, containing 13 cells,

averaged 3.2 cells/nest (range of 2-4 cells). Cells were separated by 3.3 mm (SD 1.6

mm) of fibrillose plant parts. One nest was finished and had a vestibule of fibrillose

material 20 mm long; it was capped with a 5 mm thick plug of masticated plant parts

and pebbles.

Cell size and adult weights varied considerably. Average length of cells of males

was 17.2 mm (SD 1.6 mm, n = 5) and average length of female cells was 19.0 mm (SD

2.6 mm. n = 4); these averages, however, were not significantly different (P < 0.24).

Although male bees were heavier (average weight of males was 205. 1 mg, SD 25.4

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PAN-PACIFIC ENTOMOLOGIST

mg, n = 5; females 185.2 mg, SD 34.1 mg, n = 4), weights were not significantly

different (P < 0.35). There was a significant correlation between cell length and

adult weight in both sexes (r=.91, P< 0.003 for males; r=.96, P < 0.04 for

females). Average length of cocoons from which males emerged was 14.2 mm (SD

0.8 mm) and average width was 8.7 mm (SD 0.4 mm); average length of cocoons

from females was 13.8 mm (SD 0.5 mm) and average width was 8.3 mm (SD 0.3 mm).

One nest had males below and a single female above, one contained males, and

another contained only females.

One cell contained a bee larva infected with the fungus, Ascosphaera sp., and two

other cells were destroyed by a larva of the clerid beetle, Trichodes ornatus Say.

Discussion

The placement of sexes within Callanthidium nests differed from the usual pattern

in xylophilous nesting bees in that females were at the bottom and males were at the

top (Krombein, 1979). Males of Anthidiine genera such as Callanthidium,

Anthidium and Dianthidium are larger and weigh more than females (Alcock, 1977;

Alcock et al., 1977; Frohlich and Parker, 1985). Thus, placement of the male sex first

in cell series appears to be a response to the nontypical mating system exhibited by

many anthidiine bees (Thornhill and Alcock, 1983).

Hicks (1929) reported that C. illustre females used resin and an undetermined

substance for the final cap on nests. None of the nest entrance plugs of either species

reported here contained resin. The material was masticated plant parts mixed with

pebbles, dirt, or other organic debris.

This is the first report of Ascosphaera attacking Callanthidium. Recent

trap-nesting studies have yielded many new host records (unpublished data) for this

important disease (chalkbrood), and it appears that the disease may be spreading

from commercially managed populations of the alfalfa leaf cutting bee. Megachile

rotundata (Fab.) (Parker, 1985b), to populations of native bees (Parker and

Frohlich, 1983; Youssef et al., 1985).

Acknowledgments

Thanks are due my technician, D. F. Veirs, for construction and field placement of

trap materials, and part-time assistants, R. T. Griswold and V. A. Rhea, for

recording data and rearing the specimens. Helpful manuscript suggestions were

made by V. J. Tepedino of this Laboratory and D. Mayer (Washington State

University).

I wish to dedicate this paper to E. Gorton Linsley in recognition of his

contributions to the study of bees.

Contribution from Utah Agricultural Station, Utah State University, Logan, Utah

84322-4810, Journal Paper No. 3296 and USD A-Agricultural Research Service-Bee

Biology and Systematics Laboratory, Utah State University, Logan, Utah

84322-5310.

Literature Cited

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Alcock, J., G. C. Eickwort, and K. R. Eickwort. 1977. The reproductive behavior of Anthidium

maculosum (Hymenoptera: Megachilidae) and the evolutionary significance of multiple

copulations by females. Behav. Ecol. Sociobiol. 2:385-396.

VOLUME 63, NUMBER 2

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Frohlich, D. R. and F. D. Parker. 1985. Observations on the nest-building and reproductive behavior of a

resin-gathering bee: Dianthidium ulkei. Ann. Entomol. Soc. Amer. 78:804-810.

Grigarick, A. A. and L. A. Stange. 1968. The pollen-collecting bees of the Anthidiini of California. Bull.

Calif. Insect Survey 9:1-113.

Hicks, C. H. 1929. On the nesting habits of Callanthidium illustre (Cresson). Canadian Entomol. 61:1-8.

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Burks [eds.], Catalog of Hymenoptera in America North of Mexico. Washington, D.C.:

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Krombein, K. V. 1967. Trap-nesting wasps and bees; Life histories, nests and associates. Washington,

D.C.: Smithsonian Institution Press.

Parker, F. D. 1985a. Nesting habits of Osmia grinnelli Cockerell. Pan-Pac. Entomol. 61:155-159.

Parker, F. D. 1985b. Effective fungicide treatment for controlling chalkbrood disease (Ascomycetes:

Ascosphaeraceae) of the alfalfa leafcutting bee (Hymenoptera: Megachilidae) in the field. J. Econ.

Entomol. 78:35-40.

Parker, F. D. and R. M. Bohart. 1966. Host-parasite associations in some twig-nesting Hymenoptera

from western North America. Pan-Pac. Entomol. 42:91-98.

Parker, F. D. and D. R. Frohlich. 1983. Hybrid sunflower pollination by a manageable composite

specialist: The sunflower leafcutter bee. Environ. Entomol. 12:576-581.

Stephen, W. P. and J. M. Undurraga. 1976. X-radiography, an analytical tool in population studies of the

leafcutter bee. Megachilepacifica. J. Apic. Res. 15:81-87.

Thornhill, R. and J. Alcock. 1983. The evolution of insect mating systems. Cambridge: Harvard Univ.

Press.

Torchio, P. F. and V. J. Tepedino. 1980. Sex ratio, body size and seasonality in a solitary bee, Osmia

lignariapropingua Cresson. Evolution 39:993-1003.

Youssef, N. N., W. R. McManus and P. F. Torchio. 1985. Cross-infectivity potentials of Ascosphaera

spp. (Ascomyeetes: Ascosphaera) on the bee, Osmia lignaria propingua Cresson. J. Econ.

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PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 130-134

Observations on the Prey and Nests of Podalonia occidentalis Murray

(Hymenoptera: Sphecidae)

Howard E. Evans

Department of Entomology, Colorado State University, Fort Collins, Colorado

80523.

Abstract.—Podalonia occidentalis Murray is evidently a specialist on last instar

larvae of tent caterpillars (Malacosoma spp.) (Lasiocampidae), as evidenced by

records from New Mexico, Nevada, California, Alberta, and numerous records

reported here from north central Colorado. Nests are shallow and typical of other

Podalonia species, but much variation in details of nesting behavior was noted. Four

species of miltogrammine flies were reared from nests, the four together causing a

75% destruction of the wasps’ eggs at this locality.

It is a pleasure to dedicate this paper to that avid and versatile entomologist E.

Gorton Linsley, whose 1956 paper on Cerceris californica (with J. W. MacSwain) is a

small classic of its kind. Sphecology is the worse for his fascination with wasps’ “fuzzy

relatives,” the bees, and with longhorned beetles and diverse other insects. I spent

only a few days with “Gort” in the field, but they were enough to charge my batteries

for some time to come.

The present report concerns sphecid wasps of the genus Podalonia. Most species

of this genus prey on cutworms (Noctuidae), which they exhume from the ground

and use to provision a shallow nest dug nearby (reviews in Murray, 1940; Bohart and

Menke, 1976; see also O’Brien and Kurczewski, 1982; Steiner, 1983). There are,

however, at least two species of this genus that capture hairy caterpillars that live well

above ground. The best known of these is P. valida (Cresson), which is a specialist on

“woolly bears” of the genus Estigmene (Arctiidae) (Steiner, 1974, 1975). I have

many records from north central Colorado that indicate exclusive use of saltmarsh

caterpillars, E. acrea (Drury), in this area. A second species, P. occidentalis Murray,

has been the subject of a brief report by Murray (1940). Near Santa Fe, New Mexico,

these wasps were found “working with much effectiveness on the tent caterpillar” in

the month of June. Williams (1928) reported Podalonia violaceipennis (Lepeletier)

preying upon tent caterpillars at 1980 m in the Sierras of California. He found 11

nests in close proximity and believed that all were made by one female. The species

of Podalonia have often been confused in the past, and it seems quite possible that

Williams was dealing with occidentalis rather than violaceipennis.

There are three previously unpublished records of P. occidentalis preying upon

Malacosoma. R. M. Bohart (personal communication) has collected females

carrying tent caterpillars at Sagehen Creek, Nevada, where there was a large

population of the wasps in 1974, following an outbreak of tent caterpillars in previous

years. M. F. O’Brien has sent me two additional records of P. occidentalis based on

museum specimens (personal communication). A female in the Canadian National

Collections, from Fincher, Alberta, was collected by R. W. Salt on 9 July 1941 with a

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131

Malacosoma larva; and four females in the University of Michigan Museum of

Zoology, from Crowley Lake, Mono Co., California, were collected by L. Bezark on

8 June 1976 “carrying tent caterpillars.”

The observations reported here were made between 16 and 23 June 1985 and

between 12 and 30 June 1986, at three sites 2-5 km apart, all about 23 km west of

Livermore, Larimer Co., Colorado, at an elevation of about 2300 m. In this area

western tent caterpillars, Malacosoma californicum (Packard) (Lasiocampidae) are

extremely common, especially on bitterbrush, Purshia tridentata (Pursh). The active

period of P. occidentalis appears to correspond closely with the time when tent

caterpillars reach the final instar and leave their tents to feed individually. I did not

observe prey capture in the field but several times saw females carrying prey from

areas where caterpillars were feeding on Purshia bushes. Without exception prey

carriage and nest construction occurred during the morning hours, between 0840 and

1115 Mountain Daylight Time. Both males and females were frequent visitors to the

flowers of miner’s candle, Cryptantha virgata (Porter), which blooms in abundance

during the active period of the wasps.

The three study sites were all along trails or little-used dirt roads, either in tracks or

in bare places along the roadside. In every case there were infested Purshia bushes

not far away. It was common to see males patrolling these sites, flying back and forth

in a weaving pattern 5-15 cm above the ground. Occasionally a male descended upon

a female that was active at a nest, but contact was broken off immediately. I saw only

one copulation. In this case a female walked along a road with a male astride her,

holding her in the neck region with his mandibles. He extruded his genitalia briefly

before flying off after about 20 seconds. I did not see the initial union of the pair,

which may have occurred many seconds or minutes earlier. R. M. Bohart (personal

communication) observed about 40 males forming a struggling ball around a female

along a dusty road at Sagehen Creek, Nevada. This occurred during a period of

unusual abundance of Podalonia occidentalis.

I observed stinging of the prey twice. On 12 June, at 0950, a female was seen

carrying a caterpillar along a road. She dropped the prey and disappeared for three

minutes; when she returned she stung the prey seven times (even though it appeared

already well paralyzed). She mounted the prey obliquely over its back and stung it

along the midventral line, beginning at the thorax and moving back slightly each

time, covering most of the length of the abdomen. Essentially this same behavior was

observed in a terrarium, where a female Podalonia had been placed with tent

caterpillars.

Females carry their prey forward over the ground, holding it venter up, the wasp

grasping the caterpillar with her mandibles on the first or second abdominal segment

and straddling it. Since the caterpillars are much longer than the wasp, they extend a

considerable distance in front of and behind the wasp. In four instances the wasp was

seen to proceed to a plot of more or less bare, friable soil and then deposit her prey

while she searched for a place to dig. In one instance a female, with much apparent

effort, pulled her prey into a weed 2 cm off the ground. She then walked about and

scraped the soil here and there, returning to her prey in 18 minutes and again in 29

minutes, each time moving it slightly. Finally, 33 minutes after entering the site, she

started a nest 2.5 m away from the prey. Nest construction required only 10 minutes,

and the female then returned to the prey and carried it directly to the edge of the

hole. She then entered and pulled the prey in head first. She emerged within a few

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PAN-PACIFIC ENTOMOLOGIST

seconds and began filling the burrow by scraping in soil from the edge of the hole.

Filling was complete in 3.5 minutes, and the female then rested in the shade of a plant

and cleaned herself for 3 minutes before flying off.

Similar behavior was observed on three other occasions, although in each case the

prey was left on the ground in the shadow of plants or within a grass clump. In two

instances, a wasp was seen to leave a caterpillar under a plant and not return for

several hours. In one case the caterpillar was eventually carried off by several ants

{Formica sp.).

In contrast, there were three other occasions when it was clear that the wasp

prepared her nest first, leaving it open and provisioning it much later. In each case a

female was seen completing a nest in the morning, but the nest was not provisioned

and closed until the morning of the following day. Two of these instances occurred

late in the active season (21-22 June) when most tent caterpillars had spun their

cocoons. In one case the caterpillar was undersized and bore several eggs of

Tachinidae; evidently it had been unable to attain full size and pupate. Thus it

appears that when females are unable to obtain prey during a morning hunting

period, they proceed to dig a nest, leaving it open and filling it later. I found two nests

that were never filled.

Nests are shallow and are dug with rapid thrusts of the fore legs, small pebbles

being pulled out with the mandibles. The burrow is about 1 cm in diameter and is

oblique, terminating in a horizontal cell about 2.5 cm in length at a depth of from 2 to

6 cm (mean 3.9 cm, n = 14). Since the cell is shorter than the prey, the latter is coiled

in a broadly C-shaped posture, lying on its side. The egg measures 2.5 mm in length

and is laid vertically on the uppermost side of the abdomen, its anterior end attached

firmly to the prey, the posterior and extending ventrally free from the prey. Of 11

eggs recorded, seven were attached at an intersegmental membrane, three between

A2 and A3, two between A3 and A4, and two between A4 and A5. Four others were

attached toward the middle of the segment, three on A3 and one on A4.

Following oviposition, the female fills the burrow rapidly from soil at the periphery

of the entrance. In five of nine closures observed, much of the soil was taken from a

shallow quarry close beside the entrance; in three of these cases there were two such

quarries. The quarries varied from 0.5 to 2.0 cm in depth and from 2 to 4 cm in

distance from the entrance. Presumably these quarries are homologous to the

accessory burrows that have been described in a variety of digger wasps (Evans,

1966). A portion of the mound at the nest entrance is usually left intact, and nests

(especially those with quarries) can sometimes be spotted in the absence of the wasp.

Closures are not always completed level with the soil surface; in one case the top 1.5

cm of the burrow was left unfilled.

The egg hatches in about two days and full larval development requires eight to 10

days. However, of 12 nests in which the egg was recovered, nine contained maggots

that quickly destroyed the egg and later the prey. Four different species of

Miltogramminae (Sarcophagidae) were involved, maggots numbering two to 12 per

nest. In order of abundance the flies were:

Hilarella hilarella (Zetterstedt), 14 flies from 4 nests

Sphenometopa sp. nr. nebulosa (Coquillett), 10 flies from 3 nests

Taxigramma heteroneura (Meigen), 2 flies from 1 nest

Senotainia trilineata (Wulp), 2 flies from 1 nest

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133

Flies emerged from 22 to 32 days following the date of larviposition. Oddly, I rarely

saw these flies in the field; only on one occasion did I see a satellite fly perched on a

stone 15 cm from a female that was digging a nest. Others have noted that Hilarella is

an especially prevalent parasite of species of Podalonia (Murray, 1940; O’Brien,

1983).

Podalonia occidentalis appears to be a specialist on Malacosoma, and the wasps

disappear from the field when the tent caterpillars have pupated. On one occasion I

placed a female in a terrarium with several Malacosoma larvae; she stung one of

them (as described above) but failed to lay an egg or bury the prey. On a second

occasion I placed a Podalonia female in a terrarium with two Malacosoma, two

unidentified “woolly bears” (Arctiidae), and two cutworms (Noctuidae). She stung

one of the Malacosoma but failed to attack the others. These experiments are far

from conclusive, but so far as they go they do tend to support evidence from the field,

where 17 prey records were obtained, all involving M. californicum.

Discussion

Despite the host specificity of Podalonia occidentalis, many aspects of its behavior

seem unusually variable. Both prey and nests were sometimes abandoned; nests

were sometimes incompletely filled; some nests were filled with quarries (either one

or two) and others lacked quarries; egg position varied considerably. The most

striking variation was in the prey-nest or nest-prey dichotomy. This behavioral

difference is often considered a fundamental one, most Podalonia (and many other

more generalized wasps) taking prey before they make a nest (for reviews see Evans

and West-Eberhard, 1970; Iwata, 1976). However, such variation has been reported

in at least two other species of Podalonia (Myartseva, 1963 [cited in Bohart and

Menke, 1976]; Tsuneki, 1968). P. valida females regularly dig the nest before

obtaining prey (Steiner, 1975; personal observations). P. valida is unusual in that

females make a series of nests in a restricted territory that is defended against

intrusion by other females. There was evidence that P. occidentalis females return

again and again to the same general area to nest, but nests were not closely clumped

and there was no evidence of territorial behavior. Podalonia appears to be a genus in

transition with respect to whether the prey is taken before nest building (as it is in the

genus Prionyx, a member of the same subfamily) or whether the nest is dug first (as it

is in the closely related genus Ammophila).

The incidence of parasitism by miltogrammine flies (75%) is possibly the highest

ever recorded for a digger wasp. Since P. occidentalis nests earlier in the season than

most digger wasps, its inability to avoid the attacks of miltogrammines may permit

these flies to build up populations that are able to exploit species that nest later in the

summer (species of Ammophila, Philanthus, and other genera, in fact occurred in

these same study sites). The great abundance of tent caterpillars might seem to

compensate for the high incidence of parasitism, but in fact I have no evidence that

females ever provision more than one nest a day, and they are restricted to the brief

period when tent caterpillars are in the final instar. The impact of P. occidentalis on

the natural control of these pests appeared negligible in the study area.

Acknowledgments

I thank Mary Alice Evans for assistance with the field work, Richard M. Bohart

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PAN-PACIFIC ENTOMOLOGIST

(University of California, Davis) for confirming my identification of the wasps, and

N. E. Woodley (Systematic Entomology Laboratory, USD A) for identifying the

Sphenometopa.

Literature Cited

Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the world. A generic revision. Berkeley and Los

Angeles: Univ. Calif. Press, 695 pp.

Evans, H. E. 1966. The accessory burrows of digger wasps. Science, 152:465-471.

-, and M. J. West-Eberhard. 1970. The wasps. Ann Arbor: Univ. Michigan Press, 265 pp.

Iwata, K. 1976. Evolution of instinct. Comparative ethology of Hymenoptera. New Delhi: Amerind Publ.

Co. 535 pp.

Murray, W. D. 1940. Podalonia of North and Central America. Entomol. Amer., 20:1-82.

O’Brien, M. F. 1983. Observations on the nesting behavior of Podalonia argentifrons. Southwest.

Entomol., 8:194-197.

-, and F. E. Kurczewski. 1982. Ethology and overwintering of Podalonia luctuosa (Hymenoptera:

Sphecidae). Great Lakes Entomol., 15:261-275.

Steiner, A. L. 1974. Unusual caterpillar-prey records and hunting behavior for a Podalonia digger wasp:

Podalonia valida (Cresson). Pan-Pac. Entomol., 50:73-77.

-. 1975. Description of the territorial behavior of Podalonia valida (Hymenoptera: Sphecidae)

females in southeast Arizona, with remarks on digger wasp territorial behavior. Quaest. Entomol.,

11:113-127.

-. 1983. Predatory behavior of digger wasps (Hymenoptera: Sphecidae) VI. Cutworm hunting and

stinging by the ammophiline wasp Podalonia luctuosa (Smith). Melanderia, 41:1-16.

Tsuneki, K. 1968. The biology oiAmmophila in East Asia. Etizenia, 33:1-64.

Williams, F. X. 1928. The sphecid wasp, Podalonia violaceipennis (Lep.). Proc. Haw. Entomol. Soc.,

7:163.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 135-144

Biosystematic Studies of Ceylonese Wasps, XVIII:

The Species of Trachepyris Kieffer (Hymenoptera: Bethylidae:

Epyrinae)^

Karl V. Krombein

Smithsonian Institution, Washington, DC, 20560

Abstract.—Pristobethylus Kieffer, 1905, and Acanthepyris Kieffer, 1910, are new

synonyms of Trachepyris Kieffer, 1905. Epyris serricollis Westwood, Pristobethylus

indicus Muesebeck, P. crenaticollis Kieffer, P. semiserratus Kieffer and

Acanthepyris ceresensis Turner are new combinations in Trachepyris, and T.

haemorrhoidalis Kieffer is confirmed as a member of the genus. Trachepyris indicus

and T. haemorrhoidalis occur in Sri Lanka where they parasitize larvae of

Tenebrionidae.

A major problem with Kieffer’s large works on bethylid classification (e.g.,

1904-1906, 1914) is that he relied greatly on differences and did not give equal

significance to similarities. Evans was obliged to synonymize some 20 of Kieffer’s

genus-level names in his studies of New World Bethylidae. The same pattern is

beginning to emerge in the Old World bethylid fauna. During my revisionary study

of the Ceylonese Bethylidae, I became aware that Pristobethylus Kieffer, 1905, and

Acanthepyris Kieffer, 1910, were synonymous. Later, while sorting the extensive

bethylid collection made in Egypt by Priesner, I realized that Trachepyris Kieffer,

1905 {in Kieffer and Marshall, 1904-1906), was the senior synonym of these three

names.

These relationships were apparently sensed in part by two earlier workers on

Bethylidae. Turner (1928) must have subconsciously recognized the close

relationship of Acanthepyris and Pristobethylus, for he described two species in the

former genus, one of which was later transferred, correctly in the Kiefferian sense, to

Pristobethylus by Benoit (1957). Earlier, Benoit (1952) suggested that Acanthepyris

and Trachepyris were probably synonymous but did not make the synonymy and did

not mention this surmise in the 1957 paper.

Trachepyris Kieffer

Trachepyris Kieffer. 1905 (Jan): 107 (type-species, Trachepyris spinosipes Kieffer,

original designation and monotypic).

Pristobethylus Kieffer in Kieffer and Marshall, 1905 (Nov): 248 (type-species, Epyris

serricollis Westwood, original designation and monotypic). NEW SYNONYMY.

Epyris subg. Acanthepyris Kieffer, 1910: 103 (type-species, Epyris (Acanthepyris)

hildebrandti Kieffer, monotypic). NEW SYNONYMY.

Acanthepyris Kieffer, 1914: 401 (raised to generic rank).

The preceding number in this series is “XVII: A revision of Sri Lankan and South Indian Bembix

Latreille (Hymenoptera: Sphecoidea: Nyssonidae)” with J. van der Vecht, Smithson. Contrib. Zool.,

451:1-30,36 figs., 1987.

135

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PAN-PACIFIC ENTOMOLOGIST

Trachepyris females are unusual among the Bethylidae in the structure of the

mandible and antennal scape, and in the presence of a rake of stout bristles on the

fore tarsus. The mandible (Figs. 11-13) in dorsal view is somewhat flattened,

rounded at the apex, and has along the inner margin a strong inwardly directed tooth,

beneath which are modified sensilla chaetica (Figs. 14-18). The upper surface of the

scape (Figs. 5-7) is rather flattened and smooth, margined below by a row of close

short setae and above by a row of more scattered setae. The fore tarsal rake (Figs.

1-3) consists of stout setae along the outer margin, three on the basitarsus and one

each at the apices of the second and third segments.

The sensilla chaetica adjacent to the large tooth on the inner mandibular margin

are greatly enlarged and bizarrely modified. Their shape and position suggest that

they may have a fossorial function.

The stout setae of the fore tarsal rake obviously function in excavation of soil.

Possession of a tarsal rake is rare in the bethylids, for so few genera have fossorial

habits. Disepyris Kieffer has such a rake (Fig. 4) but it is composed of much longer,

more slender bristles than the short stout setae of Trachepyris.

Kieffer separated Pristobethylus from most Bethylidae because the pronotum of

the type-species is margined anteriorly and laterally by a scalloped carina.

Essentially, this carina is the only noteworthy difference between females of

Pristobethylus MidAcanthepyris. The significance can be gauged by the fact that the

lateral carina is complete in serricollis, crenaticollis (Kieffer) and ceresensis (Turner)

but present on only the anterior half in semiserratus (Kieffer) and indicus

(Muesebeck). The posterolateral angles of the head have a short scalloped carina in

serricollis, ceresensis and indicus; this was not noted in the descriptions of

crenaticollis and semiserratus. The fore-going species of Pristobethylus are all new

combinations in Trachepyris.

The radial vein is short in Trachepyris spinosipes and the costal vein bears a row of

extraordinarily long setae. These venational characters are subject to variation in

species assigned originally to Acanthepyris and Pristobethylus and do not warrant

separation of Trachepyris from the other two genera.

Treatment of these three assemblages of species as the spinosipes, serricollis and

hildebrandti species-groups is consistent with the treatment of similar groups in

Holepyris and other genera of Epyrinae. The hildebrandti group is the most

generalized and the spinosipes group the most specialized.

The spinosipes group is known from Algeria and Egypt. The other two groups are

primarily Ethiopian but each has one species in the Indian subcontinent.

Diagnosis. —Small wasps, 2.5-6.2 mm long; body black, apex of abdomen and

appendages sometimes red or brown. Head of female flattened, posterior margin

straight to emarginate; female mandible (Figs. 11-18) somewhat flattened above,

curved, apex bluntly rounded, inner margin with large sub apical tooth and modified

sensilla chaetica beneath and a smaller median tooth or two, male mandible

relatively slender, with large apical tooth and three to four small teeth above it;

clypeus narrow, with rounded median lobe and less prominent lateral lobes,

ecarinate medially; antenna 13-segmented, arising from beneath frontal lobes,

scrobes not carinate, female scape (Figs. 5-7) above somewhat flattened, mostly

smooth, margined below by short stout setae and above by longer setae, male

antenna relatively long, first and second flagellar segments subequal in length; malar

space absent; female eye not prominent, not hairy, not extending close to vertex.

