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
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES

The Pan-Pacific Entomologist

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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.

2

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

3

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.

4

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

6

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

7

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

8

PAN-PACIFIC ENTOMOLOGIST

i

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

la

lae

bealesi

indai

VOLUME 63, NUMBER 1

9

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.

10

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

11

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

12

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

13

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

14

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

15

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

17

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.

18

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

19

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

20

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

21

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)

22

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

23

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.

24

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

25

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.

26

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

27

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

28

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

29

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.,

30

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

31

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

32

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

33

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

34

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

35

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.

36

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

38

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

39

Table 1. Characteristics of nesting trees oiApis dorsata in Sabah.

Tree

Bark

Ladder

Number of

Active Colonies

Remarks

rough

+

0

rough

+

0

rough

—

>2

possibly one or two
other hidden nests

semi-rough

—

1

smooth

+

0

dead tree

smooth

+

0

smooth

+

0

smooth

+

0

smooth

+

1

smooth

+

3

smooth

+

7

on two adjoining trees

smooth

—

0

definite comb-scar

smooth

—

4

smooth

—

4

smooth

—

7

% 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

40

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

41

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

42

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;

44

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

45

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

47

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

49

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

51

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.

54

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

55

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

57

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

58

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

59

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

61

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

3

14

18

2

2

Hippodamia

South

9

30

33

0

0

convergens

East

13

35

27

0

0

West

7

11

7

0

0

Chrysoperla

South

6

15

5

1

1

carnea

East

14

13

4

0

1

West

0

0

0

0

0

Hemerobius sp.

South

1

0

0

0

1

(prob. ovalis)

East

2

2

0

1

0

West

8

5

5

2

2

Parasitic Hymenoptera

South

7

4

22

11

3

East

5

19

19

3

9

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

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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

63

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

EA

LA LE

EA

LA

LE

Chrysoperla

carnea

36

45 20

28

26

44

Hemerobius sp.

(prob. ovalis)

6

3 26

2

1

30

®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

64

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).

66

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

67

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.

68

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

69

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.

70

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

71

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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.

72

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

73

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.

74

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.

76

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

77

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

78

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

79

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

80

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

81

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

83

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

2

0.3

0

0.0

2

0.7

Wormaldia undescr. sp. A

44

8.8

12

4.7

21

7.3

Psychomyiidae

Tinodes consuetus McL.

24

4.8

10

3.9

10

3.5

Tinodes siskiyou Denning

0

0.0

2

0.8

0

0.0

Polycentropodidae

Polycentropus halidus Milne

0

0.0

1

0.4

0

0.0

Unassociated Polycentropus

Females:

Polycentropus sp. #1

3

0.6

0

0.0

0

0.0

Polycentropus sp. #2

1

0.2

0

0.0

0

0.0

Hydropsychidae

Cheumatopsyche mickeli

Denning

3

0.6

5

2.0

0

0.0

Homoplectra oaklandensis

(Ling)

1

0.2

0

0.0

0

0.0

Hydropsyche californica Banks

0

0.0

2

0.8

0

0.0

Hydropsyche philo Ross

51

10.2

25

9.8

14

4.9

Hydropsyche occidentalis

Banks

5

1.0

7

2.7

2

0.7

Hydropsyche oslari Banks

4

0.8

2

0.8

7

2.4

Hydropsyche sp. A

1

0.2

0

0.0

1

0.3

Parapsyche almota Ross

1

0.2

0

0.0

3

1.0

Rhyacophilidae

Rhyacophila angelita Banks

1

0.2

3

1.2

12

4.2

Rhyacophila vedra Milne

0

0.0

0

0.0

1

0.3

Rhyacophila vocala Milne

1

0.2

2

0.8

4

1.4

Rhyacophila undescr. sp. A

1

0.2

0

0.0

2

0.7

Rhyacophila undescr. sp. B

0

0.0

1

0.4

1

0.3

Rhyacophila undescr. sp. C

0

0.0

1

0.4

0

0.0

Unassociated Rhyacophila

Females:

Rhyacophila sp. #1

1

0.2

2

0.8

4

1.4

Rhyacophila sp. #2

0

0.0

3

1.2

16

5.6

Rhyacophila sp. #3

0

0.0

0

0.0

1

0.3

Rhyacophila sp. #4

0

0.0

0

0.0

1

0.3

Glossosomatidae

Unassociated Agapetus

Female:

Agapetus sp. #1

0

0.0

0

0.0

1

0.3

84

PAN-PACIFIC ENTOMOLOGIST

Table 1. continued

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Glossosoma califica Denning

1

0.2

0

0.0

0

0.0

Glossosoma oregonense Ling

0

0.0

0

0.0

1

0.3

Unassociated Glossosoma

Females:

Glossosoma sp. #1

0

0.0

2

0.8

0

0.0

Glossosoma sp. #2

0

0.0

0

0.0

3

1.0

Hydroptilidae

Hydroptila arctia Ross

31

6.2

47

18.7

22

7.7

Hydroptila sp. A nr.

rono Ross

18

3.6

9

3.5

10

3.5

Unassociated Hydroptila

Females:

Hydroptila sp. #1

39

7.8

12

4.7

8

2.8

Hydroptila sp. #2

0

0.0

1

0.4

4

1.4

Ithytrichia sp. A

0

0.0

1

0.4

0

0.0

Neotrichia sp. A

1

0.2

0

0.0

0

0.0

Neotrichia undescr. sp. A

0

0.0

1

0.4

0

0.0

Unassociated Neotrichia

Female:

Neotrichia sp. #1

11

2.2

2

0.8

0

0.0

Ochrotrichia undescr. sp. B

0

0.0

0

0.0

1

0.3

Ochrotrichia undescr. sp. C

0

0.0

0

0.0

1

0.3

Ochrotrichia \md&sci. sp. D

0

0.0

0

0.0

2

0.7

Ochrotrichia undescr. sp. E

1

0.2

0

0.0

1

0.3

Unassociated Ochrotrichia

Female:

Ochrotrichia sp. #1

4

0.8

2

0.8

1

0.3

Oxyethira sp. A

0

0.0

5

2.0

0

0.0

Unassociated Hydroptilidae

Females:

Hydroptilidae sp. #1

0

0.0

3

1.2

0

0.0

Brachycentridae

Amiocentrus aspilus (Ross)

4

0.8

3

1.2

2

0.7

Unassociated Amiocentrus

Females:

Amiocentrus sp. #1

2

0.3

0

0.0

0

0.0

Amiocentrus sp. #2

5

1.0

3

1.2

14

4.9

VOLUME 63, NUMBER 1

85

Table 1. continued

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Lepidostomatidae

Lepidostoma strophis Ross

7

1.4

0

0.0

1

0.3

Unassociated Lepidostoma

Females:

Lepidostoma sp. #1

15

3.0

3

1.2

2

0.7

Lepidostoma sp. #2

0

0.0

1

0.4

0

0.0

Sericostomatidae

Gumaga nigricula (McL.)

78

15.6

20

7.9

62

21.7

Odontoceridae

Mariliaflexuosa Ulmer

0

0.0

1

0.4

0

0.0

Unassociated Marilia

Females;

Marilia sp. #1

0

0.0

1

0.4

0

0.0

Marilia sp. #2

4

0.8

0

0.0

0

0.0

Unassociated Odontoceridae

Female:

Odontoceridae sp. #1

0

0.0

0

0.0

1

0.3

Helicopsychidae

Helicopsyche borealis
(Hagen)

126

25.1

51

20.2

42

14.7

Calamoceratidae

Heteroplectron californicum

McL.

0

0.0

1

0.4

2

0.7

Leptoceridae

Mystacides alafimbriata

Hill-Griffin

0

0.0

2

0.8

1

0.3

Ylodes frontalis
(Banks)

6

1.2

2

0.8

0

0.0

Unassociated Ylodes

Females:

Ylodes sp. #1

0

0.0

0

0.0

1

0.3

Ylodes sp. #2

4

0.8

1

0.4

1

0.3

Total N

Total No. Taxa (Excluding

Unassociated Females)

No. Unassociated Females

501

23

89

252

25

36

286

26

58

86

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)

24

16.7

0

0.0

%

0.6

Wormaldia cruzensis (Ling)

0

0.0

1

0.2

0

0.0

Wormaldia\xnd&%cx. sp. A

0

0.0

2

0.4

0

0.0

Psychomyiidae

Nyctiophylax sp. A

0

0.0

0

0.0

1

0.3

Tinodes siskiyou Denning

1

0.7

1

0.2

0

0.0

Unassociated Tinodes

Female:

Tinodes sp. #1

0

0.0

2

0.4

0

0.0

Polycentropodidae

Polycentropus halidus Milne

0

0.0

2

0.4

1

0.3

Unassociated Polycentropus

Female:

Polycentropus sp. #3

0

0.0

1

0.2

0

0.0

Hydropsychidae

Hydropsyche philo Ross

0

0.0

0

0.0

2

0.6

Hydropsyche occidentalis

Banks

11

7.6

3

0.6

1

0.3

Hydropsyche oslari Banks

12

8.3

21

4.4

10

2.9

Parapsyche almota Ross

0

0.0

2

0.4

3

0.9

Rhyacophilidae

Rhyacophila narvae Navas

0

0.0

3

0.6

2

0.6

Rhyacophila pellisa Ross

0

0.0

35

7.3

22

6.4

Rhyacophila sierra Denning

0

0.0

0

0.0

1

0.3

Rhyacophila vao Milne

2

1.4

13

2.7

2

0.6

Rhyacophila vedra Milne

0

0.0

5

1.0

0

0.0

Rhyacophila vocala Milne

0

0.0

1

0.2

1

0.3

Rhyacophila undescr. sp. A

0

0.0

1

0.2

1

0.3

Rhyacophila\xr\dQS>cx. sp. C

0

0.0

1

0.2

0

0.0

Unassociated Rhyacophila

Females:

Rhyacophila sp. #2

0

0.0

1

0.2

0

0.0

Rhyacophila sp. #5

0

0.0

3

0.6

10

2.9

Rhyacophila sp. #6

0

0.0

35

7.3

27

7.9

Rhyacophila sp. #7

0

0.0

4

0.8

6

1.7

Rhyacophila sp. #8

0

0.0

0

0.0

1

0.3

Rhyacophila sp. #9

0

0.0

6

1.3

0

0.0

Rhyacophila sp. #10

3

2.1

2

0.4

0

0.0

VOLUME 63, NUMBER 1

87

Table 2. continued

No.

1979

Percent

of total

No.

1980

Percent

of total

No.

1981

Percent

of total

Glossosomatidae

Agapetus taho Ross

23

16.0

0

0.0

7

2.0

Unassociated Agapetus

Female:

Agapetus sp. #1

0

0.0

3

0.6

1

0.3

Anagapetus sp. A

1

0.7

2

0.4

0

0.0

Glossosoma ore gone ns e Ling

0

0.0

1

0.2

0

0.0

Unassociated Glossosoma

Female

Glossosoma sp. #1

0

0.0

8

1.7

4

1.2

Hydroptilidae

Hydroptila arctia Ross

9

6.3

14

2.9

15

4.4

Hydroptila sp. A nr.
rono Ross

12

8.3

1

0.2

3

0.9

Unassociated Hydroptila

Females:

Hydroptila sp. #1

8

5.6

9

1.9

5

1.5

Hydroptila sp. #2

0

0.0

1

0.2

1

0.3

Neotrichia sp. A

5

3.5

1

0.2

0

0.0

Unassociated Neotrichia

Female:

Neotrichia sp. #1

5

3.5

0

0.0

0

0.0

Ochrotrichia undescr. sp. A

1

0.7

0

0.0

0

0.0

Ochrotrichia undescr. sp. E

1

0.7

0

0.0

0

0.0

Ochrotrichia undescr. sp. F

0

0.0

0

0.0

2

0.6

Unassociated Ochrotrichia

Female:

Ochrotrichia sp. #1

1

0.7

3

0.6

1

0.3

Oxyethira sp. A

0

0.0

0

0.0

1

0.3

Unassociated Hydroptilidae

Females:

Hydroptilidae sp. #1

1

0.7

0

0.0

1

0.3

Brachycentridae

Amiocentrus aspilus (Ross)

0

0.0

9

3.0

13

3.8

Unassociated Amiocentrus

Females:

Amiocentrus sp. #1

0

0.0

95

20.1

54

15.7

Amiocentrus sp. #2

0

0.0

0

0.0

1

0.3

Amiocentrus sp. #3

0

0.0

0

0.0

8

2.3

Micrasema undescr. sp. A

0

0.0

123

25.8

38

11.1

88

PAN-PACIFIC ENTOMOLOGIST

Table 2, continued

1979

1980

1981

No,

Percent

No.

Percent

No.

Percent

of total

of total

of total

Lepidostomatidae

Lepidostoma cantha Ross

0

0.0

36

7.5

67

19.5

Lepidostomafischeri Denning

0

0.0

0

0.0

2

0.6

Lepidostoma strophis Ross

0

0.0

0

0.0

1

0.3

Unassociated Lepidostoma

Females:

Lepidostoma sp. #1

0

0.0

0

0.0

5

1.5

Lepidostoma sp. #2

0

0.0

0

0.0

1

0.3

Limnephilidae

Limnephilidae sp. A

0

0.0

1

0.2

0

0.0

Sericostomatidae

Gumaga nigricula (McL.)

1

0.7

2

0.4

2

0.6

Odontoceridae

Marilia flexuosa Ulmer

Nerophilus californicus

2

1.4

16

3.4

14

4.1

(Hagen)

0

0.0

1

0.2

0

0.0

Parthina vierra Denning

1

0.7

0

0.0

0

0.0

Helicopsychidae

Helicopsyche borealis

(Hagen)

17

11.8

2

0.4

1

0.3

Calamoceratidae

Heteroplectron

californicum McL.

1

0.7

2

0.4

2

0.6

Leptoceridae

Mystacides alafimbriata

HiU-Griffin

0

0.0

1

0.2

0

0.0

Oecetis sp. A

2

1.4

1

0.2

0

0.0

Total N

144

477

343

Total No. Taxa (Excluding

Unassociated Females)

18

30

27

No. Unassociated Females

18

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

89

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

N

Mean Temp.
(Oct.-April)

°C

SR

with BSC

SR

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

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

DF

Mean

Variance

(=AV)

Big Sulphur Creek

1979-1980

7

-0.35

0.104

Big Sulphur Creek

1980-1981

7

0.09

0.098

Big Sulphur Creek

1979-1981

7

-0.29

0.063

Big Canyon Creek

1980-1981

8

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.,

90

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.

Literature Cited

Brown, W. M. Ill, and L. E. Jackson, Jr. 1974. Sediment source and deposition sites and erosional and
depositional provinces, Marin and Sonoma counties, California. U.S. Geol. Surv. Publ. No.
MF-625,32pp.

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

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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

93

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;

94

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

95

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).

96

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

97

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,
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Vol. 63

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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

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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

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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

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THORP, R. W.—A new species of Andrena {Onagrandrena) from Utah’s San Rafael Desert

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Proceedings of the Pacific Coast Entomological Society, 1986 . 199

Corrigenda . 205

Publications received. 150

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THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is pubUshed quarterly (January, April, July,
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The Pan-Pacific Entomologist (ISSN 0031-0603)
PRODUCED BY A-R EDITIONS, MADISON, WISCONSIN 53703, U.S.A.

Dedicated

to

E. Gorton Linsley

E. Gorton Linsley

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

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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.

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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

no

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.

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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

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 but

with sparse and minute

moderately long hairs;

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

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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

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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

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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

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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

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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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Claude-Joseph, F. 1929. Le repos nocturne chez quelques Hymenopteres du Chili. Zool. Anz.,
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Friese, H. 1908. Die Apidae (Blumenwespen) von Argentina nach den Reisenergebnissen der Herren
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Friese, H. 1910. Neue Biennenarten aus Siid-Amerika. Deutsche Entom. Zeitschr., 1910:693-711.
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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
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Hurd, P. D., Jr. and J. S. Moure. 1963. A classification of the large carpenter bees (Xylocopini). Univ.
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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.
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Michener, C. D. 1944. Comparative external morphology, phylogeny, and a classification of the bees.
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155:67-240.

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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.

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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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6, Paris.

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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

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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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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

Alcock, J. 1977. Patrolling and mating by males of Callanthidium illustre. Southwest. Nat. 22:554-557.
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

129

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.
Hurd, P. D., Jr. 1979. Superfamily Apoidea, pp. 1741-2209. In\ P. D. Hurd, Jr., D. R. Smith and B. D.
Burks [eds.], Catalog of Hymenoptera in America North of Mexico. Washington, D.C.:
Smithsonian Institution Press.

Johnson, S. A. 1904. Notes and news, entomological gleanings from all quarters of the globe. Entomol.
News 15:284.

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.
Entomol. 78:227-231.

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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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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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.

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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

VOLUME 63, NUMBER 2

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).

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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.

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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

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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

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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

152

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

9

1.50

3:3=*

2

Iwata, 1955

Sphecodes japonica

—

4

0.67

3:3“

2

Iwata, 1955

Sphecodes sp. A

0.57

6

1.00

3:3

1

present study

Sphecodes sp. B

0.76

—

—

3:3

1

present study

Megachilidae

Dioxys pacificus

0.74

2

0.33

3:3

2

present study

Stelis sp.

0.61

2.67

0.44

3:3

3

present study

Euaspis basalis

0.53

3.5

0.58

3:3“'=

3

Iwata, 1955,1960

Coelioxys (Rhinocoelioxys) sp.

0.41

—

—

prob.3:3

1

present study

Coelioxys yanonis

—

2

0.33

3:3“

1

Iwata, 1955

Coelioxys fenestratus

0.54

18

3.00

3:3“

1

Iwata, 1955

Coelioxys brevis

0.40

5

0.83

3:3“

1

Iwata, 1955

Coelioxys sp. (138)

—

9

1.50

3:3“

1

Iwata, 1955

Coelioxys decipiens

—

4

0.67

prob. 3:3^*

1

Iwata, 1965

Anthophoridae

Anthophorinae

Thyreus japonicus

0.85

3.5

0.438

4:4

2

Iwata, 1955

Zacosmia maculata desertorum

0.74

6

0.75

4:4

1

present study

Mesoplia proh. rufipes

0.74

4

0.50

4:4

1

present study

Ericrocis lata

0.77

2

0.20

5:5

1

present study

Anthophoridae

Nomadinae

Neolarra (Neolarra) californica

0.33

13.75

1.325

5:5

3

present study

5:6

1

present study

Neolarra (Neolarra) sp.

—

—

—

5:5 (or 5:4)

1

present study

Neolarra (Phileremulus) vigi-

Ians

—

11

1.10

5:5

1

present study

Holcopasites calliopsidis

—

—

—

5:6

1

present study

Ammobates carinatus

0.47

6

0.50

6:6

1

present study

Oreopasites sp. A

0.49

7

0.74

5:4

1

present study

5:5 (or 6:4)

1

present study

Oreopasites vanduzeei

—

—

approx. 11 total

1

Rozen, 1986a

prob. 5:5

1

Rozen, 1986a

Kelita chilensis

0.30

7

0.70

prob. 5:5

1

Rozen,1986a

5:5

1

present study

Epeolus zonatus

0.53

8.5

0.85

5:5

2

present study

Epeolus scutellaris

0.76

6.33

0.45

7:7

4

present study

Epeolus sp.

0.56

4

0.40

5:5

1

present study

Epeolus japonicus

1.00

6.5

0.54

6:6

2

Iwata, 1955

Triepeolus pectoralis

0.62

5

0.50

5:5

1

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

2

0.20

5:5

1

present study

Centrias articulata

0.28

—

—

5:5

3

present study

Nomada banksi

0.30

15.5

1.55

5:5

3

present study

Nomada vicina

0.39

8

0.80

5:5

1

present study

Nomada illinoiensis

—

—

—

5:4

1

present study

5:5

1

present study

Nomada (“Gnathias”) sp.

—

—

—

5:5

1

present study

Nomada pyrifera

0.50

14.5

1.45

5:5

2

Iwata, 1955

Nomada sp. nr. glabella

0.38

8.67

0.90

5:5

2

Iwata, 1955

5:4

1

Iwata, 1955

Nomada japonica

0.39

21.125 2.15

5:5

6

Iwata, 1955

4:4

1

Iwata, 1955

6:5

1

Iwata, 1955

Apidae

Psithyrus barbutellus

—

—

—

10:10

1

Cumber, 1949

10:12

1

Cumber, 1949

6:7

1

Cumber, 1949

8:9

1

Cumber, 1949

Psithyrus bohemicus

—

—

—

14:13

1

Cumber, 1949

Psithyrus campestris

—

—

—

7:8

1

Cumber, 1949

6:7

1

Cumber, 1949

Psithryus rupestris

—

—

—

18:15

1

Cumber, 1949

15:13

2

Cumber, 1949

16:14

1

Cumber, 1949

12:14

1

Cumber, 1949

Psithyrus vestalis

—

—

—

10:10

1

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

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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 .

1

1

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

31

a) With Unchanged 1

Number of Ovarioles

k 5.62 V

0.90V

i 14

Us = 170

Us = 124

1

lz = 2.02 1

1 Z = 0.198

1

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

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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.

164

PAN-PACIFIC ENTOMOLOGIST

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

165

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PAN-PACIFIC ENTOMOLOGIST

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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PAN-PACIFIC ENTOMOLOGIST

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

VOLUME 63, NUMBER 2

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

VOLUME 63, NUMBER 2

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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173

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.

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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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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.

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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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PAN-PACIFIC ENTOMOLOGIST

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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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PAN-PACIFIC ENTOMOLOGIST

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.

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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

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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

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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

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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.

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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

196

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

VOLUME 63, NUMBER 2

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

200

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,

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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.

202

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
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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
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Typing.—Two copies of each manuscript must be submitted (original and one xerox copy or two xerox copies are suitable). All
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Underscore only where italics are intended in the body of the text. Number all pages consecutively and put authors name on each
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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¬
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Figure legends. —Legends should be typewritten double-spaced on separate pages headed EXPLANATION OF FIGURES and
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References. —All citations in text, e.g., Essig (1926) or (Essig 1958), must be listed alphabetically under LITERATURE CITED
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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
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Proofs, reprints, and abstracts. —Proofs and forms for the abstract and reprint order will be sent to authors. Changes in proof will
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Editing and administrative charges. —Papers by members of the Pacific Coast Entomological Society are charged at the rate of
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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.
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THE PAN-PACIFIC ENTOMOLOGIST.

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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¬
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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
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U.S. orders, or $1.70 for foreign orders). No members discount at this special
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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
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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¬
dressed to the Secretary, Mr. Vincent F. Lee, California Academy of Sciences, Golden
Gate Park, San Francisco, CA 94118-9961.

Manuscripts, proofs, and all correspondence concerning editorial matters should be
addressed to Editor, Pacific Coast Entomological Society, 201 Wellman Hall, Univer¬
sity of California, Berkeley, CA 94720. See back cover for instructions.

The annual dues, paid in advance, are $15.00 for regular members of the Society,
$7.50 for student members, $30.00 or more for sponsoring members, or $20.00 for sub¬
scription only. Members of the Society receive The Pan-Pacific Entomologist. Single
copies of recent numbers are $5.00 each or $20.00 a volume. See back cover for prices
of earlier back numbers. Make checks payable to the Pacific Coast Entomological Soci¬
ety.

Pacific Coast Entomological Society

OFFICERS FOR 1987

Stanley C. WiUiams, President Wojciech J. Pulawski, Treasurer

Alan I. Kaplan, President-Elect Vincent F. Lee, Secretary

THE PAN-PACIFIC ENTOMOLOGIST (ISSN 0031-0603) is published quarterly (January, April, July,
and October) for $20.00 per year by the Pacific Coast Entomological Society, California Academy of Sciences,
Golden Gate Park, San Francisco, California 94118-9961. Second-class postage paid at San Francisco, Califor¬
nia, and at additional mailing offices. POSTMASTER: Send address changes to THE PAN-PACIFIC EN¬
TOMOLOGIST, California Academy of Sciences, Golden Gate Park, San Francisco, California 94118-9961.

This issue mailed August 26,1987

The Pan-Pacific Entomologist (ISSN 0031-0603)
PRODUCED BY A-R EDITIONS, MADISON, WISCONSIN 53703, U.S.A.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 207-217

The Male Mating Strategy of the Bee Nomia nevadensis
(Hymenoptera: Halictidae):

Leg Structure and Mate Guarding

Kevin M. O’Neill and Louis Bjostad

Department of Entomology, Colorado State University, Fort Collins, CO 80523;
(KMO present address) Department of Entomology, Montana State University,
Bozeman, Montana 59717

Abstract. —Males of the bee Nomia nevadensis hakeri Cockerell congregate in
large numbers within the previous generation’s nesting area and mate with emerging
virgin females. Competition among males, much of which occurs below ground
before the female emerges, and the fact that females are receptive to only one male
upon emergence (confirmed experimentally) favors males that: a) search for
evidence (possibly odor) that a female is about to emerge at a specific location; b)
rendezvous with a female before she emerges from the ground; and c) upon finding a
female, use behavioral and morphological means to prevent takeover by other
males. Evidence is presented that males use flattened expansions of their hind tibia to
grip females firmly. The activities of males in the emergence area makes them
conspicuous targets for predation by robberflies (Asilidae).

Females of many species of solitary ground-nesting bees and wasps nest in dense
aggregations. Thus, adult virgin females emerging the following generation may
provide mate-searching males with a clumped source of receptive females. If these
females mate only once, or several times over long periods in their lives, there is a
selective advantage to traits that aid males to find mates as close as possible to the
time at which the females become receptive (Thornhill and Alcock, 1983). It may
even be beneficial for males to attempt to reach a female before she emerges from the
ground (e.g., Alcock et al, 1976; O’Neill and Evans, 1983; Schone and Tengo,
1981). However, in many situations, the high density of conspecific males makes it
difficult, even for those finding females early, to complete copulation without
interference from competitors. A variety of mechanisms have evolved to minimize
such interference including removing females from the vicinity of competitors or
guarding the female in a manner that prevents takeovers. In some insects, guarding
ability is enhanced by the presence of morphological structures that help the male
secure a hold upon the female (Thornhill and Alcock, 1983).

This paper reports on a study of the mating strategy of males of the bee Nomia
nevadensis hakeri Cockerell. Females of this and other species of Nomia construct
multicellular nests in extremely dense aggregations (Cross and Bohart, 1960;
Kerfoot, 1964; Johansen et al. , 1978), such that large numbers of males and females
emerge the following year within a restricted area. We studied a dense natural
aggregation of N. nevadensis bakeri in the summers of 1984 and 1985. Here we
present information on the mating strategy of males, the behavioral and

207

208

PAN-PACIFIC ENTOMOLOGIST

morphological adaptations that may help them avoid interference from competing
males, and the predation risk undertaken by mate-searching males. We also
conducted experiments to determine whether females are receptive to more than one
male upon emergence.

Materials and Methods

Nomia nevadensis bakeri was studied on five days between 28 July and 11 August,
1984 and on five days between 24 July and 2 August, 1985. The study site was located
beside a dirt road approximately 9 km northeast of Roggen, Weld County,
Colorado, U.S.A. The soil in the emergence area was sandy, with a surface crust
about 0.5 to 1.0 cm thick. The area from which female bees emerged measured
approximately 2 x 10 m in 1984 and 2 x 5 m in 1985, and contained sparse
vegetation, primarily sunflowers (Helianthus sp.) and scurfpea {Psoralea lanceolata
Pursh).