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male eye more prominent. Pronotum longer than scutum, anterior and lateral

margins of disk carinate or not, posterior submarginal groove lacking, collar

depressed; scutum with distinct notauli and parapsidal furrows; scutellum with pair

of separated pits at base; mesopleuron with small pit below hind wing and curved

sulcus near lower margin; propodeal disk margined by a carina laterally and

posteriorly and with several discal carinae; posterior propodeal surface with median

carina on at least upper half; female fore femur somewhat to moderately broadened;

female fore tarsus (Figs. 1-3) with rake of stout bristles, three on basal segment and

one each at apices of second and third, male with weak tarsal rake; female mid tibia

spinose on outer surface; tarsal claw with inner tooth (Figs. 8-10); female forewing

(Figs. 25-27) with enlarged stigma, radial vein of variable length, basal vein meeting

subcosta only slightly basad of stigma, transverse median vein sometimes with short

stub. Abdomen somewhat depressed apically; male subgenital plate (Figs. 22-24)

with apical margin rounded or lobate, base with a median stalk; aedeagus relatively

broad, shorter than digitus, cuspis biramous, paramere relatively narrow, with or

without long apical setae (Figs. 19-21).

Behavior. —Little was known previously as to the host preferences of species of

Trachepyris. When Kieffer described haemorrhoidalis, he noted that the unique type

was captured while dragging a “chenille,” 8 mm long, on the sand of a dry stream

bed. Considering our prey records discussed below, it is probable that this

“caterpillar” was actually the larva of a tenebrionid beetle.

P. B. Karunaratne captured a female haemorrhoidalis at Palatupana Tank, 22

June 1978, dragging by its head end a slender paralyzed tenebrionid larva, 13.5 mm

long, belonging to an unknown genus of Tenebrionidae.

We obtained three host records for indicus at Ma Villu near Kondachchi; all were

slender larvae of a genus and species of Tentyriinae (Tenebrionidae). T. Wijesinhe

collected two females on 19 September 1979, each with a slender paralyzed larva,

11.5 and 15.2 mm long respectively. L. Jayawickrama collected a female on 18

September 1980 walking with a paralyzed larva, 6 mm long. She held the head end of

the larva in her mouth and the posterior section of the host body was over her back.

I watched indicus females in January 1979 hunting on the beach at Palatupana

between the dunes and the high tide mark. The wasps crawled swiftly over the sand,

occasionally taking short flights just above the surface. They examined the basal

rosette of leaves of small prostrate plants. Their larval hosts were presumably on the

roots of such plants. The stout spatulate setae of the fore tarsal rake would enable the

wasp to dig readily through the friable soil to reach a host larva. However, I did not

observe digging behavior by any of the wasps. The transport of host larvae noted

above indicates that the host is probably interred in a burrow separate from the site

where it was captured.

Behavioral data are unavailable for other species of Trachepyris. Inasmuch as

females of all species have similarly shaped mandibles and fore tarsal rake, I presume

that they too have tenebrionid larvae as hosts for which they search in sand or other

friable soil.

Collection data within Sri Lanka indicate that haemorrhoidalis is more widely

distributed than indicus, occurring in both the Dry Zone and Wet Zone at altitudes of

10 to 700 m with an average annual rainfall not over 2400 mm. The latter species is

restricted to sandy areas in the Dry Zone from sea level to 100 m with an average

annual rainfall not exceeding 1100 mm.

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PAN-PACIFIC ENTOMOLOGIST

Key to Trachepyris of the Indian Subcontinent

Pronotal disk not carinate anteriorly and laterally, surface with moderate sized

punctures separated by 2-3X diameter of puncture; costa with short setae only

(Fig. 25); head not carinate posterolaterally; female fore femur broader, 2.2X as

long as wide; male legs except coxae light red; apex of paramere with several long

setae, cuspis clavate at apex (Fig. 19) .

. haemorrhoidalis Kieffer

Pronotal disk with scalloped carina anteriorly and on basal half of lateral margin,

surface with only a few scattered moderate sized punctures; costa with longer setae

interspersed among shorter (Fig. 26); head posterolaterally usually with short

scalloped carina; female fore femur more slender, 2.4X as long as wide; male legs

dark brown, tarsi lighter; apex of paramere with only short setae, cuspis slender

(Fig. 20).

. indicus (Muesebeck)

Trachepyris haemorrhoidalis Kieffer

Figures 1, 5, 8,11,14,19, 22, 25

Trachepyris haemorrhoidalis Kieffer, 1911: 230-231 ($; Karachi, Pakistan, E.

Comber; holotype in British Museum (Natural History)).

Acanthepyris haemorrhoidalis YdQitQT, 1914: 404.—Kurian, 1954: 275.

Female. —Length 4.7-5.8 mm. Black, mandible, scape, flagellum beneath, tegula,

fore and mid femora and all tibiae occasionally, tarsi, and last two or three abdominal

segments red; wings slightly infumated, stigma medium brown, veins lighter. Head

with length 0.86X width, not carinate posterolaterally, posterior margin slightly

incurved; large tooth on inner margin of mandible subapical in position (Fig. 11);

four modifled sensilla chaetica beneath subapical tooth (Fig. 14); front delicately

alutaceous, scarcely impressed anteriorly in middle, moderately punctate, those

anteriorly separated by half a puncture diameter, becoming sparser posteriorly and

separated by twice or more a puncture diameter, least interocular distance 0.7X head

width and 1.5-1.6X eye length; ocelli small, front angle 90°, posterior ocelli

separated by half a diameter from posterior margin of head, ocellocular distance

1.4-1.5X width of ocellar triangle. Thoracic dorsum delicately alutaceous,

propodeal disk glossy; pronotal disk without marginal carinae, punctures of

moderate size, separated by half a puncture’s width along anterior margin, dispersed

by two or more puncture widths elsewhere; scutum with a few small punctures in

middle; median length of propodeal disk 0.6X width, enclosed median area twice as

wide at base as at apex, quinquecarinate, median and lateral carinae stronger,

reaching discal apex, intermediate carinae weaker and sometimes not reaching apex,

surface between carinae with transverse carinules; median carina on upper

three-fourths of posterior surface; fore femur relatively broad, 2.2X as long as wide;

costa with short setae only (Fig. 25); transverse median vein with stub.

Male. —Length 4.4—4.8 mm. Coloration as in female except abdomen black and

legs except coxae light red. Head with length 0.9X width, posterior margin slightly

incurved, not carinate posterolaterally; front glossy, moderately punctate, closely so

anteriorly, punctures separated by once or twice diameter of puncture posteriorly,

least interocular distance 0.55X head width and subequal to eye length; ocellocular

distance I.IX width of ocellar triangle. Thorax shining; pronotal disk without

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139

Figures 1-10. Trachepyris and Disepyris females. 1-4. Fore tarsus: 1, T. haemorrhoidalis Kieffer (X45);

2, T. indicus (Muesebeck) (X84); 3, T. spinosipes Kieffer (X67); 4, D. rufipes Kieffer (X73). 5-7. Scape; 5,

T. haemorrhoidalis (X134); 6, T. indicus (XlOl); 7, T. spinosipes (X134). 8-10. Claws: 8, T.

haemorrhoidalis (X224); 9, T. indicus (X280); 10, T. spinosipes (X280).

140

PAN-PACIFIC ENTOMOLOGIST

anterior and lateral carinae; median length of propodeal disk 0.6X width, enclosed

median area quinquecarinate, lateral carinae converging strongly toward apex, none

of carinae reaching apex; costa and subcosta with short setae only; transverse median

vein with stub. Genitalia and subgenital plate (Figs. 19, 22); apex of paramere with

several long setae; cuspis clavate at apex.

Specimens examined .—SRI LANKA. NORTHERN PROVINCE. Mannar

District: 0.5 mi NE Kokmotte Bungalow, Wilpattu Natl. Pk., 21-25 May 1976, 6 ;

Ma Villu, Kondachchi, 16-19 Sep 1980, d. NORTH CENTRAL PROVINCE.

Anuradhapura District: Padaviya, Irrigation Bungalow or Antiquities Site, 180 ft,

2-8 Nov 1970, 6 ; 18, 21 May 1976,1 in Malaise trap, IS ; 20-23 July 1978, Malaise

trap, 9. CENTRAL PROVINCE. Kandy District: Udawattakele Sanctuary,

Kandy, 2100 ft, 16-31 Aug and 1-17 Sep 1976, 29. WESTERN PROVINCE.

Colombo District: Colombo Museum Gardens, 50 ft, 23, 28 Mar and 13-15 Apr

1977, 9,26. SABARAGAMUWA PROVINCE. Ratnapura District: Panamure,

500 ft, 15-21 Oct 1970,2 9. UVA PROVINCE. Badulla District: Ulhitiya Oya, 15 mi

NNE Mahiyangana, Malaise trap, 5-6 Sep 1980, 36. Monaragala District:

Angunakolapelessa, Malaise trap, 8-9 Oct 1980, 6. SOUTHERN PROVINCE.

Hambantota District: Palatupana Tank, 10-20 m, 27-29 Sep 1977 and 22 June 1978,

29.

Trachepyris indicus (Muesebeck), New Combination

Figures 2, 6, 9,12,15,17, 20, 23, 26

Pristobethylus indicus Muesebeck, 1934: 233-225, Fig. 1 (9; Chowghat, Malabar,

India, K. P. A. Menon; holotype inU. S. National Museum).—Kurian, 1954:273.

Female .—Length 4.0-6.2 mm. Black, mandible, scape, flagellum beneath, tegula,

occasionally fore and mid femora and all tibiae, tarsi, and last one to three abdominal

segments red, basal segments of legs usually brown; wings slightly infumated, stigma

dark brown, veins much lighter. Head with median length 0.78-0.92X width,

posterolaterally usually with short scalloped carina, posterior margin strongly

emarginate; large tooth on inner margin of mandible apical in position (Fig. 12), with

a broad cutting edge dorsally (Fig. 17); two modified sensilla chaetica beneath apical

tooth, third modified sensillum chaeticum displaced to base of apical tooth (Fig. 17)

by rounded boss along inner margin (Fig. 15); front delicately alutaceous, with short

median groove anteriorly, moderately punctate, punctures anteriorly separated by

half a puncture diameter, becoming much sparser on rest of lower half of front and

virtually absent on upper half; least interocular distance 0.7X head width and

1.5-1.8X eye length; ocelli small, front angle about 135°, posterior ocelli separated

by half a diameter from posterior margin of head, ocellocular distance 1.0-1.IX

width of ocellar triangle. Thoracic dorsum delicately alutaceous, propodeum glossy;

pronotal disk with scalloped carina anteriorly, extending half distance to apex

laterally, with widely dispersed punctures of moderate size; scutum with only a few

small punctures posteriorly in middle; median length of propodeal disk 0.6-0.7X

width, enclosed median area quinquecarinate, only median carina reaching discal

apex, sides converging strongly toward apex, area between carinae with irregular

transverse carinules; median carina on upper half of posterior surface; fore femur

2.4X as long as wide; costa with longer setae interspersed among shorter (Fig. 26);

transverse median vein with short stub.

VOLUME 63, NUMBER 2

141

Figures 11-18. Trachepyris female mandibles. 11-13. Apical half, dorsal: 11, haemorrhoidalis Kieffer

(X236); 12, indicus (Muesebeck) (X201); 13, spinosipes Kieffer (X372). 14-16. Apical half, ventral: 14,

haemorrhoidalis (X236); 15, indicus (X248); 16, spinosipes (X395). 17. Apical half, oblique, indicus

(X378). 18. Modified sensilla chaetica, ventral, spinosipes (X944). (t = tooth, s = modified sensilla

chaetica; b = mandibular boss.)

142

PAN-PACIFIC ENTOMOLOGIST

Male. —Length 2.5-4.5 mm. Coloration similar to female except abdomen entirely

black, legs and antennae rarely light red. Head with median length 0.8X width,

posterolaterally rounded or slightly irregular from a few punctures, posterior margin

not so deeply emarginate as in female; front glossy, with scattered small punctures,

somewhat more sparsely so on posterior half; least interocular distance 0.64-0.68X

head width and 1.3-1.5X eye length; ocellocular distance 1.1-1.2X width of ocellar

triangle. Thorax shining; pronotal disk with scalloped carina anteriorly and laterally

halfway to apex, weaker than in female; median length of propodeal disk 0.7-0.8X

width, enclosed median area tri- or quinquecarinate, only median carina reaching

apex, lateral carinae converging strongly toward apex, area with irregular transverse

carinules; costa with longer setae interspersed among shorter; transverse median

vein without stub. Genitalia and subgenital plate (Figs. 20, 23); apex of paramere

with short setae; cuspis slender.

Specimens examined. —SRI LANKA. NORTHERN PROVINCE. Mannar

District: Ma Villu, Kondachchi, 19 Sep 1979, 2$ , 2c?; 16-19 Sep 1980, $ , (?; 11-12

Apr 1981, c?. UVA PROVINCE. Badulla District: Ulhitiya Oya, 15 mi NNE

Mahiyangana, on or in leaf litter on sand, 5-6 Sep 1980, 6. Monaragala District:

Mau Aru, 10 mi E of Uda Walawe, 100 m, 24-26 Sep 1977, $ . SOUTHERN

PROVINCE. Hambantota, 10 ft, 28 Oct 1970, 29 . Palatupana, near Wildlife and

Nature Protection Society Bungalow, 0-50 ft, 8-10 Mar 1972, 29,2c?; 20-22 June

1978, Malaise trap, 9 ; 18-21 Jan 1979, 5 9.

INDIA. KERALA, Chowghat, Malabar, 29 May 1931, caught on sand, 9

(holotype). MADRAS. Karikal, Tanjore, 17 Aug 1951, 29.

Trachepyris spinosipes KiefFer

Figures 3, 7,10,13,16,18, 21, 24, 27

Trachepyris spinosipes Kieffer, 1905: 107.—Kieffer in Kieffer and Marshall, 1906:

413 (9; Algeria: Ismailia, Ain Sefra; type series in Paris Museum).—Kieffer,

1914: 406.

I have examined a series of 18 females and a single male from 10 localities in Egypt.

A description is withheld for a revisionary study of Egyptian Bethylidae.

Acknowledgments

I am pleased to dedicate this paper to E. Gorton Linsley, an occasional wayfarer in

entomological areas other than biosystematics of bees and long-horned beetles. Gort

is a dedicated naturalist, always interested in elucidating the complex, sometimes

arcane relationships between his solitary bees and their nest associates. His detailed

pioneer work with J. W. MacSwain (1957) on the interactions of Stylops pacifica

Bohart with its ground-nesting host bee Andrena complexa Viereck afforded

valuable insights a few years later in my observations on the trap-nested solitary

eumenid, Euodynerus foraminatus apopkensis (Robertson), and its stylopid

parasite, Pseudoxenos hookeri (Pierce).

I am grateful to P.B. Karunaratne, T. Wijesinhe and L. Jayawickrama,

technicians with the Smithsonian’s Ceylon Insect Project, for their assistance in the

field and for the host records they obtained.

I thank T. J. Spilman and J. R. Dogger, Systematic Entomology Laboratory, U. S.

Department of Agriculture, for identifying the tenebrionid larvae. M. C. Day,

VOLUME 63, NUMBER 2

143

Figures 19-27. Trachepyris species. 19-21. Male genitalia, ventral at left, dorsal at right: 19, haemor-

rhoidalis Kieffer; 20, indicus (Muesebeck); 21, spinosipes Kieffer. 22-24. Male subgenital plate, ventral;

22, haemorrhoidalis’, 23, indicus', 24, spinosipes. 25—'ll. Female right forewing: 25, haemorrhoidalis', 26,

indicus; 27, spinosipes.

144

PAN-PACIFIC ENTOMOLOGIST

British Museum (Natural History), and C. O’Toole, Oxford University, furnished

helpful information on the types of haemorrhoidalis and serricollis respectively. I am

indebted to my friend and mentor for many years, Carl F. W. Muesebeck, for his

thorough review of the manuscript.

The line drawings are by George L. Venable, Department of Entomology,

Smithsonian Institution (SI). Beth Norden (SI) made the painstaking preparations of

specimens for the scanning electron microscope and assisted in the photography;

Susan G. Braden (SI) made the micrographs.

Literature Cited

Benoit, P. L. G. 1952. Bethylidae (Hym.) Nord Africains nouveaux ou pen connus du Museo Civico di

Storia Naturale di Genova. Doriana 1, No. 27:1-7.

- 1957. Hymenoptera-Bethylidae. Parc Natl. Albert, Mission G. F. de Witte, 1933-1935, Fasc.

88:1-57.

Kieffer, J. J. 1905. Description de nouveaux Proctotrypides exotiques. Ann. Soc. Sci. Bruxelles,

29:95-142.

-, 1910. Serphidae, Cynipidae, Chalcididae, Evaniidae und Stephanidae aus Aequatorialafrika.

Wiss. Ergeb. Deut. Zent.-Afrika-Exped. 1907-1908, 3, Lief. 2:1-29.

-, 1911. Nouveaux Bethylides et Dryinides exotiques du British Museum de Londres. Ann. Soc. Sci.

Bruxelles, 35:200-233.

-, 1914. Bethylidae. Das Tierreich, 41:1-595,205 figs.

-andT. A. Marshall. 1904-1906. Proctotrypidae in E. Andre. Species des Hymenopteres d’Europe

& d’Algerie, 9:1-552,21 pis.

Kurian, C. 1954. Catalogue of Oriental Bethyloidea. Agra Univ. Jour. Res., 3:253-288.

Linsley, E. G. and J. W. MacSwain. 1957. Observations on the habits of Stylops pacifica Bohart. Univ.

Calif. Pubs. Entomol. 11:395-430.

Muesebeck, C. F. W. 1934. Seven new species of Indian Bethylidae (Hymenoptera). Rec. Indian Mus.,

36:223-232.

Turner, R. E. 1928. New Hymenoptera of the family Bethylidae. Ann. &Mag. Nat. Hist. (10) 1:129-152.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 145-146

A New Mexican Species of Linsleyella Chemsak

(Coleoptera: Cerambycidae)

John A. Chemsak

University of California, Berkeley

The purpuricenine genus Linsleyella Chemsak (1984) previously contained three

species, virens (Bates), ricei Chemsak, and michelbacheri Chemsak. The new species

described below is one of the many with longitudinal eburneous vittae, which

indicates that this characteristic spans a number of genera.

It is a pleasure to dedicate this paper to E. Gorton Linsley, friend and colleague.

Christine Jordan prepared the illustration.

Linsleyella virgulata, New Species

(Figure 1)

Male .—Form small to moderate sized; integument dark metallic blue-green,

appendages metallic; pubescence pale, long, erect. Head small, front irregularly

punctate, with numerous long, erect, dark hairs; vertex coarsely, confluently

punctate, long, erect hairs moderately dense; antennae longer than body, basal

segments shining, moderately coarsely, confluently punctate, erect and

subdepressed setae numerous, segments from sixth opaque, densely clothed with

very short, appressed pubescence, third segment longer than first, fourth shorter

than third, slightly longer than first, eleventh segment acute at apex. Pronotum

broader than long, sides usually subangulate behind middle; disk convex,

moderately coarsely, subconfluently punctate, often with a longitudinal, median,

glabrous callus; pubescence long, erect; prosternum rather finely, transversely

punctate, moderately densely clothed with long, pale, erect pubescence;

mesosternum subopaque at sides; metasternum deeply, separately punctate at

middle, sides subopaque, densely clothed with pale, depressed pubescence, long,

suberect hairs numerous. Elytra more than 2^/2 times as long as broad, sides slightly

tapering toward middle; each elytron with an eburneous longitudinal vitta near

suture, extending from basal margin to near apical margin and another, narrower

pair at sides behind humeri but not extending to apex; punctures between vittae

coarse, subconfluent, epipleura subopaque; pubescence long, erect; apices sinuate

truncate, inner angles dentate. Legs slender; femora confluently punctate,

pubescence long, erect, hind pair extending almost to elytral apices; tibiae

moderately clothed with subdepressed hairs. Abdomen finely, densely punctate at

sides, middle almost glabrous; last sternite subtruncate at apex, shallowly

emarginate at middle. Length, 8-12 mm.

Female .—Form more robust. Antennae shorter than body, segments from sixth

enlarged. Legs with femora shorter. Abdomen with last sternite broadly

subtruncate, shallowly emarginate at middle. Length, 8-13 mm.

Holotype male, allotype (California Academy of Sciences) from 4 miles SW

Morelos Canada, Puebla, Mexico, 20 September 1977 (J. Chemsak, A. & M.

Michelbacher). Paratypes include: 26 males, 17 females, same data; 14 males, 21

145

146

PAN-PACIFIC ENTOMOLOGIST

Figure 1. Lins ley ella virgulata Chemsak, $ .

females, 7 km SE Morelos Canada, 4 October 1975 (J. Chemsak, J. Powell, T.

Eichlin, T. Friedlander); 2 males, 7 km SE Morelos Canada, 4-10 July 1974 on Selloa

glutinosa flowers. (J. Chemsak, J. Powell, E. G. Linsley); 2 males, 1 female,

Tehuacan, Puebla, Mexico, 17 October 1941 (DeLong, Good, Caldwell &

Plummer).

The eburneous vittae of the elytra make this species distinctive from other known

Linsleyella. The coloration is fairly uniform within the type series although a little

variation is evident in the thickness of the yellowish vittae. In males the discal pair

tend to be broader anteriorly and in females the bands are somewhat narrow.

Adults were mostly collected on flowers of Selloa glutinosa at the type locality.

Literature Cited

Chemsak, John A. 1984. Description of a new purpuricenine genus Linsleyella (Coleoptera:

Cerambycidae). Pan-Pac. Entomol. 60:114—118.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 147-150

A New Genus and Species in the Tribe Macrotomini

(Coleoptera: Cerambycidae) from Costa Rica

Edmund F. Giesbert

9780 Drake Lane, Beverly Hills, California 90210.

Abstract. —A single new Cerambycid genus and species from Costa Rica is

described and figured: Parastrongylaspis linsleyi (Macrotomini).

The genus and species described below, and proposed in conjunction with ongoing

studies of the cerambycid fauna of Monteverde, Costa Rica, and its environs by F. T.

Hovore, is so far unique to that locality.

Parastrongylaspis, New Genus

Form stout, convex. Head with front short, concave; vertex with median line

carinate; mandibles stout, arcuate; genae usually apically produced, parallel; palpi

nearly equal, apical segment of maxillary pair slightly larger, truncate at apex; eyes

large, moderately coarsely facetted; antennal tubercles prominent, divergent;

antennae 11 segmented, moderately robust and slightly longer than body in male,

less robust and shorter than body in female, serrate, scape short, compressed, outer

segments finely longitudinally striolate, flattened, third segment about twice as long

as scape, longer than fourth to tenth segments, eleventh segment of males longest,

appendiculate. Pronotum wider than long, convex, lateral suture not expanded nor

crenulate, hind angles spinose, disk uneven; prosternum narrow, intercoxal process

slender, strongly arcuate, with apex rounded, coxal cavities open behind, strongly

angulate externally; mesosternal process moderately slender, coxal cavities open to

epimera; metasternum with episternum broad, sides subparallel. Scutellum cordate,

moderately large, convex, asperate. Elytra nearly 2^/2 times as long as width across

humeri, sides subparallel, apices widely rounded, with sutural angle dentate. Legs

moderately stout; trochanters of male deeply excavated ventrally, with excavation

densely pubescent; femora linear, tibiae feebly arcuate, distally expanded and

apically spined; tarsi with third segment moderately expanded, cleft to base,

metatarsi with first segment longer than following two together. Abdomen normally

segmented.

Type species.—Parastrongylaspis linsleyi New Species

This genus resembles Strongylaspis Thomson, and presumably bears a close

relationship to that neotropical genus. It may be easily separated by the flattened,

serrate antennae and modified trochanters of the male, and by the lack of lateral

crenulations of the pronotum. The species of Strongylaspis which occur north of

South America have received little attention from modern systematists, with the

exception of S. corticaria Erichson, which is quite abundant in collections, and has

been redescribed by Linsley (1962) and de Zayas (1975).

147

148

PAN-PACIFIC ENTOMOLOGIST

Parastrongylaspis linsleyi Giesbert, New Species

(Fig. 1)

Male .—Form moderately large, robust. Integument dark yellow brown, head,

pronotum, and appendages reddish brown. Head moderately closely granulate and

granulate-punctate, with fine, long, suberect golden pubescence on front and vertex;

median line feebly cariniform, slightly darkened; antennal tubercles moderately

prominent; antennae moderately robust, usually exceeding elytral apices by one or

two segments, scape somewhat flattened, moderately coarsely punctate, sparsely

pubescent, segments 3 to 11 serrate, somewhat flattened, finely longitudinally

striolate and glabrous, third segment nearly twice as long as scape, about 1^/4 times as

long as fourth, segments 4 to 10 subequal, eleventh segment slightly longer than

third, appendiculate. Pronotum wider than long, convex, sides straight, tapering

anteriorly, with a small stout spine at each posterior angle; disk with an indistinct

obhque cicatrix on each side before middle; surface granulate, moderately densely

clothed with long, fine, erect, golden hairs not obscuring surface. Scutellum convex,

widely rounded behind, bearing distinct, transverse, cicatrix-like asperites, and

fringed with fine golden hairs. Elytra parallel sided, strongly convex anteriorly, less

so toward apices, which are widely, separately rounded with sutural angle dentate;

surface moderately densely granulate, granules becoming less distinct toward apices,

with indistinct fine, short, subdepressed pubescence. Underside granulate, with

sternum densely clothed with fine, erect golden pubescence; abdomen with

pubescence less dense, terminal sternite widely emarginate at apex. Legs with

trochanters scaphiform, ventrally modified into a deep, cup-like excavation filled

with long pale hairs; femora sublinear, somewhat compressed, moderately sparsely

punctate and pubescent, distally asperate beneath; tibiae asperate, finely pubescent,

feebly curved, flattened, and widened distally, with outer apical angle acuminate.

Length 17-28 mm.

Female .—^Form similar to male. Head with antennal tubercles somewhat less

prominent; antennae moderately slender, subserrate, reaching at most to apical 1/3

of elytra, segments from fifth striate. Abdomen with apex of terminal sternite feebly

bilobed, deeply emarginate in middle. Legs with ventral surface of trochanters

shallowly excavated, and bearing dense, fine, erect hairs. Length 21-28 mm.

Types.—Holotype male, allotype (California Academy of Sciences), and 19

paratypes, from Monteverde, Puntarenas prov., COSTA RICA, with the following

data: 1 male, 1 female, 3-5 June 1974 (E. Giesbert); 1 male, 1-3 June 1978 (E.

Giesbert); 6 males, 4 females, 26 May-4 June 1984 (E. Riley, D. Rider, D. LeDoux);

1 male, 22-24 May 1985 (F. Hovore); 1 male, 5 May 1980 (W. A. Haber); 1 male, 5

April 1981 (Haber); 1 male, 1 female, 24-28May 1985 (Haber); 1 female, 17-20May

1985 (J. Chemsak); 3 males, 9-12 June 1986 (Hovore, Giesbert).

Remarks .—The combination of the peculiar modification of the male trochanters,

with the distinctly pubescent pronotum and serrate antennae, most noticeably in the

male, is unique among the known Central American macrotomine fauna.

I would like to thank F. T. Hovore and J. E. Wappes for providing specimen data

from their fine personal collections, and J. A. Chemsak for data from the collection

at the Essig Museum of Entomology, Berkeley, California, as well as his review of

the manuscript.

VOLUME 63, NUMBER 2

149

Figure 1. Strongylaspis linsleyi Giesbert, male

150

PAN-PACIFIC ENTOMOLOGIST

It is a pleasure to dedicate this handsome species to my friend E. Gorton Linsley in

recognition of his lifetime of devotion to the study and to the students of entomology.

Literature Cited

Linsley, E. G. 1962. The Cerambyddae of North America. Part 11. Taxonomy and Classification of the

Parandrinae, Prioninae, Spondylinae, and Aseminae. University of Cahfornia Publications in Entomology

19:1-102.

Zayas, F. de. 1975. Revision de la famiha Cerambyddae (Coleoptera: Phytophagoidea). Habana. 433 pp.

Publications Received

The Sucking Lice of North America. An Illustrated Manual for Identification. By

Ke Chung Kim, Harry D. Pratt, and Chester J. Stojanovich. The Pennsylvania State

University Press, University Park and London, xii + 241 pp., figs. 1-203, pis. 1-76.

Available from The Pennsylvania State University Press, 215 Wagner Building,

University Park, Pennsylvania 16802. Price $39.50 clothbound. ISBN 0-271-00395-2.

Publication date given as 23 December 1986. Copy received by PCES at CAS on 31

December 1986.

This volume, dedicated to the late Professor Gordon F. Ferris recognizes 76

species of Sucking Lice as occurring in North America, from a total of currently

approximately 500 described world species. The authors estimate the world

Anopluran fauna at about 1,000 species.

Chapters on Collecting and Preparation Techniques, Morphology and Diagnostic

Characters, Biology and Immature States, Public Health and Veterinary Importance

precede the Synopses of North American Anoplura, while chapters on Parasite-Host

List, Host-Parasite List, References, and Index complete the volume.

In addition to illustrations occupying a full page plate for each species, each

couplet to the keys to the families, genera and species of the North American

Anoplura is finely and helpfully illustrated (a total of 190 figures). This would

certainly meet with the approval of Ferris. The majority of the illustrations were

prepared by Mr. Stojanovich, who in 1951 collaborated with Professor Ferris in the

pubhcation of “The Sucking Lice.” This latter publication is still available from the

Pacific Coast Entomological Society for the nominal price of $10.00.

—Paul H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San

Francisco, California 94118.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 151-154

A New Genus and Species of Cerambycidae from Costa Rica

[Coleoptera]

Frank T. Hovore

Placerita Canyon Nature Center, 19152 W. Placerita Canyon Road, Newhall, CA

91321

Preparation of a faunal inventory of the Cerambycidae of the Monteverde Cloud

Forest and surrounding environs revealed a number of undescribed taxa. The

following new genus and species are presented at this time to make the name

available for the Monteverde study, and to pay tribute to E. Gorton Linsley, a friend

and source of professional guidance and inspiration to me for many years.

Gortonia , New Genus

Form elongate, cylindrical. Head with front vertical, subquadrate, with a narrow,

moderately deep, transverse impression parallel to the clypeal margin; genae broad,

slightly produced laterally; palpi very short, terminal segments elongate,

subcylindrical, bluntly rounded at apices; mandibles stout, broad at base, apices

acute, narrowly emarginate internally; eyes finely faceted, moderately deeply

emarginate, upper lobes small, widely separated on vertex, lower lobes large,

rounded; antennal tubercles slightly elevated, obtuse, divergent, directed

posteriorly; antennae 11 segmented, slender, scape stout, subconical, shorter than

third segment, segments 3 to 8 subequal in length in male, slightly decreasing in

length in female, intermediate segments feebly expanded and dentate externally at

apices. Pronotum with sides evenly rounded or with a very feeble indication of a

median lateral tubercle, basal margins narrowly expanded laterally over elytral

humeri; discal surface smooth, broadly convex, feebly depressed medially;

prosternum broad, evenly convex, prosternal process narrow, arcuately declivous

behind in male, more evenly rounded and slightly expanded apically in female, coxal

cavities feebly angulated externally, open behind by about the width of the apex of

the prosternal process; mesosternal process about twice as broad as that of procoxae,

feebly convex, evenly arcuate between coxae, occasionally tuberculate anteriorly in

females; metepisternum narrow, sides tapering posteriorly. Elytra elongate, narrow,

only slightly tapering apically, surface metallic, shining; apices evenly, separately

rounded to suture. Scutellum subchordate, apex acute. Legs with profemora

moderately stout, gradually expanded apically, mesofemora nearly twice as long as

profemora, less strongly expanded apically, feebly carinate on inner and outer

surfaces at apical one-half, metafemora very elongate, sinuate, extending to or

beyond elytral apices in both sexes; tibiae slender, straight, metatibiae longest, with

two apical spurs; posterior tarsi slender, elongate, first segment longer than

following two together, third segment cleft almost to base. Abdomen normally

segmented, female with setal brush.

151

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PAN-PACIFIC ENTOMOLOGIST

Tribal placement: Purpuricenini, near Eriphus Serville.

Type species: Gortonia linsleyi, New Species

The elongate, cylindrical form, long posterior legs, long, slender antennae,

smooth, unarmed pro thorax, and metallic elytra distinguish Gortonia from all other

purpuricenine genera. Superficially, Gortonia resembles an elongated Batyle

Thomson, but structurally appears to be more closely related to Purpuricenopsis

Zajciw, Axestoleus Bates, Zenochloris Bates, and Eriphus Serville. From

Purpuricenopsis it differs by the feebly dentate antennal segments, evenly rounded

and unarmed elytral apices (emarginate-dentate in Purpuricenopsis), unarmed

femoral apices, and longer posterior tarsi. (Comparisons with Purpuricenopsis are

based upon Zajciw’s (1969) generic key and redescription of the genus.) The genus

Eriphus may contain polyphyletic elements, with both metallic and non-metallic

species, exhibiting a variety of pronotal shapes. From Eriphus sensu stricto,

Gortonia differs immediately by the unarmed prothorax, densely pubescent,

metallic elytra, with the apices separately rounded, long, slender antennae, and

shorter, more quadrate front. Zenochloris Bates, which appears to differ from

Eriphus primarily by the metallic coloration of its species, may be distinguished from

Gortonia by the other characters enumerated above for the separation of Eriphus.

Axestoleus differs by the non-metallic coloration, stouter body form, truncate elytral

apices, shorter antennae, shorter legs, and callouses on the pronotal disk.

Eriphusprolixus Bates resembles G. linsleyi in general body form, and may belong

in Gortonia; however, I have seen only a color transparency of the type specimen,

and generic assessment must await examination of the underside and appendages.

Gortonia linsleyi, New Species

(Figure 1)

Male. —Form moderate-sized, dorsal surface of body very feebly depressed;

integument black, prothorax and basal three-fourths of anterior femora

reddish-orange, pronotal disk usually with dark infuscation medially and at sides on

the apical one-half, median marking often in shape of narrow, inverted triangle,

elytra metallic greenish-gray; body pubescence mostly pale, black on elytral apices

and portions of antennae, meso- and metatibiae. Head coarsely, sparsely,

transversely punctate on neck and vertex, genae with a few large punctures, front

densely, finely, irregularly punctate, densely clothed with short, fine,

posteriorly-appressed pubescence, longitudinal midline moderately-deeply

impressed between antennal tubercles; eyes with upper lobes separated on vertex by

about the diameter of antennal scape, lower lobes large, about one-third taller than

genae; genal apex slightly produced laterally, rounded, with narrow emargination

medially at mandibular insertion; antennae surpassing elytral apices by about four

segments, scape robust, densely, finely punctate, finely setose, second segment

moniliform, about as long as wide, segments 3 to 5 distinctly, thinly fringed internally

with black, suberect hairs, distal segments with a few scattered erect hairs internally,

all segments sparsely clothed with short, very fine, appressed pubescence, segments

3 to 7 feebly expanded and dentate externally at apices, segments 3 to 8 subequal in

length, segment 9 slightly shorter, segment 10 shortest, segment 11 subequal in

length to segment 9, sinuate, curved outward, Pronotum slightly wider than long,

base and apex constricted, base with a narrow, transverse sulcus, apex about

VOLUME 63, NUMBER 2

153

Figure 1. Gortonia linsleyi Hovore, male.

one-third narrower than base, median lateral tubercle evident only as a small,

impunctate, tumid area, discal surface nearly glabrous, microscopically

alutaceous-punctate, with scattered larger, seta-bearing punctures, sides with fine

punctures and large, irregularly-shaped or elongate punctures intermixed, thinly

clothed with fine, short, erect pubescence, a few longer hairs present on basal

two-thirds; prosternum coarsely, sparsely, irregularly punctate, finely,

moderately-densely pubescent; meso- and metasternum densely, minutely punctate.

154

PAN-PACIFIC ENTOMOLOGIST

densely clothed with fine, suberect pubescence. Elytra more than three times longer

than humeral width, surface densely, moderately to coarsely punctate, punctures

becoming shallower, less defined, confluent apically, densely clothed with pale,

recumbent pubescence which does not obscure the surface, hairs at apex of elytra

longer, coarser, black; apices separately rounded. Legs with profemora densely,

minutely punctate-pubescent, meso- and metafemora moderately to coarsely

punctate and pubescent. Abdomen densely, minutely punctate, densely clothed with

long erect pubescence, apical margins of sternites narrowly glabrous and

impunctate; second and fifth sternites subequal in length; apex of fifth sternite and

fifth tergite each broadly rounded or feebly truncate, fringed with long pale hairs.

Length: 9-14 mm.

Female .—Coloration similar to that of male, form slightly more robust; elytra very

slightly shorter; pronotal infuscation lacking; antennae surpassing elytral apices by

about two and one-half segments, segments beginning with third decreasing slightly

in length, tenth shortest, apical segment straight; pronotal disk more lightly punctate

than in male, sides nearly glabrous, punctate areas only feebly indicated by surface

irregularities; abdomen with fifth sternite nearly twice as long as second, apex of fifth

sternite and fifth tergite each broadly rotundate-truncate, feebly emarginate

medially, fringed with long pale hairs. Length: 12-15 mm.

Holotype male, allotype (California Academy of Sciences) and 19 paratypes (14

males, 5 females) from COSTA RICA, Puntarenas Province, 6 km S Santa Elena,

9-12 June 1986, on blossoms of Croton sp. (F. T. Hovore). Additional paratypes (9

males, 3 females), all topotypical: 5 males, 3 females, 5-7 June 1980 (E. F. Giesbert,

J. E. Wappes); 1 male, 6-7 June 1983 (E. F. Giesbert); 2 males, 1 female, 18 May

1984 (F. T. Hovore, R. L. Penrose); 2 males, 9 June 1986 (E. F. Giesbert). A single

male specimen, not designated as paratypical, is at hand from PANAMA, Panama

Province, Bayano district, 3-5 km W Ipeti, 19 May 1985, on blossoming tree. Aside

from its smaller size (9 mm), this specimen does not differ significantly from

topotypical material.

Variation in the type series is minimal. Some individuals have a pronounced bluish

tint to the elytra, and most males possess the dark median pronotal macula. In a few

individuals this macula is reduced to a vague apical patch, while in others it extends

posteriorly to the base of the disk. The normally reddish profemora are heavily

infuscated with black in a few males, and appendage length appears to be somewhat

allometric; larger individuals possess relatively longer antennae and legs.

It is my great pleasure to dedicate this new genus and species to E. Gorton Linsley,

Professor Emeritus, University of California, Berkeley.

Acknowledgments

I am grateful to R, L. Penrose (California Department of Food and Agriculture)

and J. A. Chemsak (University of California, Berkeley) for reviewing the

manuscript; and to E. F. Giesbert for specimen data and color slides of cerambycid

type specimens.

Literature Cited

Zajciw, D. 1969. Sinopse dos generos brasileiros da tribo purpuricenini Fair m., 1864, Rev. Brasil. Biol.,

29(1): 109-120.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 155-164

Ovaries, Ovarioles, and Oocytes in Parasitic Bees

(Hymenoptera: Apoidea)

Byron Alexander and Jerome G. Rozen, Jr.

Department of Entomology, Cornell University, Ithaca, New York 14853 and

Department of Entomology, American Museum of Natural History, Central Park

W. at 79th Street, New York, New York 10024.

Abstract .—Ovarian features of parasitic bees including the number of ovarioles

per ovary, the number and size of mature oocytes, and the morphology of mature

oocytes are explored. Included are data from 44 species representing approximately

8 separately derived cleptoparasitic lineages within the Apoidea. The number of

ovarioles of parasitic Halictidae and Megachilidae is 3 per ovary, as is characteristic

of non-parasitic Colletidae, Andrenidae, Halictidae, Melittidae, and Megachilidae.

In the anthophorid/apid lineage, which has 4 ovarioles per ovary as a plesiomorphic

condition, the Nomadinae tend to have increased numbers of ovarioles, with 5

ovarioles per ovary seeming to be the most common condition. In two other parasitic

lineages of the Anthophoridae (Melectini and Ctenioschelini), only Ericrocis lata

with 5 ovarioles has an increase from the plesiomorphic state. In the Apidae,

Psithyrus consistently exhibits an above normal number of ovarioles.

Cleptoparasitic bees in all families tend to have a larger number of mature oocytes

in their ovaries at a given time than do solitary bees, and these oocytes tend to be

smaller than are those of solitary bees. Mature oocytes of the Nomadinae show

considerable variation in structure from one another and from the oocytes of

non-nomadine bees, as illustrated by Nomada, Ammobates and Triepeolus.

Introduction

Rozen (1986a) published a survey of the number of ovarioles in various taxa of

bees, and realized, as had Iwata (1955, 1960, 1964, 1965) and Iwata and Sakagami

(1966) before, that the number of ovarioles and other ovarian characteristics of bees

have phylogenetic, taxonomic and adaptive significance. Cleptoparasitic bees

especially seem replete with variable ovarian features. The works of these authors

suggested that interesting patterns of variation exist from one parasitic lineage to

another and that at least some of the variation may be explained in terms of the mode

of life of the bees. To explore these matters more broadly, we collected, preserved in

fixative and examined as many additional parasitic taxa as possible. Using both

recently collected specimens and information from the literature, this paper reports

on the number of ovarioles of different cleptoparasitic lineages, on the number and

size of their mature oocytes and on other aspects of the morphology of the oocytes of

the Nomadinae.

With great pleasure we dedicate this article to Dr. E. Gorton Linsley, one of the

great systematists of cleptoparasitic bees (as well as many other groups of insects).

The second author remembers with fondness being required to undertake the life

155

156

PAN-PACIFIC ENTOMOLOGIST

history study of one species of bee as part of his dissertation research by his major

professor, Gort Linsley. This simple requirement led the second author to 35 years of

digging holes both shallow and deep in most of the continents of the world in search

of biological data about, and immature stages of, solitary and parasitic bees. Gort

Linsley more than anyone else taught him that the data base for systematics was far

greater than the specimen on a pin.

The work of the first author was aided by the Undergraduate/Graduate Research

Program of the American Museum of Natural History, supported by the Greenwall

Foundation.

We thank Dr. George C. Eickwort, Cornell University, for reviewing this manu¬

script.

Materials and Methods

Original observations reported in this paper are based upon dissections of

specimens that either were fixed and preserved in Kahle’s solution prior to dissection

(Epeolus zonatus from the Cornell collection and all of the AMNH specimens except

Triepeolus sp. R) or were dissected as freshly killed specimens and subsequently

fixed in Kahle’s solution (all other Cornell material). The internal organs of

specimens that have been preserved prior to dissection are somewhat brittle and

more difficult to dissect without damaging structures one wishes to observe. New

material reported in this paper deals with 25 species in 16 genera and 3 families of

cleptoparasitic bees.

Because we compare our observations with the large data set previously gathered

by Iwata (1955,1960,1965) and Iwata and Sakagami (1966), we use their definitions

of stages of oocyte development and their measurement of body size. We repeat the

definitions of their terms used in this paper: (1) EGG LENGTH (E) is the length (to

the nearest 0.1 mm) of “the largest oocyte in the ovaries”; (2) MESOSOMAL

WIDTH (M) is the “distance between the outer extremities of the tegulae” (mean

value if more than one specimen was available); (3) EGG INDEX is the ratio E/M;

and (4) MATURE OOCYTES are all oocytes in Iwata’s (1955) categories A or B,

defined as follows:

Category A: “mature oocytes, the nutritive cells of which have quite

disappeared. In certain species the mature oocytes diminish their

sizes a little to take their proper shapes.”

Category B: “nearly mature oocytes, the nutritive cells of which begin to

collapse. Generally they attain the maximum size proper to each

species, but have not yet taken their proper shapes.”

We count all oocytes tallied by Iwata as category A or B as mature oocytes because

we were not confident that we could reliably distinguish between the two categories

in our specimens. (Iwata worked primarily with fresh rather than preserved material,

and his definitions of four discrete categories of developing oocytes were formulated

in the framework of a survey of the oocytes of all Hymenoptera, rather than bees

alone. The distinctions may be more apparent and significant in other Hymenoptera

than they are in bees.)

The statistical tests used in this paper to analyze the number of mature oocytes in

parasitic bees are Mann-Whitney U Tests (where sample sizes are adequate,

P-values are based upon a standardized normal distribution of the U statistic). A

VOLUME 63, NUMBER 2

157

non-parametric statistical test is used because distributions are decidedly

non-normal. Data for solitary species are all from Iwata (1955,1960,1965) or Iwata

and Sakagami (1966). Data for parasitic species are from Iwata (13 species) and the

present study (18 species).

Results

Number of ovarioles. —In most families of bees (Colletidae, Andrenidae,

Halictidae, Melittidae, andMegachilidae; unknown for the Stenotritidae, Oxaeidae,

and Ctenoplectridae), there are 3 ovarioles per ovary (indicated by the notation 3:3

in Table 1), regardless of whether the bees are solitary, parasitic, subsocial, or social

(Iwata and Sakagami, 1966; Rozen, 1986a). The present study corroborates this

pattern for cleptoparasitic species in the Halictidae and Megachilidae (no

cleptoparasitic bees in the other families for which data are available) (Table 1). In

the Anthophoridae and Apidae, the presumed plesiomorphic condition is 4 ovarioles

per ovary (i.e., 4:4 in Table 1). A notable exception is the subfamily Nomadinae, all

of whose members are cleptoparasitic. Ovariole number appears to be more variable

within the Nomadinae than in most other bees. It differs not only from species to

species, but among individuals within a species, and sometimes even between ovaries

within an individual (Table 1). Despite this variability, the number of ovarioles is

almost always higher than the common anthophorid/apid condition of 4:4, and

within the Nomadine 5:5 seems to be the most common condition.

We have examined four species of cleptoparasitic anthophorids that are not in the

subfamily Nomadinae. Three species have the common anthophorid condition of 4

ovarioles per ovary, but Ericrocis lata has 5 ovarioles per ovary (Table 1).

Within the Apidae, an inerease in ovariole number has been reported in five

species of the socially parasitic genus Psithyrus (Cumber, 1949). The exact number of

ovarioles is extremely variable, ranging from 6 to 18 per ovary (Table 1). By contrast,

Bombus, the presumed sister group of Psithyrus, possesses the plesiomorphic

number 4. (Queens in the related advanced eusocial genus Apis are remarkable

among bees for their large number of ovarioles. Snodgrass (1956) estimated that

Apis mellifera queens have 160 to 180 ovarioles per ovary.)

The fact that both Psithyrus and Ericrocis lata have more than 4 ovarioles (the

plesiomorphic anthophorid/apid number) suggests that selection pressure for

increased number of ovarioles has independently played a role in three separate

groups in the anthophorid/apid line. We suspect, as did Iwata (1955,1964) and Iwata

and Sakagami (1966), that increased number of ovarioles functions to increase the

total number of eggs that an individual can deposit. It also permits a greater number

of mature oocytes to be ready for deposition within a short time period as pointed out

below.

Number of mature oocytes. —Cleptoparasitic bees in all families tend to have a

larger number of mature oocytes in their ovaries at a given time than do solitary bees.

The difference is statistically significant, whether one tallies the total number of

mature oocytes or the number of mature oocytes per ovariole (Table 2). The pattern

also holds up if one divides the cleptoparasitic species into two subgroups on the basis

of whether they have the same number of ovarioles as their non-parasitic relatives or

an increased number of ovarioles. In comparing these two subgroups, we find that

those species with an increased number of ovarioles have a significantly higher total

158

PAN-PACIFIC ENTOMOLOGIST

Table 1. Number of ovarioles and size and number of mature oocytes in various taxa of cleptoparasitic bees. Defini¬

tions of terms and notations are given in the text.

Taxon

Egg

Index

Total

Number

Mature

Oocytes

Mature

Oocytes

per

Ovariole

Number of

Ovarioles

Number of

Specimens

Examined Reference

Halictidae

Sphecodes esakii

0.36

1.50

3:3=*

Iwata, 1955

Sphecodes japonica

—

0.67

3:3“

Iwata, 1955

Sphecodes sp. A

0.57

1.00

3:3

present study

Sphecodes sp. B

0.76

—

3:3

present study

Megachilidae

Dioxys pacificus

0.74

0.33

3:3

present study

Stelis sp.

0.61

2.67

0.44

3:3

present study

Euaspis basalis

0.53

3.5

0.58

3:3“'=

Iwata, 1955,1960

Coelioxys (Rhinocoelioxys) sp.

0.41

—

prob.3:3

present study

Coelioxys yanonis

—

0.33

3:3“

Iwata, 1955

Coelioxys fenestratus

0.54

3.00

3:3“

Iwata, 1955

Coelioxys brevis

0.40

0.83

3:3“

Iwata, 1955

Coelioxys sp. (138)

—

1.50

3:3“

Iwata, 1955

Coelioxys decipiens

—

0.67

prob. 3:3^*

Iwata, 1965

Anthophoridae

Anthophorinae

Thyreus japonicus

0.85

3.5

0.438

4:4

Iwata, 1955

Zacosmia maculata desertorum

0.74

0.75

4:4

present study

Mesoplia proh. rufipes

0.74

0.50

4:4

present study

Ericrocis lata

0.77

0.20

5:5

present study

Anthophoridae

Nomadinae

Neolarra (Neolarra) californica

0.33

13.75

1.325

5:5

present study

5:6

present study

Neolarra (Neolarra) sp.

—

5:5 (or 5:4)

present study

Neolarra (Phileremulus) vigi-

Ians

—

1.10

5:5

present study

Holcopasites calliopsidis

—

5:6

present study

Ammobates carinatus

0.47

0.50

6:6

present study

Oreopasites sp. A

0.49

0.74

5:4

present study

5:5 (or 6:4)

present study

Oreopasites vanduzeei

—

approx. 11 total

Rozen, 1986a

prob. 5:5

Rozen, 1986a

Kelita chilensis

0.30

0.70

prob. 5:5

Rozen,1986a

5:5

present study

Epeolus zonatus

0.53

8.5

0.85

5:5

present study

Epeolus scutellaris

0.76

6.33

0.45

7:7

present study

Epeolus sp.

0.56

0.40

5:5

present study

Epeolus japonicus

1.00

6.5

0.54

6:6

Iwata, 1955

Triepeolus pectoralis

0.62

0.50

5:5

present study

continued

VOLUME 63, NUMBER 2

159

Table 1. continued

Taxon

Egg

Index

Total

Number

Mature

Oocytes

Mature

Oocytes

per

Ovariole

Number of

Ovarioles

Number of

Specimens

Examined

Reference

Triepeolus sp. R

0.49

0.20

5:5

present study

Centrias articulata

0.28

—

5:5

present study

Nomada banksi

0.30

15.5

1.55

5:5

present study

Nomada vicina

0.39

0.80

5:5

present study

Nomada illinoiensis

—

5:4

present study

5:5

present study

Nomada (“Gnathias”) sp.