This aggregation was observed for a total of 24.3 hours over the ten days of study.
Focal observations were made on interactions of males with females and conspecific
males at emergence holes. Occurrences of all observed matings and predations upon
males were recorded and mating pairs were collected and preserved in order to
determine the size of mating males and emerging females. An estimate of the body
size distribution of males in the population was made from three sweep net samples
taken in the emergence area on two days in 1984. Head widths of both males and
females were determined to the nearest 0.1 mm with a VWR Scientific Products
micrometer accurate to 0.05 mm. In 1985, 53 males were marked on the thorax with
dots of enamel paint to facilitate later identification.

We preserved mating pairs in natural positions so that they could be returned to
the laboratory to examine the posturing and positioning of the males’ legs. In the
field, pairs were immersed and stored in hquid nitrogen (boiling point: -195.8°C);
they were examined under a dissecting microscope immediately upon removal from
the liquid. This technique was possible because, once a male grasped a female, we
could usually transfer them to the hquid nitrogen without causing them to separate.
Liquid nitrogen is commonly used for rapid freezing of biological specimens (Dawes,
1979).

We conducted an experiment to determine if females were receptive to the mating
attempts of more than one male upon emergence. For each manipulation, we
recorded whether the female mated with the male to which she was presented. In one
experimental group (29 females), each was allowed to complete copulation with the
male that had found her upon emergence. Following this, each female was presented
to a second male. To determine if manipulating females in this way affected their
receptivity, each of 15 females in a control group was separated, prior to copulation,
from the male that was mounted upon her as she emerged; each was then presented
to a second male. As a second experimental group, 12 of the females from the control
group were presented to a third male after the second interaction was complete. The
protocol for this experiment is outhned in Table 1.

Results

Searching and Mating Behavior. Males were active in the emergence area from
about 900 to 1300 hours when soil surface temperatures ranged between about 25°
and 45°C. At the peak of activity, hundreds of males swarmed over the emergence

VOLUME 63, NUMBER 3

209

Table 1. Experimental protocol and results of mating experiments. Females used in #3 were those used originally in
# 2 .

Female Response to Manipulation

Protocol

Female Refused

to Mate

Female

Copulated

1)

Copulation completed; female then
paired with another male.

29

0

2)

First copulation interrupted before
coupling; female then paired with
second male.

0

15

3)

Mated female from #2 paired with third
male.

12

0

area within 10 cm of the surface. Each male flew in an irregular pattern, occasionally
landing to investigate holes, 0.5 to 1.0 cm in diameter, in the surface of the sand. The
lack of a tumulus or depression around these holes indicates that they were
emergence holes, rather than active nest entrances (Kerfoot, 1964). No nesting
females were seen in the area during the course of the study. After landing near a
hole, a male usually stood facing the entrance or entered it to remain underground
for from several seconds to over a minute. As many as four males were observed
within in a single hole at a given time with others standing near the entrance, facing
the hole, when a female was about to emerge.

Initial pairing of males and females always occurred below the surface within the
tunnels before the female had emerged from the ground. Each female emerged from
a hole with a male mounted upon her, although they had not coupled genitalia at this
time. During the study, we saw males digging at the surface only three times.
Apparently, waiting males entered holes after the female broke through to the
surface. However, this is an inference, since we never observed the exact moment
when the emergence hole was opened. Usually within 60 seconds of emerging with
the female, if other males were not in contact with the pair, the mounted male moved
back along the female’s body, probed with his genitalia, and coupled. After coupling
genitalia, the male’s abdomen began to pulsate rhythmically and he often released
the female from his leg grasp. During this entire period, a receptive female remained
quiescent and did not attempt to break free from the male. This copulatory phase
lasted from 4 to 43 seconds (mean = 19.2; SD = 8.8; N = 42). The male then broke
genitalic contact and left or, more often (90% of 42 cases), moved forward on the
female’s body to his original position and remained for 2 to 72 seconds
(mean = 22.2; SD = 16.8; N = 38). Following this post-copulatory phase, the male
broke contact and the female immediately flew away from the emergence area.
Males often reentered the swarm following copulation. On one day in 1985, six males
that mated were marked after they had copulated. Five of these males were resighted
in the swarm within 5 to 29 minutes. Males were also observed within the emergence
area up to seven days after they were marked.

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PAN-PACIFIC ENTOMOLOGIST

All mating pairs were collected within several cm of an emergence hole. Males
were never observed flying and carrying a female away from the emergence area. A
total of 271 mating pairs were seen during the course of the study. Most of the
matings (N = 246; mean = 18.1/hour of observation) were observed in the first year
of the study. During the second year, activity was much lower probably due to heavy
rains before and during the seasonal period of activity, and only 25 matings were
observed (mean = 2.3/hour of observation).

Emerging females released a strong, “sweet” odor. If a female (mated or
unmated) was held in a pair of forceps or within the end of an insect net 5 to 10 cm
above the surface of the ground within several minutes of her emergence, males
always approached upwind in a rapid zig-zag flight and landed upon her, often
causing a struggle among males for the female. This procedure never failed to attract
males to recently emerged females (alive or dead; N = 66) or to males that had
recently been in contact with a live female. The entire sequence of copulation could
be initiated in this way if the female was a virgin (see next section).

On five occasions we observed males investigating (i.e. walking in tight circles and
antennating the soil surface) localized areas (several cm^), but not in the vicinity of a
hole. All five times we scraped away the surface soil, once to a depth of 5 cm, and
discovered a female who was releasing the strong scent. Apparently, males, possibly
by orienting to the “sweet” odor, can detect females that have not yet reached the
surface. By doing so, they could wait for the females at the exact point of emergence.
The presence of the surface crust may act to delay the female just below the surface
and increase the chance that she will be discovered by a male.

Female Receptivity in the Emergence Area. We were able to demonstrate that
females will mate with only one male between the time that they emerge and the time
they leave the emergence area. Females that had already copulated always refused to
mate with other males (Table 1, #1 & #3), although the latter made vigorous mating
attempts. On the other hand, females that were separated before copulation from
the first male mounted upon them were always receptive (Table 1, #2). The latter
data indicate that our technique was not responsible for non-receptivity of the
females used in the above experiments.

When a female was not receptive to mating attempts she had means of preventing
copulation, even though the male often had a secure hold upon her. The form of
these interactions, which sometimes lasted longer than 5 minutes, was distinctly
different from those that resulted in successful copulation. To refuse a mating
attempt, a female curled her abdomen forward until her genitalic region nearly
touched her head. This prevented the male from making genitalic contact. She also
attempted to pry herself loose from the male by pushing backwards and upwards at
him with her hind legs and by beating her wings if they were free. Typically, she
walked forward during these attempts. None of these behaviors were seen in
interactions that resulted in successful copulation. Eventually, unreceptive females
were able to break free and leave the emergence area without further interference
from males. Outside of our experiments, we observed one instance of a female
refusing to mate with a male in the emergence area.

Interactions among Males. There is potential for competition among males both
below and above ground. We have noted that a number of males may enter an
emergence hole. By peering into the emergence holes, we often observed what were
apparently intense struggles for females, though we were unable to record the

VOLUME 63, NUMBER 3

211

duration of such interactions or determine whether they involved interference or
scramble competition.

Above ground, after the female emerged with a male mounted upon her back, one
to four males were frequently in contact with the pair during the pre-copulatory
phase, struggling to gain access to the female. The males were mounted either
dorsally on the first male’s back or ventrally beneath the female. Generally, the
posture and behavior, particularly of the dorsally mounted males, was similar to that
of a male mounted upon a female. Some of these interactions may simply have been
mistaken attempts by males to mount females. The pervasive odor of the female
could have been responsible for a failure of males to discriminate between the sexes.
We also observed twelve cases of isolated “homosexual” pairs (i.e. males mounted
upon other males). Odor has been implicated as the trigger for inappropriate sexual
mounts in other insect species in both natural (Tomkins et al., 1980; Tengo, 1979)
and experimental (Shimron and Hefetz, 1985) situations.

We detected no size-biased mating success among males. Males sampled from
copulating pairs (mean head width = 3.01 mm; S.D. = 0.07; N = 89) were not
significantly different in size from males in sweep net samples taken in the emergence
area (mean = 3.01 mm; S.D. = 0.08; N = 252; t-test, 1339 = 0). On the other hand,
females taken in the emergence area (mean = 3.19;S.D. = 0.09;N = 66 ) were not
only significantly larger than copulating males on average (t-test, ti 53 — 14.2;
p < 0.001), but in each mating pair were either larger than (95%) or equal to (5%)
the male in size (N = 43). There were no females present among the 252 bees taken
in the sweep samples, giving further confirmation that they do not remain in the
emergence area after mating (although they must return later to nest).

Morphological Modifications for Mate Guarding. The structure and exact
placement of the hind legs may combine to prevent the mounted male from being
supplanted by others during the pre-copulatory phase. The male’s head was just
behind that of the female’s, with his front and middle legs usually held over her
wings, preventing them from moving. His hind legs were wrapped around her
abdomen just posterior to her petiole, usually between the first and second gastral
segments. Above ground at least, males were never supplanted by others, when
positioned in this manner. In addition, when an unreceptive female was presented to
a male, it was difficult for her to break free, although the male could not induce her to
mate. In hundreds of observed interactions, a pair was disrupted during the
post-copulatory phase only twice, and during the pre-copulatory phase only once. In
the latter case, the male did not have his legs properly situated under the female’s
abdomen; she mated with the usurping male.

By immersing pairs that were in the pre-copulatory position into a Dewar flask
containing liquid nitrogen, we were able to return five pairs from the field intact, for
closer examination. Only the right side of each pair was examined, since
manipulation caused the pair to separate slightly.

A structure on the males’ hindlegs that may help them grasp a female was evident:
the tibia is expanded distally to form a pair of flattened triangular flanges (Fig. 1) that
lie flat against the female’s abdomen, pointing anteriorly, when a male has his legs
wrapped around a female. The positioning of the flanges on the males’ tibiae suggests
that they locked onto the female by sliding the larger one beneath the posterior edge
of a sclerite beneath her abdomen (Fig. 2). On one female the larger flange was
almost completely inserted beneath the ventral posterior edge of gastral tergum I

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PAN-PACIFIC ENTOMOLOGIST

Figure 1. A) Scanning electron micrograph of the left hindleg of male of N. nevadensis baked, (20X;
side of tibia shown is that which lies against the female when the male is mounted in the pre-copulatory
posture); B) Close-up view (SOX) of right hind tibia (opposite aspect from A) of male showing expansions
(arrows); F = femur; T = tibia.

VOLUME 63, NUMBER 3

213

Anterior

Figure 2. Schematic representation of the venter of the abdomen of a female N. nevadensis bakeri,
showing where the large flange of the male hind tibia was positioned on five pairs examined. One each
found in position (a) and (c) (stippled area denotes portion inserted under female’s sclerite); three found
in position (b) (black area denotes portion inserted).

where the latter curls beneath the abdomen (Fig. 2,a). The tip of the flange was so
inserted when examined on three others (Fig. 2,b), although it may have pulled out
slightly in the process of the examination. On the fifth female, approximately half of
the flange was inserted beneath the posterior edge of sternum II (Fig. 2,c). The tarsal
segments of the males’ legs were lying back along the medial line of the female’s
venter.

Predation upon Males. Males were sometimes preyed upon by robberflies of the
species Diogmites angustipennis Loew. These flies, which reach up to 22 mm in
length (Lavigne and Holland, 1969), were apparently attracted by the conspicuous
movements of male bees flying about the emergence area. Twenty-three successful

214

PAN-PACIFIC ENTOMOLOGIST

predations were observed, the majority (21) of which occurred on the one day in
which the flies were most abundant.

Discussion

Females of Nomia nevadensis bakeri in this population were receptive upon
emergence, mated with only one male while in the emergence area, were apparently
detectable prior to emergence, possibly because of the odor they emit, and emerged
within an area containing many conspeciflc males. It is also probable that some
degree of protandry occurs in this population, as it does in other species of Nomia
(Kerfoot, 1964; Cross and Bohart, 1960; Johansen etal., 1978). The combination of
these factors creates a competitive situation favoring males that reach and mate with
an emerging female before conspeciflc males do. To accomplish this, males in this
population: 1) search for evidence of females about to emerge, 2) attempt to
rendezvous with females before they emerge from the ground, and 3) upon finding a
female, use behavioral and morphological means to prevent other males from
usurping their position. Males were rarely seen digging where a female was about to
emerge. This contrasts with other species of Hymenoptera with similar mating
systems (e.g., Alcock et al., 1976; O’Neill and Evans, 1983; Schone and Tengo,
1981), where males invest much time and energy digging for pre-emergent females.

Male insects competing for females within a crowded emergence area use a variety
of means to prevent interference form conspeciflc males (Thornhill and Alcock,
1983). Males of some species of bees and wasps avoid takeover by competitors by
carrying the female away in flight from the emergence area before mating with her
(Alcock etal., 1976; O’Neill and Evans, 1983). However, it would have been difficult
for males of N. nevadensis in this population to do this, since they were never larger
than the female and would probably have had difficulty carrying her in flight. The
male is usually larger than the female in insects in which the male carries the female
during courtship or mating (O’Neill, 1985). In contrast to other species of
Hymenoptera in which larger size has been shown to aid males in their attempts to
obtain matings (e.g., Alcock e? a/., 1976; O’Neill, 1983a,b; O’Neill and Evans, 1983;
Severinghaus et al., 1981), no such effect was detected in our analysis. Males of N.
nevadensis show much less range in size than species of digger wasps (O’Neill, 1985)
and bees (Alcock, et al., 1976) for which a size advantage among competing
conspecifics has been shown, although the reason for this is unclear. We can
speculate on one factor selecting against large size in this species: males that are too
large may be unable to maneuver for position on a female within an emergence hole.
However, this hypothesis would be difficult to test.

Rather than leave the emergence area with the female, males of N. nevadensis
remained. Those males grasping females with the aid of modified leg structures were
highly successful at maintaining contact with a female and completing copulation. It
could also be hypothesized that these leg structures function to subdue the female so
that mating can take place, much as male scorpionflies use their notal organ to force
copulation upon females (Thornhill, 1980). However, this seems unlikely for several
reasons. First, virgin females were generally quiescent during mating attempts, so
need not have been subdued. Second, as demonstrated in the mating experiments,
previously mated females were capable of refusing to copulate and could break free
from the males. Therefore, it seems more likely that the behavior and leg structure of

VOLUME 63, NUMBER 3

215

males have evolved as traits that prevent interference from conspecific males, thus
assuring sole access to a receptive female.

Males of a variety of insect species utilize leg modifications to maintain a grip upon
females during courtship and mating. Males of other species of the subgenus to which
N. nevadensis belongs {Epinomia) also have flattened expansions of their hind tibia
(Cross, 1958). Males of some species of the subgenus Acunomia possess even larger
expansions of the hind tibia that are not flattened like they are in N. nevadensis and
are sometimes larger than the main part of the tibia itself (Ribble, 1965); this
indicates that these tibial modifications may be utilized in a slightly different manner
than they are in N. nevadensis, if indeed they are used at all during courtship and
mating in these other species. The hind femora are also enlarged in some species of
Nomia. We have not been able to find reference to the potential use of these leg
structures in other species of Nomia. Males of the melittid bee Meganomia binghami
Cockerell use enlargements of the hindlegs to assist in maintaining a grip upon
females in the presence of up to ten conspecific males (Rozen, 1977; Stage, 1971).
Male bees of some species of the genus Agapostemon (Halictidae) have similar
modifications of the hindlegs, although their function is unknown (Roberts, 1969,
1972). The unmodified structure of the legs of males of most species of bees contrasts
markedly with that of males of Nomia, Agapostemon, and Meganomia and with
females of most non-parasitic species (Stephen et al., 1969). Given the
morphological specialization associated with foraging evident in the hind legs of
female bees (e.g., Roberts and Vallespir, 1978), leg structure in bees appears to have
maintained a high degree of evolutionary plasticity.

Some species of beetles (Crowson, 1981) and flies (Spieth, 1952) also have leg
structures that provide a firm grasp upon mates. For example, in the fly Sepsis
cynipsea (L.), the front legs of a male are structured so that they act as a clamp
around the female’s wing bases. In this species, mating occurs at cattle droppings
where up to 500 males may be present (Parker, 1972). Males of a variety of insects
possess modifications of wings, genitalia, jaws, and antennae that are used to grip
females (Thornhill and Alcock, 1983; Rothschild and Hinton, 1968).

Along with our observations on Nomia, the comparative information suggests that
sexual selection has promoted the independent evolution of a variety of
morphological structures that enhance the mating success of individual males
through an ability to grasp females securely in the presence of competitors. The
ability to manipulate interactions among male and female Nomia and, potentially,
the leg structure of males, may provide a good system for experimental and
comparative studies of the function and efficiency of mate guarding. Our brief study
also leaves open questions concerning the below ground competition for females, the
function of the male’s remaining with the female for a short time after insemination,
and cues used by males to locate pre-emergent females.

Acknowledgments

We thank Ruth O’Neill and Howard Evans for assistance in the field and for
comments on the manuscript. John Alcock, Byron Alexander, Robert Bartelt, J.
Wayne Brewer, George Eickwort, Wendell Morrill, and David Reduker also
reviewed the manuscript. Howard E. Evans identified the bees and robberfly.
Scanning electron micrographs were taken by J. A. Blixt. This work was carried out

216

PAN-PACIFIC ENTOMOLOGIST

with support from the Colorado Agricultural Experiment Station (grant #1-53971)
and the Montana Agricultural Experiment Station (MONB-155).

Literature Cited

Alcock, J., C. E. Jones and S. L. Buchmann. 1977. Male mating strategies in the bee Centrispallida Fox
(Anthophoridae: Hymenoptera). Amer. Nat. 977:145-155.

Cross, E. A. 1958. A revision of the bees of the subgenus Epinomia in the New World (Hymenoptera:
Halictidae). Univ. Kans. Sci. Bull. 38:1261-1301.

Cross, E. A. and G. E. Bohart. 1960. The biology otNomia {Epinomia) triangulifera with comparative
notes on other species ofNomia. Univ. Kans. Sci. Bull. 41:761-792.

Crowson, R. A. 1981. The Biology of the Coleoptera. Academic Press, New York.

Dawes, C. J. 1979. Biological techniques for transmission and scanning electron microscopy.

Cryotechniques, Chapter 11. Ladd Research Industries, Burhngton, Vermont.

Johansen, C. A., D. F. Mayer, and J. D. Eves. 1978. Biology and management of the alkali bee, Nomia
melanderi Cockerell (Hymenoptera: Halictidae). Melanderia 28:25-33.

Kerfoot, W. B. 1964. Observations on the nests of Nomia nevadensis bakeri with comparative notes on
Nomia nevadensis arizonensis (Hymenoptera: Halictidae). J. Kans. Entomol. Soc. 37:152-157.
Lavigne, R. J. and F. R. Holland. 1969. Comparative behavior of eleven species of Wyoming robber flies
(Diptera: Asilidae). Univ. Wyoming Ag. Expt. Sta. Science Monograph 18:1-61.

O’Neill, K. M. 1983a. The significance of body size in territorial interactions of male beewolves
(Hymenoptera: Sphecidae, Philanthus). Anim. Behav. 31:404-411.

O’Neill, K. M. 1983b. Territoriality, body size, and spacing in males of the beewolf Philanthus basilaris
(Hymenoptera: Sphecidae). Behaviour 86:295-321.

O’Neill, K. M. 1985. Egg size, prey size, and sexual size dimorphism in digger wasps (Hymenoptera:
Sphecidae). Can. J. Zool. 63: 2187-2193.

O’Neill, K. M. and H. E. Evans. 1983. Alternative mating tactics in Bembecinus quinquespinosus
(Hymenoptera: Sphecidae): correlations with size and color variation. Behav. Ecol. Sociobiol.
14:39-46.

Parker, G. A. 1972. Reproductive behaviour of Sepsis cynipsea (Diptera: Sepsidae). 1. A preliminary
analysis of the reproductive strategy and its associated behaviour patterns. Behaviour 41:172-206.
nibble, D. W. 1965. A revision of the banded subgenera of Nomia in America (Hymenoptera:
Halictidae). Univ. Kans. Sci. Bull. 45:277-359.

Roberts, R. B. 1969. Biology of the bee genus Agapostemon (Hymenoptera: Halictidae). Univ. Kans.
Sci. BuU. 48:689-719.

Roberts, R. B. 1972. Revision of the bee genus Agapostemon (Hymenoptera: Hahctidae). Univ. Kans.
Sci. Bull. 49:437-590.

Roberts, R. B. andS. R. Vallespir. 1978. Specializationsof hairs bearing pollen and oil on the legs of bees
(Apoidea: Hymenoptera). Ann. Entomol. Soc. Amer. 71:619-627.

Rothschild, M. and H. E. Hinton. 1968. Holding organs on the antennae of male fleas. Proc. R. Ent. Soc.
Lond. 43:105-107.

Rozen, J. G. 1977. Biology and immature stages of the bee genus Meganomia (Hymenoptera:
Melittidae). Amer. Mus. Nov. 2630:1-14.

Schone, H. and J. Tengo. 1981. Competition of males, courtship behaviour and chemical communication
in the digger wasp Bembix rostrata (Hymenoptera: Sphecidae). Behaviour 77:44-66.
Severinghaus, L. L., B. H. Kurtak, and G. C. Eickwort. 1981. The reproductive behavior otAnthidium
manicatum (Hymenoptera: Megachihdae) and the significance of size for territorial males. Behav.
Ecol. Sociobiol. 9:51-58.

Shimron, O. and A. Hefetz. 1985. Mating behavior and sex attraction of Eucera palestinae Friese
(Hymenoptera: Anthophoridae). J. Kans. Ent. Soc. 58:526-531.

Spieth, H. T. 1952. Mating behavior within the g&nus Drosophila. Bull. Am. Mus. Nat. Hist. 99:395-474.
Stage, G. I. 1971. Family placement of the African genus Meganomia Cockerell with a review of the
included species (Hymenoptera: Apoidea). Proc. Enf. Soc. Wash. 73:306-313.

Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The Biology and External Morphology of Bees.
Oregon State University Ag. Expt. Sta., Corvallis, Oregon.

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Tengo, J. 1979. Odour-released behaviour in Andrena male bees (Apoidea: Hymenoptera). Zoon.
7:15-45.

Thornhill, R. 1980. Rape in Panorpa and a general rape hypothesis. Anim. Behav. 28:52-59.

Thornhill, R. and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard Univ. Press,
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Tompkins, L., J. C. Hall, and L. M. Hall. 1980. Courtship-stimulating volatile compounds from normal
and mutant Drosophila. J. Ins. Physiol. 26:689-697.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 218-223

Phenology of Douglas-Fir Tussock Moth, Orgyia pseudotsugata, Egg
Eclosion and Mortality in a Thinned and Unthinned Stand

(Lepidoptera: Lymantriidae)

Boyd E. Wickman and Torolf R. Torgersen

Forestry and Range Sciences Laboratory, Pacific Northwest Research Station,
Forest Service, U.S. Department of Agriculture, La Grande, Oregon 97850

Abstract. —Heat-unit accumulation and egg eclosion were monitored in unthinned
and thinned white fir stands in southern Oregon. Degree-day accumulation and egg
eclosion on the unthinned site were 7 to 10 days behind development in the thinned
site. Parasitism and predation were higher on egg masses in the thinned stand.

The ability to predict phenological development of insect-host systems has
important applications for pest management as well as for research on population
biology. Studies on phenology of the Douglas-fir tussock moth, Orgyia
pseudotsugata (McDunnough), in California and Oregon (Wickman 1976a, 1976b,
1981) show egg eclosion is synchronous with host budburst. During population
sampling for early instars, we observed a spread in developmental stages from 1st to
3rd instars on some sites. Phenological data suggest that egg eclosion on a given site is
related to the amount of solar radiation reaching an egg mass; eclosion usually occurs
first in the exposed tops of large trees and last in shaded areas under full forest canopy
(Wickman 1976b).

In the past, we compared tussock moth egg masses only from exposed trees but at
different elevations (Wickman 1976a, 1976b, 1981). Egg eclosion was assumed to be
early on exposed trees, and later in shaded habitats. In this study, our object was to
compare tussock moth eclosion and egg mortality in two different environments at
the same elevation—a thinned (exposed) and an adjacent unthinned (shaded) stand.

The study was limited by available sites, egg masses, and instrumentation. These
preliminary data are reported to encourage more definitive studies on the effects of
silvicultural prescriptions on insect phenology and mortality.

Methods

The study site was near Mare’s Egg Spring, 10 km southwest of Fort Klamath, in
south-central Oregon. The site, with an eastern exposure, is at 1350 m elevation and
on a level bench about 100 m above the upper Klamath basin. The site was partly on a
population-study plot used for phenological monitoring of tussock moth from
1976-80 (Wickman 1981).

One exposed and one shaded habitat were compared. The exposed habitat was in a
thinned stand of second-growth white fir {Abies concolor (Gordon and Glend.)
Lindley ex Hildebr.) with widely spaced saplings and pole-sized trees and a few
overstory ponderosa pine {Pinus ponderosa Douglas ex Lawson). The shaded
habitat, about 100 m to the east, was a dense, unthinned, second-growth white fir
stand of pole-sized trees with a closed canopy.

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219

Temperature was recorded in the thinned stand by a 31-day battery-powered,
weather-sheltered hygrothermograph from November 1976 to November 1977.
From these records, Fahrenheit degree-day (°D) accumulations were calculated for
the exposed environment. The heat-unit threshold for development selected was
5.6°C (42°F) Wickman 1976a). When budburst began on May 31 in the area, a
weather-sheltered, 7-day hygrothermograph was installed in the unthinned stand;
monitoring continued until July 5. The °D accumulations for the two stands were
compared from June 1 to July 5.

Because natural populations of Douglas-fir tussock moth in the area were low and
egg masses difficult to find, we used egg masses reared in the laboratory from larvae
collected locally in June 1976. The egg-mass stock, which was also used for other
population studies (Torgersen and Ryan 1981), showed normal fecunditv and
development. The egg masses ranged from 100 to 250 eggs (x = 150).

To acclimate the egg masses, they were randomly divided into two batches, placed
in fine-mesh nylon bags, and hung in both stands on February 9. On April 5, egg
masses were individually wired to branches (Torgersen and Mason 1979). One egg
mass was wired at about 2 m from the ground at the four cardinal directions on four
trees, for a total of 16 egg masses in each stand. In the thinned stand, the masses were
wired to the underside of live, lower crown foliage. Because not much foliage was
present at 2 m in the unthinned stand, the masses were placed on small, foliated
adventitious branches on the main trunk at that height. Egg masses occur naturally in
such locations, as well as on large foliated branches. A 30- x 60-cm fine-mesh nylon
bag was placed over each stocked branch to prevent predation, although the mesh
did not keep out parasites. Bags were left in place until June 6. Masses were checked
every 2 days from June 6-12, and then daily through July 5. Each morning, egg
eclosion was recorded and new larvae counted and removed. On July 5, the egg
masses were removed for further rearing in the laboratory at 20°C. Each egg mass
was dissected when eclosion was complete and parasites ceased to emerge. Emerging
adult parasites were counted daily, and egg numbers were recorded. Unbroken eggs
from which neither tussock moth larvae nor parasites emerged were recorded as
“unknown mortality” (Torgersen and Mason 1979).