—

5:5

present study

Nomada pyrifera

0.50

14.5

1.45

5:5

Iwata, 1955

Nomada sp. nr. glabella

0.38

8.67

0.90

5:5

Iwata, 1955

5:4

Iwata, 1955

Nomada japonica

0.39

21.125 2.15

5:5

Iwata, 1955

4:4

Iwata, 1955

6:5

Iwata, 1955

Apidae

Psithyrus barbutellus

—

10:10

Cumber, 1949

10:12

Cumber, 1949

6:7

Cumber, 1949

8:9

Cumber, 1949

Psithyrus bohemicus

—

14:13

Cumber, 1949

Psithyrus campestris

—

7:8

Cumber, 1949

6:7

Cumber, 1949

Psithryus rupestris

—

18:15

Cumber, 1949

15:13

Cumber, 1949

16:14

Cumber, 1949

12:14

Cumber, 1949

Psithyrus vestalis

—

10:10

Cumber, 1949

^Species included in Iwata’s Table IV (A), comprising “Anthophila with three pairs of ovarioles.”

'’Iwata’s Fig. 50 shows a single ovary which clearly has only 3 ovarioles.

^wata (1960) explicitly lists number of ovarioles (for a single specimen) as 3:3.

number of mature oocytes, but they do not produce more oocytes per ovariole

(Table 2).

Hence different lineages of parasitic bees have independently undergone an

increase in the number of mature oocytes in their ovaries at a given time. This may

reflect that parasitic bees produce more eggs in their life span than do solitary bees

and that as a result a larger number of eggs are ready for deposit in a short interval.

However, it is also likely that there is selective advantage for parasitic bees to be able

to oviposit in rapid succession. Unlike a female solitary bee that must construct and

provision a cell before each oviposition, a female cleptoparasite may find, and

therefore must be ready to lay eggs in, more than one host cell in a short time period.

Size of mature oocytes .—Iwata and Sakagami (1966) reported that the mature

160

PAN-PACIFIC ENTOMOLOGIST

Table 2. Comparison of numbers of mature oocytes in parasitic vs. solitary bees.

Total Mature

Number of Oocytes

Mature

Oocytes

per

Ovariole

Sample

Size

1) Solitary Bees

/ 2.17 .

Us = 320

Us = 725

Us = 678.5

Z = 3.913

lz = 5.16

1 Z = 4.41

P<0.001 <

P < 0.001

P<0.01

2) Cleptoparasitic Bees 1

7.18 i

' 0.86J

a) With Unchanged 1

Number of Ovarioles

k 5.62 V

0.90V

i 14

Us = 170

Us = 124

lz = 2.02 1

1 Z = 0.198

I P < 0.05 I

( P = 0.84

b) With Increased

Number of Ovarioles

8.46

' 0.84>/

1 17

oocytes of cleptoparasitic bees tend to be smaller than those of solitary species. Our

observations corroborate this pattern, as shown in Figure 1 (which combines data

from the present study and Iwata and Sakagami’s study). It also appears that the

distribution of egg sizes is relatively symmetrical in solitary species but decidedly

assymmetrical and weighted toward smaller egg sizes in cleptoparasitic bees.

The egg index proposed by Iwata and Sakagami is a measure of oocyte size relative

to overall body size. They suggested such a measure because they expected that

oocyte size would vary with body size in a regular manner. Indeed, when maximum

oocyte length is plotted against mesosomal width, both parasitic and non-parasitic

species show a positive linear relationship between the two variables (Fig. 2. r^ = .758

for parasites; r^= .748 for solitary species). Within the lower ranges of body sizes,

parasitic species of a given size seem consistently to have smaller oocytes than

solitary species of the same size. Such a trend is not apparent among larger bees in

our sample.

Morphology of oocytes of the Nomadinae. —As has been noted by others (Iwata,

1960; Rozen, 1986a), mature oocytes of Nomadinae show remarkable variation in

structure from one another and from the oocytes of non-nomadine bees. Figures 3-5

illustrate the oocyte features of some of the bees that we examined. The nipple-like

structure on the anterior end of the oocyte of Nomada vicina (Fig. 3) was found also

on two other Nomada seen by us but not on a fourth species; Iwata (1960) depicted

nipples in two of the three species he studied. We presume that the corrugations that

predominate on one side of the oocytes of Ammobates (Fig. 4) and Oreopasites

(Rozen, 1986a, Fig. 1) permit the eggs to bend U-shaped as they are inserted into the

cell walls of the hosts, as seems characteristic of the Ammobatini (Rozen, 1986b)

with the exception of Pseudodichroa (Rozen and Michener, 1968). The completely

annular corrugations of Triepeoluspectoralis oocytes (Fig. 5) and of the eggs of some

other species in the same genus (Bohart, 1966) may allow for the expansion of the egg

as it absorbs water in the cell wall of the host. We believe that oocytes of other

VOLUME 63, NUMBER 2

161

Figure 1. Frequency distributions for oocyte size (expressed as the EGG INDEX, or ratio of length of

mature oocyte to mesosoma width) in cleptoparasitic bees (solid line) and solitary bees (dashed line).

Nomadinae will also be distinctive in shape and that shape will be correlated with the

mode of oviposition of the taxon.

Material Studied

Sphecodes sp. A, 2 mi E of Apache, Cochise Co., Arizona, 9 May 1986 (J. G.

Rozen). Sphecodes sp. B, 3 mi SW of Rodeo, Hidalgo Co., New Mexico, 5 May 1965

(J. G. Rozen). Dioxys pacificus Cockerell, 2 mi E of Apache, Cochise Co., Arizona,

9 May 1986 (J. G. Rozen). Stelis sp. A, Cienega Ranch, Hidalgo Co., New Mexico,

13 May 1986 (J. G. Rozen). Coelioxys (Rhinocoelioxys) sp., near Chetumal,

Quintana Roo, MEXICO, 12 October 1986 (G. C. Eickwort). Zacosmia maculata

desertorum Cockerell, 2 mi E of Apache, Cochise Co., Arizona, 9 May 1986 (J. G.

Rozen). Ericrocis lata (Cresson), 1 mi E of Douglas, Cochise Co., Arizona, 21

August 1986 (J. G. Rozen). Mesoplia prob. rufipes Perty, Hollis Reservoir, near

Valencia, TRINIDAD, 3 March 1968 (J. G. Rozen). Ammobates carinatus

162

PAN-PACIFIC ENTOMOLOGIST

MESOSOMA WIDTH (mm)

Figure 2. Scatter plot of mesosomal width versus length of mature oocytes in various species of bees.

□ = solitary; ★ = cleptoparasites with same number of ovarioles as non-parasitic relatives;

# = cleptoparasites with increased number of ovarioles.

Morawitz, 10 km S of Skhirate, MOROCCO, 29 April 1968 (J. G. Rozen & E.

Suissa). Oreopasites A, 4 mi E of Willcox, Cochise Co., Arizona, 5 September

1986 (J. G. & B. L. Rozen), from nesting area of Nomadopsis meliloti (Cockerell).

Kelita chilensis (Friese), Penuelas, Valparaiso Prov., CHILE, 28 October 1969

(J. G. Rozen). Triepeolus sp. R, 11 mi S of Animas, Hidalgo Co., New Mexico, 20

August 1986 (J. G. Rozen). Triepeolus pectoralis (Robertson), Cornell Campus,

Ithaca, Tompkins Co., New York, 3 September 1985 (B. Alexander). Epeolus

scutellaris Say, Cornell Campus, Ithaca, Tompkins Co., New York, 1-5 September

1985 (B. Alexander). Epeolus sp. , 4 mi E of Willcox, Cochise Co., Arizona, 8 May

1986 (J. G. Rozen). Holcopasites calliopsidis (Linsley), Van Natta sand pit, Ithaca,

Tompkins Co., New York, 11 July 1985 (B. Alexander). Neolarra (Neolarra)

californica Michener, Cienega Ranch, Hidalgo Co., New Mexico, 10 May 1986

(J. G. Rozen). Neolarra (Phileremulus) vigilans (Cockerell), 19 mi SW of Apache,

Cochise Co., Arizona, 29 August 1986 (J. G. & B. L. Rozen), from nesting site of

Perdita lenis and Perdita sp., Neolarra (Neolarra) sp. , 4.8 mi SW of Rodeo, Hidalgo

VOLUME 63, NUMBER 2

163

Figure 3-5. Mature oocytes of Nomadinae, anterior end at top. 3. Nomada vicina. 4. Ammobates

carinatus. 5. Triepeolus pectoralis. Scale = 1.0 mm.

Co., New Mexico, 1 September 1975 (J. G. Rozen), from Perdita mentzeliae nest #7.

Centrias articulata (Smith), Cornell campus, Ithaca, Tompkins Co., New York, 11

July 1985 (B. Alexander). Nomada banksi Cockerell, Van Natta sand pit, Ithaca,

Tompkins Co., New York, 22, 29 September 1985 (B. Alexander). Nomada vicina

Cresson, van Natta sand pit, Ithaca, Tompkins Co., New York, 23 September 1985

(B. Alexander). Nomada sp. (perhaps illinoiensis Robertson), Cornell campus,

Ithaca, Tompkins Co., New York, 15 May 1986 (B. Alexander). Nomada

{'‘Gnathias”) sp., near Junius Ponds, 6miNW of Waterloo, Seneca Co., New York,

18 May 1986 (B. Alexander).

Literature Cited

Bohart, G. E. 1966. Notes on Triepeolus remigatus (Fabricius), a “cuckoo bee” parasite of the squash

bee, Xenoglossa strenua (Cresson). Pan-Pacific Ent., 42:255-262.

Cumber, R. A. 1949. Humble-bees and commensals found within a thirty mile radius of London. Proc.

Royal Ent. Soc. London (A), 24:119-127.

Iwata, K. 1955. The comparative anatomy of the ovary in Hymenoptera. Part 1. Aculeata. Mushi,

29:17-34.

-. 1960. The comparative anatomy of the ovary in Hymenoptera. Supplement on Aculeata with

descriptions of ovarian eggs of certain species. Acta Hymenopterologica, 1:205-211.

-. 1964. Egg giantism in subsocial Hymenoptera, with ethological discussion on tropical bamboo

carpenter bees. Nature and Life in Southeast Asia, 3:399-435.

-. 1965. The comparative anatomy of the ovary in Hymenoptera (records on 64 species of Aculeata

in Thailand with descriptions of ovarian eggs). Mushi, 38:101-111.

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Iwata, K. & S. F. Sakagami. 1966. Giantism and dwarfism in bee eggs in relation to the modes of life, with

notes on the number of ovarioles. Japanese Jour. Ecology, 16:4-16.

Rozen, J. G. 1986a. Survey of the number of ovarioles in various taxa of bees (Hymenoptera, Apoidea).

Proc. ent. Soc. Washington, 88:707-710.

-. 1986b. The natural history of the Old World nomadine parasitic bee Pasites maculatus

(Anthophoridae: Nomadinae) and its host Pseudapis diversipes (Halictidae: Nomiinae). Amer.

Museum Novitates, no. 2861, 8 pp.

Rozen, J. G. and C. D. Michener. 1968. The biology of Scrapter and its cuckoo bee, Pseudodichroa

(Hymenoptera: Colletidae and Anthophoridae). Amer. Museum Novitates, no. 2335,13 pp.

Snodgrass, R. E. 1956. Anatomy of the Honey Bee. Cornell University Press, Ithaca, New York.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 165-171

A Revision of the Bee Genus Aztecanthidium

(Hymenoptera: Megachilidae)

Roy R. Snelling

Natural History Museum of Los Angeles County, 900 Exposition Blvd., Los

Angeles, California 90007.

Abstract. —The exclusively Mexican bee genus Aztecanthidium is revised and the

three known species separated in a key. One new species, A tenochtitlanicum, is

described from the State of Jalisco. Distribution data for the two previously known

species are provided. Distinctive morphological features of each species are

illustrated.

This paper is dedicated to my friend and colleague, E. Gorton Linsley, the first

live-bee systematist I was privileged to meet, more than 35 years ago. Thank you,

Gort, for your kindness and encouragement!

Introduction

The higher classification of the New World anthidiine bees was reviewed by

Michener (1948) and the component genera separated by a key. The Mexican genus

Aztecanthidium was subsequently described by Michener and Ordway (1964) for two

previously undescribed species. A third species, heretofore undescribed, has

prompted the present study.

Material Examined

Specimens used in this study are from the following collections: Estacion Biologica

Chamela, Universidad Autonoma de Mexico (CHAM); Natural History Museum of

Los Angeles County (LACM); Snow Entomological Collection, University of

Kansas (UKAN); USD A Bee Biology and Systematics Laboratory, Utah State

University (UTSU).

Systematics

Aztecanthidium has been adequately described by Michener and Ordway (1964),

and there is no need to repeat that description. The few known species are

moderate-sized, sparsely hairy bees, and the body is more or less elongate and

parallel-sided. Two of the species are principally reddish with more or less defined

yellowish areas on the head and body; they resemble the species of Paranthidium,

subgenus Mecanthidium Michener (1942). The third species, described below, is

black and has sharply contrasting yellow markings.

Aztecanthidium differs from Mecanthidium in possessing a sharply carinate

(almost flange-like) preoccipital ridge across the top of the head and down the side to

the hypostomal carina; the mandibles of both sexes have one or two teeth on the

cutting margin; the clypeus, especially of the female, is protuberant; the pronotal

lobes are carinate; the last metasomal tergum of the male is bilobed. In

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Mecanthidium the preoccipital ridge is weak and present only across the top of the

head; the mandibles of both sexes are without teeth on the cutting margin (except in

the male of one species, which has a single tooth); the clypeus, in profile, is weakly

convex; the pronotal lobes are lamellate; the last metasomal tergum of the male is

protuberant and blunt.

In Michener’s (1948) key to the New World anthidiine genera, Aztecanthidium

runs to Allanthidium, from which it differs by the depressed tergal margins, the

completely carinate preoccipital ridge, the presence of an anterior mesepisternal

Carina, and the lack of a row of pits across the base of the propodeum.

The type species ot Aztecanthidium is^. xochipillium Michener and Ordway.

Key to Species of Aztecanthidium

la. Male tergum 4 with short lateral tooth or angle and emargination of tergum 7 as

wide as deep or wider (Figs. 4, 6); apex of female labrum concave and clypeus

moderately protuberant and rounded in profile (Figs. 3,5).2

b. Male tergum 4 without lateral tooth or angle and emargination of tergum 7

deeper than wide (Fig. 2); apex of female labrum tridentate and clypeus strongly

protuberant and angulate in profile, with biarcuate carina (Fig. 1) .

. cuauhtemocum

2a. Male tergum 7 with lateral margins mainly convex and apical lobes broad (Fig.

6 ); female clypeal profile evenly convex from base to summit of apical declivity

(Fig. 5); both sexes principally reddish, with limited blackish areas and obscure

yellowish marks . xochipillium

b. Male tergum 7 with lateral margins mainly concave and apical lobes narrow (Fig.

4); female clypeal profile convex at base, concave toward apical declivity and

distinctly elevated at declivity (Fig. 3); both sexes black with yellow maculations

. tenochtitlanicum

Aztecanthidium cuauhtemocum Michener and Ordway

Figures 1-2, 7-9

Aztecanthidium cuauhtemocum Michener and Ordway, 1964: 75; S $ .

The type series was collected 2 mi S Tzitzio, 4450 feet elevation, Michoacan, 29

July 1962 (M. G. Naumann), on Leguminosae, and all four specimens are in the

UKAN collection. I have seen a paratype of each sex. No additional specimens have

been available for study.

Both sexes of this species are easily recognized by the figures and the

characteristics cited in the key. The female is especially characterized by the

biarcuate, strongly protuberant clypeal carina and the tridentate labral apex. The

male is less obviously distinct, but lacks a lateral tooth on metasomal tergum 4 and

the emargination of tergum 7 is deeper than wide. Both sexes are largely ferruginous

bees, with obscure yellowish marks, especially on the face and the metasomal terga.

Aztecanthidium tenochtitlanicum, New Species

Figures 3-4,10-12

Diagnosis. —Both sexes black and yellow, scutellum weakly bilobed. Female

labrum emarginate and clypeus moderately protuberant and with small preapical

tubercle in profile (Fig. 3). Male metasomal tergum 4 angulate or subdentate at side,

emargination of tergum 7 wider than deep and apical lobes relatively narrow (Fig. 4).

Fig. 1-6. Lateral view of female head and apical male metasomal terga, respectively, of : 1-2,

Aztecanthidium cuauhtemocum; 3-4, A. tenochtitlanicum; 5-6, A. xochipillium.

Description male holotype .—Measurements (mm): head width 4.67; head length

3.85; wing length 11.3; total length 16.1.

Paratypes .—head width 4.29-4.68; head length 3.33-3.87; wing length 10.2-11.3;

total length 12.3-16.3.

Head and body black, the following yellow: clypeus, except narrow black band

along apical margin; paraocular area, ending broadly at about mid-level between

base of clypeus and antennal socket; spot adjacent to inner upper eye margin at level

of anterior ocellus; broad stripe on either side of anterior margin of mesoscutum and

short, narrow stripe adjacent to tegula; broad posterior stripe on scutellum; small

anterior spot on tegula; outer apical spot on metacoxa; large lateral spot on tergum 1;

tergum 2 with preapical bar across middle one-third and with very narrow preapical

stripe extending to large lateral spot; terga 3 and 4 similar to 2, but with preapical

stripe only slightly broadened at sides; tergum 5 with only very narrow preapical

stripe on lateral one-third; sterna 2-5 with irregular lateral blotches.

Head about 1.2 times as broad as long, inner eye margins slightly divergent below.

Clypeus rugosopunctate. Paraocular and supraclypeal areas and frons

rugosopunctate to subcontiguously punctate, interspaces moderately shiny; vertex

similar but punctures subcontiguous and slightly larger. Ocellocular and interocellar

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Fig. 7-15. Male genitalia and metasomal sterna 7-8 of: 7-9, Aztecanthidium cuauhtemocum.-, 10-12, A.

tenochtitlanicum; 13-15, A. xochipilium.

distances about equal; ocelloccipital distance about 1.6 times interocellar distance.

Greatest width of gena slightly less than width of eye. Labrum slightly depressed

along midline. Apical tooth of mandible long, acute; second and third teeth obtuse;

inner tooth short, acute.

Mesoscutal interspaces moderately shiny between subcontiguous, moderate

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169

punctures. Propodeum dull, interspaces distinctly roughened, with broad median

impunctate area. Dorsal face of scutellum with disc weakly depressed in middle,

more broadly so distad, apical margin slightly concave between weak sublateral

tubercles. Punctures of mesepisternum very coarse (about 0.08 mm), contiguous to

subcontiguous; punctures of metepisternum moderate to coarse, mostly contiguous,

but with some irregular interspaces up to a puncture diameter; interspaces of meso-

and metepisterna shiny; side of propodeum roughened and slightly shiny between

fine, subcontiguous punctures.

Profemur more than three times as long as broad or thick; meso- and metafemora

less than three times as long as broad or thick. Pro- and mesobasitarsi each shorter

than combined lengths of following segments; metabasitarsus longer than following

segments combined.

Punctures of metasomal terga fine to moderate, subcontiguous to dense across

middle, becoming almost uniformly subcontiguous at sides, interspaces moderately

shiny. Tergum 4 with low, obtuse preapical tubercle at side in dorsal view; tergum 5

with conspicuous lateral spine; tergum 6 with a pair of spines on each side, inner

spine longer and stouter; apical margin of tergum 6 slightly produced and

subtruncate across middle one-third; tergum 7 bilobed, emargination broader than a

semicircle, lobes narrow and subtruncate at apex. Sterna shiny, segments 3-5 slightly

raised and impunctate along midline; apical margins straight, except 6 strongly

convex medially; sterna 7 and 8 and genitalia as illustrated (Figs. 10-12).

Vestiture whitish, generally sparse and inconspicuous, longer and dense on gular

area, side of mesosoma (especially on propodeum), and on side and apical margins of

metasomal sterna.

Female .—Measurements (mm): head width 4.51-4.93; head length 3.72-4.09;

wing length 10.6-11.5; total length 15.2-17.0.

Agrees generally with description of male except as follows. Inner eye margins

strongly divergent below, lower interocular distance (at level of base of clypeus)

1.20-1.26 times minimum interocular distance. Clypeal profile convex toward base,

concave before preapical declivity and summit of declivity slightly protuberant (Fig.

3); summit of declivity marked by more or less definite median tubercle and pair of

smaller submedian tubercles (latter sometimes absent); disc of clypeus roughened

and coarsely rugosopunctate in middle, becoming finely and subcontiguously

punctate toward sides, usually without traces of median impunctate line. Ocellocular

distance about 1.2 times interocellar distance; ocelloccipital distance 2.0-2.2 times

interocellar distance. Gena, in profile, about as wide as eye. Mandible with four

approximately equidistant teeth, second and third obtuse; setae of ventral brush

sparse, none as much as one-half as long as mandible length. Flagellar segments

about as broad as long.

Metasomal terga 4 and 5 without lateral spines or teeth; tergum 6 with short,

obtuse spine on each side of broadly convex apical margin; terga 5 and 6 without

short, stout, brown setae; sterna densely punctate and with long, yellowish white

scopal hairs.

Color about as described for male, but clypeus with transverse yellow band across

basal one-half or less, paraocular area black, metasomal tergum 2 with lateral mark

only, tergum 5 wholly black, and sterna without lateral spots.

Type material .—Holotype male: Chamela, Jalisco, MEXICO, 10 June 1983 (S. H.

Bullock, 1953), in LA CM. Paratypes, all from same locality: 18 (3 <3,14 9 ?, various

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PAN-PACIFIC ENTOMOLOGIST

dates from 9 June to 3 December, various years (S. H. Bullock, C. D. Michener,

F. D. Parker, and T. L. Griswold); paratypes in CHAM, LACM, UKAN, and

UTSU.

Etymology. —The specific name is derived from that of the former Aztec empire,

Tenochtitlan.

Discussion. —In addition to the type material cited above, I have seen 3 d c?, 4 $ $

from 17 km W Tehuantepec, Oaxaca, MEXICO, 8 September 1965 (D. H. Janzen;

UKAN), on Caesalpinia sclerocarpa. The only additional floral data are on Chamela

specimens of both sexes collected by Michener at flowers of Longchocarpus sp.

The specimens examined are generally uniform in their morphological features.

One male is unusually small, total length 11 mm, with a head width of 3.72 mm.

Variation among the males otherwise consists of minor fluctuations in the extent of

the yellow markings; many specimens lack tegular spots and lateral stripes on the

mesoscutum, and most males are without a median preapical band on metasomal

tergum 2. A few specimens possess small yellow maculations on the axillae.

The available females exhibit even less variety. The anterior and lateral

mesoscutal maculations are more commonly united than in the males. The axilla is

immaculate in all specimens and metasomal tergum 2 is consistently without a

median preapical band. In many females the submedian tubercles at the summit of

the clypeal declivity are reduced or absent; the median tubercle is consistently

present.

This species is morphologically most similar to A. xochipillium, with which the

female shares the emarginate labrum, clypeal shape, and quadridentate mandible.

Males of these two species are similar in that both possess a lateral spine or tubercle

on tergum 4 and the emargination of tergum 7 is wider than deep. These features

separate both from A. cuauhtemocum.

The most conspicuous difference between A. tenochtitlanicum and A. xochipillium

is color. The former is conspicuously black and yellow and the latter is red with

obscure yellowish marks. The genitalia and associated sterna of A. tenochtitlanicum

(Figs. 10-12) and A. xochipillium (Figs. 13-15) are distinctive for each species, but

more similar to each other than either is the A. cuauhtemocum (Figs. 7-9).

Aztecanthidium xochipillium Michener and Ordway

Figures 5-6,13-15

Aztecanthidium xochipillium Michener and Ordway, 1964: 72-73; $ $.

The type locality of A. xochipillium is 17 mi N Chilpancingo, 2550 feet elevation,

Guerrero; the holotype and allotype were collected by Ordway and Roberts on 13

August 1962. The primary type and most paratypes are in UKAN. In addition to a

paratype pair, and the specimens from Ahuacatlan, Nayarit, recorded by Michener

and Ordway, I have seen the following specimens.

MEXICO, Jalisco: 1 $, 3 SS, Guadalajara, no date (Crawford; LACM).

Michoacan: 1 9 , Cojumatlan, 9 August 1964 (R. E. Ryckman, D. Spencer, C. P.

Christianson: LACM). Nayarit: 2 d d, 8 mi NW Ixtlan del Rio, 10 September 1970

(E. M. Fisher; LACM).

Like A. cuauhtemocum, both sexes are largely reddish bees with obscure yellowish

markings on the head and body. In both sexes the scutellum is distinctly bilobed and

the lobes extend over the metanotum. Males of A. xochipillium hstvQ a distinct

preapical spine or tooth laterad on metasomal tergum 4. The female clypeus is only

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171

moderately protuberant, lacks a biarcuate carina, and the apex of the labrum is

emarginate. In these characters the female is similar to that of A. tenochtitlanicum,

but the clypeal profile is evenly convex and is without a definite preapical tubercle

(Fig. 15).

Acknowledgments

For the opportunity to examine material in their care I am indebted to S. H.

Bullock (CHAM), T. L. Griswold (UTSU), and C. D. Michener (UKAN).

Literature Cited

Michener, C. D. 1942. Taxonomic observations on bees with descriptions of new genera and species

(Hymenoptera; Apoidea). J. New York Entomol. Soc., 50:273-282.

-. 1948. The generic classification of the anthidiine bees (Hymenoptera, Megachilidae). Amer.

Mus. Novit. 1381,29 pp.

Michener, C. D., and E. Ordway, 1964. Some anthidiine bees from Mexico (Hymenoptera:

Megachilidae). J. New York Entomol. Soc., 72:70-78.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 172-177

Host Records and Nest Entry by Dolichostelis, a Kleptoparasitic

Anthidiine Bee (Hymenoptera: Megachilidae)

Frank D. Parker, James H. Cane, Gordon W. Frankie and S. B. Vinson

(FDP) USD A, Agricultural Research Service, Bee Biology and Systematics

Laboratory, Logan, Utah; (JHC) Department of Entomology, Auburn University,

Auburn, Alabama; (GWF) University of California, Berkeley, California; (SBV)

Texas A&M University, College Station, Texas.

Abstract. —Host-parasite associations were recorded for three species of

Dolichostelis, a kleptoparasitic anthidiine genus. D. louisae (Cockerell) was reared

from nests of the megachild bees, Chalicodoma angelarum (Cockerell) and C.

campanulae (Robertson). D. costaricensis Friese was reared from cells made by C.

otonita Cresson. D. rudheckiarum (Cockerell) was observed parasitizing nests of C.

subexelis (Cockerell). These host records are the first confirmations for any of these

parasites. Cocoons of Dolichostelis are described and illustrated, sex ratios are

calculated, and observations on nest entry by D. rudbeckiarum are described and

illustrated.

The biologies and taxonomic relations of the parasitic bees in the tribe Anthidiini

are relatively unknown. Many undescribed species exist (Parker, unpublished data),

and only a few parasites have been associated with their host bees. Anthidine bees

are found in all zoogeographic regions and some genera are globally distributed; four

genera (Stelis, Odontostelis, Dolichostelis, and Heterostelis) occur in North and

Central America (Hurd et al., 1979; Parker and Bohart, 1979). Hosts of the parasitic

American genera are megachilids such as Osmia, Proteriades, Hoplitis,

Ashmeadiella, Anthidium, and Megachile (Hurd et al., 1979, unpublished records).

Odontostelis parasitizes Euglossa, a Neotropical apid (Bennett, 1966).

Dolichostelis is a newly proposed genus from North and Central America.

Previously, no host associations were confirmed for any of the six species included

(Parker and Bohart, 1979) although Krombein (1967) reared one (D. louisae

(Cockerell)) from a nest of an unidentified resin-using bee. In this paper, host

associations for Dolichostelis resulted from three separate studies. First, wooden

block traps were placed at several locations near Auburn, Alabama, for a

collaborative study of trap-nesting aculeates in cooperation with J. Cane. The design

of the traps and methods of rearing the specimens resembled those described by

Parker (1985). In the Alabama study, the traps were opened and their contents

individually isolated and reared. Adults were weighed after emergence, killed,

mounted, and identified. The Costa Rican study was done in cooperation with G.

Frankie and S. Vinson, who placed traps for us in the field, employing the same

design used in Alabama. In addition, Frankie and Vinson deployed many units of

individual stick traps (pine) that had been taped together into bundles bearing

several holes sizes/unit (borings of approximately 4.5, 6, 7.5, 10, and 11 mm in

diameter). Nests from the stick traps were not opened initially, but they were

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individually isolated and all emerging insects were killed and labeled. Adult weights

and placement of cells within the nest were not recorded. Observations on D.

rudbeckiarum were recorded by the senior author from nesting materials placed in

his yard in Logan, Utah.

Dolichostelis louisae (Cockerell)

Nests of four species of Chalicodoma (Chelostomoides) were recovered from traps

placed near Auburn, Alabama, during 1985. Twelve specimens of D. louisae

(Cockerell) emerged from nests of two species of Chalicodoma. An eight-celled nest

of C. angelarum (Cockerell) had six cocoons of D. louisae in the outer-most cells. In

one of six recovered nests of C. campanulae (Robertson), a single cell was parasitized

by this same bee. Three additional nests of Chalicodoma were parasitized by D.

louisae, but since no hosts emerged, specific associations could not be confirmed.

The single nest obtained by Krombein (1976) also contained only Dolichostelis, but

he believed, correctly, that the host bee was a species of Chalicodoma.

Cocoons of D. louisae bear a nipple dorsally, a feature that characterizes

Anthidiini cocoons. Cocoons of D. louisae differed from typical Stelis cocoons (Fig.

1) by their barrel-shape and lesser amount of silk. The short fecal pellets formed by

D. louisae larvae differed from the typical ribbon-like strands made by many Stelis

larvae. Fecal pellets of D. louisae were woven into the outermost layer of the cocoon.

Such pellets are not incorporated in cocoon formation by Stelis larvae. D. louisae

cocoons were made from three layers of coarse, white silk strands. Inside the first

layer, which bore an anterior nipple, a second layer had a conical and hollow nipple;

the second layer was made from an amber-colored substance. The third layer was

similar in texture and color to the first layer and it covered all the inner surfaces

except beneath the nipple. Cocoons averaged 8 mm long and 5 mm wide.

The observed sex ratio was 1.4 females to 1 male and the calculated sex ratio

(Torchio and Tepedino, 1980) was 1.06 males to 1 female. Females were only slightly

heavier (22.5 mg, SD 7.1 mm, range 14.5-32.7 mg, n = 7) than males (21.2 mg, SD

4.2 mg, range 15.2-26.4 mg, n = 5). Average adult weights of the parasites and their

two hosts were compared; average weight of the parasites was 60.7% of C.

angelarum and 73.3% of C. campanulae. All pollen and nectar provisions were

consumed in parasitized cells. Thus, differences in weights between host and parasite

were attributed to differences in relative proportions of fecal and silken materials

produced by their respective larvae. Similar differences in allotments of resources

have been recorded for a related parasitic bee, Stelis depressa Timberlake, and such

behavior may be important in survival of these parasites (Parker, 1984).

Dolichostelis costaricensis Friese

Sixteen nests of Chalicodoma otonita Cresson were obtained from the stick traps

placed at Lomas Barbudal Biological Reserve, Guanacaste, in Costa Rica. Stick

traps were placed in shaded forest locations during the extended season, from

December to May. A male of C. otonita and a male of D. coastaricensis emerged

from one of these isolated nests. When the nest was examined, the first cell made by

the host contained an empty cocoon of Dolichostelis. Apparently, the male emerged

from the second cell. In a three-celled nest of C. otonita, two empty cocoons of D.

costaricensis were found in the same host cell and the cell above contained an empty

parasite cocoon. Another two-celled Chalicodoma nest produced two Dolichostelis

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PAN-PACIFIC ENTOMOLOGIST

adults. Five more Dolichostelis emerged from traps with no host emergence, but

these nests were probably made by C. otonita. One dead female of this parasite was

found in a wooden block trap, but no cells of C. otonita were successfully parasitized.

The observed sex ratio was 1:1; since adults were not weighed, calculated sex

ratios could not be estimated. Cocoons of D. costaricensis were similar in formation,

size, and color to those of D. louisae.

Dolichostelis rudheckiarum (Cockerell)

For the past several years, a population of Chalicodoma subexilis (Cockerell) has

nested in several sizes (4, 6, 8 mm in diameter) of borings in pine wood placed on

window sills and in the garage of the senior author’s home. In 1986, D. Broemling, a

graduate student at Utah State University, observed a female of D. rudheckiarum

chewing at an entrance plug of a C. subexilis nest; it was captured and identified.

During the next several weeks in August, other females were noted and the following

observations recorded.

Often during the day, females of Dolichostelis were seen as they inspected nests of

Chalicodoma. These small bees were unusually rapid fliers for bees, and darted

among the layers of stacked borings. They darted rapidly back and forth horizontally

before the faces of the borings from 5-10 cm. The females landed only to inspect

resin nests of Chalicodoma. They never inspected active nests of other aculeates that

used the same sites such as Megachile, Osmia, Eumegachile, Euodynerus, Isodontia,

and Trypoxylon, none of which use resin in nest construction. During nest

inspection, Dolichostelis females either entered opened nests or briefly examined the

entrance plug.

At 8:00 p.M. (MDT, 8 August 1986), a female was observed chewing on a resin

entrance plug, and F. Parker recorded and photographed its activities. The parasite

worked at the entrance plug for several hours, removing tiny pieces of resin which it

then stuck on the wood surrounding the boring. As the parasite removed small

pieces, it worked most of the resin into an extended lip (Fig. 2). After about two

hours, the parasite bent the lip down with the weight of its body while chewing at the

top of the plug. During the entire process, the parasite deposited glistening droplets

from the tip of its abdomen onto the surface of the resin plug. After each deposition,

it then turned around and chewed the area where the droplet was deposited.

Apparently, this substance, acting as a solvent, enabled the parasite to soften and

mold the resin. Also, this liquid may aid in preventing resin from sticking to the

mouthparts, since the parasite frequently groomed and cleaned its head. After

sunset, a small lamp was placed near the glass to illuminate the nest surface. The

parasite seemed undisturbed by the light since it continued to work. After the

Fig. 1. Portion of a nest of Chalicodoma angelarum (Cockerell) made in a trap block and with two cells

containing cocoons of Dolichostelis louisae (Cockerell). Fig. 2. Female of Dolichostelis rudheckiarum

(Cockerell) opening sealed entrance of a nest of Chalicodoma subexilis. About 9:00 p.m. (MDT). Fig. 3.

Same nest at 8:00 a.m. the following day and with the female parasitic bee resting just inside the entrance.

Fig. 4. Same nest being closed by the parasite. Fig. 5. Same nest after parasite had finished closing the

entrance. Note the small pellets of resin adjacent to the plugged opening; such pellets indicate nest parasit-

ization by Dolichostelis. Fig. 6. Cocoon of Dolichostelis rudheckiarum. Fig. 7. Female of Chalicodoma

subexilis inspecting a previously capped nest. Fig. 8. Female of Chalicodoma subexilis depositing drop¬

lets from its abdomen around rim of nest plug.

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PAN-PACIFIC ENTOMOLOGIST

parasite finally gained entrance to the nest, it proceeded to remove more resin from

within the nest. Later, it removed what appeared to be a small amount of host

provision. The female continued its back-and-forth entry into the boring until after

12:00 A.M., when observations were discontinued.

The next morning, at 8:00 a.m. , the nest was checked and the female was resting

just inside the entrance (Fig. 3). As soon as sunlight struck the nest, about 9:00 a.m. ,

the parasite resumed its activities. Soon, it commenced refilling the entrance with the

small pieces of resin it had previously stuck adjacent to the opening (Fig. 4). By

hanging vertically from above the hole, the parasite grasped the resin lip protruding

from the hole and pulled it up and into place as if it had been hinged at the base.

Then, the nest plug was smoothed across with more resin (Fig. 5). The parasite

finished working at 9:55 a.m. and flew away. The nest was left on the ledge to be

opened in the laboratory on Monday, but before removal, the original owner

returned to the nest, opened the entrance, and began removing pollen and nectar.

Sometime later, it again plugged the nest, but this time using masticated leaf pulp

mixed with resin for the final closure. Upon opening in the laboratory, the nest

contained a single cell with a large egg lying across the pool of nectar and pollen; this

egg hatched into a Chalicodoma larva. Twenty other Chalicodoma nests were

opened, including several with evidence of parasite entry, but only one contained

parasite cocoons.

Cocoons of D. rudbeckiarum (Fig. 6) resembled Stelis cocoons; both had a small

nipple. The amber-colored inner layer of D. rudbeckiarum cocoons was less dense

and the overwintering prepupal larvae were visible. The barrel-shaped cocoons

averaged 8 mm long and 5 mm wide.

During the course of these observations, many females of Chalicodoma were seen

landing at nests they had previously capped. The females would inspect the cap (Fig.

7), turn around, and deposit droplets of liquid from the tip of the abdomen onto the

inner rim of the nest cap (Fig. 8). Females of Dolichostelis were not observed

inspecting such nests.

These limited observations suggest that unique behavioral traits may exist in these

parasites and their host. Such behavior has not been reported previously although

Bennett (1966) observed that a related parasite, Odontostelis, that invaded nests of

Euglossa, drove the nest owner away, opened cells of its host, removed and killed

eggs or early larval instars of Euglossa, deposited its own egg on the provision, and

resealed the cells. It appears that Chalicodoma females can detect parasitized nests

and neutralize parasitized cells. It was not uncommon to observe females of the host

bee examining, opening, and removing pollen and nectar (along with the parasite

egg?) from a previously finished nest and then recapping it. One such female

remained in the entrance of a nest for two days before the nest was finally recapped.

Nest usurping among host females probably does not explain this kind of nesting

behavior (since females were not marked) because there were few nesting females

and a surplus of available nesting sites.

During the past 15 years of collecting and observing bee nests in the vicinity of

Logan, nests of C. subexilis have been commonly found in units provided for the

alfalfa leafcutting bee. Megachile rotundata (F.). Not a single nest contained a cell

parasitized by the Dolichostelis, nor have any specimens of this parasite been

observed or net-collected this far north. It appears that this parasite may have

extended its range and/or its available hosts recently.

VOLUME 63, NUMBER 2

177

Acknowledgments

We are dedicating this manuscript to E. Gorton Linsley, who has contributed

many papers on the biology and taxonomy of bees, including the discovery of several

new species of Stelis.

We would like to thank D. Veirs, R. Griswold, and V. Rhea (USDA Bee Lab,

Logan, Utah) for making the trap blocks, recording the biological information, and

curating the specimens.

Partial support for this research came from Hatch Project ALA 00648 of the

Alabama Agricultural Experiment Station, Auburn.

Contribution from Utah Agricultural Experiment Station, Utah State University,

Logan, Utah 84322-4810, Journal Paper No. 3326, and USD A-Agricultural

Research Service-Bee Biology and Systematics Laboratory, Utah State University,

Logan, Utah 84322-5310.

Literature Cited

Bennett, F. D. 1966. Notes on the biology of Stelis (Odontostelis) bilineolata (Spinola), a parasite of

Euglossa cordata (Linnaeus). J. N.Y. Entomol. Soc., 74:72-79.

Hurd, P. D., Jr. 1979. Superfamily Apoidea. In: K. V. Krombein, P. D. Hurd, Jr., D. R. Smith, and

B. D. Burks (eds.). Catalog of Hymenoptera in America North of Mexico, vol. 2, Apocrita, pp.

1741-2209. Washington, D.C.: Smithsonian Institution Press.

Krombein, K. V. 1967. Trap-nesting wasps and bees: life histories, nests, and associates. Smithsonian

Publ. 4670,570 pp.

Parker, F. D. 1984. The nesting biology of Osmia (Trichinosmia) latisulcata Michener. J. Kansas

Entomol. Soc., 57:430-436.

Parker, F. D. 1985. Nesting habits of Osmia grinnelli Cockerell. Pan-Pac. Entomol., 61:155-159.

Parker, F. D. and G. E. Bohart. 1979. Dolichostelis, a new genus of parasitic bees. J. Kansas Entomol.

Soc., 52:138-153.

Torchio, P. F. and V. J. Tepedino 1980 Sex ratio, body size, and seasonality in a sohtary bee, Osmia

lignariapropinqua Cresson. Evolution, 34:993-1003.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 17&-193

A Revision of the Dufoureine Genus Micralictoides Timberlake

(Hymenoptera: Halictidae)

George E. Bohart and Terry L. Griswold

USD A, ARS, Bee Biology and Systematics Laboratory, Utah State University,

Logan, Utah 84322-5310.

Abstract. —The southwestern North American genus, Micralictoides, is reviewed,

a key to species presented, five new species described (M. chaenactidis, M. dinoceps,

M. grossus, M. linsleyi, and M. quadriceps), and new records provided for the three

previously included species.

Timberlake (1939) established Micralictoides as a subgenus of Dufourea for two

species, D. altadenae (Michener) and D. ruficaudus (Michener), described under

Halictoides (Michener 1937). Bohart (1942) later gave Micralictoides generic status

and described a third species, M. mojavensis. Michener (1965), in a generic review of

the New World Dufoureinae, briefly characterized Micralictoides and provided a key

separating it from the other six New World genera.

Genus Micralictoides Timberlake

Micralictoides Timberlake 1939: 397 (as subgenus). Type: Halictoides ruficaudus

Michener.

Generic Diagnosis. —Size small, body length 3.5 to 6 mm; abdomen of male broad

as in female; punctures of head and mesonotum shallow; tergal punctures fine and

shallow; body without metallic reflections; legs with tarsi (and usually tibiae) paler

than femora; clypeal margin of female truncate or nearly so between tubercles;

labrum very short, nearly truncate; maxillary palpus of female nearly as long as

flagellum; flagellum of male with segments not or scarcely longer than broad, without

modified hairs; male without modified legs; forewing with two submarginal cells, first

submarginal twice as long as wide, marginal cell from apex of stigma to apex of cell no

longer than distance from apex of cell to wing tip; propodeum with weak, irregular

striae becoming obsolescent laterally and along posterior margin; sternum VI of

male without distinct grooves, carinae, or strong protuberances, but sometimes

gently swollen toward apex; sternum VII with pair of slender, depressed, apical

lobes, each with terminal, incurved appendage; sternum VIII medially with broad

articulating lobe, apically with slender median lobe about one-third total segment

length; genital capsule swollen basally, with simple, strap-like gonostylus, distinct

from gonobase, with short, strongly incurved sagitta, and knob-like, polished

volsella.

Micralictoides can be distinguished from other dufoureine genera by a

combination of the two submarginal cells in the forewing and the hind basitarsus

which is distinctly paler than the hind femur. The configuration of metasomal sterna

VII and VIII in the male is unique in the subfamily. Superficially, Micralictoides

resembles some of the smaller species of Dufourea such as the D. leachi Timberlake

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VOLUME 63, NUMBER 2

group. Both have relatively simple antennae, legs, and visible sterna in the male, and

both have a swollen basal portion of the genital capsule and a shortened volsella.

Systematics. —The eight known species of Micralictoides can be separated into

three groups based on the structure of the metasomal sterna (especially VII) and the

genitalia of the males. The altadenae group includes four species: M. altadenae

(Michener) with a broad face and short mouthparts, M. chaenactidis n. sp. and M.

linsleyi n. sp. with face and mouthparts moderate in length, and M. quadriceps n. sp.

with a long face and long mouthparts. The grossus group, which contains only M.

grossus n. sp., may be related to M. quadriceps of the altadenae group on the basis of

a somewhat similar development of sternum VI. The ruficaudus group includes M.

ruficaudus (Michener), M. mojavensis Bohart, and M. dinoceps n. sp. The

relationship between these three species is most apparent in the configuration of

male sternum VII. This group probably developed from the altadenae group; in fact,

the female of M. mojavensis is difficult to distinguish from that of M. chaenactidis.

Distribution.—Micralictoides is restricted to the southwestern United States where

it is known only from California, Nevada, and Arizona. All eight species are found in

California and only one of them, M. chaenactidis, ranges much beyond its borders

(into north-central Nevada and west-central Arizona). In California, the various

species are distributed as follows: M. chaenactidis in the central and southern coastal

ranges and the southern deserts, M. mojavensis in the Mojave Desert and Los

Angeles Basin, M. dinoceps in the San Bernardino Mountains, M. ruficaudus and M.

altadenae from the central and southern coastal ranges, M. quadriceps and M.

grossus in the central Sierra Nevada foothills, and M. linsleyi on the eastern side of

the northern Sierras.

Biology. —The nesting habits of Micralictoides are unknown. Collection records

indicate that members of the genus are remarkably oligolectic. Apparent pollen

sources for Micralictoides are: Allium (M. dinoceps), Chaenactis (M. chaenactidis),

Eriophyllum (M. altadenae), Gilia (M. grossus), Eschscholtzia (M. ruficaudus), and

Navarretia (M. quadriceps). Pollen preference in M. mojavensis is unclear. Floral

records for females include Gilia, Salvia, Eschscholtzia, Phacelia, Layia, Baeria, and

Malacothrix. There are no floral associations for M. linsleyi.

Since species of Micralictoides are strikingly oligolectic, we deemed it appropriate

to dedicate this paper to Dr. E. G. Linsley, whose important contributions to bee

systematics and biology include special studies on problems of oligolecty, and to

name one of the included new species after him.

Key to Males of Micralictoides

1. Head distinctly longer than wide. quadriceps n. sp.

Head at least slightly wider than long .2

2. Abdomen largely red or reddish-brown . ruficaudus (Michener)

Abdomen dark brown to black, segments sometimes paler apically.3

3. Sternum VI in profile with low but distinct bulge near middle, surface nearly

obscured by moderately long, dense pubescence (Fig. 18), lateral arm of

sternum VII short, stout basally, hammer-shaped apically (Fig. 16) .. grossus

n. sp.

Sternum VI nearly straight in profile, with short, sparse pubescence not at all

obscuring surface (Fig. 19), lateral arm of sternum VII long, slender basally,

clubbed apically (Figs. 10-12,14,15) .4

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PAN-PACIFIC ENTOMOLOGIST

4. Sternum VIII with lateral arm (preceding apical “foot”) with abrupt expansion

near middle (Figs. 14,15).5

Sternum VII with lateral arm not distinctly expanded preceding apical “foot”

(Figs. 10,11,12) .6

5. Head width at most 1.2 times length; lateral arm of sternum VII beyond expansion

slender then widened apically (Fig. 15) . mojavensis Bohart

Head width 1.3 times length; lateral arm of sternum VII beyond expansion

uniformly wide (Fig. 14) . dinoceps n. sp.

6. Sternum VIII with median apical projection short, slender, tapering toward apex

throughout (Fig. 2); sternum VII with basal flaps nearly touching broadly

triangular apical “foot” (Fig. 10) . linsleyi n. sp.

Sternum VIII with median apical projection long, parallel sided or slightly

narrowed sub-basally (Figs. 3,4); sternum VII with basal flaps remote from

oval apical “foot” (Figs. 11,12) .7

7. Prementum shorter than eye; propodeal enclosure with well-defined but slightly

irregular striae even when viewed from directly above or from slightly in front

altadenae (Michener)

Prementum longer than eye; propodeal enclosure appearing rugose or with striae

highly irregular and difficult to trace separately, even when viewed from

directly above or from shghtly in front . chaenactidis n. sp.

Key to Females of Micralictoides

1. Head distinctly longer than wide (Fig. 38). quadriceps n. sp.

Head at least slightly broader than long (Figs. 33-37, 39, 40) .2

2. Tergum II with punctures of mid-line area mostly less than one puncture width

apart (except on apical impunctate border), if punctures fine and indistinct,

interpunctural areas shagreened .3

Tergum II with punctures of mid-line area mostly more than one puncture width

apart (except sometimes on basal half) and with interpunctural areas shiny . 5

3. Head more then nine tenths as long as wide (Fig. 40); distance between antennal

scrobes less than median clypeal length; anterior wing (measured from apex of

tegula) at least 3.5 mm long . grossus n. sp.

Head less than nine tenths as long as wide; distance between antennal scrobes

greater than median clypeal width; anterior wing not more than 3.4 mm long 4

4. Prementum and stipes shorter than eye; head not more than 0.82 times as long as

wide (Fig. 39) . altadenae (Michener)

Prementum and stipes longer than eye; head at least 0.85 times as long as wide

(Fig. 35). chaenactidis n. sp.

5. Frons with surface between punctures strongly shagreened . linsleyi n. sp.

Frons with surface between punctures polished.6

6. Abdomen red; stipes and prementum shiny, not shagreened . ruficaudus

(Michener)

Abdomen black; stipes and prementum shagreened.7

7. Maxillary palpus elongate, as long or longer than stipes (Fig. 26) .. . mojavensis

Bohart

Maxillary palpus not elongate, shorter than stipes (Fig. 27). dinoceps n. sp.

VOLUME 63, NUMBER 2

181

Micralictoides chaenactidis, New Species

(Figs. 4,12, 30, 35)

Holotype male .—Length about 4.5 mm, forewing length 2.9 mm; body moderately

shining, black except mid and fore tarsi, tip of hind tibia, abdomen dark brown;

tegula, hind tarsi, sternum VI light brown; pubescence white, sparse, rather short,

not concealing integument.

Head. Head slightly broader than length from vertex to clypeal margin (9:8);

clypeus about half as long as breadth of apical truncation, its surface with rather

numerous, coarse punctures; lower margin of median ocellus lower than upper eye

margin; distance from median ocellus to antennal scrobe more than twice that

between scrobes; mandible moderately long but not slender, apical tooth less than

one-third length of mandible and slightly darker than middle third; punctures of

frons close and broad except sparser between and close to ocelli; antennal scape

nearly half as broad apically as long, with scattered large punctures; flagellomeres I

and II distinctly, III slightly broader than long, IV to X ranging from about as long as

broad to about one-fifth longer than broad; length of prementum, maxillary palpus

each greater than eye length.

Thorax. Punctures of central half of scutum averaging about one puncture width

apart, interpunctural areas not roughened; scutellum more irregularly punctate but

average distance between punctures as on scutum; propodeal enclosure strongly,

rather closely, somewhat irregularly striate throughout.

Abdomen. Impunctate margins of terga translucent, not much lighter than

remainder of segments; punctures of first ter gum about half as large, nearly as close

as those on scutellum, those on succeeding terga successively smaller but not minute;

sternum VI practically straight in profile, uniformly sparsely haired; sternum VII as

in Fig. 12; sternum VIII as in Fig. 4.

Female .—Differs from male in having the face much more polished, punctation of

frons sparse (usually separated by two or more puncture widths) and with an

additional scattering of much smaller punctures. Scutum and scutellum rather

polished, with fine punctures generally more than one puncture width apart

submedially. Abdominal terga more finely punctate, generally more polished. Hind

tibia nearly as light as hind basitarsus, dorsal scopal hairs longer than apical width of

tibia. Elevated portion of clypeus slightly rounded apically, marginal truncation

about twice as broad as long, head and mouthparts proportioned as in Figs. 30, 35.