Results and Discussion

A steady divergence of °D accumulation is evident between the thinned and
unthinned stands (Fig. 1). On June 20 when eclosion began in the thinned stand, °D
reached 432 (from June 1), and 282 °D had accumulated in the unthinned stand—a
difference of 150 °D. By July 5, thermal development on the two sites had diverged
by 285 °D. These °D differences between thinned and unthinned stands at the same
elevation were greater than those found between two exposed sites with a 100 m
difference in elevation (Wickman 1981). Hopkins’ (1918) “bioclimatic law” states
that natural development is delayed 4 days for each 400 ft (122 m) rise in elevation at
a given latitude. Thus, local variation in °D development resulting from thinning may
surpass elevational differences of as much as 100 m.

The mean °D accumulation in the unthinned stand from June 1 to 20 was 14
°D/day. Therefore, °D accumulation in the thinned stand was about 11 days ahead of
the unthinned stand. Egg masses in the unthinned stand were exposed to predation
for nearly 2 weeks longer than eggs in the thinned stand.

Egg eclosion differed on the two sites (Fig. 1), consistent with the previous finding

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June July

Figure 1. Accumulated degree-days in thinned and unthinned stands in south-central Oregon, 1977.

that hatch of tussock moth eggs is influenced by exposure to solar radiation
(Wickman 1976a). Egg eclosion began on June 20 in the thinned stand and was 99
percent complete on July 5. Eclosion did not begin in the unthinned stand until June
27 and was only 61 percent complete on July 5. Eclosion from eggs in the unthinned
stand continued for 10 days in the laboratory.

The mesh bags around the egg masses did not prevent heavy attack by the egg
parasite Telenomus californicus Ashmead in the early spring. In another study,

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Table 1. Douglas-fir tussock moth egg eclosion, Telenomus californicus emergence, and unknown mortality in a
thinned and unthinned stand, south-central Oregon, 1977.

Eggs

per

mass

Douglas-fir
tussock moth

eclosion

Telenomus

californicus

emergence

Unknown

mortality

Parasitism

Unknown

mortality

Number

Percent

Thinned

Total

298

943

392

X

101.9

61.9

15.9

S.D.

47.83

22.06

22.16

Unthinned

Total

781

1072

362

x"

138.4

48.9

17.8

S.D.

52.75

14.75

17.44

females of this tiny parasite were first observed on egg masses at nearby Mare’s Egg
Spring plots on April 1, and parasite oviposition was essentially completed by April
20 (Torgersen and Ryan 1981).

In our study, egg masses were also heavily parasitized, averaging 61.9 percent in
the thinned stand and 48.9 percent in the unthinned stand (Table 1). Parasites began
to emerge from egg masses in the unthinned stand 6 °D later, lagging behind the
thinned stand; emergence ceased on the same day in both stands, however (Fig. 2).
Emergence of T. californicus can be roughly predicted from heat-unit accumulation.
In the combined field and laboratory rearing, emergence was completed at 1,635 °D,
accumulated from April 1 to July 24. This included 468 °D accumulated in the
laboratory—26 °D daily for 18 days.

Higher parasitism in the thinned stand (Table 1) suggests either that the parasites
do more searching in exposed tree crowns than in closed canopies or that some other
environmental condition draws them there.

The egg masses were exposed to predators from June 6 to July 5. Torgersen and
Mason (1979, 1985) found that birds and ants can partially destroy or completely
remove egg masses, and about half the eggs can be lost from predation by birds
alone. They also found significantly greater egg parasitism on xeric than on mesic
plots in California, Oregon, and Idaho.

In our study, egg masses appeared to be about the same size when we placed them
in the field. On July 5, masses from the thinned stand had one-third fewer eggs than
those from the unthinned stand and looked broken and ragged, which is typical of
predation.

A t-test showed that the differences in egg survival, after parasitism and apparent
predation between thinned and unthinned stands, were significant (p = < 0.05). The
difference between the two sites in unknown egg mortality was not significant,
however. As expected, egg viability was similar (p = > 0.05) because the egg masses
came from the same natural population.

Thinning increases the exposure of host trees to solar radiation and affects °D
accumulation, which speeds tussock moth egg development and eclosion. Thus,

222

PAN-PACIFIC ENTOMOLOGIST

Figure 2. Accumulated percentage emergence of Telenomus californicus, from Douglas-fir tussock
moth egg masses in thinned and unthinned stands in south-central Oregon, 1977.

some survival advantage could be gained by egg deposition in open stands, where
eggs might be exposed to predation for 14 fewer days. Thinning also apparently
enhances predation by birds and ants and parasitism by T, californicus. These limited
data leave unclear whether the advantages of earlier egg eclosion and reduced length
of exposure to predation can be offset by greater intensity of predation and higher
parasitism. More study is needed to measure precisely the effects of thinning on the
population dynamics of the tussock moth and other defoliating insects.

Acknowledgments

This study was funded in part by the USDA Expanded Douglas-fir Tussock Moth
Research and Development Program, Portland, Oregon. We thank Sherry Gollob
and Lee Roe for their assistance in the field and laboratory.

Literature Cited

Hopkins, A. D. 1918. Periodic events and natural laws as guides to agricultural research and practice.
USDA Mon. Weather Rev. Suppl. 9:42.

Torgersen, T. R., and R. R. Mason. 1979. Predation and parasitization of Douglas-fir tussock moth egg
masses. USDA Handb. 549.11 pp.

VOLUME 63 , NUMBER 3

223

Torgersen, T. R., and R. B. Ryan. 1981. Field biology of Telenomus californicus Ashmead, an important
egg parasite of Douglas-fir tussock moth. Ann. Entomol. Soc. Am. 79:185-186.

Torgersen, T. R., and R. R. Mason. 1985. Characteristics of egg parasitization of Douglas-fir tussock
moth, Orgyia pseudotsugata (McD.) (Lepidoptera: Lymantriidae), by Telenomus californicus
Ash. (Hymenoptera: Scelionidae). Environ. Entomol. 14:323-328.

Wickman, B. E. 1976a. Phenology of white fir and Douglas-fir tussock moth egg hatch and larval
development in California. Environ. Entomol. 5:316-320.

Wickman, B. E. 1976b. Douglas-fir tussock moth egg hatch and larval development in relation to
phenology of grand fir and Douglas-fir in northeastern Oregon. USDA For. Serv. Res. Pap.
PNW-206. 13 pp.

Wickman, B. E. 1981. Degree-day accumulation related to the phenology of Douglas-fir tussock moth
and white fir over five seasons of monitoring in southern Oregon. USDA For. Serv. Res. Note
PNW-392. 13 pp.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 224-235

A Morphometric Analysis of Mordellistena Costa in the
Southwestern United States (Coleoptera: Mordellidae)

Jeffrey C. Burne

University of Wyoming, Laramie, Wyoming

Abstract. —Beetles of the genus Mordellistena Costa in the southwestern United
States are submitted to a morphometric analysis in order to obtain more reliable
taxonomic characters and to establish the foundations for construction of a
phytogeny. Thirty external characters were chosen for measurement in a preliminary
statistical analysis of four well-defined species groups. These results were submitted
to a discriminant analysis which reduced the number of significant characters to 14.
Over 500 specimens were then measured for these characters, and the results then
submitted to a cluster analysis. The results of the analysis are discussed as to their
taxonomic and classificatory possibilities. Accomplishments of the study include the
generation of a list of the species of Mordellistena for Arizona, the discovery of
several undescribed species, a more confident method of identification, and the
foundation of a phytogeny for the genus. Shortcomings of the analysis were the
failure to construct species specific groups and inability to group 100%.

Beetles of the genus Mordellistena Costa have long been a problem to taxonomists
in that they are difficult to identify because the characters used to separate species are
indecisive. Despite the extensive work of Liljeblad (1945), the use of leg ridges as the
major identification characters has remained unsatisfactory. A morphometric
analysis of the genus Mordellistena was undertaken in the hope of attaining 1) more
reliable characters, 2) an easier and more confident taxonomy, and 3) the basis for a
phytogeny.

Since Liljeblad’s (1945) monograph on the entire family, little work has been done
on New World Mordellidae. Ray (1944, 1946, and 1947) dealt with neotropical
species and Khalaf (1970 and 1971) did some studies on wing venation of
southeastern species. Recently, much work has been done in Europe and Asia on the
Mordellidae, but there has been a complete lack of work on members of the
Mordellidae of the southwestern United States. Liljeblad (1945) lists 22 species of
Mordellistena from the southwest, the majority from California only. Individuals of
the genus from the southwest (New Mexico, Arizona, southern California, and
western Texas) are even more uniform than the mordellids in general, being
relatively slender and largely of a unicolorous brown or black body color. Because of
this uniformity and in consideration of the past difficulties with Mordellidae in
general, it was thought that a morphometric analysis would prove especially useful in
a study of the group. While the immediate goal of the study was the discovery of
reliable taxonomic characters and the generation of a list of the species of
Mordellistena present in Arizona, the long term goal is to use such an analysis for a
complete revision of the North American Mordellistena.

224

VOLUME 63, NUMBER 3

225

Materials and Methods

The University of Arizona insect collection contains over 1,500 specimens of
Mordellistena from a variety of localities throughout the Southwest. Along with
approximately 200 specimens borrowed from the Arizona State University
collection, a large base was present for a morphometric analysis of the Southwestern
species.

All of the approximately 1,700 specimens were first tagged with individual
identification numbers. Next came the selection of morphologic characters to be
measured. This was based primarily on which characters were consistently
observable in the genus. This selection resulted in the choice of 30 external
morphological characters which are listed in Table 1 and illustrated in Figures 1-4.

Following character selection, four groups of specimens, all identified to species
through the use of Liljeblad’s key (1945), all very distinct in appearance, and all
represented by a large series from a single collection locality were selected. The four
selected were: M. scapularis (Say), M. nunenmacheri\A\]Qh\did, M. sericans'PdM, and
M. tosta LeConte. It was on these groups that the preliminary set of 32 characters was
tested in order to determine their validity for taxonomic identification.

Measurements for the characters were accomplished using a Lasico Auto-Scaler
device attached to the right ocular tube of a Wild M-5 stereomicroscope.
Adjustments were made which allowed results to be recorded in millimeters in rapid
fashion.

Following measurement of specimens in the four groups for the 30 characters, a
discriminant analysis (SAS program) was run on the results. From these data were
chosen only those morphological characters which separated all of the specimens
back into their four original species groups. Only raw measurements were utilized in
the study due to the problems associated with the use of ratios (Blacklith and
Reyment, 1971). A series of 80 specimens were measured and 14 of the characters
accomplished the desired separation. These 14 characters are listed in Table 2.

Next came the measurement of as many specimens as possible for the 14 characters
with subsequent analysis of the results. In doing this, it was assumed that no
additional character variation was present. This was necessary in order to establish
some starting point for the cluster analysis. Thus the discriminant analysis was
utilized in order to select only those characters useful in taxonomic separation. Prior
to selection of specimens for measurement, all 1,700 Mordellistena were grouped
according to locality. From each resulting locality group, a random selection of
specimens, up to a maximum of 25, was made. For localities in which there were
fewer than 25 examples, all members from that locality were measured. In this way, a
total of 650 beetles were selected and then measured. Results of the measurements
were then transposed to key-punch computer cards with one card per beetle. The
identification number (i.e. label) of each insect and results of measurement of each
character on that insect were thus present on cards and ready for analysis. This
analysis was accomplished through the use of BMDP Computer Program P2M
written by Lazio Engelman (1979).

The BMDP P2M clustering program is written so that each case (beetle) is read
and considered as a single cluster to begin with. By then comparing the results of each
case for each variable (morphological character), the process of grouping is initiated
with each case placed into ever-enlarging groups until ultimately all cases are united

226

PAN-PACIFIC ENTOMOLOGIST

Table 1.

Original 30 External Morphological Characters

1. length of ultimate abdominal sternite (a)

2. length of antenna (b)

3. combined length of 3rd and 4th antennal segments (c)

4. length of 5th antennal segment (d)

5. length of metatrochanter (e)

6. length of metepisternum (f)

7. eye height (g)

8. eye length (g)

9. ocular width (h)

10. length of metafemora (i)

11. length of large metatibial spur (j)

12. length of small metatibial spur (k)

13. height of mandible (1)

14. width of mandible (1)

15. length of metatarsal claw (m)

16. length of head capsule (n)

17. length of total body

18. length of ultimate tergite (o)

19. width of metasternum (p)

20. width of pronotum (q)

21. length of 1st metatibial ridge (r)

22. length of 2nd metatibial ridge (r)

23. length of 3rd metatibial ridge (if present) (r)

24. width of clypeus (s)

25. length of metatarsus (t)

26. length of mesofermora (u)

27. length of mesotibia (v)

28. length of mesotarsus (w)

29. length of ultimate segment of maxillary palp (x)

30. width of ultimate segment of maxillary palp (x)

in a single cluster. As might be expected, there are limits to the number of cases that
can be analyzed in a program of this type, dependent on the number of variables
utilized. Indeed, it was found that the needed computer capacity for 650 specimens
and 14 characters exceeded the core memory of the University of Arizona CYBER.
For this reason, a reduction in the number of cases was necessary. This was done
through the elimination of cases from localities with larger groups in that all localities
with 25 specimens were reduced to 20 until the total sample set numbered 400. This
was the maximum number of cases that the computer was able to handle with 14
variables. In this way, the number of cases was reduced with no sacrifice of samples
from localities with few representatives.

VOLUME 63, NUMBER 3

227

Figure 1. Scanning electron microscope photograph, lateral view of a specimen of the genus
Mordellistena; see Table 1 for explanation of letters.

The results of the BMDP P2M program were given in several different graphical
displays. These included 1) a vertical tree showing the sequence of cluster formation,
2) a table listing the amalgamation distance and the mean for each variable as each
new cluster is formed, 3) a shaded distance matrix graphical display, 4) a listing of the
data, 5) a matrix of the distances between all cases, and 6) a histogram of the
distances. Of these the most useful for determination of relationships were found to
be the vertical tree, the matrix of distances, and the shaded distance matrix.

The shaded distance matrix is made by the computer via the overprinting of
characters beyond each specimen number (which run along the x-axis) with darker
characters, or more overprinting, indicating closer relatives. By following the line
out (along the horizontal) from any specimen number, it is possible to observe the
extent of relationship of any specimen to any other. Through examination of the

228

PAN-PACIFIC ENTOMOLOGIST

Figure 2. Scanning electron microscope photograph, ventral view; see Table 1 for explanation of letters.

shaded matrix for large, dark triangles, it is an easy matter to observe the location of
groups and discern the members of these. Dark triangles of this type may be seen by
referring to the sample shaded matrix in Figure 5.

The vertical tree is formed by the computer through the process of step-by-step
clustering and reflects precisely this process. The 400 beetles are first arranged across
the top of the display and at this point each case represents a separate cluster.
Proceeding down the page, then, it is possible to follow a line from any specimen and
determine at what point that beetle is grouped with another and when these two are
joined with another and so on. Along the y-axis of this tree are listed the
amalgamation distances so that it is possible to determine at what mathematical
distance any cluster is formed. These amalgamation distances ranged from 0.00 to
10.734 and increased down the display in increments less than or equal to 0.500 with
the exception of the final step, which went from 3.522 to 10.374. These numbers
reflected the sum of squares of the variables measured. It was at the amalgamation
distance of 10.374 that all 400 cases finally came to rest in a single cluster. The
diagram was very useful in the rapid discovery of odd specimens that had few
relatives. In these examples, odd specimens were seen to join the clustering process
near the bottom of the page. A sample vertical tree diagram is illustrated in Figure 6.

Finally, there is the actual distances matrix. In this printout, any specimen may be
compared to any other and thus the matrix of distances is simply a 400-by-400 chart

VOLUME 63, NUMBER 3

229

Figure 3. Scanning electron microscope photograph, close-up view of frons; see Table 1 for explanation
of letters.

with separation distances ranging from 0.00 (comparison of any case to itself) to
distances over 13.00.

All three of the preceding printouts discussed were used to determine what groups
had been constructed by the clustering program. Following this all 400 individual
specimens were examined and compared with Liljeblad’s key and descriptions
(1945) to determine what species had been found and overall concurrency with the
grouping process of the computer.

Results and Discussion

The cluster analysis delineated 14 major groups (those with five or more members)
comprising a total of 223 specimens, 25 minor groups (those with four or few
members) comprising a total of 73 specimens, and 104 specimens not definitely
aligned with any of the 39 total groups. All 104 of these non-grouped specimens were

230

PAN-PACIFIC ENTOMOLOGIST

Table 2.

Final 14 Characters Used in Cluster Analysis

1. length of ultimate abdominal sternite

2. length of metatrochanter

3. length of metepisternum

4. eye height

5. eye length

6. length of large metatibial spur

7. length of small metatibial spur

8. length of metatarsal claw

9. length of head capsule

10. length of metafemora

11. length of mesotibia

12. length of mesofemora

13. length of ultimate segment of maxillary palp

14. ocular width

Figure 4. Scanning electron microscope photograph, close-up view of metatarsal claw; see Table 1 for
explanation of letters.

VOLUME 63, NUMBER 3

231

DISTANCtS BETWEEN CASES RbPRcScNTED IN SHADED FORM.
HEAVY SHADING INDICATES SMALL DISTANCES.

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however easily grouped by hand, based on having at least one close relative among
the remaining 399 specimens.

Following visual examination of all 400 beetles, it became evident that while the
minor groups were all single species, only two of the major groups were single
species, while the other 12 all contained more than one species. However, these were
easily separated on the basis of ground color and color patterns. The 25 minor groups
all consisted of but a single species, so at least in some cases the 14 characters selected

232

PAN-PACIFIC ENTOMOLOGIST

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through the discriminant analysis were species specific. It is evident that in further
analyses a coding system for color and color patterns will be necessary to ensure
species specific grouping throughout. In the large groups the separation on the basis
of color was generally very easy, often involving nothing more than the identification
of solid brown specimens, solid black specimens and bicolorous specimens.

The study yielded a total of 58 different species present in Arizona and the
surrounding Sonoran desert. Of these, 14 were either unidentifiable in Liljeblad or

VOLUME 63, NUMBER 3

233

represented new species. A list of these species is presented in Table 2. It should be
noted that Liljeblad had 10 species recorded as occurring in Arizona. The generation
of such a species list for Arizona was one of the major goals of this project.

The other goal of this study was the identification of reliable taxonomic characters
and the validation of the leg ridges. It was clearly evident that the ridges of the
metathoracic legs were unreliable as shown by their failure to separate the four
groups of the discriminant analysis. The tibial ridges did seem to demonstrate more
constancy in form and number and may yet prove useful as identification characters.
The tarsal ridges, however, are far too transient, even varying from one side to the
other of a single insect, to be of any use. Liljeblad attempted no conclusions on
phylogeny. Rather, he merely grouped species according to ridge pattern. There
were several examples where the cluster analysis grouped different species together
that were in close proximity in Liljeblad’s work. This is an indication that there may
indeed be some phylogenetic significance to the ridges.

While ridges proved unreliable, the examination of the members of the groups
showed that members of the same group possessed very similarly structured ultimate
maxillary palp segments. This was true even in the larger groups made up of several
species. Liljeblad suggested the possible importance of this character and this
importance was reinforced by the fact that it was shown to be successful in the
separation of the four species in the discriminant analysis phase of the study.
Especially noticeable was the shape of the ultimate maxillary palp segment in two of
the major groups in which the shape exhibited was long, narrow and parallel-sided in
contrast to the usual scalene shape found in Mordellistena. As the pollen-feeding
habits of the adults would appear to indicate an important food-gathering function
for these segments, it is possible that the related morphology may be species specific.
The use of this small character (usually under .300 mm in length and often somewhat
hidden) in a taxonomic scheme is indeed much easier where numerical valves are
attached rather than general descriptions of the shape.

Perhaps the most noticeable problem encountered was the limitation in the
number of cases that the CYBER was able to handle, as previously mentioned.
Considering the case-by-case comparison method of the analysis, it is perhaps not
too surprising that there is indeed a limit to the number of cases that any computer
can handle. The point here is that there are finite and attainable limits restricting a
numerical study.

The occurrence of one apparent species in more than one group occurred in two
cases in the study and involves specimens which key in Liljeblad to M. tosta LeConte
and M. comata (LeConte). In each case, specimens keying to these species were
found in five groups formed by the cluster analysis. There are several possible
reasons for this placement. First, there is the possibility that one or both of these
species exhibit a wide range of variation in size and characters seemingly not usual for
Mordellistena. This would mean that the 14 characters which work for the rest of the
group are not suitable for these two species and this seems unlikely. A more plausible
explanation is separation based on sexual dimorphism. There is a definite wide
separation between small specimens keying to M. tosta and M. comata and larger
specimens keying to these two species. In addition, there is a noticeable difference in
the morphology of the ultimate maxillary palp segment with it being scalene in the
smaller specimens and elongate-securiform in the larger specimens. This difference
and these shapes were noted by Liljeblad in the descriptions of both species with the

234

PAN-PACIFIC ENTOMOLOGIST

males having the scalene, and the females the elongate-securiform, morphology.
Finally, there is the explanation that several species are represented and cannot be
distinguished except by the exacting method of a morphometric analysis. Whatever
the answer, all specimens keying to M. tosta and M. comata warrant further study.

The grouping of several apparent species in a single cluster questions the
discriminatory power of the analysis. As previously mentioned, in all instances little
or no difficulty was encountered in separating these based on color and color
patterns. The addition, or coding, of these characters is evidently a necessary one in a
future revision of the entire group.

Finally, there is the inability of the analysis to group 104 specimens. The 104 all had
at least one close relative and the majority had several among the rest of the
specimens. In addition, all but two specimens fell just outside the limits of a group.
The reason for these specimens not fitting into any group may be something as simple
as an inaccurate measurement of one or more characters or something more complex
involving character abberation. The relative of these specimens as indicated by the
analysis should, however, provide rapid clues as to their identity. The ideal
procedure, in keeping with the idea behind a study such as this, would be to add more
characters until these obstacles are overcome and grouping is accomplished for all
104 specimens. Investigation of such possibilities, which again may simply involve a
color coding, is something to be accomplished in an expanded study of the entire
group.

Summary

The question of usefulness of a morphometric analysis such as this in forming a
phylogeny for Mordellistena is now addressed. Two pieces of evidence indicate that
indeed this study can form an important part of such a phylogenetic reconstruction.
The first is the placement of the two groups with unusually elongate ultimate
maxillary palpal segments next to each other in the analytical results. It seems likely
that this elongate segment represents some sort of offshoot of a section of the
Mordellistena and the close clustering indicates phylogenetic significance to the
character. The second concerns the character of an elongated upper tibial ridge. In
cases where this striking character was present, it was seen in adjacent or solitary
groups, again indicating possible phylogenetic significance.

The cluster analysis formed 39 groups of specimens comprising a total of 296 of the
400 specimens submitted for analysis. This is a 74% successful grouping with 102 of
the remaining 104 specimens just outside the limits of the 39 groups. Thus the
establishment of valid characters for taxonomic use, the 14 delineated plus those of
color and color patterns, is seen as having been achieved. For a more detailed
explanation of the groups, the reader is referred to Burne (1985). Forty-four species
recognizable in Liljeblad, as well as 14 species either unrecognizable or undescribed,
were found through the analysis generating the list of Mordellistena of Arizona seen
in Table 3.

Acknowledgments

I would like to express gratitude to the following: Dr. Floyd G. Werner for serving
as advisor during this research and Carl A. Olson for assistance with photography,
both of the University of Arizona, Tucson; and Dr. C. Ferris, Dr. C. C. Burkhardt
and Dr. R. Pfadt for critical analysis, all of the University of Wyoming, Laramie.

VOLUME 63, NUMBER 3

235

Table 3. Species List from Study

M. vapida

M. conformis

M. husseyi

M. intermixta

M. picipennis

M. unicolor

M. tantula

M. Smithi

M. pustulata

M. aspersa

M. divisa

M. sp. 1

M. sp. prob. testacea

M. texana

M. sp. 2

M. paradisa

M. marginalis

M. sp. 3

M. pullata

M. calignosa

M. sp.4

M. aethiops

M. palmi

M. sp. 5

M. tosta

M. lutea

M. sp. 6

M. morula

M. ruficeps

M. sp. 7

M. nebulas a

M. rufa

M. sp. 8

M. comata

M. aemula

M. sp. 9

M. rubrifascia

M. splendens

M. sp.10

M. nigricans

M. blandula

M. sp.11

M. sericans

M. nunenmacheri

M. sp. 12

M. nubila

M. wickhami

M. sp.13

M. knausa

M. parva

M. sp.14

M. ambusta

M. nigella

M. subfucus

M. militaris

M. indistincta

M. pallens

Literature Cited

Burne, J. C. 1985. A Morphometric Analysis of Mordellistena in the Southwestern United States
(Coleptera: Mordellidae), Ph. D. Dissertation, University of Arizona, Tucson, 70 pp.

Engelman, L. P. 1979. Cluster Analysis of Cases. In BMDP Biomedical Computer Programs, P-Series,
1979. W. J. Dixon and M. B. Brown, eds.. University of California Press, Berkeley, 880 pp.
Khalaf, K. T. 1970. Wing venation and phylogenetic relationships in Mordellidae. Fla. Entomol.
53(3):153-160.

Khalaf, K. T. 1971. Mordellidae from Louisiana and Mississippi. J. Kans. Entomol. So. 44(4);441-445.
Khalaf, K. T. 1971. Five new species of Mordellidae from Louisiana and Mississippi. Pan-Pacific Ent.
47(2): 140-145.

Liljeblad, E. 1945. Monograph of the Family Mordellidae (Coleoptera) of North America North of
Mexico. Misc. Publ. Mus. Zool. Univ. Michigan, no. 62,7 pis., 229pp.

Ray, E. 1944. New Mordellidae Beetles from the Western Hemisphere. Zool. Serv. Field Mus. Nat. Hist.,
29:117-133.

Ray, E. 1946. Studies on North American Mordellidae, II, III, and IV. Pan-Pacific Entomol. 22:41-50,
90-99, and 121-132.

Ray, E. 1947. Studies on North American Mordellidae, V. Pan-Pacific Entomol. 23:121-131.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 236

Acanthochalcis nigricans Cameron—

New distribution information, including Central America

(Hymenoptera: Chalcididae)

Jeffrey A. Halstead

2110 N. Hayes, Fresno, California 93722

Abstract. —New distributional information for Acanthochalcis nigricans in
Nevada, United States; Baja California Sur, Mexico; Quezaltepeque, El Salvador;
and Guanacaste, Costa Rica is presented. Rubinistic color variation for nigricans is
noted.

Acanthochalcis nigricans Cameron is the second largest chalcidid wasp in America
north of Mexico. Its body length (males 10 mm, 8 to 11 mm; females 17 mm, 7 to 24
mm) is slightly shorter than its North American congener, unispinosa Girault.
Acanthochalcis are predominantly black with reddish brown (or black) labrum,
clypeus, legs, tegulae, sternites, and tergites (ventrally). Females are unique among
all North American Chalcididae in having the ovipositor project posteriorly a
distance equal to or greater than the length of the abdomen.