Type Material .—Holotype male: ARIZONA, Yavapai Co., 30 miles (48.3 km)

NW Wickenburg 16-IV-1965, Chaenactis, G. E. Bohart, P. F. Torchio, N. Youssef.

Paratypes: Thirty-nine females and four males, same data as holotype; Pima County:

3 females, Marana, 26-IV-1973, Chaenactis douglasii, G. E. Bohart; 3 males, 9

females. Silver Bell Bajada, J. L. Neff; Pinal County: 1 male. Mammoth, 29-III-68,

Phacelia, Torchio & Youssef. Holotype deposited at U.S. National Museum,

paratypes at BBSL and LACM.

Additional Material. —CALIFORNIA. Los Angeles County: 1 male, Aliso

Canyon, 3-V-1934, C. D. Michener; 1 female. Little Rock, Mojave Desert,

20-V-1937, E. P. VanDuzee; 1 female, Lovejoy Butte, ll-V-1944,

Acamptopappus, P. H. Timberlake; 1 female, Mint Canyon, 3-V-1936, E. G.

Linsley; 2 males, Newhall, 20-IV-1940, R. M. Bohart; 1 female, Palmdale, 2 miles

(3.2 km) E, 17-V-1937, Astragalus, E. P. VanDuzee; 1 male, Piute Butte, Mojave

Desert, 12-V-1944, A. L. Melander; 6 males, 15 females, Piute Butte, S of.

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183

12-V-1944, Chaenactis brachypappa, P. H. Timberlake; 1 male, 1 female, Tapia

Park, 18-IV-1953; 3 males. Three Points, 2-V-1968, D. Veirs, Riverside County; 8

males, 3 females, Joshua Tree Natl. Mon., 20-V-1970, Chaenactis, P. F. & D. M.

Torchio. San Benito County: 1 male, 1 female, Pinnacles Natl. Mon. vicinity,

5-V-1979, Chaenactis glabriuscula, A. R. Moldenke; 1 female. Pinnacles, 3 miles

(4.8 km) NE, 27-IV-1962, J. K. Drew; 1 female, same except Chaenactis

glabriuscula, R. W. Thorp. San Bernardino County: 2 males, Morongo Valley,

21-IV-1957, R. R. Snelling & M. D. Stage; 3 females, same except Malacothrix; 1

male, Wildwood Cyn., 3 miles (4.8 km) E Yucaipa, 29-V-1977, T. Griswold. San

Luis Obispo County: 1 male. Black Mt., 6 air miles (9.7 km) NE Pozo,

3300'-3600' (1006m-1097m), l-V-1962, Eriophyllum confertiflorum, C. A.

Toschi; 2 males. Black Mt., same except J. Powell; 2 males, 2 females, Creston, 10

miles (16.1 km) SE, 25-IV-1968, J. Powell; 1 male, Creston, 10 miles (16.1 km) SE,

25-IV-1968, D. Veirs; 2 males, same except Malacothrix, J. Doyen; 1 male,

Creston, 2.5 miles (4.0 km) S, 4-V-1962, C. A. Toschi; 4 males. La Panza Camp, 12

miles (19.3 km) NE Pozo, 29-IV-1962, Eriophyllum confertiflorum, P. H.

Timberlake; 2 males, same except P. D. Hurd; 27 females, Santa Margarita, 6 miles

(9.7 km) NE, 5-V-1962, Chaenactis glabriuscula, P. H. Timberlake; 5 males, 1

female, Shandon, 7 miles (11.3 km) SW, 24—IV-1960, J. W. MacSwain. Ventura

County: 1 male, Chuchupate Ranger Station, base Frazier Mt., lO-V-1959, Baeria

chrysostoma, P. H. Timberlake; 1 male, same except 8-V-1959, P. D. Hurd; 1

female. Hungry Valley, 5 miles (8.0 km) S Gorman, 4-V-1959, Chaenactis

glabriuscula, J. Powell; 1 male, same except 6-V-1959, J. R. Powers; 26 females,

same except P. H. Timberlake; 8 males, 29 females, same except 9-V-1959, P. H.

Timberlake; 20 males, 51 females, Quatal Canyon, NW corner Ventura Co.,

9-V-1959, Chaenactis glabriuscula, P. H. Timberlake; 1 male, same except P. D.

Hurd; 1 male, same except Eriophyllum confertiflorum, C. W. O’Brien; 1 female,

same except J. Powell; 1 female, same except no floral data, J. R. Powers; 1 female,

same except G, 1. Stage. NEVADA. Lander County: 3 males, 2 females. Battle

Mountain, 12-VI-1962, Chaenactis, G. E. Bohart; 1 female, same except on

Cruciferae. Specimens in the collections of BBSL, CAS, CIS, LA CM, and UCR.

Distribution. —This is the widest ranging of the Micralictoides species, ranging

from the desert regions of Nevada and Arizona across the Sonoran and southern

Mojave Deserts of California and into drier parts of the coastal ranges of California.

Discussion. — M. chaenactidis is probably closer to M. altadenae than to other

species on the basis of its almost identical male sterna and genitalia. It differs

principally in having longer mouthparts and face, and sparser punctation on the

female frons. The female of M. chaenactidis is also similar to that of M. mojavensis,

but the basal zone of the propodeum is more strongly sculptured and the dorsal

scopal hairs of the hind tibia are longer than the apical width of the tibia. The male

differs from M. mojavensis most obviously in sternum VII (Fig. 12), but it also has a

more strongly punctate clypeus and more roughened frons.

Figs. 1-24. Males. Figs. 1-8: Metasomal sternum VIII. Fig. 1, M. quadriceps', Fig. 2, M. linsleyi', Fig. 3,

M. altadenae', Fig. 4, M. chaenactidis'. Fig. 5, M. ruficaudus'. Fig. 6, M. dinoceps'. Fig. 7, M. mojavensis-.

Fig. 8, M. grossus. Figs. 9-16: Metasomal sternum VII. Fig. 9, M. quadriceps'. Fig. 10, M. linsleyi'. Fig. 11,

M. altadenae'. Fig. 12, M. chaenactidis'. Fig. 13, M. ruficaudus-, Fig. 14, M. dinoceps; Fig. 15, M.

mojavensis'. Fig. 16, M. grossus. Fig. 17: Metasomal sternum VII, lateral view, M. quadriceps. Figs.

18-19: Metasomal sternum VI. Fig. 18, M. grossus-, Fig. 19, M. ruficaudus. Figs. 20-24: Genital capsule.

Fig. 20, M. quadriceps', Figs. 21, 22, M. grossus'. Figs. 23, 24, M. dinoceps. (Illustrations not drawn to

scale.)

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185

Specimens from California localities differ from the type series in having the

integument less intensely black, the frons of the female more uniformly and usually a

little more densely punctured, and the abdominal terga slightly more coarsely and

closely punctate.

Micralictoides altadenae (Michener)

(Figs. 3,11, 29, 39)

Halictoides altadenae Michener 1937: 395 (Holotype female: SMEK).

Dufourea (Micralictoides) altadenae', Timberlake 1939: 397.

Micralictoides altadenae', Bohart 1942: 123.

Male. —Length about 4.5 mm, forewing length 3 mm; very similar to M.

chaenactidis, but differs in having a wider head (8:7) and shorter mouthparts. It also

has denser pubescence on the foreparts of the face (partially obscuring the clypeus),

somewhat paler integument of the abdomen (dark brown instead of largely glossy

black), and partially brownish (instead of entirely whitish) pubescence on terga IV

and V. The mid and hind basitarsi are somewhat paler in contrast to the tibiae and the

abdominal pubescence is generally more abundant. Sterna VI, VII (Fig. 11), VIII

(Fig. 3) and the genital capsule are similar to those of M. chaenactidis.

Female. —Much like M. chaenactidis but differs in having a wider face and shorter

mouthparts (Fig. 29, 39). Punctures of the frons are generally larger and closer

together (from less than one to slightly over one puncture width apart) and those of

the scutum are denser and more irregular in size (averaging about one puncture

width apart). The integument is somewhat paler (mostly dark brown on the abdomen

instead of mostly black) and the hind tibia is almost the same color as the basitarsus

instead of distinctly darker brown.

Distribution. —Known only from two widely separated localities in cismontane

California, the type series from Altadena, Los Angeles County, California and a new

record from Yolo County. (The record from Aliso Canyon, Los Angeles County, is

an error and actually represents M. chaenactidis .)

New Records. —CALIFORNIA. Yolo County: 2 males, 4 females, 30-V-1956,

R. M. Bohart.

Discussion. —This species is very close to M. chaenactidis (for differences, see

discussion under the latter). The female resembles M. mojavensis as well, but has

shorter mouthparts (Fig. 29) and browner abdominal pubescence. The face is

proportionately broader than that of M. mojavensis (compare Fig. 39 and Fig. 34)

but the contrast is less pronounced than in comparison with M. chaenactidis. The

male is easily distinguished from M. mojavensis on the basis of sternum VII.

Figs. 25-40. Females. Mouthparts in lateral view. Fig. 25, M. linsleyi; Fig. 26, M. mojavensis; Fig. 27,

M. dinoceps; Fig. 28, M. ruficaudus; Fig. 29, M. altadenae; Fig. 30, M. chaenactidis; Fig. 31, M.

quadriceps; Fig. 32, M. grossus. Figs. 33-40: Head in frontal view. Fig. 33, M. linsleyi; Fig. 34, M.

mojavensis; Fig. 35, M. chaenactidis; Fig. 36, M. dinoceps; Fig. 37, M. ruficaudus; Fig. 38, M. quadriceps;

Fig. 39, M. altadenae; Fig. 40, M. grossus. (Illustrations not drawn to scale.)

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Micralictoides linsleyi, New Species

(Figs. 2,10, 25, 33)

Holotype male. —Length about 3.5 mm, forewing length 2.5 mm; body black

except antenna, tegula, tarsi brown; abdomen slightly reddish. Pubescence white,

sparse, rather short, not concealing integument.

Head. Head length less than width (0.80); clypeus one-half as long as width of

truncation, its surface with dense, coarse punctures; lower margin of midocellus well

below upper eye margin; distance from median ocellus to antennal scrobe

approximately twice that between antennal scrobes; mandible moderately long but

not slender, apical tooth less than one-third length of mandible, slightly darker than

middle part; punctures of frons close and broad; antennal scape one-third as broad

apically as long, with scattered large punctures; flagellomeres I-III distinctly broader

than long, IV-X ranging from as long as broad to one and one-half times as long as

broad; prementum and stipes slightly shagreened, length of prementum slightly

greater than eye length, maxillary palpus approximately equal to eye length.

Thorax. Punctures of disk of scutum and scutellum more than a puncture width

apart; interspaces shiny; propodeal enclosure with fine, close, irregular carinae not

reaching posterior margin.

Abdomen. Impunctate margins of terga translucent, not much lighter than

remainder of segments; punctures of terga as large as those on scutum and scutellum

but much sparser; sternum VI practically straight in profile, with a few long hairs

medially; sternum VII as in Fig. 10, sternum VIII as in Fig. 2.

Female. —Head proportions and mouthparts as in Figs. 25, 33; elevated preapical

margin of clypeus nearly straight; vertex in frontal view well elevated behind eye;

scutum and scutellum rather polished, punctures finer and much closer than in male;

hind tibia with dorsal scopal hairs not as long as apical width of tibia. Abdominal

punctation as in male.

Type Material. —Holotype male: NEVADA, Washoe Co., 4 miles (6.4 km) N

Sparks, 18-VI-1959, G. I. Stage. Paratypes: 7 females, same data as holotype; 1

female same except 3.4 miles (5.5 km) N of Sparks. Holotype deposited at CAS,

paratypes at BBSL, LA CM, and UCB.

Additional Material. —CALIFORNIA. Lassen County: 1 male, Litchfield, 10.6

miles (17.1 km) N, 19-VI-1959; 1 male, Ravendale, 10 miles (16.1 km) S,

5-VI-1964, R. C. Dickson; 1 male, Termo, 9-VI-1960, G. 1. Stage.

Distribution. —Eastern side of the northern Sierra Nevada Mountains.

Discussion. —^Females of M. linsleyi are similar to those of M. ruficaudus.

Differences include frons roughened between punctures, vertex elevated above

eyes, and preapical margin of clypeus less arcuate.

The males are distinct from those of M. ruficaudus in having the abdominal reddish

coloration nearly confined to the apical tergal margins. From other Micralictoides

males, they can be distinguished by the contrastingly reddish brown color of the

apical tergal margins and by the configuration of sterna VII and VIII.

Micralictoides quadriceps, New Species

(Figs. 1,9,17,20,31,38)

Holotype male. —Length about 4 mm, forewing length 2.6 mm; body dull black,

with close, broad, rather shallow punctures and short, pale, sparse pubescence.

VOLUME 63, NUMBER 2

187

Head. Face from vertex to clypeal margin subquadrate, broadened apically, head

slightly longer than broad (7:6.5); lower margin of median ocellus above upper eye

margin; distance from median ocellus to antennal scrobe about three times that

between scrobes or between lateral ocelli; mandibles unusually long, slender, with

middle one-third yellowish; punctures of frons separated by much less than one

puncture width except a little sparser near ocelli and medially on clypeus; clypeal

integument only slightly obscured by pubescence; antennal scape shining between

punctures and three times as long as broad; flagellomeres I and III to XI longer than

broad; mouthparts (including palpi) over twice as long as mesonotum, metanotum,

dorsal face of propodeum combined.

Thorax. Scutal punctures moderately strong, uniform, averaging less than one

puncture width apart; median impressed line well developed on scutum and

scutellum; scutal width (between tegulae) less than length of scutum and scutellum

combined (4:5); hind basitarsus, tip of hind femur, yellowish brown, other basitarsi,

all femoro-tibial areas brown; dorsal propodeal enclosure with moderately regular

striae laterally, weak, irregular ones medially.

Abdomen. Transparent marginal zones of terga conspicuously lighter than

remainder; discs of terga with fine, rather shallow punctures from one to two or more

puncture widths apart; sternum VI with shallow median impression and rather

abrupt bend in dorsal direction at apical one-fifth, the sternum not obscured in

ventral view by rather abundant preapical zone of pubescence; sternum VII as in

Figs. 9,17; sternum VIII as in Fig. 1.

Female. —Similar to male, but head longer, more parallel sided (Fig. 38), labrum

longer, rounded apically; mouthparts as in Fig. 31; clypeus with seven or eight large,

well separated punctures on flattened portion; hind basitarsus, tibia, tip of femur

orange-yellow; discs of terga more closely punctured and minutely roughened.

Type Material. —Holotype male: CALIFORNIA, Amador County, Daffodil Hill,

3-VI-1963, R. M. Bohart. Paratypes: 48 males, 3 females, same as holotype; 34

males, 1 female, same except F. D. Parker; 55 males, 2 females, same except

M. E. Irwin; 8 males, 4 females. Volcano, Amador County, California, 4-VI-1961,

Navarretia, R. M. Bohart. Holotype deposited at UCD; paratypes at BBSL and

UCD.

Additional Material. —1 female, Dunlap, Fresno County, California, 25-VI-1963,

R. R. Snelling; 1 female. Bear Valley, Placer County, California, 4-VII-1956, R. R.

Snelling. Specimens at LA CM.

Distribution. —Known only from the western foothills of the Sierra Nevada

Mountains.

Discussion. —Variation is slight in this species. Some specimens are smaller than

those described above and some of the males have the face nearly as parallel-sided as

the females.

The elongated, nearly parallel-sided face is unique in the genus. The male genitalia

and sterna closely resemble those of M. chaenactidis and M. altadenae. On the other

hand, the reflexed tip of sternum VI bears some resemblance to that of M. grossus.

Micralictoides grossus, New Species

(Figs. 8,16,18,21,22,32,40)

Holotype male. —Length about 6 mm, forewing length 4 mm; body dull black,

closely punctured on head but only moderately so elsewhere, the punctures deeper

188

PAN-PACIFIC ENTOMOLOGIST

than usual for the genus; pubescence pale brownish, moderately profuse, partially

obscuring lower parts of face, side of mesepisternum, tip of abdomen.

Head. Head slightly broader than long (9:8.5); lower margin of median ocellus

shghtly below upper eye margin; distance form median ocellus to antennal scrobe less

than three times that between scrobes (6:2.5) and about twice that between lateral

ocelli; punctures of face, including clypeus, less than one puncture width apart

except close to ocelli, in subantennal area; scape dull, rough, less than twice as long

as broad; flagellomeres mostly broader than long; mandible moderately long, black

basally, reddish apically, the apical tooth less than one-third as long as mandible;

tongue with total length slightly less than twice that of mesonotum, metanotum,

dorsal enclosure of propodeum combined (17:9).

Thorax. Punctures of scutum averaging about one puncture width apart, not

unusually broad or shallow, those of scutellum sparser sublaterally; median,

impressed line of scutum visible on scutellum only as dense row of punctures; width

of scutum between tegulae as great as length of scutum and scutellum combined;

basitarsi, apical portion of posterior tibia yellowish; middle two-thirds of posterior

tibia as dark as femur; dorsal propodeal enclosure appearing finely granular with

superimposed, weak, irregular striae.

Abdomen. Dorsally longer than broad (14:10); apical impunctate margins of terga

transparent, rather inconspicuous; discs of terga with moderately fine punctures,

mostly ranging from one to two puncture widths apart; sternum VI conspicuously

bulging in profile, the bulge densely covered with reddish-brown pubescence (Fig.

18); sternum VII as in Fig. 16; sternum VIII as in Fig. 8.

Female. —^As in male but larger (length 6 mm) with pubescence shorter, generally

sparser, punctation closer, stronger on scutum, clypeus with only a few large

punctures on anteromedian flattened area, propodeum more finely seulptured,

tergal discs less polished, a little more closely punctate, hind tibia about same color of

brown as hind basitarsus, somewhat paler than hind femur. Head proportions as in

Fig. 40; mouthparts as in Fig. 32.

Type Material. —Holotype male: CALIFORNIA, Tuolumne Co., Tuolumne

City, 3-VI-1953, Gilia capitata, B. L. Rozen. Paratypes: CALIFORNIA. Tuolumne

County: 1 female, same data as holotype; 2 males, 18 females, same except J. G.

Rozen; 1 female, same except 22-VI-1953, J. G. Rozen; 3 males, same except

27-V-1953; 2 males, same except 29-V-1953; 1 male. Buck Meadows-Mather site,

#50982, A. R. Moldenke; 1 female, Early Intake Power House, 3 miles (4.8 km) S,

Stanislaus Natl. For., 1-VI-1977, #119159, P. Lincoln; 1 female, same except

2-VI-1977, Gilia capitata, P. Lincoln; 1 female, same except 4-VI-1977. Madera

County: 1 female. North Fork, lO-V-1936, F. T. Scott. Sierra County: 1 male, Yuba

River, 9 miles (14.5 km) W Goodyears Bar, 29-V-1965, C. D. MacNeill. Holotype

deposited at CAS, paratypes at UCB, LA CM, and BBSL.

Distribution. —Known only from the western foothills of the Sierra Nevada

Mountains.

Discussion. —Size is somewhat variable in this species, the smallest male being

about 4.5 mm and the smallest female 5 mm in length. Some of the males have the

propodeal striae completely irregular.

This species is easily distinguished by its large size, strong, deep punctures, and

dull first tergum. The male is very distinctive in sterna VI, VII and VIII and the

genital capsule as illustrated. It is not closely related to any other species, although

VOLUME 63, NUMBER 2

189

the male shows a possible relationship to M. quadriceps in the bulging and pubescent

profile of sternum VI.

Micralictoides ruficaudus (Michener)

(Figs. 5,13,19,28,37)

Halictoides ruficaudus Michener 1937; 397 (Holotype female: SMEK).

Dufourea (Micralictoides) ruficauda; Timberlake 1939: 397.

Micralictoides ruficaudus; Bohart 1942: 121.

Male. —Length about 4.5 mm, forewing length 3 mm; body black except

metasomal terga I-III, and frequently IV, mostly orange (dark areas basally on I and

laterally on II) and legs varying from dark brown on coxae and femora to lighter

brown (but only slightly) on tarsi. Punctures of head and thorax mostly closer than

one puncture width apart except about one puncture width apart on middle of

scutum and sides of mesepisternum and sparse and fine subgenally; pubescence

off-white, rather sparse, only slightly obscuring lower parts of face, sides of

mesepisternum; that of abdominal apex yellowish.

Head. Head slightly broader than long (7:6.5); lower margin of median ocellus in

line with upper eye margin; distance from median ocellus to antennal scrobe a little

over twice that between scrobes or between lateral ocelli; clypeus at least half as long

as breadth of truncate apical margin, surface strongly, rather closely, punctate;

mandibles rather short, the apical third orange to red; antennal scape nearly half as

broad as long, strongly roughened; flagellomeres II and III about twice as broad as

long, much shorter than either I or IV, both of which (and all succeeding segments

except the last) are broader than long; mouthparts (including palpi) about I 1/2 times

as long as mesonotum, metanotum, and dorsal enclosure of propodeum combined;

terminal maxillary palpal segment longer than either of preceding segments.

Thorax. Punctures rather broad and shallow, those of median portion of scutum

about one puncture width apart, those of lower portions of mesepisternum broad but

very shallow, leaving its surface rather polished; tarsi only slightly paler than femora,

central portion of tibiae; propodeal enclosure strongly, rather irregularly striate

throughout; total propodeal length considerably greater than that of metanotum and

scutellum combined.

Abdomen. Tergal discs polished but punctures rather coarse (about as large as

scutal), shallow, and mostly a little more than one diameter apart; sterna completely

orange in some specimens, but parts or all of sterna III-VI dark brown in others;

sternum VI approximately straight in profile, with moderately dense, dark

pubescence medially (Fig. 19); sternum VII (Fig. 13) similar to that of M. mojavensis;

sternum VIII (Fig. 5) and genital capsule similar to those of M. altadenae.

Female. —Differs from male in having face more shining but only slightly more

sparsely punctate (generally less than one diameter apart) except clypeus which has

punctures two or more diameters apart; head unusually convex and with inner eye

margins nearly parallel (Fig. 37); mouthparts as in Fig. 28; low portions of

mesepisternum with punctures somewhat finer and scutum with punctures coarse

and slightly less than one diameter apart, even medially; tarsi about unicolorous with

tibiae, scarcely paler than femora; metasomal terga I to III generally orange except

for large dark spot laterally on II; terga IV and V becoming darker (especially on

190

PAN-PACIFIC ENTOMOLOGIST

some specimens); tergal punctures coarse, ranging from less than one to about two

diameters apart.

Distribution. —^Found in the coastal ranges of California from San Diego to Marin

Counties. This spring species appears to be oligolectic on Eschscholtzia californica.

New Records. —CALIFORNIA. Alameda County: 1 male, 8 females, Oakland,

Skyline Drive, 12-V-1953, E. Gilbert. Kern County: 1 male, Kern River Pres., 1.5

miles (2.4 km) W Weldon, 21-IV-1983, J. D. Pinto & R. K. Velten. Los Angeles

County: 1 female, Altadena, 2-V-1936, Eschscholtzia calif ornica, C. D. Michener.

Marin County: 1 female. Mill Valley, 12-V-1950, F. X. Williams; 1 female, Novato,

6-V-1962, D. C. Rentz. Monterey County: 2 males, Hastings Reserve, near

Jamesburg, 3-V-1958, J. Powell; 9 males, 8females, Hollister, 22miles (35.4 km) S,

3-V-1982, Eschscholtzia calif ornica, P. F. Torchio; 2 females, Jolon, 14 miles (22.5

km) W, 2-IV-1959, C. W. O’Brien. Riverside County: 2 males. Corona, 12 miles

(19.3 km) S, 1-15 & Indian Truck Trail, 14-IV-1985, Cryptantha intermedia, R. R.

Snelling; 2 males. Riverside, lO-IV-1935, Eschscholtzia californica, P. H.

Timberlake; 1 male, same except 12-IV-1935; 1 male, same except Cryptantha

intermedia', 1 female, same except 12-IV-1933, Eschscholtzia californica, S. C.

Dorman; 1 female, same except 15-IV-1929, P. H. Timberlake; 1 male, same except

15-IV-1932, Cryptantha intermedia', 1 male, same except 15-IV-1933, H. L.

McKenzie; 1 female, same except 15-V-1934, Cryptantha intermedia, P. H.

Timberlake; 1 female, same except 2-IV-1929, Eschscholtzia calif ornica', 1 female,

same except 24-IV-1928; 1 female, same except 29-HI-1933, Amsinckia

douglasiana, H. L. McKenzie; 1 male, same except 5-IV-1933, Cryptantha

intermedia, P. H. Timberlake; 1 male, same except 9-V-1935. San Bernardino

County: 1 female, Devore, 21-IV-1974, J. C. & E. M. Hall. San Luis Obispo

County: 3 females. Black Mt., 6 air miles (9.7 km) NE Pozo, 3300-3600'

(1006-1097m), l-V-1962, J. K. Drew.

Discussion. —This is a very distinctive species, apparently related to M.

mojavensis on the basis of sternum VII in the male, but with many unusual features

including the largely orange abdomen, the short male flagellum, and the coarse

abdominal punctation. Except that the abdomen varies in the amount of black

beyond tergum III, there is no evidence of geographical variation. This red abdomen

is distinctive among Micralictoides except for some females of M. linsleyi which also

exhibit orange coloration on the basal terga but differ from M. ruficaudus in having

roughened interpunctural areas on the frons.

Micralictoides mojavensis Bohart

(Figs. 7,15,26,34)

Micralictoides mojavensis Bohart 1942:120 (Holotype male: CAS).