The gQmxs Acanthochalcis contains three species: nigricans —Kansas, Oklahoma,
Texas, New Mexico, Arizona, California, and Mexico; unispinosa —Texas, Arizona,
and California; gigas Steffan—Madagascar (Burks 1979, Catalog of Hymenoptera in
America north of Mexico, Smith, Instit. Press I: 860-874; Steffan 1959, Acta
Entomol. Mus. Natl. Prague 33(549):306-309). Both North American species are
presumed to be parasitoids of buprestid beetle larvae which inhabit oak trees
(Halstead and Haines 1985, Pan Pacific Entomol. 61(3):227-228).

Recently I examined specimens of nigricans which include new range records for
Nevada (United States), Baja California Sur (Mexico), El Salvador, and Costa Rica.
DeSantis (1979, Comision de Investigacioines Cientificas de La Provincia de Buenos
Aires, La Plata, Argentina, p. 66) listed nigricans from Mexico but no state
information was presented. Type material is from the state of Sonora (Cameron
1884, Biol. Cent. Amer., Hym., I: p. 100). These new records represent a
considerable southward range extension.

Locality data is as follows: UNITED STATES. NEVADA, Clark Co.: Las Vegas,
20.vi.39, on Cersium, P. Timberlake, 1$, (UCR). MEXICO. BAJA
CALIFORNIA SUR, Los Medanos, 10.3 mi. SW, elevation 0.4 m., 28.iii.64, on
Laria, sand dune association, M. E. Irwin, Id, (UCR); Santiago, 11 km W, Canon
de La Zorra, 285 m, 4-5.ix.77, E. Fisher and R. Westcott, 1$, (CAS). EL
SALVADOR. QUEZALTEPEQUE, 12.vii.61, 23.viii.61, M. E. Irwin, 3d,
(UCD). COSTA RICA, GUANACASTE, Palo Verde Station, 29 km WSW Canas,
10° 21'N, 85° 21'W, 10.vii.76, H. E. Hespenheide, 1$, (Hespenheide personal
collection).

The three specimens from El Salvador are rubinistic color forms (i.e., typically
black colored areas are reddish brown). No color variation has previously been noted
for Acanthochalcis.

236

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 237-241

Ecology of Rhopalomyia californica Felt at Jasper Ridge

(Diptera: Cecidomyiidae)

L. E. Ehler

Department of Entomology, University of California, Davis, California 95616

Abstract. —An ecological study of Rhopalomyia calif ornica Felt (Diptera:
Cecidomyiidae) was conducted at the Jasper Ridge Biological Preserve (Stanford
University) during 1982-83. The midge, which develops in terminal galls on
Baccharis pilularis DC, was relatively rare throughout the course of the study.
Analysis of life-table data suggested that its predators and parasites played a major
role in maintaining population density at comparatively low levels. The parasite
guild consisted of seven hymenopteran species: Torymus koebelei (Huber) & T.
baccharidis (Huber) (Torymidae), Zatropis capitis Burks & Mesopolobus sp.
(Pteromalidae), Tetrastichus sp. (Eulophidae), Eupelmus inyoensis Girault
(Eupelmidae), and Platygaster calif ornica (Ashmead) (Platygastridae). Malathion-
bait sprays applied to an adjacent area (Woodside) not only resulted in a massive
midge outbreak, but also indirectly altered the spatial structure of the midge
population. It is suggested that R. calif ornica, when introduced without its natural
enemies, may be an important biological-control agent against weedy Baccharis spp.
in Texas and Australia.

Rhopalomyia californica Felt is a native cecidomyiid midge which develops in
multi-chambered, terminal galls on Baccharis pilularis DC in California. Tilden
(1951b) described the natural history of the midge and most subsequent
investigations have dealt largely with the midge and its parasites as a model system
for addressing basic issues in community ecology and applied biological control
(Doutt, 1961; Force, 1974; Ehler 1982, 1985; and Hopper, 1984). The latest
development concerns the use of R. californica as a biological-control agent for
related Baccharis spp. which are rangeland weeds. In this regard, the midge is now
well established in Queensland, Australia, where it shows considerable promise for
eventually controlling Baccharis halimifolia L. in many parts of its range (McFadyen,
1985; W. A. Palmer, pers. comm.). It was also introduced into Texas during 1985
and 1986 (for control of several Baccharis spp.) and is now established at certain
release sites (P. Boldt, pers. comm.).

The purpose of the present paper is to summarize available data on the ecology of
R. californica at one site in its native range (Jasper Ridge Biological Preserve) in
order to facilitate eventual comparative studies in Texas and Queensland. Details of
the study sites, materials, and methods were given by Ehler et al. (1984).

Analysis of Life-Table Data

From 30 March 1982 to 9 March 1983, the midge population at Jasper Ridge was
relatively stable—i.e., density of galls never exceeded 2 per 100 terminals and larval
density did not exceed 20 per 100 terminals (Ehler et al., 1984). Life tables were

237

238

PAN-PACIFIC ENTOMOLOGIST

constructed for six cohorts of R. californica galls collected on the following dates in
1982: 30 March (n = 144), 3 May (n = 47), 1 June (n = 85), 14 July (n = 124), 12
Aug. (n = 41) and 7 Oct. (n = 83). Because major trends in mortality/survival in
each of the life tables were very similar, an average (n = 6) life table, based on 524
galls (5680 midge larvae), was calculated for a hypothetical cohort of 1000 individuals
(Table 1). Three major sources of mortality were apparent: predation of eggs and
neonate larvae, parasitization of endophagous larvae (i.e., inside the gall) and
residual mortality of larvae. Midge survival from egg to adult was ca. 5% and the sex
ratio was in favor of females.

The numbers of eggs and neonate larvae plus attendant mortality rates were
estimated from figures given by Ehler et al. (1984). In this case, the value for lx was
derived from the average gall size (assuming 100% survival from egg to endophagous
larva) in an adjacent area (Woodside) which was heavily sprayed with malathion bait
during the medfiy eradication campaign. These sprays presumably destroyed most of
the predators of these host stages and this apparently resulted in the consistently
larger galls found in this zone (see next section). The value for dx was determined by
subtraction. Because of the particular methods employed, the estimates obtained
should not be viewed as giving a complete picture of events during this age interval.
For example, predation was the only mortality factor estimated, even though other
factors (e.g., infertility) are probably involved. In view of this, the value for lx should
be regarded as an underestimate and that for dx as an overestimate. Nevertheless,
the data do suggest that predation on eggs and neonate larvae (i.e., before gall
formation) is of major importance in the population dynamics of the midge (see also
Ehler et al., 1984). Although numerous predatory insects occur on the plant (Tilden,
1951a), it was not possible to determine which species were responsible for the
mortality detected during this phase of the study.

Parasites destroyed ca. 38% of the endophagous larvae (25% real mortality). This
overall parasitization rate is somewhat low and is not necessarily characteristic of the
parasite guild in other parts of the host’s range. Seven species of solitary parasites
were reared from R. californica galls. Torymus koebelei (Huber) (Torymidae) is a
primary ectoparasite; parasitization by this species ranged from 1 to 13%. A second
torymid primary ectoparasite, T. baccharidis (Huber), was generally rare and never
parasitized >2% of the host larvae. Zatropis capitis Burks (Pteromalidae) is a
facultative secondary ectoparasite which parasitized from 0.9 to 6.5% of the hosts.
The dominant species in the parasite guild was Platygaster californica (Ashmead)
(Platygastridae), an egg-larval, primary endoparasite. This species parasitized from
1 to 36.2% of the larvae. Mesopolobus sp., another pteromalid facultative secondary
ectoparasite, parasitized > 10% of the hosts in March, but seldom parasitized > 1%
thereafter. The two remaining species were consistently rare: Tetrastichus sp.
(Eulophidae), a primary endoparasite, never parasitized >1.1% of the hosts
whereas Eupelmus inyoensis Girault (Eupelmidae), another facultative secondary
ectoparasite, was collected on only one date (Oct. 7). Over 5% of the larvae (range:
0.5-9.5) were parasitized, but due to the condition of the material, it was not possible
to attribute host mortality to any particular species listed above. Finally, total
parasitization rate and rates for individual species were relatively constant over time;
there was no convincing evidence for temporal density dependence.

The third major category of mortality was the residual—i.e., mortality of larvae
which could not be directly attributed to parasites. The residual mortality rate was
relatively high and probably has several components. Some hosts are presumably

VOLUME 63, NUMBER 3

239

Table 1. Average life table for Rhopalomyia californica at the Jasper Ridge Biological Preserve from March to Octo¬
ber, 1982.*

X

lx

dxF

dx

lOOqx

lOOrx

Egg + neonate larva

1000

Predation

355

35.5

35.5

Endophagous larva

645

Parasitization

T. koebelei

41.9

6.5

4.2

T. baccharidis

9

1.4

0.9

Z. capitis

27.7

4.3

2.8

P. californica

106.4

16.5

10.6

Mesopolobus sp.

20

3.1

2.0

Tetrastichus sp.

2.6

0.4

0.3

E. inyoensis

3.2

0.5

0.3

Undetermined

34.8

5.4

3.5

Total

245.7

38.1

24.6

Residual

341

52.9

34.1

Pupa

58.3

Residual

6.2

10.6

0.6

Adult

52.1

(0.61$ 9:0.39dc?)

947.9

—

94.8

*Key to symbols: x = developmental stage, lx = number entering stage x, dxF = mortality factor,
dx = number dying during stage X, lOOqx = dx lx for that stage (apparent mortality) and lOOrx = dx/lx
for the initial cohort (real mortality). See Ehler (1982) and Ehler et al. (1984) for details of materials and
methods.

killed by adult parasites—e.g., by host feeding, probing with the ovipositor (see also
Force, 1974)—and this would not be detected by the methods employed in this study.
Supernumerary parasitization is common at times and this could possibly result in
death of all occupants (host and parasites) in a given chamber. For these reasons, the
data for parasitization in Table 1 are probably a conservative reflection of the actual
impact of parasites. I also suspect that many gall inhabitants (both hosts and
parasites) were inadvertently killed during collecting and processing of galls. Recent
dissections of fresh galls (prior to midge emergence) collected in Yolo and Solano
Counties revealed that up to 10% of the chambers were empty. In other words, some
mortality had occurred prior to sampling, reinforcing the belief that residual
mortality has numerous components, as opposed to being strictly an artifact of
sampling technique.

Impact of Malathion-Bait Sprays

A massive outbreak of R. californica occurred at Woodside following 24
applications of malathion bait. At the height of the outbreak (Oct. 1982), midge
population levels were ca. 90X greater than those observed at Jasper Ridge. This
classic secondary outbreak evidently resulted from the wholesale destruction of
natural enemies of the midge (Ehler et al., 1984). I report here on a more subtle or
indirect effect of the sprays. The destruction of natural enemies (probably predators)
evidently allowed greater survival of eggs and neonate larvae which in turn allowed
more larvae to enter available buds and this presumably resulted in comparatively
larger galls in the spray zone. Although this effect was evident on all sample dates in
1982, the data for October were particularly striking (Figure 1). At Jasper Ridge, the

240

PAN-PACIFIC ENTOMOLOGIST

10 -

5 -

>-

C-3

a

LU

cc

JASPER RIDGE

••

• •

• •

10 -

WOODSIDE

• • •
••• ••••••

••••••

• •

•••

••••••••••••••••• •• • •

••

• •

10

20 30

CHAMBERS/GALL

40

50

Figure 1. Frequency distribution of galls according to size (chambers per gall) at Jasper Ridge (control)
and Woodside (medfly spray zone). Galls were collected on 7 October 1982; each dot represents one gall
(n = 83 for both sites).

median number of chambers per gall was 5 compared to 19 at Woodside; the average
number of chambers per gall at Woodside (21.9) was > 2X that at Jasper Ridge (8.1).
In other words, the malathion-bait sprays altered the spatial structure of the midge
population. Conversely, the data suggest that natural enemies can play a major role
in determining the spatial structure of a host population in nature. In view of these
findings, the value of gall size as an index of host-plant quality, while perhaps
intuitively appealing, must be questioned. This may also apply to midge populations
in Texas and Australia because native generalist predators on related Baccharis spp.
could produce a similar effect. Finally, the influence of an altered spatial structure of
a host population on the behavior and performance of a natural enemy should be
investigated.

Rhopalomyia As a Biological-Control Agent

The results from Woodside suggest that R. californica, when introduced without
its natural enemies, has considerable potential as a biological-control agent against
weedy Baccharis spp. in Texas and Australia. The host plants at Woodside were so
devastated (by the midge and other phytophagous species) that sampling had to be
discontinued after October 1982. This can be taken as an indication of the kind of
impact the midge might have in biological control of weeds. In future projects,
deliberate disruption of herbivore populations in the native home of a target weed
might serve as an aid in selecting candidate natural enemies for further study.

The selection of effective natural enemies for biological control of weeds has
received considerable attention in recent years (Harris, 1973; Goeden, 1983). In this

VOLUME 63, NUMBER 3

241

context, the case of R. californica is relevant to a current controversy over the use of
coevolved versus “new”’ exploiter-victim associations. Hokkanen and Pimentel
(1984) argued that new associations should be the preferred method of selecting
natural enemies; this recommendation was based on the hypothesized lack of
coevolved, interspecific balance which attends such associations. However, Goeden
and Kok (1986) challenged these findings and suggested that new exploiter-victim
associations offer limited opportunities for biological control, especially for
non-cactaceous weeds. Although controversial, I believe that the Hokkanen-
Pimentel thesis should be carefully considered and empirically tested whenever
possible. The introduction of R. calif ornica, derived from B. pilularis in California,
into Australia and Texas for control of related Baccharis spp. should provide some
key information on this question.

Acknowledgments

I thank A. Grundmann and associates at Stanford University for the opportunity
to conduct research at the Jasper Ridge Biological Preserve and B.
Alvarado-Rodriquez, P. C. Endicott, M. B. Hertlein, H. P. Sauter and K.
Thorarinsson for assistance in collecting and analyzing the data. K. Thorarinsson,
W. A. Palmer and P. Boldt reviewed an earlier version of the manuscript. Portions of
this research were supported by the California Department of Food and Agriculture
(Standard Agreement 1779).

Literature Cited

Doutt, R. L. 1961. The dimensions of endemism. Ann. Entomol. Soc. Am. 54:46-53.

Ehler, L. E. 1982. Ecology of Rhopalomyia californica Felt (Diptera: Cecidomyiidae) and its parasites in
an urban environment. Hilgardia 50:1-32.

Ehler, L. E. 1985. Species-dependent mortality in a parasite guild and its relevance to biological control.
Environ. Entomol. 14:1-6.

Ehler, L. E., P. C. Endicott, M. B. Hertlein, and B. Alvarado-Rodriquez. 1984. Medfly eradication in
California: impact of malathion-bait sprays on an endemic gall midge and its parasitoids. Entomol.
Exp. Appl. 36:201-208.

Force, D. C. 1974. Ecology of insect host-parasitoid communities. Science 184: 624-632.

Goeden, R. D. 1983. Critique and revision of Harris’ scoring system for selection of insect agents in
biological control of weeds. Prot. Ecol. 5:287-301.

Goeden, R. D., and L. T. Kok. 1986. Comments on a proposed “new” approach for selecting agents for
the biological control of weeds. Can. Entomol. 118:51-58.

Harris, P. 1973. The selection of effective agents for the biological control of weeds. Can. Entomol.
105:1495-1503.

Hokkanen, H., and D. Pimentel. 1984. New approach for selecting biological control agents. Can.
Entomol. 116:1109-1121.

Hopper, K. R. 1984. The effects of host-finding and colonization rates on abundances of parasitoids of a
gall midge. Ecology 65:20-27.

McFadyen, P. J. 1985. Introduction of the gall fly Rhopalomyia californica from the USA into Australia
for the control of the weed Baccharis halimifolia. pp. 779-787. In: E. S. Del fosse (ed.), Proc. VI
Intern. Symp. Biol. Control Weeds. Agric. Canada, Ottawa.

Tilden, J. W. 1951a. The insect associates of Baccharis pilularis De Candolle. Microentomology
16:149-188.

Tilden, J. W. 1951b. Observations on Rhopalomyia californica Felt. Pan-Pac. Entomol. 27:20-22.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 242-246

Male Swarms Discovered in Chalcidoidea
(Hymenoptera: Encyrtidae, Pteromalidae)

H. Nadel

Department of Entomology, University of California, Riverside, California 92521

Abstract. —Male swarms have been discovered in three chalcidoid species:
Bothriothorax nigripes, Copidosoma sp. (Encyrtidae), and Pachyneuron sp.
(Pteromalidae). The swarms occurred around boulders at the top of a small ridge in
southern California. B. nigripes males, and possibly the others, swarmed for the
purpose of mating. The mating behavior of B. nigripes is described. This may be the
first report of male mating swarms in the Chalcidoidea.

Males of three chalcidoid species were discovered swarming on and around five
hilltop boulders near the University of California at Riverside campus. The
aggregations were discovered on March 7,1984 and were observed at least biweekly
until their disappearance in mid-April of the same year. Examination of samples
taken by aspiration and aerial sweepnetting from the same location on three different
days revealed the following species and numbers (males:females): Bothriothorax
nigripes Howard (Encyrtidae), 865:23; Copidosoma sp. (Encyrtidae), 291:14;
Pachy neuron sp. (Pteromalidae), 522:0. The swarms daily contained several
thousand individuals of each species. They formed anew each day after disappearing
entirely from the sites during the night.

This report includes a brief description of the swarming and mating behavior of B.
nigripes, Copidosoma sp., and of the swarming behavior of Pachy neuron sp. Greater
emphasis is given to B. nigripes. A study was made to determine whether the B.
nigripes aggregations were true mating swarms. The known literature on swarming in
the parasitic Hymenoptera is included in the discussion.

Swarming and Mating Behavior

I chose to closely observe B. nigripes to ensure that its aggregations were true
mating swarms. This species is the largest of the three and thus more easily observed
in the field. Males began arriving near the boulders at around 7:00 a.m. and landed
during intervals of low wind velocity. Throughout the day, many swarmed in flight
while others congregated on the shaded north- and northwest-facing rock surfaces.
The males’ behavior on the rocks consisted of walking in irregular paths with
frequent turns, and beating the substrate with the antennae in rapid, alternating
strokes. Landing females remained relatively motionless with their antennae tucked
close to their faces, or walked slowly for a short distance ( < 10 cm) before becoming
still. Males encountered them within a minute or two and began courting. A
schematic summary of courtship and mating behaviors is presented in Fig. 1. Males
appeared to sense a female’s presence from a distance of 1 cm away, as they increased
walking speed and headed directly toward her from that point. Uninterrupted
courtship and copulation lasted about 40 seconds. In some instances other males

242

VOLUME 63, NUMBER 3

243

Male

Female

walks, turning frequently,
antennae alternately
beating up and down

lands on substrate

accelerates pace towards
female

walks slowly in + linear
path, antennae alternately
tapping substrate

motionless, antennae 4r
tucked close to face

faces female,
waggles body
8-10 times

mounts female from behind

places pro- and mesothoracic
legs on female's abdomen or
wings; rocks rapidly side to
side, metathoracic legs
alternately touching sub¬
strate

begins copulation

exposes genital pore

motionless

motionless ca. 10 seconds

dismounts

faces female and resumes
waggling __

springs away

Figure 1. Schematic representation of uninterrupted courtship and mating in Bothriothorax nigripes.

244

PAN-PACIFIC ENTOMOLOGIST

approached a pair in apparent attempts to copulate with the female, in which case the
female usually sprang away, often carrying one mounted male with her.

Females were never observed to spring away before mating or being mobbed by
several males. They seemed, therefore, to arrive at the swarms specifically for the
purpose of mating. To gain further evidence for this, I caught and dissected six
newly-landed females (before male contact) and found that their spermathecae did
not contain sperm. I paired four other newly-landed females with males in glass vials
and found them to mate readily. Two of these females were dissected and found to
have full spermathecae, and the remainder were paired with other males and found
to reject a second mating during 9- and 18-hour video-recorded observation periods.
Four additional females which were caught after they had mated naturally in the
swarms also rejected subsequent mating attempts when paired with males in glass
vials. Because newly-landed females mated readily both in the swarms and in the
laboratory, while mated females rejected subsequent matings, B. nigripes females
apparently join the male swarms in an uninseminated condition for the purpose of
mating.

It was unlikely that females arrived in search of food; some swarm boulders were
devoid of vegetation, and no feeding by either sex was observed on the few flowering
plants that grew on and around the rest. It was also unlikely that females appeared in
the area to search for their hosts, syrphid larvae, because none were detected on
ridgetop vegetation.

I sporadically observed the other swarming species. Mating pairs of Copidosoma
sp. were evident, often on rock-top vegetation, hence I also regarded the
aggregations of this species as mating swarms. The males generally flew above or
walked on the upper surface of vegetated boulders. Pachyneuron females, however,
were neither seen nor collected. The males flew and walked around on the upper
boulder surfaces, sometimes mixing with Copidosoma, but they also swarmed apart
on the north- and northeast-facing surfaces. Although I did not observe mating
within the Pachyneuron swarms, I suggest that, on the basis of the similarity in their
aggregative behavior to the other species, the Pachyneuron assemblages were also
mating swarms.

The Swarm Site

The most obvious factor distinguishing the swarm site. Coyote Ridge, is its
situation as the only low yet abrupt peak within a 7.5 km radius. It rises 134 m above a
base altitude of 335 m, and is surrounded by flat or gently sloping land. The Box
Springs Mountains nearby peak at 920 m at 2.5 km to the east. A brief examination of
Box Springs peaks at various heights, and of the nearest low, abrupt, hill in the area.
Mount Rubidoux (165 m), revealed no chalcidoid swarms on March 24, 1984,
although swarms were active that day on Coyote Ridge. Continuous and sometimes
gusty winds may have contributed to the paucity of insects on the unpreferred peaks.
Winds around Coyote Ridge were low and occasionally still, and perhaps more
suited to controlled locomotion by the wasps.

Discussion

Male mating swarms commonly occur in insects, especially in Diptera. They are
generally believed to result from 1) males searching at sites with high probability of

VOLUME 63, NUMBER 3

245

encounter with receptive females, such as emergence, feeding, and oviposition sites,
2) active male aggregation for the purpose of enhancing attractivity toward females,
or 3) localization of mating activity at landmarks (Thornhill and Alcock 1983).
Among the parasitic Hymenoptera, male swarms have been reported in the
Ichneumonoidea (Ichneumonidae; Rotheray 1981; Braconidae: Donisthorpe 1936,
Stelfox 1944, Southwood 1957; Aphidiidae: Stary 1970) and possibly in the
Bethyloidea (Dryinidae: Jervis 1979), but not in the Chalcidoidea. Large swarms of
female Cyclogastrella (= Pteromalus) deplanata (Pteromalidae) have been reported
from buildings in England (Scott 1919), and of female Chrysocharis centralis
(Eulophidae) on vegetation in Madeira (Graham 1983), but there is no indication
that these served a mating purpose. Recently, however, pteromalid mating swarms
have been observed on citrus trees in China (J. K. Waage, personal communication).

The male chalcidoids on Coyote Ridge appeared to use either a landmark-based
swarm site and/or to aggregate to increase their power to attract females. There was
no indication that either sex arrived at the ridgetop to feed. Similarly, at least for B.
nigripes, there was no indication that females arrived in search of hosts. Certainly,
there was no indication that the ridgetop was in any way more profitable than the
surrounding areas in terms of food or oviposition sites for any of the swarming
chalcidoids. The ridgetop, however, offers a distinctive landmark which is also
accompanied by slower winds than those which prevail on the nearest peaks. It is,
therefore, not only conspicuous, but also allows controlled locomotion by the
insects. These attributes of the ridge may have set the stage for the evolution or
maintenance of a landmark-based mating system in the observed chalcidoids. It is
also possible, however, that the males of each species aggregate because they thus
have a greater chance of mating than if they remained solitary. Male aggregation
pheromones are commonly implicated in the formation of insect swarms and in the
attraction of females (Thornhill and Alcock 1983). It is not impossible that the
aggregation pheromones, if they exist, of the three studied chalcidoid species are
similar; this would explain why these presumably rare swarms occurred together.
Much more work is necessary to reveal why and how the swarms occur, but at this
point the observations indicate that they are part of a landmark-based mating system
or the result of a male tendency to aggregate, or both.

Chalcidoid mating swarms have been reported here for the first time, but is this
because little attention has been given to parasitoid mating behavior in nature, or is it
because the swarms are truly rare? As far as I know, there have been no focused
studies on the mating systems of non-inbreeding parasitic Hymenoptera beyond the
confines of the laboratory. This is unfortunate from both a practical and theoretical
standpoint. Such studies may aid the evaluation of biological control agents;
introduced exotic chalcidoids, for example, may fail to become established because
of the absence of proper mating sites. As I showed in this study, readiness to mate in
the laboratory gives no indication of the species’ mating system: B. nigripes mated
readily in glass vials. If chalcidoid mating swarms are truly rare, however, we should
search for a general tendency in the Chalcidoidea to mate at the emergence site
rather than to mate after dispersal. Many chalcidoid species have a preponderance of
females (Gordh 1979), and this has been linked, in theory, to pre-dispersal mating
(Hamilton 1967, Charnov 1982, Waage 1982). The discovery of chalcidoids which
mate away from their natal site paves the way for testing the relationship between

246

PAN-PACIFIC ENTOMOLOGIST

mating system and sex ratio in this group. In any event, the study of chalcidoid
mating systems should provide a rich store of information for both practical and
theoretical work.

Acknowledgments

I owe foremost thanks to Dr. G. Gordh for urging me to write about the swarms,
and for his critical review of earlier versions of the manuscript. Many thanks to the
following people for their help: Mr. G. Scriven forced the discovery through his
aerobics training; Drs. L. LeBeck and T. Unruh, Ms. T. McUnruh, and Mr. D.
Goodward helped with the observations; Dr. J. LaSalle identified the encyrtids, and
Mr. H. Anderson identified the pteromalid; Drs. J. Pinto and T. Unruh, Mr. R.
Stouthammer and Mr. D. Goodward reviewed an earlier version of the manuscript.

Literature Cited

Charnov, E. L. 1982. The Theory of Sex Allocation. Princeton University Press, Princeton, New Jersey,
355 pp.

Donisthorpe, H. 1936. The dancing habits of some Braconidae. Entomol. Rec. 78:84.

Gordh, G. 1979. Superfamily Chalcidoidea. pp. 743-748. In: Catalog of Hymenoptera in America North
of Mexico (K. V. Krombein, P. D. Hurd, Jr., D. R. Smith, and B. D. Burks, eds.). Smithsonian
Institution Press, Washington, D.C., 1:1-1198.

Graham, M. W. R. de V. 1983. Madeira insects: Faunal notes, additions and descriptions of new species
of Chalcidoidea (Hymenoptera). Bol. Mus. Mun. Funchal 35:5-40.

Hamilton, W. D. 1967. Extraordinary sex ratios. Science 156:477-488.

Jervis, M. A. 1979. Courtship, mating and “swarming” in Aphelopus melalucas (Dalman)
(Hymenoptera: Dryinidae). Entomol. Gaz. 30:191-193.

Rotheray, G. E. 1981. Courtship, male swarms and a sex pheromone of Diplazon pactoratorius
(Thunberg) (Hymenoptera: Ichneumonidae). Entomol. Gaz. 32:193-196.

Scott, H. 1919. Swarming of the chalcid Pteromalus deplanata Nees in buildings. Entomol. Monthly Mag.
55:13-16.

Southwood, T. R. E. 1957. Observations on swarming in Braconidae (Hymenoptera) and
Coniopterygidae (Neuroptera). Proc. R. Entomol. Soc. London (A) 32:80-82.

Stary, P. 1970. The biology of aphid parasites. Dr. W. Junk N. V. The Hague, 643 pp.

Stelfox, A. W. 1944. The swarming flights of Blacus (Hymenoptera: Braconidae). Entomol. Monthly
Mag. 80:208.

Thornhill, R. and J. Alcock. 1983. The Evolution of Insect Mating Systems. Harvard University Press,
Cambridge and London, 547 pp.

Waage, J. K. 1982. Sib-mating and sex ratio strategies in Scelionid wasps. Ecol. Entomol. 7:103-112.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 247-255

The Drosophila Fauna of a Native California Forest
(Diptera: Drosophilidae)

Herman T. Spieth
University of California, Davis, California 95616

Introduction

During the years since A.D. 1492 and especially during the past 150 years the
Drosophila fauna of North America has undergone considerable change. Two major
factors have contributed to this change: (1) a number of Drosophila species from
other continents have become introduced into the fauna, e.g. D. melanogaster and
simulans and recently D. subobscura; and (2) larval substrates have changed both
qualitatively and quantitatively. Numerous fruits and vegetables from other
continents have been introduced, e.g. European grapes, apples, and citrus, all of
which provide substrates for native polyphagous species, e.g. obscura species group.
In contrast urbanization, agriculture and lumbering have reduced the volume of
substrates provided by forest fungi, slime fluxes, cacti and fermenting bark. It is
reasonable to assume that the population sizes of most oligophagic and monophagic
species have been reduced. The virilis species group provides a clear example of such
a reduction in population size (Spieth 1979).

Blodgett Forest

The Blodgett Forest Research Station of the University of California Department
of Forestry and Conservation is located in the Eldorado National Forest on the
western slope of the Sierra Nevada Mountains. Consisting of 2,961 acres (1198
hectares) located between 4,100 and 4,600 feet (1250-1400 m) elevation, it is a
“mixed conifer” forest which supports ponderosa pine, sugar pine, white fir, incense
cedar, douglas fir, black oak, tan oak and native shrubs and forbs. The forest is
totally surrounded by the Eldorado National Forest. The sole access to Blodgett is
via a narrow two lane road that originates at Georgetown, a small town located 10 air
miles west of Blodgett. Continuous forest exists from Georgetown to Blodgett. A
number of dwellings, mostly summer homes plus camping areas, exist along the
access road, mainly along the western portion. Agricultural activities are absent
along the road and in the surrounding forest.

The Blodgett Station was established in 1933, and since its inception all introduced
plants have been systematically eliminated. Thus, as a result of the Blodgett Station’s
location within the National Forest and the efforts of the foresters, it can reasonably
be assumed that the forest approaehes closely the habitat that existed when
Columbus reached America.

Systematic collecting of the Drosophila fauna was undertaken during the years
1981-84 in order to determine both the species composition and size of the
Drosophila fauna of the forest.

Collecting Procedures

Blodgett Forest is an irregularly shaped rectangle with a north-south axis of circa
4.5 mi (7.2 km.). A network of unpaved dirt roads facilitates access to various

247

248

PAN-PACIFIC ENTOMOLOGIST

portions of the forests. Five sites along a north-northwest to south-southeast
transect were selected.

Two sites, 1 and lA, at the northern end of the forest, with elevations of 1310 m
and 1320 m, are located 375 m apart. The forest is of medium density with numerous
individuals of the native black oak, Quercus kelloggii, which are scattered among the
conifers.

Site 2, elevation 1365 m, is located 2.6 km southward of sites 1 and lA. It slopes
westward and has a grove of Q. kelloggii surrounded by a coniferous forest amongst
which are numerous oaks and shrubs.

Site 3, elevation 1292 m, located 840 m south-eastward of site 2, is beside a stream
in a dense forest of mature conifers. Oaks are totally absent and daily temperatures
are consistently 2-3°C lower than at any other area of the forest.

Site 4, elevation 1310 m, is located 1.31 km. south of site 3. The terrain slopes
south-eastward. Soil depths in the western part of the area are shallow with
numerous rocky outcrops. Numerous young Q. kelloggii oaks and Ceanothus sp.
dominate this upper, drier and rocky western portion of the area while old mature
oaks and a mixture of young oaks and conifers are present in the lower eastern
portion.

Plastic, red colored buckets, height 26 cm, dia. 21.5 cm, were used as bait traps,
and fermenting, yeasted bananas as bait. Three buckets spaced 100-200 m apart
were used at each of the five sites. The bait buckets were suspended from tree
branches 0.5-1.5 m above the forest floor. Collections were scheduled to be
accomplished at two-week intervals from mid-April to early October during 1981,
’82, ’83 and ’84. Each collection consisted of an “evening” collection followed by a
“morning” collection on the following day. The bait buckets and baits were removed
at the termination of the “morning” collections. Occasionally one or both collections
were nonproductive due to environmental conditions such as low temperatures, high
temperatures or the rare passage of a weather front through the area.

All specimens collected at each of the 15 baits were returned to the laboratory for
identification. Since the four species of the D. obscura species group cannot be
effectively separated visually, individuals belonging to the group were identified by
electrophoretic techniques.

Blodgett Forest Species

During the four years, 1981-84 inclusive, adults of 13 species belonging to three
subgenera, i.e. Dorsilopha, Drosophila and Sophophora, were collected (Table 1).

Subgenus Dorsilopha Sturtevant

A single male specimen of D. busckii Sturtevant was collected 6.vi.82 (Table 1).
D. busckii is a cosmopolitan “garbage” species. This single individual can reasonably
be considered to have been a migrant into the Blodgett Forest.

Subgenus Sophophora Sturtevant

Adults of two species of the melanogaster species group and four species of the
obscura species group were collected during each of the four years (Table 1).

VOLUME 63, NUMBER 3

249

Summary of Species

Table 1

Subgenus Species

1981

d

2

1982

d

2

1983

d

2

1984

d

2

Dorsilopha D. busckii

0

0

1

0

0

0

0

0

Sophophora

D. melanogaster

SS

5

-

4

-

5

-

38

-

D. simulans

S6

53

-

6

-

104

-

165

-

D. melanogaster

D. simulans

118

32

262

1387

D. azteca

D. miranda

1290

1410

1049

1040

1500

2170

1075

973

D. persimilis

D. pseudoobscura

Drosophila

D. californica

7

7

11

14

378

282

84

47

D. immigrans

2

35

0

5

9

53

0

3

D. montana

0

0

0

1

0

1

0

1

D. occidentalis

143

129

51

34

162

143

167

138

D. pinicola

249

20

56

67

60

50

38

45

D. subfunebris

4

13

0

1

3

10

3

14

Melanogaster species group

D, melanogaster yiQigQn and D. simulans Sturtevant

During all four years adults of both species were absent from the collections during
the months of April, May, June and July (Table 2). Presumably they were also absent
during the winter and early spring. Only two males, one of each species, plus 32
females were collected during August and 30 of these females during August of 1984
(Table 2). In each of the four years the number of females greatly exceeded the
number of males collected, i.e., 4.7 females for each male. Also the number of
simulans males greatly exceeded the number of melanogaster males.

Blodgett Forest is located in the drainage of the Sacramento Valley. At the lower
elevations vast acreages of the valley are devoted to vineyards, orchards and
vegetable crops, especially tomatoes. Each year huge populations of both D.
melanogaster and D. simulans develop in these agricultural areas. The simulans
populations greatly exceed in size the populations. Areas located about

15-20 miles southwest and west of Blodgett are known to produce large populations
of both of these species. After the fruits and vegetables are harvested the adults must
seek other food and ovipositional sites. The appearance of adults of the two species
in the Blodgett Forest in September and October during each of the four years
suggests that these individuals were migrants from lower elevations and that they
must have migrated at least 15-20 miles. Further, the lack of individuals of both

250

PAN-PACIFIC ENTOMOLOGIST

Table 2

Species

Year

Apr

May

June

July

Aug

Sept

Oct

Totals

D. melanogaster

1981

0

0

0

0

0

0

5

5

1982

0

0

0

0

0

4

0

4

1983

0

0

0

0

0

2

3

5

1984

0

0

0

0

1

37

0

38

52

D. simulans

1981

0

0

0

0

0

38

15

53

1982

0

0

0

0

0

6

0

6

1983

0

0

0

0

0

8

96

104

1984

0

0

0

0

1

162

2

165

328

D. melanogaster

9$

1981

0

0

0

0

2

94

22

118

D. simulans

1982

0

0

0

0

0

32

0

32

1983

0

0

0

0

0

95

167

262

1984

0

0

0

0

30

1352

5

1387

1749

species during the months of April through July indicates that the migrants that reach
Blodgett during the months of August through October of the preceding year were
not able to survive in the forests during the winter months.

Obscura species group

Four species of the obscura group are residents of the Blodgett Forest; i.e. D.
azteca Sturtevant and Dobzhansky, D. miranda Dobzhansky, D. persimilis
Dobzhansky and Epling and D. pseudoobscura Frolowa. Numerically they are the
dominant species of the forest (Table 1).

Morphologically, individuals of D. persimilis and D. pseudoobscura are essentially
indistinguishable. Undersized individuals of D. miranda can be confused with large
sized individuals of persimilis and pseudoobscura. Only the males of azteca can be
confidently identified. The electrophoretic patterns of each of the four species,
however, are species specific and therefore were utilized to determine the specific
identity of the flies (Table 3). D. pseudoobscura and D. persimilis are numerically the
dominant members of the obscura species constituting more than 70% of the obscura
flies collected during 1981, ’82 and ’83 but receding to 61% in 1984. In comparison D.
miranda increased from 6.88% in 1981 to 23.26% in 1984. D. miranda is a
“northern” species whose range extends from Canada to the southern portion of
California. In comparison D. azteca is a “southern” species whose range extends
from Guatemala to northern California. From 1981 to 1983 the azteca populations
decreased sharply but recovered in 1984 (Table 3).

No specimens of the recently introduced D. subobscura were collected during the
years 1981-84. During 1985 a few collections were made in the forest and D.
subobscura appeared in one of these.

VOLUME 63, NUMBER 3

251

Table 3

D. obscura group species

1981

1982

1983

1984

D. pseudoobscura

36.28%

27.16%

44.35%

30.85%

D. persimilis

34.64%

47.83%

32.66%

30.29%

D. miranda

6.88%

13.74%

20.80%

23.26%

D. azteca

22.20%

11.27%

2.19%

15.60%

Subgenus Drosophila

Six species of the subgenus were collected (Table 1). Each of these is a member of a
different species group.

D. californica Sturtevant

D. californica, a member of the repleta species group, was described by Sturtevant
(1923) on the basis of 1 d and 4 9 9 collected at Pacific Grove, California.
Subsequent collecting has indicated that the species is apparently restricted to
California. Its larval substrate is unknown and it is considered to be a “rare” species
since it appears infrequently and usually in small numbers in Drosophila collections.

The volume of the larval substrate present in any area is positively correlated with
the size of adult populations of the species that breeds in the substrate; i.e., with rare
exceptions Drosophila females are capable of producing and ovipositing large
numbers of fertile eggs during their normal life spans. Thus the major factor
controlling population size is the availability of suitable larval substrate(s).

During 1981 only 14 individuals (Table 4) were collected, 11 of which were
collected during July and August. In 1982 25 individuals were collected, but, in 1983,
660 specimens of californica were collected. Clearly the amount of larval substrate
suitable for D. californica increased dramatically in 1983. In 1984 the population
receded and only 131 individuals were collected.

D. immigrans Sturtevant

This cosmopolitan species is similar to D. melanogaster and D. simulans in that it
cannot overwinter in the forest. During the four years 107 specimens, 11 d c? and 96
9 9 , were collected. Of these 1 6 and 3 9 9 were collected during the months of
April through August while the remaining 103 were captured during September and
October.

The yearly variation was great: e.g., only three specimens, all females, were
captured in 1984,62 individuals of which nine were males in 1983, five females and no
males in 1982 and 35 females and 2 males in 1981. It can reasonably be concluded that
all of these individuals were migrants from lower elevation.

D. montana Patterson and Wheeler

Only three females, one each in 1982,1983 and 1984, were captured. This species
utilizes the fermenting bark of the aspen of its larval substrate. Since aspen is lacking

252

Table 4

D. californica Sturtevant

PAN-PACIFIC ENTOMOLOGIST

Apr

6

9

May

6

9

June

cJ

9

July

(J

9

Aug

3 9

Sept

d 9

Oct

6

9

Totals

36 99

T

1981

1

0

0

0

1

0

3

3

1

4

0

0

0

0

7

7

14

1982

0

0

0

0

3

3

5

6

1

3

2

2

0

0

11

14

25

1983

0

1

3

1

11

19

54

51

88

77

148

117

74

16

378

282

660

1984

1

0

10

9

5

2

11

7

34

22

24

7

1

0

84

47

131

480

350

830

D.

Table 5

immigrans Sturtevant

July

3 9

Aug

3 9

Sept

3 9

Oct

(J 9

Totals

33 99

T

1981

0

1

0

0

0

33

2

0

2

34

37*

1982

0

0

0

0

0

5

0

0

0

5

5

1983

1

0

0

1

6

46

2

6

9

53

62

1984

0

0

0

0

0

2

0

1

0

3

3

*one 9 coll. 4 April 81

from the Blodgett Forest these three specimens appear to be migrants from higher
elevations where aspen and D. montana are both common.

D. occidentalis Spencer and D. pinicola Sturtevant

Both species utilize fungi as larval substrates. The Blodgett Forest has a
Mediterranean type climate with only scant and unpredictable precipitation during
the summer months. As a result the area becomes increasingly desiccated during the
summer and early fall. The fungal populations reach their peak during April and May
and steadily decline during the remainder of the year. Fungi do exist during the
summer months along stream banks and at the outflows of a few permanent springs.

D. occidentalis populations peak during June and July. Except for 1982 its
populations were similar in size during each of the other three years (Table 6). In
contrast the pinicola populations reach their peak during May. D. pinicola females
become sterile when subjected to temperatures above 18°C (Spieth and Heed 1975).
In the Blodgett Forest the pinicola populations were largest during May (Table 7)
and declined rapidly during the following months, and this decline may be due to
temperature constraints. Further, the pinicola populations, for unknown reasons,
declined drastically during each of the following years after 1981.

D. subfunebris Stalker and Spencer

During the four years only 48 specimens of subfunebris were collected (Table 1)
and it appears to be the rarest of the native species in the Blodgett Forest. Since,

VOLUME 63, NUMBER 3

253

Table 6

D. occidentalis Spencer

Apr

(J 2

May
i 2

June

6 2

July

6 2

Aug

S 2

Sept
<J 2

Oct

cJ 2

Totals

66 22

T

1981

19

10

36

16

31

25

44

39

3

8

4

9

10

12

147

119

266

1982

1

0

15

5

22

10

10

12

3

3

0

4

0

0

51

54

105

1983

0

0

8

17

34

28

62

51

9

18

4

23

15

2

132

139

271

1984

0

0

6

47

39

27

39

46

22

17

4

3

3

4

113

138

251

Table 7

D. pinicola Sturtevant

Apr

6 2

May

6 2

June

6 2

July

6 2

Aug

6 2

Sept

6 2

Oct

6 2

Totals

66 22

T

1981

20

18

205

136

6

21

12

18

0

2

3

3

3

8

249

206

455

1982

22

32

28

14

2

5

2

6

1

2

0

0

0

0

55

59

114

1983

3

10

29

11

8

9

16

3

0

3

0

2

7

2

63

40

103

1984

0

1

15

12

12

6

5

14

1

8

0

4

3

0

36

45

81

however, only 10 of the specimens were males it is possible that adult individuals,
especially males, are only marginally attracted to fermenting bananas and that the
populations may have been considerably larger than the present data indicate.

Larval Substrates

As the summer progresses the forest becomes increasingly desiccated. Thus the
stream beside which collecting site 3 is located has sufficient volume to support trout
during the months of April and May but ceases to flow during August and
September. The stream bed remains somewhat moist and D. occidentalis and D.
pinicola tend to congregate in the area but other species such as D. californica, D.
subfunebris and the four obscura species, i.e. azteca, miranda, persimilis and
pseudoobscura, do not. These species consistently occupy all parts of the forest and
this indicates that their larval substrates and food sources are distributed throughout
the forest. Clearly the adults of these species are able to find adult nutrients and
larval substrates in a desiccated environment which appears to lack any free liquid. It
should be noted that climatic conditions in the forest are such that no dew forms
during the summer and fall months and none of the plants exhibits guttation.

Carson (1951), Carson et al. (1956) investigated sixty-four slime fluxes on the
California black oak Quercus kelloggii at Mather, California, which is located 90
miles southeast of and at approximately the same altitude as Blodgett. Twenty-two
of the sixty-four fluxes they examined were utilized by D. persimilis and/or D.
pseudoobscura and four by D. californica. Slime fluxes are relatively rare at
Blodgett, a fact which was confirmed by J. Powell who visited Blodgett. The Blodgett
fluxes were consistently inspected during the four years but neither adults nor larvae
of any species were found on or in the fluxes.

254

PAN-PACIFIC ENTOMOLOGIST

Quercus kelloggii produces vast numbers of acorns. The majority of these are
parasitized by either the coleopteran Curculio occidentalis or the lepidopteran
Melissopus latiferreanus. The larvae of these two species feed on the cotyledons and
embryo of the acorn. Shortly after the acorn is shed from the tree the larvae reach
maturity and they then cut a small circular escape hole through the shell of acorn,
escape and pupate in the soil. Remaining in the parasitized acorn is a mass of soft
frass. Acorns, both parasitized and unparasitized, quickly desiccate. Foresters in
rearing oaks find it necessary to pluck the mature acorns from the trees, in order to
insure that the oak embryo is viable.

In the laboratory if the parasitized acorns are kept moistened D. melanogaster, D.
immigrans and D. pseudoohscura females readily oviposit into the acorn, via the
parasite’s exit hole and/or the base of the acorn where the vascular elements of the
acorn are located. Normal adult flies eventually emerge from such acorns.

D. pseudoohscura and persimilis are widespread in California. At lower elevations
(e.g. Davis, elevation 50 ft.), they can be collected throughout the year but the
populations are small during the summer and early fall months. Both species reach
peak population size during the late winter and spring, i.e., during the time of major
precipitation.

Dr. William Marshall provided me with a liter of acorns which he had collected
during February and March 1983 at Oakmont, California, elevation 250 ft. From
these 20 S S and 12 9 9 obscura spp. and 4 d d and 8 9 9 of Z). immigrans emerged.
This suggests that at the lower elevations in California acorns during the late winter
and early spring probably serve as ovipositional sites.

Basically, Drosophila adults have pigmentary patterns that blend with the
background of their feeding-ovipositional substrates, e.g. D. virilis group species on
fermenting bark, D. mulleri group species on fermenting cacti. The melanistic
coloration of the obscura species suggested that perhaps their ovipositional
substrates might be on or in the forest soil. Dobzhansky once remarked that he could
regularly sweep individuals from the forest floor under oak trees at Mather. A
number of species of subterranean fungi exist in the California forests but the
number of fruiting bodies are few in comparison to the fungi (mushrooms) whose
fruiting bodies are above the surface of the soil. Rodents avidly feed on such fungi
and can detect the presence of the hidden fruiting bodies, but human collectors must
find them by digging up areas of the forest floor. Such digging at various sites in the
forest, totalling more than 100 sq. meters, produced six fruiting bodies representing
two specimens each of three different species. Five of the fungal bodies were bisected
to determine that they did not contain larvae when collected. These were then
immersed in moist sand in one-liter glass jars and allowed to decay. Adults of either
D. pseudoohscura or D. persimilis were then added to the jars. One species of fungus
was totally ignored by the females of both species. D. pseudoohscura females readily
oviposited into the other two species of fungi and more than 150 adults were reared
from the fungi. If the fungal material was completely immersed in the sand the
females then oviposited their eggs into the sand overlying the fungal body and the
emerging larvae then migrated to the fungal material.

D. persimilis essentially ignored the fungi. No Fi persimilis emerged from any of
the fungi although dissection of one fungal mass showed a few larvae to be present
but none matured to adulthood.

VOLUME 63, NUMBER 3

255

In sum, fungi, acorns and slime fluxes can serve as larval substrates for species of
the obscura group, but the size of the populations in the Blodgett Forest and
elsewhere indicates that there must be other, as yet undiscovered, endemic larval
substrates.

Summary

Adults of thirteen species of Drosophila were collected in the Blodgett Forest
during the years 1981-84. Eight of these species, i.e. D. azteca, miranda, persimilis,
pseudoobscura, californica, occidentalis, pinicola and subfunebris are residents.
Adults of the other five species, i.e. D. busckii, melanogaster, simulans, immigrans,
and montana are migrants into the forest.

Numerically the four species of the obscura group are the dominant species in the
forest. Intensive search was made for the larval substrates of these species. A few
slime fluxes on Quercus kelloggii were found but none was used by the flies. Most of
the acorns of Q. kelloggi are parasitized by either a coleopteran or a lepidopteran.
Such acorns, if they are kept moist after the parasite larva has emerged from the
acorn, are readily used by D. persimilis and D. pseudoobscura as larval substrates.
Further, some subterranean fungi also can serve as larval substrates for D.
pseudoobscura but these fungi were not attractive to D. persimilis.

Acknowledgments

This study was supported in part by grant GM2222-1. Forest Manager Robert C.
Heald and Robert Timoni of the Blodgett Forest and Ms. Laurie Barr of the
Department of Genetics, provided invaluable assistance and advice.

Literature Cited

Carson, H. L. 1951. Breeding sites of Drosophila pseudoobscura and D. persimilis in the transition zone
of the Sierra Nevada. Evolution 5:91-96.

Carson, H. L., E. P. Knapp, and H. J. Phaff. 1956. The yeast flora of the natural breeding sites of some
species of Drosophila. Ecology 37:538-543.

Spieth, H. T. and W. B. Heed. 1975. The Drosophila pinicola species group. The Pan-Pacific Entomol.
51:287-295.

Stalker, H. D. and W. P. Spencer. 1939. Four new species of Drosophila, with notes on the funebris
group. Ann. Ent. Soc. Amer. 32:105-112.

Sturtevant, A. H. 1923. New species and notes on synonymy and distribution of Muscidae acalypteratae
(Diptera). Amer. Mus. Novitates 76:1-12.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 256-257

On the Rearing of Microchridium minutum and Its Probable
Host —Ammoplanellus (Ammoplanellus) umatilla
(Hymenoptera: Chrysididae, Sphecidae)

Jeffrey A. Halstead

Department of Biology, California State University Fresno, Fresno, California
93740.1

Abstract. —Rearing information for Microchridium minutum and Ammoplanellus
subg. Ammoplanellus umatilla from old Andricus quercuscalifornicus (Cynipidae)
galls on valley oak is presented. The probable parasite-host relationship between M.
minutum and A. umatilla is noted.

During a study of insects inhabiting old Andricus quercuscalifornicus (Bassett)
galls (Hymenoptera: Cynipidae), more commonly referred to as “oak apples,”
fourteen specimens of Ammoplanellus subg. Ammoplanellus umatilla Pate and a
female specimen of the monotypic Microchridium minutum Bohart were reared. The
purpose of this paper is to present this rearing information and to note the probable
parasite-host association between M. minutum and A. umatilla.

The wasps emerged from 250 A. quercuscalifornicus galls which were collected
from a large, isolated valley oak (Quercus lobata Nee) in a residential backyard
located at 4557 East Dakota, Fresno, Fresno County, California. The galls ranged
from 2.5 to 10 cm (1 to 4 inches) in diameter, most possessed a few to many insect
emergence holes of various sizes, and were located at a height of 2 to 3.3 m (6 to 10
feet) upon branches near the trunk of the oak. The galls were collected on 18 January
1982 and held outdoors in a sealed cardboard box until 27 December 1982, then the
box was opened and examined for insects that failed to emerge. The Microchridium
minutum, named so because of its small size (about 2 mm), and A umatilla (about 3
mm) were dead lying amongst the galls on the bottom of the box. This data
represents the first rearing records for both M. minutum and A. umatilla.

Bohart and Kimsey (1980) indicated the habitat of minutum as desert areas of
Arizona and California. However, it was also recorded from Nebraska, Nevada, and
Baja California, Mexico and thought possibly to be widespread west of the 100th
meridian (Bohart 1980, Bohart and Kimsey 1982). The only biological information
for minutum involves collection data on mats of Euphorbia (Bohart and Kimsey
1982).

No biological information is recorded for the five North American species of
Ammoplanellus, though Pate (1945) proposed that the two species in the subgenus
Ammoplanellus may nest in pre-existing holes or crannies and provision their
burrows with Thysanoptera (thrips) as do Ammoplanus.

Over 40 years ago, Pate (1945) noted an undescribed genus of minute chrysidid

Tresent address: 2110 N. Hayes, Fresno, California 93722.

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257

wasp that was flying about holes in a chair from which Hesperorhipis mirabills Knull
(Coleoptera: Buprestidae) had emerged. Many A. xila Pate were also flying about
these holes and presumably were nesting within the buprestid burrows. Though no
specimens were reared, the observer indicated that the chrysidid was possibly a
parasite of A. xila. This chrysidid was not described because the specimen Pate
received was in poor condition (Pate 1939), but it is possible that it was a specimen of
the recently described M. minutum.

It is postulated that A umatilla nests in old A. quercuscalifornicus galls in
pre-existing holes and M. minutum enters active nesting holes to parasitize it.
Additional rearing and observation will be needed to substantiate this association.

Acknowledgments

I wish to thank R. M. Bohart, Department of Entomology, University of
California Davis, for determining Microchridium minutum and N. J. Smith, Fresno
County Agricultural Commissioner’s Office, Fresno, California, for determining
Ammoplanellus umatilla. I am also grateful to N. J. Smith and D.J. Burdick,
Department of Biology, California State University Fresno, for editorial comments
on this paper. I especially thank Mrs. Hindsburger for allowing access onto her
property.

Literature Cited

Bohart, R. M. 1980. New genera and species of North American Chrysididae. J. Kansas Ent. Soc.
53:132-136.

Bohart, R. M. and L. S. Kimsey. 1982. A synopsis of the Chrysididae in America north of Mexico. Mem.
Amer. Entomol. Instit. No. 33, 266 pp.

-and-. 1980. A generic synopsis of the Chrysididae of America north of Mexico. J. Kansas

Ent. Soc. 53:137-148.

Pate, V. S. L. 1945. 'HoX.qs on Ammoplanus. Pan-Pacific Ent. 21:81-88.