Male. —Length about 4.5 mm, forewing length 3 mm, head width slightly greater

than length (1.1 or 1.2); similar in most respects to M. chaenactidis, but differs in

having the clypeus and supra-clypeal area more polished, the clypeus more nearly

impunctate (except basolaterally) and the punctures of the supra-clypeal area mostly

separated by three or four times their diameters. The facial pubescence is somewhat

sparser, not at all concealing the clypeus and supraclypeal area. The mouthparts are

about as long as those of M. chaenactidis and distinctly longer than those of M.

altadenae. The propodeum has the sculptured area of the enclosure more restricted

VOLUME 63, NUMBER 2

191

than that of M. chaenactidis and has a broad, nearly smooth zone laterally within the

enclosural boundaries. Sternum VI resembles that of M. chaenactidis, but sternum

VII is strikingly different (Fig. 15). Sternum VIII is similar to that of M. chaenactidis,

but its apical process is a little broader (Fig. 7).

Female. —Very similar to M. altadenae and to the widespread California form of

M. chaenactidis. Head (Fig. 34) length to breadth ratio varies from 0.88 to 0.99.

Mouthparts (Fig. 26) long as in M. chaenactidis. Length of dorsal scopal hair on hind

tibia slightly less than apical width of tibia. Tergum II with punctation of disk sparse,

especially in middle.

Distribution. —Mojave Desert and Los Angeles Basin. All collection records are

from mid-April to late May. Males were collected on Gilia multicaulis and

Platystemon californicus.

New Records. —CALIFORNIA. Inyo County: 1 female, Darwin, 12-V-1969, J.

Doyen; 2 females, same except P. Wells. Riverside County: 2 males, 1 female,

Gavilan, 4-IV-1952, Salvia columbariae, P. H. Timberlake; 1 female, same except

22-IV-1952, Eschscholtzia californica; 1 female, same except lO-IV-1956; 1 female,

same except 16-IV-1956, Gilia multicaulis. San Bernardino County: 3 females,

Adelanto, 20 miles (32.2 km) N, 18-IV-1962, Malacothrix glabrata, P. H.

Timberlake; 1 male, Atolia, l-V-1952, R. M. Bohart; 3 males, 5 females. Mid Hills,

9 miles (14.5 km) SSE Cima, 26-V-1975, T. Griswold; 1 male. Mid Hills, Sec. 12

T12N R14E, 5620' (1713m), 19-V-1980, T. Griswold; 1 male, Sacaton Spr., New

YorkMts., 4100' (1250m), lO-V-1978, T. Griswold; 1 female, Twentynine Palms,

4 miles (6.4 km) S, 14-IV-1935, Malacothrix calif ornica, P. H. Timberlake; 1

female, Victorville, 12 miles (19.3 km) W, 14-IV-1963, Phacelia fremontii, M.

Dickson.

Discussion. —This species is difficult to distinguish from M. chaenactidis in its

externally visible characteristics even though sternum VII of the male resembles that

of M. ruficaudus more than it does that of M. chaenactidis. The female appears to be

separable from M. chaenactidis only by the more sparse punctation of tergum II and

the shorter scopal hair. Since the distribution of these two species overlaps, it is

possible that our association of the sexes in M. mojavensis (based primarily on

similarity in headshape) is not entirely correct. Longer series from single localities

will probably clarify this matter in the future. The punctation and coloration of M.

mojavensis, in general, appears to be intermediate between those of the Coast Range

and more interior forms of M. chaenactidis. Like M. chaenactidis, the species differs

from altadenae in both sexes by its narrower face (but only slightly so), longer

clypeus, and longer mouthparts.

Four females from Red Rock Canyon, Kern County, California are a composite of

characteristics of M. mojavensis and M. dinoceps. The head shape and propodeal

sculpturing are as in M. dinoceps, while the scutal punctation and length of the

maxillary palpi are as in M. mojavensis. It is possible that these females represent

intermediate conditions in a polymorphic species and, consequently, that M.

dinoceps is not a valid species. However, the available material of both M.

mojavensis and M. dinoceps is quite homogeneous with respect to these two

characters. It, therefore, seems possible that the Red Rock Canyon females

represent a third species. Resolution of this problem must await discovery of the

corresponding male.

192

PAN-PACIFIC ENTOMOLOGIST

Micralictoides dinoceps, New Species

(Figs. 6,14, 23, 24, 27, 36)

Holotype male. —Length about 4.5 mm, forewing length 3 mm; body black except

tegula, tarsi, sterna brown. Pubescence white, sparse, rather short, not concealing

integument.

Head. Head distinctly broader than long (1.3); clypeus less than one-third as long

as breadth of truncation, its surface with sparse, coarse punctures; lower margin of

median ocellus lower than upper eye margin; distance from median ocellus to

antellal scrobe less than twice that between antennal scrobes; mandible moderately

long but not slender, apical tooth less than one-third length of mandible, slightly

darker than middle third; punctures of frons close and broad; antennal scape

one-third as broad apically as long, with scattered large punctures; flagellomeres

I-III distinctly broader than long, IV-X ranging from as long as broad to one and

one-half times as long as broad; prementum and stipes shagreened, length of

prementum slightly greater than eye length, maxillary palpus shorter than eye

length.

Thorax. Punctures of scutum and scutellum less than a puncture width apart,

interspaces not shagreened; propodeal enclosure with strong, rather close, regular

longitudinal carinae throughout.

Abdomen. Impunctate margins of terga translucent, not much lighter than

remainder of segments; punctures of terga smaller and much sparser than those on

scutum and scutellum; sternum VI practically straight in profile, uniformly sparsely

haired; sternum VII as in Fig. 14, sternum VIII as in Fig. 6, genital capsule as in Figs.

23,24.

Female. —Differs from male in having the face more polished, punctation of frons

slightly more sparse; head proportioned as in Fig. 36; mouthparts as in Fig. 27;

elevated portion of clypeus nearly straight apically, width of marginal truncation

more than twice length; scutum and scutellum rather polished, with fine punctures

less than a puncture width apart; abdominal terga with punctures as coarse, but much

more sparse; hind tibia nearly as light as hind basitarsus, dorsal scopal hair no longer

than apical width of tibia.

Type Material. —Holotype male. CALIFORNIA, San Bernardino Co., Arrastre

Flat, 7,450 ft (2,271m), 26-VI-1919, Alliumfimbriatum, M. H. O’Brien. Paratypes:

California, San Bernardino County: 7 males and 16 females, same data as holotype; 1

female. Mill Creek, 7400' (2,256m), 29-VI-1942, R. M. Bohart. Los Angeles

County: 1 male, Valyermo, 4 miles (6.4 km) SE, 13-IV-1960, R. R. Snelling.

Holotype deposited at LA CM; paratypes at LA CM, BBSL, and UCD.

Distribution. —Apparently restricted to the San Bernardino Mountains of

southern California.

Discussion. —Males of M. dinoceps are close to those of M. mojavensis and both

can be distinguished from other Micralictoides by the abrupt expansion near the

middle of the lateral arm of sternum VII. M. dinoceps can be separated from M.

mojavensis by the wider head and differences in sternum VII. In M. dinoceps, the

lateral arms of sternum VII are in a tight arc so that together, they form an area

longer than wide; in M. mojavensis, these arms are in loose curves so that they form

an area wider than long. The lateral arm of M. dinoceps is of uniform thickness

VOLUME 63, NUMBER 2

193

beyond the medial expansion whereas in M. mojavensis, it is narrow then apically

expanded.

Females of M. dinoceps are easily confused with those of M. altadenae, M.

chaenactidis and M. mojavensis. From these, they differ by the shortened maxillary

palpi, the fine and more dense scutal punctation, and in the sculpturing of the basal

zone of the propodeum. They further differ from M. mojavensis by the broader head,

and from M. altadenae and M. chaenactidis by the shorter scopal hairs and the sparser

punctation of tergum IF

Acknowledgments

We are grateful to the following individuals for providing the majority of the

approximately 650 specimens used in this study: P. Timberlake and S. Frommer,

University of California at Riverside (UCR), R. Bohart, University of California at

Davis (UCD), P. Hurd, University of California at Berkeley (UCB), R. Snelling,

Los Angeles County Museum (LACM), C. Michener, University of Kansas

(SMEK), W. Pulawski, California Academy of Sciences (CAS), J. Rozen and M.

Favreau, American Museum of Natural History (AMNH), D. Faulkner, San Diego

Natural History Museum (SDNH), and U. Lanham, University of Colorado (UC).

This is a contribution from the Utah Agricultural Experiment Station, Utah State

University, Logan, Utah 84322-4810, Journal Paper No. 3342, and

USD A-Agricultural Research Service-Bee Biology and Systematics Laboratory,

Utah State University, Logan, Utah 84322-5310.

Literature Cited

Bohart, G. E. 1942. A synopsis of the genus Micralictoides. Pan-Pac. Entomol., 18:119-123.

Michener, C. D. 1937. Records and descriptions of North American bees. Ann. Mag. Nat. Hist., (10)

19:313-329,393-410.

-. 1965. A generic review of the Dufoureinae of the Western Hemisphere. Ann. Amer. Entomol.

Soc., 58:321-326.

Timberlake, P. H. 1939. New species of the genus Dufourea from California. Ann. Entomol. Soc. Amer.,

32:395-414.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 194-198

A new species of Andrena (Onagrandrena) from Utah’s

San Rafael Desert (Hymenoptera: Andrenidae)

Robbin W. Thorp

Department of Entomology, University of California, Davis, CA 95616.

Abstract .—A series of a large, distinctive, undescribed species of black Andrena

belonging to the subgenus Onagrandrena was collected by Dr. Frank Parker, USD A

Bee Biology and Systematics Laboratory, Logan, Utah during a survey of aculeate

Hymenoptera from Utah’s San Rafael Desert. It is related to a complex of species

described by E. G. Linsley and J. W. MacSwain for their ecological studies of bees

and Onagraceae of the Great Basin (Linsley, et al. 1963b). I describe the species at

this time to include it in this volume dedicated to E. Gorton Linsley and to make the

name available for a forthcoming review of the subgenus. The format and

abbreviations used in the description follow those used by LaBerge (1967), with the

exception that UCD is now RMB (R. M. Bohart Museum of Entomology).

Andrena (Onagrandrena) linsleyana Thorp, New Species

Female.—Measurements and Ratios .—N = 20; length, 14-15.5 mm; width, 4.8-5

mm; wing length, M = 8.7±0.06 mm; FL/FW, M = l± 0.129; FOVL/FOVW,

M = 2.8± 0.007.

Integumental Color.—Black except as follows: mandible tips mahogany; antennal

flagellomeres 3-10 brownish beneath; tegulae posteriorly mahogany; wing

membrane infuscated, veins light yellowish brown; legs and basal areas of metasomal

sternites mahogany.

Structure .—^Antennal scape equal to flagellomeres 1 to 3; flagellomere 1 slightly

shorter that 2 plus 3 combined (1:1.25); flagellomeres 2, 3 and 4 each as wide as long

and equal to each other in length. Eyes 3 times as long as wide, inner margins

essentially parallel. Malar space short, linear. Mandibles somewhat elongate, when

closed outer mandible extends to end of labrum with subapical dorsal tooth

surpassing midlabrum by two fifths length, without tooth or angle on basal inferior

margin. Galea with outer margin angled outward from apex in straight line, then

bending rearward to insertion of palpus; surface moderately dull, shagreened.

Maxillary palpus nearly twice as long as galea; segmental ratio about

(1.2:2.0:1.5:2.0:1.7:1.7). Labial palpus with first segment curved, somewhat

flattened; segmental ratio about (2.0:1.2:1.0:1.5). Labral process short, tumid,

converging and shallowly emarginate apically with two, very short, blunt teeth,

longer than apical width, without basal depression (Fig. 2); labrum apical to process

without transverse sulcus, with median longitudinal crista, with coarse,

longitudinally elongate, nearly contiguous punctures. Clypeus convex; punctures

coarse, separated by less than one diameter centrally, finer and nearly contiguous

laterally and dorsally; surface shiny, with narrow median longitudinal impunctate

line. Supraclypeal area dull, punctures nearly contiguous and finer than on clypeus.

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195

Face above antennal fossae with coarse, contiguous punctures between fine

longitudinal rugulae. Facial fovea long, extending below level of antennal bases and

2.8 times maximum width, width at top twice width at bottom and separated from

lateral ocellus by 0.4 ocellar diameter, area between fovea and ocellus finely

striate-punctate. Lateral ocelli 2.4 diameters from inner margins of compound eyes

and about one diameter from the top of the vertex. Vertex above lateral ocellus dull,

densely, moderately coarsely punctate. Genal area in profile broader than eye

(about as 8:5), punctures finer than on clypeus, sparse, separated by 1-2 diameters

centrally, closer, and separated by less than one diameter peripherally, surface dull,

shagreened, shiniest centrally.

Pronotum with rounded humeral angle and without dorsoventral ridge, dull,

finely, moderately sparsely punctate, shagreened posteriorly, shiny anteriorly.

Mesoscutum dull with coarse, close punctures separated by less than one diameter,

interspaces shagreened. Scutellum dull, punctures coarse, close (<1 diameter)

becoming contiguous posteriorly. Metanotum dull, punctures coarse and

contiguous. Propodeum with dorsal enclosure moderately rugose, irregular, without

longitudinal rugae (Fig. 1); dorsolaterally and posteriorly dull, with coarse, close and

vertically long-ovate punctures, laterally sparsely punctate in center of corbicular

area. Fore wing with base of vein M ending about 2 vein widths anterior to cu-v.

Metasomal tergum 1 shiny with anterior face impunctate, rounded to dorsal

sparsely punctate disk. Terga 2-4 shiny, finely, sparsely, punctate with punctures

separated by 1-2 diameters, impunctate margin shiny, about one-fifth length of

apical impressed area. Pygidial plate V-shaped with apex truncate, margins curved

upwards, with distinct, raised internal triangular area grading apically and

basolaterally toward margins, mediolaterally sharply declivous with fine striae

extending onto depressed submarginal, impunctate, dull, shagreened area, surface

of raised internal area finely, irregularly striate. Sterna 2-5 with basal areas sparsely,

moderately finely punctate, separated by 3-4 diameters on sternum 2, becoming

closer on succeeding sterna to 1-2 diameters on sternum 5, narrow apical areas

impunctate, shiny, moderately transparent.

Vestiture.—Black. Propodeal corbiculum incomplete, dorsal hairs moderately

long, straight, similar to other hairs above, with many internal hairs and only a

ventral tuft of hairs anteriorly; trochanteral flocculus complete with moderately long

curved hairs; tibial scopal hairs sparse, long (nearly twice tibial width), simple.

Tergal hairs erect, long on tergum 1 (7 times as long as on tergum 2).

Male.—Measurements and Ratios :—N = 20; length, 9.5-14.5 mm; width, 3-4.5

mm; wing length, M = 7.4±0.37; FL/FW, M = 1.12 ± 0.001; FS1/FS2,

M = 1.25± 0.003.

Integumental Color.—Black with exceptions as in female.

Structure.—Antennal scape equal to first two and one fourth flagellomeres;

flagellomere 1 longer than 2 (as 1.3:1.0), equal to 3; segment 2 as wide as long;

segment 3 slightly longer than wide (as 1.3:1.0). Mandibles moderately long, when

closed, outer mandible extends slightly beyond end of labrum. Malar space and galea

as in female. Maxillary palpus as in female, but segmental ratio

(1.0:1.5:1.3:1.7:1.3:1.7). Labial palpus as in female, but ratio about

(1.5:1.0:1.0:1.3). Labral process wider than long (as 2.5:1.5), deeply emarginate

with two teeth (Fig. 4); labrum apical to process convex, finely, sparsely punctate,

shiny, without crista. Clypeus with sculpture as in female, except median impunctate

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PAN-PACIFIC ENTOMOLOGIST

line more difficult to detect, especially above; shiny centrally. Supraclypeal area and

face above antennal fossae as in female. Lateral ocelli separated from inner margin

of compound eye by 3 diameters and from vertex by one diameter. Vertex sculptured

as in female. Genal area in profile broader than eye (as about 7.5:5), sculpture as in

female.

Pronotum with rounded humeral angle and without dorsoventral ridge, surface

sculpture as in female. Mesoscutum and scutellum sculptured as in female.

Propodeum sculptured as in female (Fig. 3). Fore wing with base of vein M as in

female.

Metasomal terga 1-5 sculptured as in female. Tergum 7 with pseudopygidial area

concealed by long hairs at lateral margin, V-shaped and broadly truncate apically.

Sterna 2-5 punctate as in female. Sternum 6 with apical margin turned down slightly,

without emargination. Sternum 7 emarginate apically. Sternum 8 with terminal shaft

equal in length to that of broad base and with apex slightly emarginate. Penis valves

moderately broadened medially, tips narrow, not exceeding tips of gonostyli; dorsal

lobe of gonocoxite relatively narrow, acute apically shorter than base of gonocoxite

(as 3:5); gonostyli enlarged nearly axe-shaped apically with concave outer face,

longer than distance to apex of dorsal lobe of gonocoxite (as 3.5:1.5) (Figs. 5-6).

Vestiture.—^Black.

Type Material. —The holotype female (USNM) and allotype (USNM), 8 female

and 57 male paratypes (USU; CAS; CIS; RMB) were collected at Bullfrog

Campground, Kane County, Utah, 21 April 1983 by F. D. and J. H. Parker. The

type locality is near the end of the road at Lake Powell about 120 km south of

Hanksville, Wayne County (confirmed by F. D. Parker, personal communication).

An additional 11 female paratypes (USU) were collected as follows: UTAH: Emery

Co.: 5,100', 4 air mi. N. Gilson Butte, 29 May 1981, F. D. Parker, 1 female on

Oenothera and 6 females on Compositae; 4,900', Wildhorse Creek, N of Goblin

Valley, 3 June 1982, F. D. Parker, T. L. Griswold, 4 females on Oenothera.

Variation.—Males vary considerably in size with their lengths ranging from

9.5-14.5 mm (M = 12 ± 1.263) while females vary only slightly with lengths ranging

from 14-15.5 mm (M = 14.9 ± 0.326). The median longitudinal impunctate line of

the clypeus of some males is reduced to the lower portion only.

Systematics.— Andrena linsleyana belongs to the species complex that includes: A.

(O.) chylismae-nevadae-thorpi-stagei all described by Linsley and MacSwain. Its

females can be separated from others of this complex by their larger size, the less

pronounced and more irregularly rugose propodeal enclosure, the presence of a

median longitudinal impunctate line on the clypeus, the shorter and more

emarginate labral process. Males of A. linsleyana can be separated from those of A.

chlyismae, the only species of this complex for which males have been described, by

their larger size, the less pronounced and more irregularly rugose propodeal

enclosure, and the presence of a median longitudinal impunctate line on the clypeus.

Males of both have all black pubescence. This is a unique characteristic among

Onagrandrena of the Intermountain Region and will likely be true for other

members of this species group as the males become known.

Flower and Colleciton Records. —According to Dr. F. D. Parker (personal

communication) the specific floral records for the above are: Oenothera pallida

Lindley and Hymenopappus filifolius Hook, and the bees were foraging as late as 11

A.M. I examined the specimens for pollen. All the bees collected on Oenothera bore

small amounts of Oenothera pollen in their scopae. All the females collected on

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197

Figures 1-6. Andrena linsleyana, paratypes. 1. Propodeal enclosure of female. 2. Labral process of

female. 3. Propodeal enclosure of male. 4. Labral process of male. 5,6. Male genital capsule lateral and

apical views.

Hymenopappus had moderate amounts of pollen from Compositae on their faces

below their antennae, but none in their scopae indicating that this was a nectar

source. Its flight season is from late April into June. The specimens from Bullfrog

Campground on 21 April 1981 are all very fresh. The collections made at the other

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PAN-PACIFIC ENTOMOLOGIST

two sites in late May and early June show wear of the mandibles, hairs of the clypeus

and labrum and of the wing tips. The presence of pollen in the scopae of the females

and the lack of males in the latter collections also suggests that these were made near

the end of the season for the bees.

Discussion. —Based on the flower records and pollen analyses, females of A.

linsleyana presumably collect residual pollen in the morning from the evening

opening O. pallida when the nectar supply in the deep hypanthium is low and forage

for nectar on flowers of compositae. This is similar to the behavior of females of

(O.) linsleyi Timberlake which forage in the morning for residual pollen from the

nocturnal O. deltoides Torrey & Fremont and for nectar on Geraea canescens Torrey

& Gray in the Colorado Desert of California (Linsley, et al. 1963a). Females from

Bullfrog Campground were collected at a nest site and males were found sleeping in

shallow burrows under rocks (F. D. Parker, personal communication).

The Canyon Lands subdivision of the Colorado Plateau Province which contains

the San Rafael Desert of Utah has the greatest number of endemic plant species of

any part of the Intermountain Region (Cronquist, et al. 1972). The San Rafael

Desert has produced several endemic bee species (F. D. Parker, personal

communication) as well, so it is not surprising that A. linsleyana with the largest

females of any Onagrandrena represents another endemic species.

Etymology. —I take great pleasure in naming this species in honor of Dr. E.

Gorton Linsley who has contributed greatly to the understanding of the systematics

and ecology of this subgenus of Andrena and who had a great influence on my career

as my major professor and mentor during my graduate years at Berkeley.

Acknowledgments

I thank Dr. Frank Parker for bringing this species to my attention and for

providing additional biological information. I thank Drs. Parker and W. E. LaBerge

for reviewing the manuscript. I thank Mrs. Mary Ann Tenorio, Entomology,

California Academy of Sciences, for assistance with the scanning electron

microscope photos.

Literature Cited

Cronquist, A., A. H. Holmgren, N. H. Holmgren and J. L. Reveal. 1972. Intermountain flora. Vol. 1.

Hafner Publ. Co., NY. 270 pp.

LaBerge, W. E. 1967. Revision of the bees of the genus Andrena of the Western Hemisphere. Part 1.

Callandrena. (Hymenoptera: Andrenidae). Bull. Univ. Nebraska State Mus. 7:1-318.

Linsley, E. G., J. W. MacSwain and P. H. Raven. 1963a. Comparative behavior of bees and Onagraceae.

1. Oenothera bees of the Colorado Desert. Univ. Cahf. Pubis. Entomol. 33:1-24.

-. 1963b. Comparative behavior of bees and Onagraceae. 11. Oenothera bees of the Great Basin.

Univ. Calif. Pubis. Entomol. 33:25-58.

PAN-PACIFIC ENTOMOLOGIST

63(2), 1987, pp. 199-204

Proceedings of the Pacific Coast Engomological Society, 1986

FOUR HUNDRED AND FORTY-FOURTH MEETING

The 444th meeting was held Friday, 21 February 1986, at 8:15 p.m., in the Morrison Auditorium,

California Academy of Sciences, Golden Gate Park, San Francisco, with President Mr. Larry G. Bezark

presiding.

The minutes of the meeting held 13 December 1985 were read and accepted. Nine persons were

proposed and elected as new members: Mr. Richard L. Bottorff and Mr. Tim A. Christiansen as student

members; Mr. Richard P. and Mrs. Louise H. Fall as regular family members; and Mr. Robert D. Haines,

Dr. Patricia G. Lincoln, Mr. Terry D. Miller, Dr. Frank J. Radovsky, and Dr. Jose A. Ramos as regular

members.

Mr. Bezark announced the topics for some of the forthcoming meetings. Several guests were

introduced. Dr. Kirby A. Brown showed slides of bronze medallions casted in 1911 by the Paris mint in

honor of the famous entomologist Jean-Henri Fabre.

The featured speaker. Dr. Jarmila Kukalova-Peck, Department of Geology, Carleton University,

Ottawa, presented a lecture entitled “Paleozoic Insects, the Origin of Wings and the Evolution of Insect

Flight.” She showed slides of these monstrous and bizarre-looking insects from the Carboniferous and

Permian Periods, a majority of which have highly modified beaks for penetrating and sucking the strobili

of primitive plants. She discussed the old paranotal theory which proposes evolution of flight by gradual

modifications of the paranotal lobes from rigid gliding structures to foldable, then flapping structures, and

finally to wings that can power flight. Combining knowledge from paleontology, embryology, and

comparative anatomy, she presented her own “exite-ting” theory which proposes wing evolution from

basipodite exites that migrated from the ventral to the dorso-lateral position. The presence of musculature

in the articulations of the wings supports her theory that any prewing structures were already flexible

before true wings evolved. Nearly all of the very early insects also had segmented abdominal legs. Her

research also led her to conclude that the Arthropoda is a monophyletic group.

The social hour was held in the entomology conference room following adjournment of the meeting.

A total of 102 persons was present, of which 51 signed as members and 29 as guests.—^V. F. Lee,

Secretary.

FOUR HUNDRED AND FORTY-FIFTH MEETING

The 445th meeting was held Friday, 21 March 1986, at 8:15 p.m., in Mulford Hall, University of

California, Berkeley, with President Mr. Larry G. Bezark presiding.

The minutes of the meeting held 21 February 1986 were read and accepted. Five persons were proposed

and elected as new members: Mr. Marc L. Utheim as a student member; and Dr. Edward I. Coher, Prof.

Hua Lizhong, Dr. Daniel Udovic, and Dr. Floyd G. Werner as regular members.

Mr. Bezark mentioned a few papers that appeared in some recent journals and the recent publication of

the 1986 volume of Annual Review of Entomology. Dr. Edward L. Smith presented his recent findings on a

proposed new classification of the hexapodan groups.