-. 1939. Studies in the Pemphredonine Wasps II. Records and descriptions of new forms in the

Ammoplanoid complex from Southwestern United States. Trans. Amer. Ent. Soc. 64:373-420.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 258-264

Probing Behavior of Aphis helianthi (Homoptera: Aphididae) and Its
Preference for Pittosporum tobira Leaves of Different Ages

G. P. Walker

Department of Entomology, University of California, Riverside, California 92521
USA

Abstract. —A natural infestation of Aphis helianthi Monell exhibited a bimodal
distribution on actively growing shoots of Pittosporum tobira Aiton. Peaks of
abundance occurred on the youngest and oldest foliage. Aphids were also common
on the mature leaves of year-old shoots that were producing new growth at their
apices. Preference tests on leaf disks indicated a strong attraction to young leaves but
not to mature or senescent leaves. Probing behavior was observed on young and
mature leaves and duration of test probes was significantly longer on mature than on
young leaves (median = 44.0s vs. 20.5s; p = 0.0002). The cuticle and outer
epidermal cell walls of P. tobira were significantly thicker on mature than on young
leaves (cuticle: 3.14pL vs. I.OSjjl; cell walls: 6.60|jl vs. 1.75|jl; p < 0.0001). The results
of the probing behavior experiments were not consistent with previous reports of
probing behavior on Citrus and probable causes for the difference are discussed.

Although there have been many studies comparing aphid probing behavior among
host and non-host or resistant and non-resistant plants, there have been few studies
comparing probing behavior among different aged leaves of the same plant species.
In one such study on Citrus sinensis (Linnaeus) Osbeck (Zettler et al. 1969), the
duration of test probes by four species of aphids {Aphis gossypii Glover, A.
spiraecola Patch, A craccivora Koch, and Myzus persicae [Sulzer]) was extremely
brief on mature leaves (over 60% of the probes were only 1-15 sec. in duration) in
contrast to longer probes on succulent, immature leaves. Because of the extreme
brevity of these probes, the authors concluded that some quality of mature citrus
cuticle and/or outer cell wall of the epidermis was repellent to probing aphids and
that this quality was either chemical in nature or merely an impenetrability of the
thick mature cuticle. This differs from the conventional explanation that leaf age
preference by aphids is primarily a function of nutritional quality of phloem sap
(Kennedy 1958). Since there is a paucity of probing behavior studies on leaves of
different ages, it was of interest to determine if aphid preference of young over
mature leaves could be associated with repellency by mature cuticle and/or outer
epidermal cell walls in other plant species. This would be of particular interest in
plants with traits similar to citrus that might be associated with thick, tough mature
leaf cuticles: evergreen perennials with long-lived leaves that thrive in xeric
conditions of southern California. Pittosporum tobira Aiton, a common ornamental
plant in southern California, meets these criteria. An infestation ot Aphis helianthi
Monell on P. tobira in Riverside, California, provided an opportunity to determine
1) if this aphid exhibited leaf age preference on P. tobira’, 2) if nonpreference of a leaf

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259

age was associated with very brief test probes; and 3) if the duration of test probes
was correlated with cuticle thickness.

Materials and Methods

Natural distribution. —A single P. tobira plant infested with A. helianthi was used
for examining the distribution and leaf age preference of this aphid. This particular
plant, unlike most ornamental P. tobira, was not shaped into a dense hedge and was
therefore ideal for this study because it had long, actively growing terminals with a
variety of leaf ages present on the same terminal. Effects of leaf age on the natural
distribution of A. helianthi were evaluated during March 1985. Aphids were counted
on the upper and lower surfaces from all leaves, from base to apex, on each of 6
actively growing terminals and each of 3 terminals from the previous year’s growth.
These 9 terminals comprised the bulk of the infestation on this plant.

Leaf age preference. —Aphids from this natural infestation were used to evaluate
their leaf age preference. A cork borer (1.3 cm diameter) was used to cut leaf disks
from leaves that were young (Y) (light green; not fully expanded; from near the apex
of an actively growing terminal), mature (M) (fully expanded; dark green;
hardened), or senescent (S) (mostly green but beginning to turn yellow on the upper
leaf surface; lighter green on lower surface than mature leaves). Only one disk was
cut per leaf. In each test, two leaf disks from each of two leaf ages were placed adaxial
side down on moistened filter paper in a 5 cm diameter glass Petri dish. The 4-leaf
disks were arranged in a square pattern so that leaf ages alternated and the edges of
adjacent disks were in contact. Using an aspirator, 4 apterous aphids of various
instars were carefully placed on each of the 4-leaf disks per Petri dish. There were
four such dishes per test for a total of 64 aphids/test. The tests were conducted at
room temperature and after 3.5-4.5 hrs. (except in one test: ca. 8.5 hrs.) numbers of
aphids on each leaf disk were recorded.

Probe duration. —The durations of probes by A. helianthi on young and mature
leaves were recorded using leaves collected from several field grown P. tobira plants.
Leaves with intact petioles were collected by removing them from twigs and
immediately immersing them in water. All tests were completed within 12 hrs. of
picking the leaves. Leaves were placed individually in 1-dram shell vials filled with
water and saturated cotton. The petioles were immersed in the saturated cotton and
the blades extended erect above the vials. Parafilm was used to seal the opening of
the vials around the protruding leaves such that the vials were water tight. The vials
were held by a test-tube holder on a ring stand and could be easily rotated to facilitate
microscopic viewing of the aphids on both leaf surfaces. Immediately prior to use, a
thin barrier of petroleum jelly/mineral oil was placed on the base of the leaves to
confine the aphids on the leaves.

Aphids were collected from a field plant in the morning (ca. 9-11 a.m.) and were
held in glass Petri dishes without a food source until they were used in the experiment
(two 10-min. periods between 1 p.m. and 10 p.m.). Using an aspirator, aphids
(apterous adults and final nymphal instars) were carefully placed on the leaves and
were observed for 10 min. with a Wild M5A stereomicroscope. A Volpi Intralux 5000
fiber optic ringlight provided a uniform and cool light source. The duration of each
probe on the abaxial surface was recorded. If a probe started prior to the end of the
10-min. observation period and extended beyond it, the full duration of the probe

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PAN-PACIFIC ENTOMOLOGIST

was recorded. The duration of a probe was defined as the time which the aphid was in
characteristic “probing posture”; labium extended perpendicular to the body plane
and its tip touching the leaf surface. After the 10-min. testing period, each aphid was
returned to a glass Petri dish without food. The next aphid was then tested on the
alternate leaf age. After all aphids were tested once, each aphid was retested using a
leaf age that was opposite to that used in its first test. There were 8 replicates, each
conducted on a different day with different aphids. The first 4 replicates were
conducted in April-May 1985 and the second 4 replicates in February-March 1986.
The time involved in setting up the experiments resulted in only 5-9 aphids being
used in any one replicate.

Duration of aphid probes (abaxial surfaces only) were analysed two ways. First,
for each replicate, the duration of probes recorded on each leaf age was compared by
the Mann-Whitney U test. Second, data from the 8 replicates were pooled and the
median probe duration for each aphid on each leaf age was calculated. Median probe
durations on young and mature leaves were paired for each aphid and were
compared using the Fisher distribution-free sign test (Hollander and Wolfe 1973).
Since test probes and not prolonged feeding probes were of major interest in the
study, probe durations > 5 min. were truncated and given values of 5 min. prior to all
analyses (such probes accounted for only 3% of all probes and had negligible effect
on the medians).

Cuticle thickness. —Young and mature P. tobira leaves were collected for
histological sectioning on 18,19, and 23 April and 22 May 1985 (the same time period
that replicates 1-4 of the probe duration experiment were conducted). These were
fixed in FAA, embedded in paraffin, sectioned at 10|jl, and stained with hemalum
and safranin. The thickness of the cuticle and outer epidermal cell walls on the lower
leaf surfaces were measured with an ocular micrometer at lOOOX.

Results

Natural distribution. —On actively growing terminals, A. helianthi tended to be
bimodally distributed with respect to leaf position (leaf age) on the terminals. Peaks
of aphid abundance occurred at the basal (oldest) and/or apical (youngest) leaf
positions. Fig. 1 illustrates the aphid distribution on an actively growing terminal
where the bimodality was pronounced. The growth pattern of P. tobira is such that in
the spring, groups of several new actively growing terminals tend to rise from the
apex of a terminal from the previous year’s growth. In addition to being present on
actively growing terminals as described above, A. helianthi was also found in large
numbers on some of the previous year’s terminals that had actively growing terminals
arising from their apices. The leaves on the previous year’s terminals ranged from
senescent at the base to mature at the apex. The hmited number (3) of the previous
year’s terminals that were found infested with aphids prevents conclusions from
being made, but it is important to note that aphids were found in large numbers on
mature foliage of these terminals. In each of the 9 terminals that were examined, A.
helianthi was found in greater numbers on lower leaf surfaces than on upper surfaces.
In 7 of these 9 terminals the difference was significant (p < 0.05, Wilcoxon signed
rank test).

VOLUME 63, NUMBER 3

261

Figure 1. Numbers of A. helianthi found on leaves along the length of a long actively growing P. tobira
terminal. Leaves are numbered from base to apex; number of aphids is plotted in a log scale (#
aphids +1).

Leaf age preference .—Table 1 shows that leaf disks cut from young leaves were
preferred over those from mature or senescent leaves. In tests where no young leaf
disks were included (M vs. S, Table 1), few aphids settled on any disk; most were on
the moist filter paper or on the sides and top of the glass Petri dish. There was no
consistent preference observed in the M vs. S tests although significantly more aphids
were found on the mature disk than on the senescent disk after 4 hours in replicate 2
of the M vs. S comparison (Table 1). However, after 11 hours in this replicate there
was no significant difference between the two ages of leaf disks (5 aphids were on
each). The results presented in Table 1 did not significantly change over time in any
of the other tests.

Probe duration .—Median probe duration of A. helianthi was greater on mature
leaves than on young leaves in all 8 replicates (Table 2). The difference was
significant at the p = 0.10 level in 7 of the 8 replicates and significant at the p = 0.05
level in 5 of the 8 replicates. When median probe durations on young and mature
leaves were calculated for each aphid and analysed by paired analysis (pairing by
aphid), probes on mature leaves were significantly longer (p < 0.05) than those on
young leaves for the pooled data of each year of testing and the pooled data of both
years combined (Table 2).

Cuticle thickness .—Histological examination of young and mature P. tobira leaves
revealed that on lower leaf surfaces, the cuticle was significantly thicker on mature
than on young leaves (3.14|jl ± 0.13 vs. 1.08(x ± 0.20; mean ± SE; p < 0.0001, t test).
In addition, there was also a significantly thicker outer cell wall of the lower
epidermal cells on mature leaves as compared to young leaves (6.60fx ± 0.21 vs.
1.75|jl ± 0.47; p< 0.0001, t test). The differences between lower leaf surfaces of
young and mature leaves is illustrated in Fig. 2.

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PAN-PACIFIC ENTOMOLOGIST

Table 1. Aphis helianthi preference for Pittosporum leaf disks cut from young (Y), mature (M), and senescent (S)
leaves

Number of settled aphids^

Comparison

Leafage Rep. 1

Rep. 2

Rep. 3

Yvs.M

Y 42

36

29

* * *

* * *

* *

M 5

3

2

Yvs. S

Y 42

48

-

* * *

* * *

—

S 8

6

-

M vs. S

M 9

15

-

n.s.

-

S 6

4

-

^Total of 64 aphids (assorted instars) in each replicate; only those that settled on a leaf disk were used

for analysis. Significance levels based on the binomial distribution and Hq = Pq = 0.5 are:
* 0.01 < p < 0.005; n.s. not significant.

*** p < 0.0002;

Table 2. Median probe duration of A. helianthi on young and mature leaves of P. tobira.

Median probe duration (sc

;c.) and (N)^

Significance

Young leaf

Mature leaf

level

Replicate 1

20.0 (22)

54.0 (25)

. 0087b

Replicate 2

16.0 (26)

40.0 (23)

.0003b

Replicate 3

32.0 (43)

48.5 (32)

.0002b

Replicate 4

24.5 (32)

39.0(30)

. 0275 b

Pooled rep. 1-4

23.5 (29)

52.0 (29)

.0207=

(1985 tests)

Replicate 5

22.0 (29)

23.5 (26)

. 9194 b

Replicate 6

14.5 (34)

35.0(14)

.OOOP

Replicate 7

18.5 (46)

36.0 (30)

.0607b

Replicate 8

23.0 (43)

42.0 (36)

.0982b

Pooled rep. 5-8

19.0(31)

35.0(31)

.0011=

(1986 tests)

Pooled rep. 1-8

20.5 (60)

44.0 (60)

.0002=

“N = number of probes for Mann-Whitney comparisons and N = number of aphids for Fisher
distribution-free paired comparisons.

•^Mann-Whitney U-test.

Tisher distribution-free sign test.

Discussion

Natural distribution .—The bimodal distribution of A. helianthi on actively
growing terminals of P. tobira (Fig. 1) was not unexpected since this distribution
pattern is commonly seen in the Aphididae as a group (Kennedy 1958). However, it
was unexpected to find large numbers of aphids on mature leaves from some of the

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263

Figure 2. Lower leaf surface of P. tobira leaves. A; mature leaf. B: young leaf, c; cuticle, w: cell wall.

previous year’s terminals that bore new, actively growing terminals at their apices.
Perhaps these leaves were mobilizing nitrogen to supply the new, growing terminals
and were consequently more nutritious for the aphids.

Leaf age preference. — A. helianthi showed a distinct preference for leaf disks cut
from young P. tobira leaves vs. mature or senescent leaf disks (Table 1). This finding
offers at least partial explanation for the great abundance of A. helianthi observed on
young P. tobira leaves in nature (Fig. 1). Aphids did not readily settle on senescent
leaves even though aphid abundance appeared to increase on senescent leaves in the
field. However, Kennedy and Booth (1951) demonstrated that as leaves progress
from the beginning of senescence to complete senescence, the preference of aphids
for these leaves changes. Thus, the failure of the preference tests to detect preference
of senescent over mature leaves may be a result of my criterion for choosing
senescent leaves (slightly yellowed) not corresponding to the criterion that aphids
presumably was (high amino acid concentration in the phloem). Alternatively,
preferences observed on intact plants are not always detectable when using excised
plant parts (Risch 1985) and may explain the discrepancy between the natural
distribution and the preference tests. Nonetheless, both the natural distribution and
preference tests suggested preference for young over mature leaves. Therefore, the
probing behavior was studied in detail on these two age classes.

Probe duration and cuticle thickness. —The duration of probes by A. helianthi was
significantly greater on mature P. tobira leaves than on young leaves. In contrast,
Zettler et al. (1969), observed that probe duration of four species of aphids on Citrus
sinensis was shorter on mature leaves than on young leaves. It is important to
emphasize that most of the probes in both this study and in Zettler et al. (1969) were
“test probes” and not phloem feeding probes since most were less than one minute
and aphids require approximately a minute to penetrate the epidermis and
considerably longer to reach the phloem (Pollard 1977). Both plant species are
characterized by possessing a thin immature leaf cuticle and a thick mature leaf
cuticle {P. tobira: Fig. 2, Citrus sinensis: Walker, unpublished observations).

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PAN-PACIFIC ENTOMOLOGIST

Therefore, the mere greater thickness of mature cuticle compared to immature
cuticle is not likely to be, by itself, the cause for the extremely brief probes observed
on mature Citrus. This contention also is supported by Zettler et al. (1969) who
observed that fully expanded Crotalaria spectabilis Roth leaves (which had cuticles
similar in thickness to mature Citrus leaves) did not elicit the large proportion of brief
probes seen on mature Citrus leaves. They did not compare young and mature
Crotalaria leaves.

Clearly, the process of discrimination between young and mature leaves differs
between aphids feeding on P. tobira and those on C. sinensis. I hypothesize that A.
helianthi does not discriminate between young and mature P. tobira leaves until its
stylets penetrate the cuticle and it can sample the internal constituents of the leaf,
and that the greater duration of test probes on mature leaves is a consequence of the
extra time required to penetrate the extremely thick mature cuticle and outer
epidermal cell wall. This hypothesis is congruous with the widely held belief that
aphid leaf age preference is primarily a function of the nutritional quality of the leaf.
However, in the case of citrus, aphids probably discriminate between young and
mature leaves before their stylets penetrate the cuticle and hence mature leaves are
rejected after very brief test probes. In citrus, the epicuticular wax or other surface
components may contain probing repellents or lack required probing stimulants
(Klingauf et al. 1978, Jordens-Rottger 1979).

Acknowledgments

I thank J. D. Hare, M. P. Parrella, and J. T. Trumble for their helpful review of
this manuscript and T. Kono for his identification of the aphids.

Literature Cited

Hollander, M. and D. A. Wolfe. 1973. Nonparametric Statistical Methods. John Wiley and Sons, New
York. 503 pp.

Jordens-Rottger, D. 1979. The role of phenolic substances for host-selection behavior of the black bean
Aphis fabac. Ent. exp. andappl. 26:49-54.

Kennedy, J. S. 1958. Physiological condition of the host-plant and susceptibility to aphid attack. Ent. exp.
and appl. 1:50-65.

Kennedy, J. S. and C. O. Booth. 1951. Host alternation and fecundity in Aphis fabae Scop. I. Feeding
preferences and fecundity in relation to the age and kind of leaves. Ann. Appl. Biol. 38:25-64.
Klingauf, F., K. Nocker-Wenzel, and U. Rdttger. 1978. The role of cuticle waxes in insect infestation
behavior. (In German with English summary.) Z. Pflanzenkr. Pflanzenschutz 85:228-237.

Pollard, D. G. 1977. Aphid penetration of plant tissues. In K. F. Harris and K. Maramorosch (eds.).
Aphids as Virus Vectors. Academic Press.

Risch, S. J. 1985. Effects of induced chemical changes on interpretation of feeding preference tests.
Entomol. exp. and appl. 39:81-84.

Zettler, F. M., M. O. Smyly, and I. R. Evans. 1969. The repellancy of mature citrus leaves to probing
aphids. Ann. Ent. Soc. Amer. 62:399-402.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 265-269

A New Genus of Naucoridae (Hemiptera) from the Philippines,

with Comments on Zoogeography

Dan a. Polhemus and John T. Polhemus

University of Colorado Museum, 3115 S. York St., Englewood, Colo. 80110

Abstract. —The genus Sagocoris Montandon is endemic to New Guinea with the
supposed exception of a single species, S. usingeri, described by La Rivers from
Luzon. A recent examination of the type of this species reveals that it is not
congeneric with S. biroi, the type-species of Sagocoris. A new genus,
Philippinocoris, is thus proposed to hold the Philippine taxon, and notes are given on
the relationships of the Philippine naucorid fauna to that of New Guinea and Asia.

We thank Dr. R. T. Schuh of the American Museum of Natural History (AMNH)
for the loan of the types of S. usingeri, and Dr. Wayne Gagne for the loan of material
held in the Bishop Museum, Honolulu (BPBM). All additional material is held in the
J. T. Polhemus collection, Englewood, Colorado (JTPC). Measurements are given
in millimeters. This research was supported in part by a grant from the National
Geographic Society, Washington, D.C.

Philippinocoris n. gen.

Description. —Form oblong with sides subparallel, widest across abdomen (Fig.

1 ).

Head broad, produced ahead of eyes for over .25 the length of an eye, anteclypeal
margin gently rounded, vertex produced for short distance behind eyes, posterior
margin gently curving; eyes twice as long as wide, with small lateral flange; labrum
large, well developed, arising behind anterior margin of anteclypeus; maxillary
plates vertically oriented, tips pointed, exceeding base of labrum; rostrum arising
well behind anterior margins of eyes when viewed laterally, occupying cavity formed
by maxillary plates and carinate margins of tectiform gula; antennae with segment I
short, segment II enlarged, segments III and IV slender, covered with long setae.

Pronotum evenly domed, not sulcate medially behind head, lateral margins
broadly rounded, posterolateral angles obtuse. Scutellum shallowly and transversely
sulcate basally, with several backwardly angling shallow sulci apically to either side
of smooth medial surface. Hemelytra complete, attaining base of genital segment,
surface shining, finely rugose, bearing granular white microstructure; embolium,
clavus and claval vein well defined, embolar margin not expanded, nearly straight,
lacking setae.

Abdomen with connexivum evenly rounded, posterolateral angles not prominent,
posterior margin of tergite V in males weakly sinuate, posterolateral angles of tergite
VI in females angled sharply downward.

Ventral surface with prosternum carinate medially, exposed for entire length,
anteromedial portion angled forward over base of gula (Fig. 2); propleural plates not
touching medially, widely separated by exposed prosternum, posterior margins and

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PAN-PACIFIC ENTOMOLOGIST

Figures 1-6. Philippinocoris usingeri (La Rivers). 1. Male, dorsal habitus. 2. Prosternal keel, lateral
view. 3. Prothorax, right side as viewed ventrally. 4. Male left paramere. 5. Male right paramere. 6. Fe¬
male subgenital plate (setiferation omitted).

flanges bordering fore coxal cavities fringed with gold setae (Fig. 3), inner portions
adjoining fore coxal cavities darkened, pruinose; mesosternal plate reflexed
anteromedially, tumescent posteromedially, tumescence bearing long gold setae;
metasternal plate glabrous, cruciform, with sharp longitudinal carina medially;
abdominal venter thickly clothed with gold hydrofuge pile, paratergites bearing
paired elongate glabrous depressions adjacent to spiracles.

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267

Legs with fore tarsus single segmented, bearing single minute claw, middle and
hind tibia bearing numerous short stout spines along anterior margins, long fine
swimming hairs present on middle and hind femora, tibiae and tarsi, parempodia
setiform.

Male genitalia with asymmetrical parameres of roughly equal size (Figs. 4, 5);
vesica slender, tapering, asymmetrical, sclerotized, coming to acute point apically.

Discussion

Philippinocoris n. gen. is most closely related to Stalocoris and Asthenocoris, all
three genera being endemic to the Philippines. These genera share a common plan in
the male genitalia, possessing a slender, tapering, asymmetrical sclerotized vesica
and distinctively shaped left parameres of much the same form as those found in the
genus Naucoris. In the Papuan Sagocoris, by contrast, the vesica, though still
asymmetrical, bears a broadly expanded, partially membranous lobe apically and the
left paramere is reduced to a small truncate stub, character states also found in the
closely related Papuan genus Aptinocoris. Additional characters present in
Philippinocoris and absent in Sagocoris are the large dark pruinose area on the inner
portion of the popleura adjoining the fore coxae, the fringes of gold setae along the
posterior margin of the propleura and on the flange adjoining the fore coxal cavity,
the mesosternal tumescence covered with long gold setae and the strongly carinate
cruciform metasternal plate. In Sagocoris the rostrum arises ahead of the anterior
margin of the eye when viewed laterally, while in Philippinocoris the rostrum is set
farther back, arising behind the anterior eye margin. Abdominal asymmetry is
present in both sexes of Sagocoris, the males having a distinct offset notch in the
posterior margin of tergite V and the females of most species having the
posterolateral angles of tergites VI and VII prolonged on the left side when viewed
from above. In Philippinocoris the posterior margin of tergite V in males is only
weakly sinuate, lacking a deep notch, and the female tergites are symmetrical.

Philippinocoris is quite closely allied to Stalocoris, from which it may be separated
by the greater degree of prolongation of the head in front of the eyes. In Stalocoris
the head is produced beyond the eyes for only .11 the length of an eye while in
Philippinocoris it extends anteriorly for .27 of the eye length. In addition the apices
of the maxillary plates in Stalocoris do not exceed the base of the labrum as in
Philippinocoris, due to the labrum being recessed less deeply beneath the
anteclypeus, and the posterior margins of the propleurae lack a fringe of gold setae.
Stalocoris species are also much smaller, averaging 6 mm. in length while specimens
of Philippinocoris, at over 10 mm., are nearly twice as large.

As noted above, the three endemic genera of Philippine Naucoridae appear to be a
closely related monophyletic group derived from the Naucorinae. One can trace a
progressive modification of head structures from the primitive state in Naucoris, in
which the labrum and rostrum arise essentially at the front of the head, to Stalocoris,
in which both are slightly recessed, and on through Philippinocoris to Asthenocoris’,
in this latter taxon the labrum is recessed well under the anteclypeus and the rostrum
has moved far back into a cavity on the underside of the head. The head structure in
Asthenocoris is similar to that encountered in the Asian Cheirochelini and was
considered evidence for a relationship between the Philippine and Indochinese taxa
by Usinger (1938), but this resemblance is more likely the result of convergent
evolution since the Cheirochelini have a distinctive symmetrical vesica and
parameres quite unlike anything seen in the Philippine genera, as well as numerous

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PAN-PACIFIC ENTOMOLOGIST

other synapomorphies distancing them from any Philippine or New Guinea species
(D. Polhemus, in press). The endemic Philippine taxa are distinguished by the
anterior projection of the prosternum, which angles forward over the base of the gula
when viewed laterally (Fig. 2); this character state is not shared with the New Guinea
taxa, in which the prosternal keel and gula meet evenly or show a weak projection of
the gula posteriorly over the prosternum.

Although the naucorid faunas of the Philippines and New Guinea are allied, they
appear to represent independent insular radiations from a common ancestral taxon
near the present day Naucoris. Neither fauna shows an exceptionally strong
relationship with either continental Asia or Australia. New Guinea is geologically
younger than the Philippines but the degree of generic differentiation there has been
far higher, producing seven endemic genera as compared to only three in the
Philippines. This may be in part attributable to competition from other naucorid
lineages in the latter region. Aphelocheirus has diversified extensively on Luzon but
is represented by but a single species in New Guinea. In the southern Philippines
laccocorine naucorids, the dominant subfamily on the Asian continent and through
the Greater Sunda Islands, have managed to invade Mindanao via the Sulu
Archipelago but occur no farther north than Leyte, and have not differentiated,
being represented only by the abundant Laccocoris hoogstraali, a species closely
related to taxa present in north Borneo; this subfamily does not occur on New
Guinea.

Philippinocoris is known only from Luzon, which harbors an extremely rich and
entirely endemic naucorid fauna; most of the distinctive Philippine elements appear
to have arisen here and dispersed southward through the archipelago, but their
influence is not felt beyond Mindanao. The large island of Celebes, lying between the
Philippines and New Guinea, has a surprisingly impoverished naucorid fauna
containing only the widespread genera Naucoris and Aphelocheirus despite an
abundance of favorable habitats, a situation also pertaining in Australia. Why these
two areas should not have developed radiative faunas similar to those found in the
Philippines and New Guinea is puzzling and indicates that the latter two areas may
have had a geological association predating the present juxtaposition of land masses
in Wallacea.

Philippinocoris usingeri (La Rivers) New Combination

Sagocoris usingeri La Rivers, 1970:167.

The original description of the species by La Rivers (1970) is quite complete, but
was based only on male specimens. We now have female examples, which are quite
similar to the males in general structure and coloration. The subgenital plate is
trapezoidal with a small, deep notch apically (Fig. 6), and the posterolateral angles of
abdominal tergite VI are sharply downturned, usually to a greater degree on the left
side when viewed from above. All specimens so far examined, of both sexes, have
complete hemelytra but only partially developed hind wings and would appear to be
incapable of flight. At present P. usingeri has been found only in the Cordillera
Central of northern Luzon, where it inhabits cold, crashing, unpolluted rocky
streams. The insects were found only in a few of the many streams sampled, but then
in great numbers clinging to submerged vertical rock faces or root tangles along the
margins of protected pools sheltered from the full force of the current, a habitat
similar to that preferred by the related genus Stalocoris.

VOLUME 63, NUMBER 3

269

Material examined. —PHILIPPINES, Luzon, Benguet Prov.: 13 males, 20
females, 3 nymphs, rocky stream nr. km. 230 on Kennon Rd., 20 km. S. of Baguio
City, 613 m. (2000 ft.), vii.8.85, CL 1966, J. T. and D. A. Polhemus (JTPC); 52
males, 33 females, 1 nymph, stream below Camp John Hay hydro plant, S. of Baguio
City, 920 m. (3000 ft.), vii.8.85, CL 1968, J. T. and D.A. Polhemus (JTPC).
Mountain Prov.: 2 males (holotype and paratype), Benguet, viil5.46, D. G. Frey
(AMNH). Ifugao Prov.: 1 male, 1 immature, Jacmal Bunhian, 24 km. E. of
Mayoyao, 800-1000 m. (2624-3280 ft.), iv.25.67, H. M. Torrevillas (BPBM).

Literature Cited

La Rivers, 1. 1970. A new Philippine Sagocoris (Hemiptera: Naucoridae). Pan-Pac. Entomol.
46:167-169.

Polhemus, D. A. (in press). A review of the genus Coptocatus Montandon (Hemiptera: Naucoridae).
Pan-Pac. Entomol.

Usinger, R. L. 1938. The Naucoridae of the Philippine Islands (Hemiptera). Phil. Jour. Sci. 64:299-311.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 270-275

A Review of the Genus Nesostethus Kirkaldy
(Hemiptera: Lygaeaidae: Lygaeinae)

G. G. E. ScUDDER

Department of Zoology, University of British Columbia, Vancouver, B.C.,
Canada V6T2A9.

Abstract. —The genus Nesostethus Kirkaldy is reviewed, and the known species,
N. niger China and N. ornatus Kirkaldy, are redescribed. Four new species are
described: N. bipartitus and N. variegatus from Fiji, N. fuscus from the New
Hebrides, and N. lunatus from Tonga.

Kirkaldy (1908) described Stalagmosthethus ornatus as a new species from Fiji
(Rewa) and placed it in a new subgenus that he named Nesostethus. He characterized
this by color and by the following structural features: fourth labial segment longer
than third; scutellum with T-shaped carina; hemelytra with strong veins; pronotum
with keel, but this obsolete anteriorly.

China (1930) elevated Nesostethus to generic rank and described a new species N.
niger from Samoa. However, he was unsure of the generic placement, stating “the
present Samoan species has been referred to it, in spite of the fact that there are
certain differences which Stal and Kirkaldy have regarded as of subgeneric
importance. This course seems to be preferable (for the time being at any rate) to
that of creating still another genus.” Since this time, there has been no reference to
the genus, except for the citation in Slater (1964).

In my ongoing study of the Pacific Lygaeinae, I have needed to clarify the identity
of the genus. In the study of available material, I have found four new species. I
herein redescribe the genus, the known species and the new taxa.

Nesostethus Kirkaldy

Stalagmostethus sg. Nesostethus Kirkaldy 1908, Proc. Linn. Soc. N.S.W. 33:355.
Nesostethus', China 1930, Ins. Samoa, Pt. 2, Hemipt. 3:115.

Moderate to large (7.7-11.1 mm), variously colored orange, red and black insects;
macropterous, body matt with head, lateral margins of pronotum, costal margin of
corium, legs and abdomen shiny. Dorsum at most with short, curved, decumbent
pubescence.

Head (Fig. 1) moderately declined, and in line with pronotum; wider than long;
eyes not stylate; eyes close to anterior margin of pronotum; vertex gently convex and
smooth; ocelli closer to eyes than to each other; antennal tubercles visible from
above; bucculae prominent anteriorly, tapering posteriorly, and extending to level of
middle of eye. First antennal segment surpassing apex of head by about half of
length; third segment gradually thickening apically; fourth antennal segment usually
slightly longer than second. Rostum reaching to or just beyond hind coxae; first
segment slightly thicker than second and just surpassing anterior margin of
prosternum; fourth segment longer than third.

270

VOLUME 63, NUMBER 3

271

Figures 1-5. Nesostethus ornatus. 1. Head and anterior part of pronotum. 2. Caudal view of male genital
capsule. 3. Right clasper. 4. Spermatheca. 5. Second gonocoxa and gonopophysis. Scale line = 0.50 mm
for Fig. 1, 0.33 mm for Fig. 2 and 0.165 for Figs. 3, 4 and 5.

Pronotum at most weakly punctate anteriorly; anterior margin without collar,
concave in centre and appearing rather sinuate; posterior margin straight or slightly
concave, with shallow impression on either side of scutellum; lateral margins
rounded, straight and somewhat convergent anteriorly; disc gently convex, with
median longitudinal carina behind callal area, and slight transverse impression

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PAN-PACIFIC ENTOMOLOGIST

one-third way from anterior; callal area flat or gently convex, the calli sinuate and
slightly oblique.

Scutellum flat with tumid T-shaped carina; laterally impunctate and fovea not
distinct; apex roundly pointed. Ostiolar peritreme auriculate.

Hemelyta reaching just beyond end of abdomen; costal margin straight or gently
sinuate; clavus and corium with veins strongly raised; basal third of costal margin and
claval commissure hirsute.

Legs unarmed; coxae hirsute. Abdomen without laterotergites, but with sternum
II exposed; sterna III-VI with anterior dorso-lateral “apertures.” Male genital
capsule and parameres as in Figs. 2-3; spermatheca as in Fig. 4; second gonapophysis
with upturned tip (Fig. 5).

Type species: Stalagmostethus (Nesostethus) ornatus Kirkaldy 1908 (Monobasic).

Structurally the head and pronotum are similar to Melanerythrus Stal, but
Nesostethus differs by having less elevated bucculae, strongly raised venation on the
corium and clavus, the fourth rostral segment longer than the third, and the
scutellum with a T-shaped tumid elevation rather than a longitudinal central carina.

Although nothing is known about the biology, specimens of N. ornatus have been
found to contain cardenolides in very high concentrations (Scudder and Duffey
1972). The genus may thus be associated with Asclepiadaceae and the abdomen
appears structurally capable of releasing cardenolides as in Oncopeltus fasciatus
(Scudder and Meredith 1982).

Nesostethus bipartitus New Species

Male. —Head, dorsally and ventrally, red with apex of clypeus black; antennae
black; rostrum black with first segment and base of second, orange-red. Pronotum
orange-red with humeral angles and median longitudinal streak on hind margin,
black. Scutellum orange-red. Clavus black; corium black with costal half of apical
two-thirds red; membrane black with apical margin narrowly white. Prosternum,
thoracic pleura and ostiolar peritreme, orange-red. Coxae orange-red; rest of legs
black. Abdomen red. Head width 1.60 mm; antennal measurements 0.77:2.13:1.67:
? mm; rostrum reaching to posterior margin of hind coxae. Pronotal width 2.47 mm,
pronotal length 1.58 mm. Anterior half of pronotum punctate. Corium with very
sparse, short decumbent pubescence; costal margin of corium sinuate. Thoracic
pleura with scattered, short, semi-erect hairs. Total length 8.8 mm.

Holotype. — S , FIJI, Matuku I., 5.vii.24 (E. H. Bryan, Jr.) (Bishop Museum).

Similar to the species herein described as N. variegatus, but differing in the color
pattern of the corium and membrane.

Nesostethus fuscus New Species

Female. —Insects completely black, except for ostiolar peritreme and abdomen
which are yellow; anterior dorso-lateral corners of sterna III-VI with small fuscous
spots. Head width 1.70 mm; antennal measurements 0.83: 2.17: 1.60: 2.23 mm;
rostrum attaining hind coxae. Pronotal width 3.07 (2.95-3.07) mm; pronotal length
2.00 (1.90-2.00) mm. Pronotum anteriorly vaguely punctate; corium with scattered,
short, decumbent pubescence; costal margin of corium sinuate. Thoracic pleura with
distinct decumbent, sericeous pubescence. Total length 10.5 (9.8-10.5) mm.

Male. —Color and structure as in female. Head width 1.60 (1.53-1.70) mm;

VOLUME 63, NUMBER 3

273

antennal measurements 0.74 (0.67-0.83): 2.05 (1.93-2.20); 1.53 (1.50-1.60): 2.18
(2.10-2.74) mm. Pronotal width 2.58 (2.37-2.80) mm; pronotal length 1.71
(1.57-1.90) mm. Total length 8.5 (8.3-9.3) mm.

Holotype. —$ , NEW HEBRIDES, Maewo I. Sounwari, 15°23'S 168°07'E, 0-400
m, 4-5.ix.79 (W. C. Gagne) (Bishop Museum).

Paratypes. — 26 1 9 , NEW HEBRIDES (but labelled Solomon Is.), Epi (H. W.
Simmonds); IM, NEW HEBRIDES, Malekula, Ounua, ii.29 (L. E. Cheesman);
IM, NEW HEBRIDES, Espirito Santo, 12.iii.43; 1 $, NEW HEBRIDES, Epi I.,
Valmali, 80-150 m, ll-18.viii.67 (J. & M. Sedlacek); (Bishop Museum; British
Museum [Nat. Hist.]; Scudder Coll.).

This species can be separated from N. niger by the moderately dense, decumbent
pubescence to the corium and thoracic pleura, the sinuate costal margin, and the
yellowish ostiolar peritreme and abdomen.

Nesostethus lunatus New Species

Male. —Head, dorsally and ventrally orange-red, with apex of clypeus and
V-shaped mark between ocelli, black; antennae black; rostrum fuscous with most of
first segment orange-red. Pronotum and scutellum orange-red. Corium and clavus
completely black; membrane black with apical margin narrowly white, and basal
angle with white area. Prosternum, thoracic pleura and ostiolar peritreme,
orange-red. Legs black with coxae orange-yellow. Abdomen orange-yellow.

Head width 1.50 mm; antennal measurements 0.67: 1.90: 1.53: 2.03 mm; rostrum
reaching to middle of hind coxae. Pronotal width 2.23 mm, pronotal length 1.42 mm.
Pronotum vaguely punctate. Corium with sparse,'short, decumbent pubescence;
costal margin of corium almost straight. Thoracic pleura with scattered, short,
semi-erect hairs. Total length 7.7 mm.

Holotype. — 6, TONGA IS., Vavua I., Neiafu, ii.56 (N. Krauss) (Bishop
Museum).

Similar to N. ornatus, but differing in the coloration of the membrane.

Nesostethus niger China

Nesostethus niger China 1930, Ins. Samoa, Pt. 2, Hemipt. 3:113.

Female. —Insects completely black, except for posterior and dorsal areas of
metapleuron and ostiolar peritreme, which are dusky orange, and abdomen, which is
orange-red.

Head width 9 1.75 mm; antennal measurements 9 0.34: 2.50: 1.80: 2.30 mm;
rostrum reaching to hind coxae. Pronotal width 9 2.65 mm, pronotal length 9 2.00
mm. Pronotum anteriorly vaguely punctate. Corium almost glabrous, with at most
scattered, short, decumbent pubescence, the costal margin straight. Thoracic pleura
virtually glabrous. Total length 9 9.3 mm.

Material examined: 19 (type) SAMOA, iii-viii.21 (F. W. O’Connor) (British
Museum [Nat. Hist.]).

N. niger can be recognized by the predominantly black color and orange-red
abdomen, with the corium rather glabrous and the costal margin straight. The
species herein described as N. fuscus differs in having the costal margin of the
hemelytra sinuate and surface of corium and thoracic pleura with a moderately
dense, short, decumbent pubescence.

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PAN-PACIFIC ENTOMOLOGIST

Nesostethus ornatus Kirkaldy

Stalagmosthethus (Nesostethus) ornatus Kirkaldy 1908, Proc. Linn. Soc. N.S.W.
33:355.

Nesostethus ornatus; China 1930, Ins. Samoa, Pt. 2, Hemipt. 3:115.

Head, dorsally and ventrally orange-red, with apex of clypeus and v-shaped area
between ocelli, black; antennae black; rostrum black with basal half of first segment
orange-red. Pronotum and scutellum orange-red, the humeral angles of pronotum
sometimes slightly dusky. Hemelytra, including membrane, completely black.
Prosternum, thoracic pleura and ostiolar peritreme, orange-red. Legs black, with
coxae orange-yellow. Abdomen pale greenish.

Head width S 1.65 (1.60-1.70) mm, $ 1.79 (1.70-1.85) mm; antennal
measurements d 0.73 (0.70-0.75): 2.08 (2.00-2.15): 1.60 (1.55-1.70): 2.33
(2.20-2.45) mm, $ 0.78 (0.75-0.80): 2.32 (2.25-2.35): 1.73 (1.70-1.75): 2.15 mm;
rostrum reaching to middle of hind coxae. Pronotal width S 2.50 (2.30-2.65) mm, 9
3.06 (2.80-3.30) mm; pronotal length 6 1.65 (1.55-1.70) mm, $ 2.01 (1.85-2.15)
mm. Pronotum anterior vaguely punctate; corium almost glabrous; costal margin of
corium sinuate. Thoracic pleura with scattered, short, semi-erect hairs. Total length
d 8.4 (8.2-8.5) mm, 9 10.8 (10.5-11.1) mm.

Material examined. —19, FIJI, Cuvu, l.i.l7 (R. Veitch) (BM); Id, FIJI IS.,
Levuka, v.21 (H. W. Simmonds) (British Museum); Id 19, FIJI, Ovalau, v.22
(H. W. Simmonds) (BM); Id, FIJI IS., Viti Levu, Colo-i-Suva, 28.vi.24 (E. H.
Bryan, Jr.) (Bishop Mus.); 19, FIJI, Colo. Emb., 10.iii.42 (R. A. Lever) (British
Museum); 19, Ins. Viti (Stockholm).

Similar to the species herein described as N. lunatus, but with a completely black
membrane.

Nesostethus variegatus New Species

Male. —Head, dorsally and ventrally, orange-red with apex of clypeus black;
antennae black; rostrum black, the first segment orange-red with extreme apex
black. Pronotum and scutellum orange-red. Clavus black; corium with basal half
black and apical half red; membrane black with apical third fading through
ochraceous to pale apical margin, the basal angle with a pale white triangular mark.
Prosternum, thoracic sterna and ostiolar peritreme orange. Coxae orange;
trochanters dusky orange; rest of legs black. Abdomen red.

Head width 1.53 mm; antennal measurements 0.67: 1.87: ?: ? mm; rostrum
reaching to sternum 11. Pronotal width 2.00 mm, pronotal length 1.33 mm. Pronotum
anteriorly vaguely punctate; thoracic pleura with scattered, short, semi-erect hairs;
corium with scattered, short, decumbent pubescence; costal margin of corium
sinuate. Total length 7.3 mm.

Holotype. —d, FIJI, Lau, Fulanga, 5.viii.24 (E. H. Bryan, Jr.) (BishopMuseum).
Museum).

Similar to the species herein described as N. bipartitus, but differing in color
pattern of the corium and membrane.

Key to Species of Nesostethus

1. Head, pronotum, scutellum, thoracic pleura and coxae, black . 2

— Head, pronotum, scutellum, thoracic pleura and coxae, orange-red . 3

VOLUME 63, NUMBER 3

275

2. Dorsum more or less glabrous; costal margin of corium straight; abdomen

red. niger China

— Dorsum with distinct, decumbent, sericeous pubescence; costal margin of

corium sinuate; abdomen pale yellow . fuscus Scudder

3. Corium completely black . 4

— Corium not completely black . 5

4. Membrane completely black . ornatus Kirkaldy

— Membrane black with margin narrowly pale, and with white triangular area at

basal angle . lunatus Scudder

5. Corium with basal half black, apical half red . variegatus Scudder

— Corium black with costal half of apical two-thirds red ... bipartitus Scudder

Acknowledgments

The research for this paper was supported by the Natural Sciences and
Engineering Research Council of Canada. I am indebted to the late E. Kjellander
(Naturhistoriska Rijksmuseum, Stockholm), G. Nishida (B.P. Bishop Museum,
Honolulu) and W. R. Dolling and the Trustees of the British Museum (Nat. Hist.)
for the loan of material.

Literature Cited

China, W. E. 1930. Insects of Samoa and other Samoan terrestrial Arthropoda. Pt. II, Hemiptera. Ease.
3:81-162. British Museum (Nat. Hist.), London.

Kirkaldy, G. W. 1908. A catalogue of the Hemiptera of Fiji. Proc. Linn. Soc. N.S.W. 33:345-391.
Scudder, G. G. E. and S. S. Duffey. 1972. Cardiac glycosides in the Lygaeinae (Hemiptera: Lygaeidae).
Can. J. Zool. 50:35-42.

Scudder, G. G. E. and J. Meredith. 1982. Morphological basis of cardiac glycoside sequestration by
Oncopeltus fasciatus (Dallas) (Hemiptera: Lygaeidae). Zoomorphology 99:87-101.

Slater, J. A. 1964. A Catalogue of the Lygaeidae of the World. University of Connecticut, Storrs. 2 vols.,

1668 pp.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 276-277

Rhyacionia zozana (Lepidoptera: Tortricidae), Host of Hockeria
tenuicornis (Hymenoptera: Chalcididae) in Oregon

Jeffrey A. Halstead and Christine G. Niwa

(JAH) 2110 N. Hayes, Fresno, California 93722; (CGN) Pacific Northwest
Research Station, Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis,
Oregon 97331.

Abstract. —A new parasite-host record of Hockeria tenuicornis (Girault) from
Rhyacionia zozana (Kearfott), a pest of pines, is presented. This information
represents the first host report and a new state record (Oregon) for Hockeria
tenuicornis.

Hockeria tenuicornis (Girault) is one of ten species of Hockeria Walker that occur
in North America (Halstead, unpublished). This wasp, described by Girault, (1918)
is known from Arizona, California, and Montana (Burks 1979). The hosts only three
North American species of Hockeria have been recorded: larvae of Myrmeleon
immaculatus DeGeer and Myrmeleon sp. (Neuroptera: Myrmeleontidae) for
Hockeria eriensis (Wallace); pupae of Neodiprion excitans Rohwer and Neodiprion
sp. (Hymenoptera: Diprionidae) for Hockeria unipunctatipennis (Girault); and
Harrisina brillians Barnes and McDunnough (Lepidoptera: Zygaenidae) for
Hockeria rubra (Ashmead) (Burks 1979).

The genus Rhyacionia contains several species that are economically important
pests of pines (Pinus spp.) in nurseries, ornamental plantings, and in natural and
planted reforestation projects (Powell and Miller 1978). One such species,
Rhyacionia zozana (Kearfott), attacks various species of pines {Pinus ponderosa,
jeffreyi, contorta, edulis, monophylla, monticola, and flexilis) in the western United
States (Stevens et al. 1980, Powell and Miller 1978). The parasites of Rhyacionia
have been the subject of many studies (see Harman and Kulman 1973, Yates 1967).

During studies on the parasite complex of Rhyacionia zozana, five female
specimens of Hockeria tenuicornis were reared. One wasp was obtained from a
collection of 729 third and fourth instars and the other four specimens from a
collection of about 7,000 cocoons. All wasps emerged from Rhyacionia zozana
cocoons the spring after their collection.

One voucher specimen (#001) of Hockeria tenuicornis is deposited in the United
States Museum of Natural History, Washington, D.C. The rearing data are as
follows: Cave Mountain, Klamath County, Oregon, exit Rhyacionia zozana on
ponderosa pine, Pinus ponderosa, 18.V.85, Christine Niwa collector.

These data represent two new records for Hockeria tenuicornis, a host in North
America, and its occurrence in Oregon (new state record).

Acknowledgments

The authors wish to thank D. J. Burdick, Department of Biology, California State
University Fresno, Fresno, California; N. J. Smith, Fresno County Agricultural

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Commissioner’s Office, Fresno, California; and R. D. Haines, Tulare County
Agricultural Commissioner’s Office, Visalia, California, for editorial comments on
this paper.

Literature Cited

Burks, B. D. 1979. Chalcididae. Pp. 860-874. In K. V. Krombeinet al. (eds.). Catalog of Hymenopterain
America North of Mexico. Vol. I. Smithsonian Inst. Press, Washington, D.C., 1198 pp.

Girault, A. A. 1918. New and old West Indian and North America chalcid-flies (Hym.). Ent. News
29:125-131.

Harman, D. M., and H. M. Kulman (eds.). 1973. A world survey of the parasites and predators of the
genus Rhyacionia, Partsito IV. Univ. of Maryland Nat. Resour ces Inst., Contrib. No. 527,179 pp.
Powell, J. A., and W. E. Miller. 1978. Nearctic pine tip moths of the genus Rhyacionia: biosystematic
review (Lepidoptera: Tortricidae, Olethreutinae). USDA For. Serv., Agric. Hdbk. No. 514, 51

pp.

Stevens, R. E., C. Sartwell, T. W. Koerber, G. E. Daterman, L. L. Sower, and J. A. Powell. 1980.
Western Rhyacionia (Lepidoptera: Tortricidae, Olethreutinae) pine tip moths trapped using
synthetic sex attractants. Can. Entomol. 112:591-603.

Yates, H. O. 1967. Key to nearctic parasites of the genus Rhyacionia: with species annotations. USDA
For. Serv., Southeast For. Exp. Stn., Asheville, N.C., 127 pp.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 278-283

A New Microcalyptris Species from California
(Lepidoptera: Nepticulidae)

David L. Wagner

Department of Entomology, California Academy of Sciences, Golden Gate Park,
San Francisco, California 94118

Abstract. —A new species of Microcalyptris Braun, is described from Antioch,
California. The adult, larva and mine are illustrated. The larval chaetotaxy of a
Microcalyptris species is figured for the first time. The larva forms a serpentine mine
in the stem epidermis of Lotus scoparius (Nutt, in T. & G.) Ottley. Stem mining has
not been reported for Microcalyptris.

The eight described species of Microcalyptris are restricted to North America
(Wilkinson 1979). A new species, Microcalyptris lotella is described from Antioch
Dunes National Wildlife Refuge, which is a remnant of a once extensive dune system
along the San Joaquin River near its confluence with the Sacramento River. The new
species shares most characters of the genus as defined by Davis (1978) and Wilkinson
(1979). Unlike other Microcalyptris species, M. lotella is a stem epidermis miner.

Microcalyptris lotella New Species

Description. —External features for male and female (Fig. 1): Forewing length
2.0-3.3 mm. Vertex and frons whitish to tan, scales occasionally dark-tipped; eyecap
large, nearly concolorous with vertex. Antenna 42-45 segmented; flagellum shiny
dark brown. Dorsum of thorax and forewing irrorated with grayish scales, darkened
apically; fringe tan. Hindwing shiny whitish gray; basal third covered with
ochraceous androconia; fringe tan. Forewing venter uniformly silver gray-brown,
basal third with ochraceous androconia between costal and anal veins. Abdomen
with shiny grayish buff scaling. Legs smooth, grayish. Venation (Fig. 2): Forewing
with six veins reaching the wing margin (nomenclature follows Wilkinson 1979);
radius three-branched (Rl, R2 + 3 and R4-h5), R4 + 5 ending at wing tip; median
unbranched, arising from R4 + 5; cubitus absent; anal vein bent basally. Hindwing
narrow; five veins reaching wing margin.

Male Genitalia (Figs. 3-5): Tegumen (pseuduncus) hoodlike enclosing the uncus,
posterior margin truncate to shallowly emarginate, bearing 20-26 curved seate.
Uncus with dorsal knob; gnathos broadly V-shaped with a large medial, dorsally
projecting spine. Lateral support rods (Davis 1978) elongate triangular, obliquely
truncate posteriorly. Valva simple, elongate, length ca. 4X width; inner surface
shallowly concave bearing numerous setae, setae thickened distally. Juxta (Fig. 4)
simple, distal portion very lightly sclerotized. Vinculum narrow medially; lobes of
saccus about as long as broad. Transtilla well developed, fused, mesad. Aedeagus
(Fig. 5) large, elongate, slender; distal end with two subequal, strongly pigmented,
curving processes; cornuti increasing in size toward insertion of larger distal process;

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Figure 1. Adult male of Microcalyptris lotella.

lateral tooth arising at about bearing numerous minute spines at its base;
phallobase elongate, broadly rounded.

Female Genitalia (Fig. 6): posterior apophyses long, nearly 2X length of anterior
apophyses. Corpus bursae narrowly ellipsoid, with two large signa that run entire
length of bursa, composed of quadrangular spinose plates. Vesicle large, elongate,
broadest anteriorad; distinct vestibule intercalated between vesicle and helical
ductus spermathecae.

Diagnosis .—Fore wings uniformly gray, irrorate. Male with androconia over basal
third of DHW and VFW. Saccus lobes as broad as long; juxta weak, simple; tegumen
(pseuduncus) with 20-26 curved hairs; valva bearing thickened apical setae;
aedeagus with two large apical processes, a lateral tooth and large cornuti. Female
with two very large signa running length of corpus bursa.

Holotype. —Male, California, Contra Costa Co., Antioch Dunes National
Wildlife Refuge, 15.hi.85, D.L. Wagner, reared from Lotus scoparius, imago
issuing 14.iv.85, male genitalic slide DLW 86-01, in the Essig Museum of
Entomology, University of California, Berkeley.

Paratypes. —California, Contra Costa Co., Antioch Dunes National Wildlife
Refuge, 15.hi.85, D. L. Wagner, J. A. Powell and J. W. Brown collectors, reared
from Lotus scoparius, imagos issuing 7.iv-13.v.85, 42 males and 57 females. DLW
Slide Numbers: larva: 85-22 and 85-23; wing: 85-27, 85-33, 85-40,85-41 and 85-46;
male: 85-45, 86-02 and 86-23; and female: 85-47, 86-06, 86-08, 86-39 and 86-40.

Ovum: Egg laid on epidermis of upper shoots of Lotus scoparius. Ovum oval,
packed with reddish purple to reddish brown frass, elevated above the epidermal
tissue.

Larva (Figs. 7,8): Fully mature larva elongate, 6.5-7.5 mm in mine, contracting
upon removal; yellow with dark brown head capsule. Cranial nomenclature and
chaetotaxy follows Gustafsson (1981). Caudal margin of head capsule deeply

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PAN-PACIFIC ENTOMOLOGIST

Figures 2-6. Microcalyptris lotella: 2. Wing venation. 3. Male genitalia. 4. Aedeagus. 5. Juxta. 6.
Female genitalia.

emarginate, lateral lobe extending posteriorad > 1.5 x basal width at tentorial
bridge, caudal margin of lobe distinctly thickened; epistomal ridge quadrangular,
bulging anteriorad, posterior margin convex; tentorial arms reduced, anterior arms
< 0.5 width of bridge; labial palpus 2-segmented, basal segment subequal to apical.
Prothorax with two weak dorsal sclerites, all setae present; mesothorax with two SV

^ An

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281

Figure 7. Head capsule of Microcalyptns lotella, ventral aspect.

1 .Ocm

\ —

9

Figures 8, 9. Microcalyptris lotella. 7. Larva. 8. Mine.

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PAN-PACIFIC ENTOMOLOGIST

setae, L3 absent; ambulatory warts on segments T2, T3 and A2-6. Abdominal
segments 1-8 with six setae; A9 with three setae and AlO with two (Fig. 8).