The featured speaker. Dr. David H. Kistner, Chico State University, presented a lecture entitled

“Wallace, Wallace’s Lines, and the Wallace Expedition.” He gave a brief biographical sketch of Alfred

Russell Wallace and discussed the Wallace’s Lines, which separate the Oriental from the Australian

Zoogeographical Regions. He suggested that the fauna of Sulawesi (or Celebes) might have been

influenced by plate tectonics since the island is a composite of land masses from each of these regions. In

1985, the Royal Entomological Society of London celebrated its 150th anniversary by sponsoring the

year-long Wallace Expedition, which is the largest expedition in the last 50 years, involving 102 scientists,

including Dr. Kistner and his wife, Alzada. She discussed the logistics of the expedition, in which

volunteers from the British military helped by carrying supplies and setting up camp, etc. for the scientists.

Dr. Kistner discussed his research on the termitophilic staphylinids taken from termite nests in the

Oriental part of the island.

The social hour was held in Wellman Hall following adjournment of the meeting.

A total of 63 persons was present, of which 34 signed as members and 14 as guests.—V. F. Lee,

Secretary.

199

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PAN-PACIFIC ENTOMOLOGIST

FOUR HUNDRED AND FORTY-SIXTH MEETING

The 446th meeting was held Friday, 18 April 1986, at 8:15 p.m., in Morrison Auditorium, Cahfornia

Academy of Sciences, Golden Gate Park, San Francisco, with President Mr. Larry G. Bezark presiding.

The minutes of the meeting held 21 March 1986 were read and accepted. Two persons were proposed

and elected as new regular members: Dr. Jeffrey C. Burne and Dr. Robert V. Dowell.

The featured speaker. Dr. Kirby W. Brown, entomologist with the San Joaquin County Department of

Agriculture, Stockton, presented a lecture entitled “Paper Beetles or Adventures in Literature.” He

showed slides of some old entomological books, some of which had very decorative title pages, and

discussed the evolution of colored illustrations from copper to steel engravings with hand-painted colors to

chromohthographs to photographs. He also discussed the importance of literature to taxonomists. He

summarized the history of entomological literature from the perspective of expeditions, taxonomic

monographs, faunistic studies, and book catalogs. He emphasized that books produced today cannot

duphcate the efforts that went into the older ones because of higher production costs. However, the older

books are being destroyed for their colored plates, which are often removed, framed, and sold separately

to command higher prices than if the books were to be sold intact. Their scientific value is therefore much

reduced.

The social hour was held in the entomology conference room following adjournment of the meeting.

A total of 36 persons was present, of which 23 signed as members and 11 as guest.—^V. F. Lee,

Secretary.

FOUR HUNDRED AND FORTY-SEVENTH MEETING

The 447th meeting was held Friday, 17 October 1986, at 8:15 p.m. , in Morrison Auditorium, California

Academy of Sciences, Golden Gate Park, San Francisco, with President Mr. Larry G. Bezark presiding.

The minutes of the meeting held 18 April 1986 were read and accepted. Twelve persons were proposed

and elected as new members: Mr. Timothy G. Myles, Mr. Felipe N. Soto Adames, and Mr. Wilham P.

Weaver Jr. as student members for 1986, and Mr. Gregory S. Forbes, Mr. Elgin E. Huckstep, Mr.

Timothy A. Kellogg, Mr. Stephen A. Manweiler, Mr. Ronald G. Robertson, Dr. Geoffrey G. E.

Scudder, and Dr. Wilham D. Shepard as regular members for 1986, and Mr. Craig Sondergaard and Dr.

Gregory R. Walker as regular members for 1987.

Mr. Bezark announced two forthcoming entomological conferences and presented shdes of his summer

collecting trip to the Rockies. Dr. Edward L. Smith presented slides of the Eagle Lake Biological Field

Station, operated by California State University, Chico, that is being threatened with closure due to lack

of funding, and make an appeal for letters of support from the audience.

The featured speaker, Dr. Harry Greene, professor of zoology at the University of California,

Berkeley, presented a lecture entitled “Behavioral Ecology of Tropical Predators.” He showed shdes of

predatory vertebrates and invertebrates of the Finca La Selva station in Costa Rica and discussed why

there is an increase in the diversity of vertebrates when one compares the polar areas to the tropics. He

also suggested that larger vertebrate predators in the tropics should not be eliminated because they can

check the smaller predators which are able to more significantly affect avian populations.

The social hour was held in the entomology conference room f oho wing adjournment of the meeting.

A total of 71 persons was present, of which 44 signed as members and 20 as guests.—V. F. Lee,

Secretary.

FOUR HUNDRED AND FORTY-EIGHTH MEETING

The 448th meeting was held as a joint meeting with the Northern California Spider Society on Friday,

21 November 1986, at 8:15 p.m., in Mulford HaU, University of Cahfornia, Berkeley, with President Mr.

Larry G. Bezark presiding.

The minutes of the meeting held 17 October 1986 were read and accepted. Two persons were proposed

and elected as new student members for 1987: Mr. Denn is W. Gray and Mr. Larry J. Orsak.

Mr. Bezark appointed members of the auditing committee, consisting of Mr. H. Vannoy Davis (as

chair). Dr. Paul H. Arnaud Jr., and Mrs. Helen K. Court, and the nominating committee, consisting of

Dr. Clifford Y. Kitayama (as chair). Dr. Barry M. Wilk, and Dr. Robbin W. Thorp. He reminded the

audience of the forthcoming Entomological Society of American meeting in Reno in December. He

demonstrated a way to conserve space when pinning specimens into schmitt boxes. Dr. J. Gordon

Edwards showed slides of his recent trip to New Zealand. Mr. Bezark then introduced Dr. Jack B. Fraser,

VOLUME 63, NUMBER 2

201

president of the Northern California Spider Society, who invited members of the audience to join the

spider society and offered their tee shirts for sale.

Dr. Fraser introduced the featured speaker, Ms. Teresa Meikle-Griswold, Natal Museum,

Pietermaritzburg, South Africa, who presented a slide lecture entitled “The Natural History of Two

Species of Group-living Eresid Spiders in Southern Africa.” She discussed the natural history of Magunia

dumicola (Pocock) and Stegodyphus mimosarum Pavesi, commonly called community spiders. These

spiders form group living cooperatives. She showed that the eresid colonies are very complex biological

communities, with predators, parasitoids, and kleptoparasites living among the eresids. The Zulus utilize

these spiders in the control of flies that live around cattle pens.

The social hour was held in Wellman Hall following adjournment of the meeting.

A total of 55 persons was present, of which 28 signed as members and 22 as guests.— W. F. Lee,

Secretary.

FOUR HUNDRED AND FORTY-NINTH MEETING

The 449th meeting was held on Friday, 12 December 1986, at 8:20 p.m., in Morrison Auditorium,

California Academy of Sciences, San Francisco, with President Mr. Larry G. Bezark presiding.

The minutes of the meeting held 21 November 1986 were read and accepted. Three persons were

proposed and elected as new regular members: for 1986, Dr. Allen M. Young, and for 1987, Mr. E.

Penryn Flemyng and Dr. Paul K. Lago.

Mr. Bezark introduced his parents. Bud and Betty. Dr. Stanley C. Williams introduced Dr. Donald K.

Fletcher. Mr. Bezark announced that he recently returned from the Entomological Society of American

meetings where he saw Christmas cards with depictions of insects, a new book with keys to braconids, and

a new book on Pacific Island names. He then called for committee reports. Dr. Paul H. Arnaud Jr., chair

of the historical committee, reported that there was considerable use of the Albert Koebele notes. Mr.

Daniel F. Gross, chair of the membership committee, reported that the 1986 membership rolls consisted

of 433 members, including 306 regular members, 60 student members, 44 sponsoring members and

sponsoring family members. The year saw an addition of 38 new members, 27 regular and 11 student

members. He then read the names of the members who continue to support the journal by being

sponsoring members for 1986: Phillip A. Adams, Robert P. Allen, Richard K. Allen, William F. Barr,

Richard M. Bohart, Paula and Robert Buickerood, Donald J. Burdick, Leopoldo E. Caltagirone, Arthur

L. Chan, Kenneth W. Cooper, J. Gordon and Alice Edwards, George R. Ferguson, William E. and

Stephenie S. Ferguson, Wayne C. Field Jr., Eric M. Fisher, John G. Franclemont, E. Eric Grissell, John

E. Hafernik Jr., Kenneth S. Hagen, Alice S. Hunter, Johannes L. Joos, Benjamin Keh, Dennis M. Kubly,

Robert J. Lyon, Robert L. Mangan, David G. Marqua, Gordon A. Marsh, Woodrow W. Middlekauff,

Robert B. Miller, Calvert E. Norland, Harry W. Oswald, Richard L. Penrose, Jacqueline L. Robertson,

Leslie S. Saul, Evert I. Schlinger, David B. Scott, Harvey 1. Scudder, Terry N. Seeno, Frank E. Skinner,

Edward L. Smith, Roy R. Snelling, Marius S. and Joanne S. Wasbauer, and David B. Weissman. Dr.

Arnaud read notes from Mr. H. Vannoy Davis, chair of the auditing committee, stating that the financial

books of the Society were in good order. He also read a report from Dr. Wojciech J. Pulawski, Treasurer,

on the fund balances of the Society as of 30 September 1986, and thanked Mrs. L. Gail Freihofer and Mr.

Davis for their help in maintaining the books. He mentioned that page charges for Dr. Hua Lizhong’s

paper on new Chinese cerambycids, published in vol. 62 of The Pan-Pacific Entomologist, was supported

by a grant from the Charles P. Alexander Publication Fund. Mr. Bezark raffled a book, California Insects

by Powell and Hogue, to a member of the audience.

Dr. Clifford Y. Kitayama, chair of the nominating committee, proposed the 1987 slate of candidates for

officers: Dr. Stanley C. Williams, as president, Dr. Wojciech J. Pulawski, as treasurer, Mr. Vincent F.

Lee, as secretary, and Mr. Alan I. Kaplan, as president-elect. The members of the Society then voted in

the slate as officers for the new year. Mr. Bezark handed the gavel over to the president. Dr. Williams,

who then presided over the remainder of the meeting.

Mr. Lee announced that, by the action of the publication committee, the Society has changed the

printer for the journal: A-R Editions of Madison, Wisconsin will replace Allen Press, starting in 1987. Dr.

Ronald E. Stecker mentioned that a former entomology student of San Jose State University, Mr. Robert

Anderson, died tragically in an auto accident a few months ago. Dr. Williams announced with regret that

the Society will not participate in the annual meeting of the American Association for the Advancement of

Science-Pacific Division in San Diego in June 1987. However, he suggested that the Society put on an

all-day seminar and meeting in place of it at the end of the year.

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PAN-PACIFIC ENTOMOLOGIST

Dr. Edward L. Smith announced two new books: Spider: Webs, Behavior, and Evolution, edited by

W. A. Shear, and E. N. Kjellesvig-Waering’s posthumous monograph on the fossil scorpions of the

world, published in Palaeontologica Americana. He also showed drawings of Carboniferous thysanurans,

japygids, and monurans, and a rediscovered Protodonata, which was previously misidentified as

belonging to another insect order, executed by Dr. Jarmila Kukalova-Peck. Mr. Benjamin Keh showed a

book on insect studies in China, written in Esperanto. Mr. Dean W. Jamieson mentioned that insect and

arachnid specimens from Iran collected by Mr. Bruce Sanford will be on display at the social hour. Mr.

Larry J. Orsak showed slides of possible spider mimics by tephritids and possibly by otitids, possible spider

images on wings of geometrids in Wau, New Guinea, and noctuid caterpillars which resemble

teratological flowers in Arizona.

Dr. Williams introduced the featured speaker, Mr. Larry G. Bezark, Biological Control Services,

California Department of Food Agriculture, Sacramento, who presented the presidential address entitled

“Biological Control of Water Hyacinth in California.” He discussed the introduction and spread of this

aquatic weed, the world’s most prohfie plant, and the attempts to control it in California with two species

of weevils {Neochetina bruchi Hustache and N. eichorniae Warner) and one species of pyrahd moth

{Sameodes albiguttalis (Warren)). He talked about the hfe cycles of the host and the biological control

agents. The insects are now well estabhshed in California and are keeping the weed in check.

The social hour was held in the entomology conference room following adjournment of the meeting.

A total of 42 persons was present, of which 30 signed as members and 12 as guests.— V. F. Lee,

Secretary.

VOLUME 63, NUMBER 2

203

PACIFIC COAST ENTOMOLOGICAL SOCIETY

STATEMENT OF INCOME, EXPENDITURES AND

CHANGES IN FUND BALANCES

Year Ended September 30,1986 and 1985

1986 1985

Income

Dues and subscriptions . $11,450 $9,714

Reprints and miscellaneous . 12,179 20,504

Sales of Memoirs . 74 47

Interest . 5,013 5,708

Dividends . 485 463

Increase in value of capital stock: American Telephone & Telegraph Com¬

pany and Pacific Telesis Group. 2,477 402

$31,678 $36,838

Expenditures

Publication costs—^Pan-Pacific Entomologist. $18,343 $25,165

Reprints, postage and miscellaneous. 1,087 514

IBM Personal Computer. . 2,400

$19,430 $28,079

Increase (Decrease) in fund balances . $12,248 $ 8,759

Fund balances October 1,1985 and 1984 . 87,230 78,471

Fund balances September 30,1986 and 1985 . $99,478 $87,230

STATEMENT OF ASSETS

September 30,1986 and 1985

1986 1985

Cash in bank

Commercial account . $ 9,258 $ 3,941

Savings accounts & Certificates of Deposit

General Fund . 18,863 17,832

Charles P. Alexander Fund. 36,264 34,282

Fall Memoir Fund . 26,359 24,918

Total cash in bank . $90,744 $80,973

Investment in 80 shares of American Telephone & Telegraph Co. common

stock and 132 shares of Pacific Telesis Group at market value. $ 8,734 $ 6,257

$99,478 $87,230

See accompanying notes to the financial statements on following page.

204

PAN-PACIFIC ENTOMOLOGIST

PACIFIC COAST ENTOMOLOGICAL SOCIETY

NOTES TO THE FINANCIAL STATEMENTS

Year Ended September 30,1986

Summary of Significant Accounting Policies

Accounting Method: Income and expenses are recorded by using the cash basis of accounting. Capital

Expenditures: Annual capital expenditures of $5,000 or less are charged to expense. Marketable Securities:

American Telephone & Telegraph Co. and Pacific Telesis Group common stock are carried at market

value. Increases and decreases in value are reflected in income. Income Tax: The Society is exempt from

Federal income and Cafifornia franchise tax. Undeposited Receipts —$525. Accounts Receivable —$3,322.

Accounts Payable —$7,042.

As Chairman of the Auditing Committee, and in accordance with its bylaws, I have reviewed the financial

records of the Society.

During the course of this review nothing was noted which indicated any material inaccuracy in the forego¬

ing statements.

H. Vannoy Davis

Chairman of the Auditing Committee

VOLUME 63, NUMBER 2

205

Corrigenda

We regret that figures 1-9 in C. Dennis Hyne’s article, “New Species of the Genus

Stringomyia from the South Pacific and Southeast Asia (Diptera, Tipulidae),” Vol. 63/

1 (January 1987), were omitted. The omitted figures appear below.

Figures 1-9. 1. Styringomyia bidentata n. sp. 2. S. bidens n. sp. 3. S. digitostylus n. sp. 4. S.

rostrostylus n. sp. 5. S. vietnamensis n. sp. 6. S. labuanae n. sp. 7. S. ysabellae n. sp. 8. S. dilinhi n.

sp. 9. 5. idioformosa n. sp. (b = basistyle, t = ninth tergite, s = ninth sternite, od = outer dististyle,

id = inner dististyle, il = inner lobe of inner dististyle, ol = outer lobe of inner dististyle, p = phallosome).

THE PAN-PACIFIC ENTOMOLOGIST

Information for Contributors

Members are invited to submit manuscripts on the systematic and biological phases of entomology, including short notes or artic¬

les on insect taxonomy, morphology, ecology, behavior, life history, and distribution. Non-members may submit manuscripts for

pubhcation, but they should read the information below regarding editing and administrative charges. Manuscripts of less than a

printed page will be published as space is available, in Scientific Notes. All manuscripts will be reviewed before acceptance. Manu¬

scripts for publication, proofs, and all editorial matters should be addressed to the editor.

General. —The metric system is to be used exclusively in manuscripts, except when citing label data on type material, or in direct

quotations when cited as such. Equivalents in other systems may be placed in parentheses following the metric, i.e. “1370 m

(4500ft.) elevation.”

Typing.—Two copies of each manuscript must be submitted (original and one xerox copy or two xerox copies are suitable). All

manuscripts must be typewritten, double-spaced throughout, with ample margins, and be on bond paper or an equivalent

weight. Carbon copies or copies on paper larger than 8V2 X 11 inches are not acceptable.

Underscore only where italics are intended in the body of the text. Number all pages consecutively and put authors name on each

sheet. References to footnotes in text should be numbered consecutively. Footnotes must be typed on a separate sheet.

Manuscripts with extensive corrections or revisions will be returned to the author for retyping.

First Page. —^The page preceding the text of the manuscript must include (1) the complete title, (2) the order and family in paren¬

theses, (3) the author’s name or names, (4) the institution with city and state or the author’s home city and state if not affiliated

(5) the complete name and address to which proof is to be sent.

Names and descriptions of organisms. —The first mention of a plant or animal should include the full scientific name with the

author of a zoological name not abbreviated. Do not abbreviate generic names. Descriptions of taxa should be in telegraphic

style. The International Code of Zoological Nomenclature must be followed.

Tables. —Tables are expensive and should be kept to a minimum. Each table should be prepared as a line drawing or typed on a

separate page with heading at top and footnotes below. Number tables with Arabic numerals. Number footnotes consecu¬

tively for each table. Use only horizontal rules. Extensive use of tabular material requiring typesetting may result in increased

charges to the author.

Illustrations. —No extra charge is made for line drawings or halftones. Submit only photographs on glossy paper and original

drawings. Authors must plan their illustrations for reduction to the dimension of the printed page (117 x 181 mm; 45/8 x 71/8

inches). If possible, allowance should be made for the legend to be placed beneath the illustration. Photographs should not be

less than the width of the printed page. Photographs should be mounted on stiff card stock, and bear the illustration number

on the face.

Loose photographs or drawings which need mounting and/or numbering are not acceptable. Photographs to be placed to¬

gether should be trimmed and abut when mounted. Drawings should be in India Ink, or equivalent, and at least twice as large

as the printed illustration. Excessively large illustrations are awkward to handle and may be damaged in transit. It is recom¬

mended that a metric scale be placed on the drawing or the magnification of the printed illustration be stated in the legend

where apphcable. Arrange figures to use space efficiently. Lettering should reduce to no less than 1 mm. On the back of each

illustration should be stated (1) the title of the paper, (2) the author’s complete name and address, and (3) whether he wishes

the illustration returned to him. Illustrations not specifically requested will be destroyed. Improperly prepared illustrations

will be returned to the author for correction prior to acceptance of the manuscript.

Figure legends. —Legends should be typewritten double-spaced on separate pages headed EXPLANATION OF FIGURES and

placed following LITERATURE CITED. Do not attach legends to illustrations.

References. —All citations in text, e.g., Essig (1926) or (Essig 1958), must be listed alphabetically under LITERATURE CITED

in the following format:

Essig, E. O. 1926. A butterfly migration. Pan-Pac. Entomol., 2:211-212.

Essig, E. O. 1958. Insects and mites of western North America. Rev. ed. The Macmillan Co., New York, 1050 pp.

Abbreviations for titles of journals should follow a recent volume of Serial Sources for the Biosis Data Base, BioSciences

Information Service. For Scientific Notes the citations to articles will appear within the text, i.e. . . .“Essig (1926, Pan-Pac.

Entomol., 2:211-212) noted. . . .”

Proofs, reprints, and abstracts. —Proofs and forms for the abstract and reprint order will be sent to authors. Changes in proof will

be charged to the author.

Editing and administrative charges. —Papers by members of the Pacific Coast Entomological Society are charged at the rate of

$30.00 per page. Members without institutional or grant funds may apply for a society grant to cover a maximum of one-half of

these charges. Non-members will be charged at the rate of $60.00 per page. Editing and administrative charges are in addition

to the charge for reprints and do not include the possible charges for author’s changes after the manuscript has been sent to the

printer.

PUBLICATIONS

OF THE

PACIFIC COAST ENTOMOLOGICAL SOCIETY

PROCEEDINGS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.

Vol. 1 (16 numbers, 179 pages) and Vol. 2 (9 numbers, 131 pages). 1901-1930.

Price $5.00 per volume.

THE PAN-PACIFIC ENTOMOLOGIST.

Vol. 1 (1924) to Vol. 51 (1975), price $10.00 per volume of 4 numbers, or $2.50

per single issue. Vol. 52 (1976) to Vol. 57 (1981), price $15.00 per volume, or

$3.75 per single issue, except for Vol. 57, no. 1, $10.00. Vol. 58 (1982) and subse¬

quent issues, $20.00 per volume or $5.00 per single issue.

MEMOIRS OF THE PACIFIC COAST ENTOMOLOGICAL SOCIETY.

Volume 1. The Sucking Lice by G. F. Ferris. 320 pages. Published October 1951.

Price $10.00 (plus $1.00 postage and handling).*

Volume 2. The Spider Mite Family Tetranychidae by A. Earl Pritchard and

Edward W. Baker. 472 pages. Published July 1955. Price $10.00 (plus $1.25 post¬

age and handling).*

Volume 3. Revisionary Studies in the Nearctic Decticinae by David C. Rentz and

James D. Birchim. 173 pages. Published July 1968. Price $4.00 (plus $0.75 post¬

age and handling).*

Volume 4. Autobiography of an Entomologist hy Robert L. Usinger. 343 pages.

Published August 1972. SPECIAL PRICE $5.00 (plus $1.00 tax, postage, and

handling for California orders, $0.70 postage and handling for non-California

U.S. orders, or $1.70 for foreign orders). No members discount at this special

price.

Volume 5. Revision of the Millipede Family Andrognathidae in the Nearctic Re¬

gion by Michael R. Gardner. 61 pages. Published January 21, 1975. Price $3.00

(plus $0.75 postage and handling).*

*(Add 6% sales tax on all California orders (residents of Alameda, Contra Costa, San Francisco, Santa

Clara and Santa Cruz counties add 6 1 / 2 %). Members of the Society will receive a 20% discount on the

price of the memoirs,)

Send orders to:

Pacific Coast Entomological Society

% California Academy of Sciences

Golden Gate Park

San Francisco, California 94118-9961

U.S.A.

Vol. 63

July 1987

THE

No. 3

Pan-Pacific Entomologist

O’NEILL, K. M., and L. BJOSTAD—The male mating strategy of the bee Nomia nevadensis

(Hymenoptera: Halictidae); Leg structure and mate guarding . 207

WICKMAN, B. E. and T. R. TORGERSEN—Phenology of Douglas-fir tussock moth Orgyia

pseudotsugata, egg eclosion and mortality in a thinned and unthinned stand (Lepidoptera;

Lymantriidae). 218

BURNE, J. C.—A morphometric analysis of Mordellistena Costa in Southwestern United States

(Coleoptera: Mordellidae) . 224

HALSTEAD J. A .—Acanthochalcis nigricans Cameron—New distributional information,

including South America (Hymenoptera: Chalcididae) . 236

EHLER, L. E.—Ecology of Rhopalomyia californica Felt at Jasper Ridge (Diptera:

Cecidomyiidae) . 237

NADEL, H.—Male swarms discovered in Chalcidoidea (Hymenoptera: Encyrtidae,

Pteromalidae). 242

SPIETH, H. T.—^The Drosophila fauna of a native California forest (Diptera: Drosophilidae) .. 247

HALSTEAD, J. A. — On the rearing of Michrochridium minutum and its probable

host —Ammoplanellus (Ammoplanellus) umatilla (Hymnoptera: Chrysididae, Sphecidae) 256

WALKER, G. P.—Probing behavior of Aphis helianthi (Homoptera: Aphididae) and its

preference fo Pittosporum tobira leaves of different ages. 258

POLHEMUS,D. A.,andJ. T. POLHEMUS—AnewgenusofNaucoridae (Hemiptera) from the

Philippines, with comments on zoogeography . 265

SCUDDER, G. G. E.—A review of the genus Nosostethus Kirkaldy (Hemiptera: Lygaeaidae:

Lygaeinae) . 270

HALSTEAD, J. A. and C. G. NIWA —Rhyacionia zozana (Lepidoptera: Tortricidae), host of

Hockeria tenuicornis (Hymenoptera: Chalcididae) in Oregon. 276

WAGNER, D. L.-—A new Microcalyptris species from California (Lepidoptera: Nepticulidae) . 278

GOEDEN, R. D.—Life history of Trupanea conjuncta (Adams) on Trixus californica Kellogg in

Southern California (Diptera: Tephritide) . 284

FORBES, G. S.—The status of Efferia similis (Williston), with descriptions of three new Nearctic

species in the albibarbis group (Diptera: Asilidae). 292

SAN FRANCISCO, CALIFORNIA • 1987

Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY

in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

EDITORIAL BOARD

J. A. Chemsak, Editor

R. S. Lane, Associate Editor

W. J. Pulawski, Treasurer J.T. Doyen

R.M. Bohart J. A. Powell J. E. Hafernik, Jr.

Published quarterly in January, April, July, and October with Society Proceedings

appearing in the October number. All communications regarding nonreceipt of num¬

bers, requests for sample copies, and financial communications should be addressed to

the Treasurer, Dr. Wojciech J. Pulawski, California Academy of Sciences, Golden

Gate Park, San Francisco, CA 94118-9961.

Application for membership in the Society and changes of address should be ad¬