Mine (Fig. 9): Early portion densely packed with dark frass; later frass deposited
as broad central band, lateral areas remaining frass-free. Mine elongate, 3.5-7.3 cm
in length, making one to three switchbacks; track rarely anastomosing except in very
narrow shoots. Larva mines in green bark, usually in shoots one to three years of age;
rare or absent from areas of reddish bark or the most recently formed shoots. Larva
fully visible beneath epidermis, leaving the mine through a crescent-shaped slit to
pupate in surface litter.

Cocoon: Cream to fuscous, loose, broadly pyriform.

Discussion

Microcalyptris lotella appears to be most closely related M. punctulata (Braun),
the only other member of the genus recorded from California. Shared features
include the comparatively large signa and the uniformly pale irrorate forewings (the
male genitalia of M. punctulata have not been described or illustrated).

The wing venation of M. lotella differs from that of M. scirpi and other
Microcalyptris species in lacking a vestigial Cu vein in the forewing. The juxta is
simple and very weakly developed in M. lotella, unlike some members of the genus
which possess a large, relatively complex juxta (e.g., M. bicornutus Davis). The
lateral support rods characteristic of the genus are well developed in this species.

Little is known about the life history of any Microcalyptris species. All apparently
form serpentine tracks, and leave the mine prior to pupation. Microcalyptris scirpi
Braun is a leaf miner in the blades of Scirpuspaludosus A. Nels. (Braun 1925). Braun
(1910) reared M. punctulata (Braun) from mines in the leaves of Ceanothus cuneatus
(Hook.) Nutt. M. lotella is unusual in that the larva is a stem miner, feeding in the
green epidermal layers of Lotus scoparius. Among the Nearctic nepticulids only one
other genus, Ectoedemia Busck, is known to mine in bark. Curiously, in the
Palearctic region, the stem epidermis of Lotus is mined by unrelated nepticulids in
the genus Trifurcula Zeller.

Presumably this species is single brooded as I have been unable to locate active
mines from April through July.

The type locality, Antioch, California, is a unique sand dune habitat along the San
Joaquin River. The once extensive dune system at Antioch was home to many
endemic plants and insects (Powell 1981). What remains of this threatened habitat is
now protected as a National Wildlife Refuge managed by the United States Fish and
Wildlife Service. Although M. lotella is presently known only from this unique
locality, abandoned mines which may represent this species were found near Briones
Reservoir in Contra Costa County and near Virner, El Dorado County, California.

Stilbosis extensa Hodges (Cosmopterigidae) was reared from the same collections
that produced Microcalyptris lotella. The larvae of S. extensa were frequently found
boring in the pith of the upper shoots, beneath mines of the Microcalyptris.

Acknowledgments

I am grateful to Dr. J. A. Powell for reviewing an earlier draft of this paper.

VOLUME 63, NUMBER 3

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Literature Cited

Braun, A. F. 1910. New species of Tineina from California. Entomol. News 21:171-179.

-1925. Microlepidoptera of northern Utah. Tran. Am. Entomol. Soc. 51:13-17.

Davis, D. R. 1978. New leaf-mining moths of the family Nepticulidae from Florida. Florida Entomol.
61:209-224.

Gustafsson, B. 1981. Characters of systematic importance in European Nepticulidae larvae
(Lepidoptera). Entomol. Scand. 12:109-116.

Powell, J. A. 1981. Endangered habitats for insects: California coastal sand dunes. Atala 6:41-55 [1978].
Wilkinson, C. 1979. A taxonomic study of the micro-lepidopteran genera Microcalyptris Braun and
Fomoria Beirne occurring in the United States of America (Lepidoptera, Nepticulidae). Tijdschr.
V. Entomol. 122:59-90.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 284-291

Life History of Trupanea conjuncta (Adams) on Trixus californica
Kellogg in Southern California (Diptera: Tephritidae)

Richard D. Goeden

Department of Entomology, University of California Riverside, Riverside,
California 92521.

Abstract.—Trupanea conjuncta is monophagous on the desert shrub, Trixus
calif ornica (Asteraceae), in southern California. Each female oviposits an average of
five eggs in a single cluster in a single immature flower head each day. The larvae feed
and develop gregariously and pupariate clustered in central cavities in flower heads.
This tephritid may be either univoltine or bivoltine; consequently, the adults may
live as long as a year. Unique among Trupanea spp. and other flower head-infesting
Tephritidae as known to date, this species is a facultative gall former. If flower heads
are absent or rare from a lack of local rainfall, gravid females may oviposit in apical
buds and the larvae develop to maturity gregariously in the galls so induced. Thus,
another reason for gall formation by insects has been discovered, i.e., as an
alternative mode of reproduction and development by a flower head-infesting
species. Egg resorption also may have evolved as a mechanism for extending the
ovipositional period and host-searching capacity of this fly. Eurytoma vernonia
Bugbee (Eurytomidae) and Pteromalus purpurieventrus (Ashmead) (Pteromalidae)
are reported as probable, primary, hymenopterous parasites of the larvae and
pupae.

Little heretofore was known about Trupanea conjuncta (Adams), except
taxonomically. I recently studied this distinctive tephritid in conjunction with a
faunistic survey of the desert shrub, Trixus calif ornica Kellogg (Asteraceae), in
southern California (Goeden and Ricker, unpubhshed data).

Taxonomy. —First described as Urellia conjuncta by Adams (1904). T. conjuncta
additionally was described and illustrated in part by Malloch (1942) (as a Trypanea),
Foote (196()a) and Foote and Blanc (1963) (as a Trupanea).

Distribution and hosts. —Before publication of the initial host-plant rearing record
for this tephritid from flower heads of T. calif ornica (Goeden 1983), T. conjuncta was
described as “rarely collected” from only a few locations in Arizona and California
(Foote 1960, Foote and Blanc 1963). This fly presumably also ranges into Mexico like
its host-plant (Shreve and Wiggens 1964), the sole representative of the genus Trixis
(Tribe Mutisieae) in the Sonoran Desert flora of Arizona (Kearney and Pebbles
1964) and California (Munz and Keck 1959, Munz 1974). Additional species of Trixis
in the Sonoran Desert in Mexico (Shreve and Wiggens 1964) represent potential
hosts.

I have reared T. conjuncta only from flower heads (and terminal-bud galls, as
described below) of T. calif ornica, among 77 genera and 182 species of California
Asteraceae sampled to date. Thus, T. conjuncta apparently is monophagous in
California (Goeden 1985). My rearing records from T. calif ornica flower heads

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collected on the dates indicated include the following: Imperial County: Indian Well,
one male and two females, 12.hi.86; Mountain Spring, two males, 20.hi.86;
Riverside County: Desert Center, 108 males and 108 females, 14.hi.84; Graham
Pass, 47 males and 41 females, 21.hi.84; San Bernardino County: Coxcomb
Mountains, 40 males and 39 females, 20.hi.84; Sheephole Mountains, 14 males and
19 females, 14.hi.84; San Diego County: Coyote Canyon, one male and one female,
29.ii.84; Yaqui Pass, three males and three females, ll.i.84. Goeden (1983)
recorded T. conjuncta from Chino Canyon, ca. 1 km NW of Palm Springs, Riverside
County, where most field observations reported herein were made and flower head
samples collected for dissection during 1983-86.

Biology

Egg. —Newly laid eggs (Fig. la) are smooth, shiny, white, and elongate
ellipsoidal, with a reduced, button-like, anterior pedicel, like those of T. bisetosa
(Coquillett) (Cavender and Goeden 1982). The posterior end narrows to a smoothly
rounded point and usually is covered and partly obscured by a whitish secretion that
glues together this end of several eggs deposited in a packet (Fig. la). Thirty-four
field-collected eggs averaged 0.77 ± 0.008 (± SE) mm in length and 0.18 ± 0.003
mm in greatest width. The pedicels averaged 0.03 mm in length and width. The eggs
of T. conjuncta are slightly longer and narrower, the pedicels slightly shorter than
those of r. bisetosa (Cavender and Goeden 1982).

Unlike the eggs of T. nigricornis (Coquillett) and T. bisetosa, which are deposited
singly (unpublished data, and Cavender and Goeden 1982, respectively), the eggs of
T. conjuncta usually are deposited in groups glued together posteriorly and along
part of their lengths. The egg clusters were easily lifted as units from the tips of the
floral tubes among the pappus hairs where the eggs usually are oviposited with their
long axes perpendicular to the receptacle surface. The “glue” loses its stickiness with
time. One or more outer eggs in a packet may also be glued to the inner curve of a
receptacle bract or to the adaxial surface of an inner phyllary. As these bracts
subsequently elongate, the attached cluster of egg chorions (by then usually empty)
is lifted upwards and away from the receptacle. All eggs in a packet are deposited
together by one female at a single insertion of her ovipositor, which usually
penetrated one or more of the leaves surrounding the young flower head and one or
two of the enveloping, alternately arranged phyllaries. The path of the ovipositor
was marked by circular punctures ca. 0.15 mm in diameter ringed by brown necrotic
tissue. Some oviposition occurred through the opening at the apex of a young flower
head where the tips of the phyllaries met. Only five (8%) of 60 infested heads
examined contained two egg packets (Fig. la), apparently laid by different females
or at least at different times, judging from the numbers and patterns of the egg
punctures as well as the different stages of development of the eggs in each pair of egg
clusters. The number of eggs in 65 clusters from field-collected heads averaged
4.5 ±0.1 (range: 3 to 7). The most eggs found in an infested head was 12. Females
preferred early-stage flower heads only 3 to 5 mm long for oviposition; this stage
lasted only two or three days in the field. Oviposition did not always follow probing,
as evidenced by punctured phyllaries of uninfested heads. Caged females were not
deterred from laying a superabundance of eggs in single buds. At least one egg mass
was observed that had been deposited atop another mass in nature. This behavior

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Figure 1. Life stages of Trupanea conjuncta. (a) Two egg clusters in young flower head, 16x. (b)
Subspheroidal gall containing puparia, 4.3x. (c) Third instars feeding gregariously in flower head, 6.4x.
(d) Cluster of empty puparia from flower head, 9x. (e) Elhpsoidal gall containing puparia, 4.3x. (f) Adult
female at rest on flower head of Trixus californica, 6.6x.

suggested that the short duration of the attractive stage of development of the young
heads, not the use of a marking pheromone as occurs with certain frugivorous
Tephritidae (Prokopy 1972), limits egg deposition in heads. Females of Tephritis
dilacerata Loew (Berube 1978) and Trupanea bisetosa (Cavender and Goeden 1972)
also apparently did not recognize or avoid immature flower heads that already
contained eggs.

The clustering of eggs by T. conjuncta apparently is an adaptation for a unique
feature of its life history, i.e., the capacity for facultative gall formation, a feature
never before reported for any flower head-infesting tephritid. No other species of
Trupanea is known to form galls; however, this behavior provides additional
ecological evidence linking this genus to the closely related genus Tephritis, several

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species of which are gallicolous (Quisenberry 1951, Foote 1959,1960a, 1960b). This
behavior may have gone undetected if an associate and I had not concurrently
surveyed the insect fauna of T. californica at various desert locations (Goeden and
Ricker, unpublished data) while I studied the life history of this fly. When surveying
plants in Grapevine Canyon, NE San Diego County, in early February, 1984,1 found
terminal bud galls on branches of plants that had not yet experienced rainfall that
winter, and probably not since the previous winter. Field and subsequent laboratory
examinations showed that these galls contained tephritid puparia from which T.
conjuncta adults subsequently were reared (Fig. lb). If gravid mature females cannot
find young flower heads generated on host plants in response to winter or late
summer rainfall, they apparently will oviposit small groups of eggs in the apical buds.
Facultative gall formation sems to be a mechanism that this flower head-infesting
tephritid has evolved to insure its reproduction for at least another generation under
conditions of inadequate rainfall or flower head availability. No galls were formed on
Trixis which flowered in February through April, 1984, in response to rainfall
received at the Chino Canyon site in January and the preceding December. Thus,
gall formation was a localized, facultative activity. However, several old, woody galls
were observed on the stems at Chino Canyon, which indicated that gall formation
also had occurred there about two growth periods beforehand. No other species of
insect produces galls on stems of T. californica in southern California (Goeden and
Ricker, unpublished data).

Twenty-four eggs hatched after six to eight days’ incubation at 27 ± 2°C and 100%
relative humidity in the laboratory. When denied their usual ovipositional sites
during solitary greenhouse cagings on non-flowering branches of Trixis, individual,
fleld-collected, sexually mature and mated females stored their eggs internally for as
long as a week. This denial of ovipositional sites occasionally resulted in the insertion
of some eggs in apical buds, the prerequisite for gall formation in nature. If immature
flower heads were resupplied to these caged gravid females after several days, eggs
were laid within the heads in large clutches, e.g., 18, 22, and 30 eggs, and various
stages of embryony, including those that hatched only one or two days after
deposition. Thus, embryony continued within the oviduct, as apparently did egg
resorption, as indicated by the oviposition of empty egg chorions together with partly
reabsorbed, flaccid eggs and viable, turgid eggs. Whether egg resorption occurs in
gravid females of T. conjuncta in nature is unknown. Conserving egg metabolites in
this manner would be another mechanism for extending the reproductive life and
host-searching capacity of this desert-inhabiting tephritid, which is solely dependent
for its reproduction on a localized, fairly uncommon, single species of host plant
growing suboptimally at the northernmost extension of its range. In January, 1985, a
severe winter frost killed the top growth and delayed or prevented flowering of many
T. californica in southern California, an event of the type which egg resorption and
the resultant extended female ovipositional period may have evolved to counter.

In flower heads, the embryo rotated 180° just before hatching, so that the
mouthparts of the first instar usually were immediately in contact with a floral tube
upon leaving the egg through a posterior-longitudinal slit in the chorion. The
duration between eclosion of the first and last individuals from a single egg clutch was
ca. one day.

Larva. —The larvae of T. conjuncta usually fed gregariously. I observed only one
instance in flower heads in which the newly hatched larvae from a single cluster had

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Split into two feeding groups that attacked separate florets. All larvae newly hatched
from an egg cluster usually entered the elongating floral tube of a single floret
together and tunneled basipetally into the immature achene. After consuming the
contents of this first floret, the larvae together transferred to an adjacent floret which
they entered through the base of its floral tube. Feeding within a series of florets
continued in this manner through the second stadium; however, most feeding and
larval growth occurred during the third stadium. The third instars fed in a central
ellipsoidal cavity (6 to 9 mm long) ca. 3 mm wide, formed among the stumps of
achenes, the scored receptacle, and the distal remnants of the pappus hairs of the
central florets excised 2 to 3 mm from their tips, all surrounded by a ring of scored or
undamaged florets (Fig. Ic). As many as 12 third instars fed in a compact mass within
a moist, central cavity, the walls of which were covered with yellowish, liquid feces
(Fig. Ic). When fully grown, the larvae pointed their mouthparts acropetally, ceased
movement, and pupariated in a compact cluster (Fig. Id).

The galls of T. conjuncta are initiated when a female oviposits in a terminal bud.
The round ovipositional scar was seen on the surface of some galls. Upon hatching,
the larvae feed on the surrounding tissues and extend the gall cavity into the pith of
the branch tip. If this feeding killed the apical meristem, the branch ceased to
elongate and a subspheroidal gall resulted (Fig. lb). If the apical meristems
remained intact, the gall assumed a spindle shape as the branch continued apical
growth (Fig. le). Twenty-one field-collected, current season’s galls measured
9.7 ± 0.6 (range: 6 to 16) mm in length and 4.6 ±0.1 (range: 3.6 to 5.5) mm in
greatest width. The larvae fed on the parenchymatous pith tissue, expanding the gall
cavity in length and width until, eventually, two to five shortened internodes were
incorporated in the fully formed galls. The central cavities of 13 fully formed galls
averaged 6.3 ± 0.4 (range: 3.3 to 9.0) mm in length and 2.6 ± 0.2 (range: 1.0 to 3.5)
mm in width. The cavities were ellipsoidal (Fig. le) or subspheroidal (Fig. lb),
smooth-walled, and free of frass. none, several, or all of the axillary buds along the
length of the gall, instead of or as well as the terminal bud, may break dormancy and
grow into branches. The vascular cylinder is incorporated in the wall of the gall,
which remains green and photosynthetic while the gall remains occupied. The fully
grown larvae cut one or two short emergence tunnels, usually laterally in the distal
half of the galls, through the 0.75-1.25 mm thick walls, leaving a thin flap of
epidermis covering each future exit hole. The larvae usually pupariate with their
head directed acropetally towards an exit tunnel. Some larvae pupariated in a
partially overlapping linear row and formed only a single, common exit hole (Fig.
le).

It is but one evolutionary step for a species to gall axillary buds in addition to
terminal buds as occurs with other genera and species of Tephritidae, e.g., Aciurina,
Procecidochares, and some Tephritis (Foote 1960b, Silverman and Goeden 1980,
Steyskal 1984). Moreover, facultative gall formation, as expressed by T. conjuncta,
indicates one means by which a flower head-infesting species of Tephritidae may
have evolved sympatrically into a gallicolous species occupying a different niche on
the same host plant, or vice versa. In a separate paper, I will describe an apparent
example of facultative gall formation in the genus Tephritis. Whether oviposition of
egg clusters and gregarious larval and pupal development are adaptations reflecting a
gall-forming ancestry or facilitating reproduction in galls by a flower head-infesting

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species, or are prerequisites thereof, I leave to evolutionary ecologists and
systematists to interpret.

Pupa .—Pupation occurs in a puparium within the central cavity in a flower head or
gall (Fig. le). The puparium (Fig. Id) is black, ellipsoidal, smoothly rounded at both
ends, but with a flattened, posterior, perispiracular plate, superficially smooth, but
very finely punctate, and slightly flattened or concave ventrally. Twenty-one
field-collected puparia measured 3.1 ± 0.1 (range: 2.3 to 3.8) mm in length by
1.3 ± 0.04 (range: 1.0 to 1.6) mm in greatest width. Adults emerged through an
anterior, two- or three-part fracture of the puparium. A very thin, whitish,
translucent pupal exuvium, left behind within the puparium, also helps to distinguish
unparasitized individuals. Adults emerged by pushing through the loose plug of
excised tips of pappus hairs of the central florets, or by breaking through the
epidermal windows covering the exit holes of galls.

Adult. —The adult (Fig. If) of both sexes is readily distinguished from other
species of Trupanea by the unbroken dark to light-brown area in the distal anterior
quarter and proximal half of the wing (Foote 1960a, Foote and Blanc 1963). Newly
emerged adults are sexually immature and apparently do not mate. Judging from the
rearing records reported above and my recovery of 394 males and 315 females
(1.25:1) from 22 flower head samples collected at Chino Canyon during 1983-85, the
sex ratio of T. conjuncta appears to be slightly male biased (X^ = 8.79, p < .005, 1
df). Males outnumbered females in 15 of the 22 samples, whereas females
outnumbered males in only five samples. T. conjuncta males were not observed to
emerge from flower heads before females, as Silverman and Goeden (1980) reported
for Procecidochares sp. from bud galls on the desert ragweed. Ambrosia dumosa
(Gray) Payne. The latter tephritid also showed a male-biased sex ratio.

The adults of T. conjuncta probably are long-lived. They apparently pass the
summer at higher elevations, moving upward along streams and washes into the
surrounding mountains as flowering ceases and their perennial hosts become
dormant. A portion may reach mountain meadows by midsummer, as occurs with
the more common, also monophagous, desert species, Trupanea imperfecta
(Coquillett) (unpublished data). As fall and colder weather approaches, the adults
migrate towards lower and warmer elevations, where they eventually rendezvous on
their host plants. I have not observed mating, which may occur in the field at this time
or earlier at higher elevations, if summer rainfall stimulates a second bloom in the
fall, these flower heads are used as oviposition sites. If their host plants remain
dormant for lack of rainfall, I suspect that rather than form galls at this time, the flies
remain inactive through the fall and early winter, awaiting the more dependable
occurrence of winter rainfall and the resulting main flowering period. Consequently,
mating either occurs in the fall in response to unknown stimuli to host flowering, or if
no rain falls, is delayed until winter, or recurs then. Both sexes were swept from
actively growing hosts before bud formation and oviposition commenced.

Oviposition was observed only once in the field at 10:21 a.m. on February 17,1984,
and lasted only 35 seconds. Oviposition is not commonly observed with this tephritid
because (1) the adults per se are uncommonly observed or swept (Foote and Blanc
1963), even in close proximity to or directly from their host plants (personal
observation); (2) most of that portion of their adult lives that is spent on or near their
hosts appears to involve resting motionless (characteristically with wings slightly

290

PAN-PACIFIC ENTOMOLOGIST

parted but mostly overlapped atop the abdominal dorsum [Fig. If]) while hidden and
sheltered within the crowns (especially when it is windy, as so commonly is the case in
the winter and early spring on the Colorado Desert); and (3) each female apparently
oviposits on the average only once a day in a single flower head. The last-named
behavior is extrapolated from the results of greenhouse cagings of field-collected,
gravid females mentioned above. Five females individually caged between March 6
to April 25, 1986, for one to 11 days each on single, flower bud-bearing branches of
potted plants laid a total 387 eggs over a total of 75 days of cagings for an average of
5.2 eggs laid per female per day. This daily oviposition rate approximated the mean
number of 4.5 eggs per cluster found in field-collected heads as reported above. The
most eggs laid by a single caged female were 137. She apparently died prematurely
and produced fertile eggs up to her death. As she probably had oviposited before
being collected, this number of eggs underestimated her fecundity.

Seasonal history. —Munz (1974) described the flowering period of T. californica as
February to April in southern California. I have reared T. conjuncta from mature
flower heads collected on May 13 and 25, June 7, and October 3,1983, in addition to
February to April, 1983, 1984, and 1985, at Chino Canyon. Thus, this tephritid and
its host plant had two reproductive periods in 1983, but only one in 1984 and 1985.

On March 7, 1985, 50, 50, 30, and 50 flower heads on four plants, respectively,
were tagged individually by slipping rubber bands attached to number tags over the
heads in the early stage favored for oviposition by T. conjuncta. The development of
each of these heads subsequently was followed and correlated with tephritid adult
emergence. From oviposition in young heads to adult emergence from mature flower
heads with faded yellow or white florets that contained fully formed achenes, the
immature stages of T. conjuncta and flower head development lasted about five
weeks under field conditions. As discussed above, the adults appear capable of living
as long as 12 or 13 months under field conditions in southern California .

Mortality factors. —The larvae of an apparently undescribed microlepidopteran,
Homeosoma sp. (Pyralidae), each destroyed the contents of several young flower
heads during the course of their development, occasionally including heads infested
by T. conjuncta. Jumping spiders (Araneida: Salticidae) and crab spiders (Araneida:
Thomisidae) appeared to be the most common potential predators of adults
observed on preblossom and flowering host plants. Two species of Hymenoptera,
both reared from flower heads of T. californica, were probable parasites of immature
T. conjuncta and two other synphagous species of Tephritidae, Tomoplagia cressoni
Aczel and Trupanea actinobola (Loew) (unpublished data). They were Eurytoma
vernonia Bugbee (Eurytomidae), probably a solitary, primary, larval or larval-pupal
endoparasite, and Pteromalus (Habrocytus) purpurieventrus (Ashmead)
(Pteromalidae), probably a solitary, primary, larval ectoparasite. Eurytoma
vernonia also were reared from puparia in apical bud galls, so adoption of this
facultative mode of reproduction by T. conjuncta does not confer “enemy-free
space” (Zwolfer 1983, Price et al. 1986); however, this could provide a measure of
relative protection from natural enemies. Both chalcidoids parasitize Trupanea
imperfecta in flower heads of Bebbia juncea (Bentham) Greene, another shrubby,
perennial, desert Asteraceae that often grows in association with T. californica. The
life history of T. imperfecta will be described in a separate paper.

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291

Acknowledgments

My thanks to D.W. Ricker for technical assistance, including the insect
photography involved in Fig. 1. Thanks also to Louie Blanc, Dick Foote, Gordon
Gordh, Earl Oatman, and John Pinto for their comments on early drafts of the
manuscript. The parasites were identified by John LaSalle, Division of Biological
Control, Department of Entomology, University of California, Riverside, and E. E.
Grissell, Systematic Entomology Laboratory, BBII, USDA, ARS, Beltsville,
Maryland. The pyralid moth was identified by D. C. Ferguson, also located at the
Systematic Entomology Laboratory.

Literature Cited

Adams, C. F. 1904. Notes on and descriptions of North American Diptera. Kans. Univ. Sci. Bull.
2:433-455.

Berube, D. E. 1978. Larval descriptions and biology at Tephritis dilacerata (Diptera: Tephritidae), a
candidate for the biocontrol of Sonchus arvensis in Canada. Entomophaga 23:69-82.

Cavender, G. L., and R. D. Goeden. 1982. Life history of Trupanea bisetosa (Diptera: Tephritidae) on
wild sunflower in southern California. Ann. Entomol. Soc. Am. 75:400-406.

Foote, R. H. 1959. A few North American species of Tephritis, with some observations on its generic
position (Diptera: Tephritidae). Bull. Brooklyn Entomol. Soc. 54:13-17.

-. 1960a. A revision of the genus Trupanea in America north of Mexico. USDA Tech. Bull. 1214,29

pp.

-. 1960b. The genus Tephritis Latreille in the Nearctic Region north of Mexico: descriptions of four

new species and notes on others. J. Kans. Entomol. Soc. 33:71-85.

-, and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect Survey

7:1-117.

Goeden, R. D. 1983. Initial host-plant records for five species of fruit flies from southern California
(Diptera: Tephritidae). Proc. Entomol. Soc. Wash. 85:399-400.

-. 1985. Host-plant relations of Trupanea spp. (Diptera: Tephritidae) in southern California. Proc.

Entomol. Soc. Wash. 87:564-571.

Kearney, T. H., and R. H. Peebles. 1964. Arizona flora. Univ. of Calif. Press, Berkeley and Los Angeles,
1085 pp.

Malloch, J. R. 1942. Notes on two genera of American flies of the family Trypetidae. U.S. Nat. Mus.
Proc. 92:1-20.

Munz, P. 1974. A flora of southern California. Univ. of Calif. Press, Berkeley and Los Angeles, 1681 pp.
Price, P. W., G. L. Waring, and G. W. Fernandes. 1986. Hypotheses on the adaptive nature of galls.
Proc. Entomol. Soc. Wash. 88:361-363.

Prokopy, R. J. 1972. Evidence for marking pheromone deterring repeated oviposition in apple maggot
flies. Environ. Entomol. 1:326-332.

Quisenberry, B. F. 1951. Astudyofthe genus Tep/inYw Latreille in the Nearctic Region north of Mexico.
J. Kans. Entomol. Soc. 24:56-72.

Shreve, F., and I. L. Wiggens. 1964. Vegetation and Flora of the Sonoran Desert. Vol. 2. Stanford Univ.
Press, 1740 pp.

Silverman, J., and R. D. Goeden. 1980. Life history of a fruit fly, Procecidochares sp., on the ragweed.
Ambrosia dumosa (Gray) Payne, in southern California (Diptera: Tephritidae). Pan-Pac.
Entomol. 56:283-288.

Steyskal, G. C. 1984. A synoptic revision of the genus Aciurina Curran, 1932 (Diptera, Tephritidae).
Proc. Entomol. Soc. Wash. 86:582-598.

Zwolfer, H. 1983. Life systems and strategies of resource exploitation in tephritids, pp. 16-30. In
Cavalloro, R., ed.. Fruit flies of economic importance. Proc. CEC/IOBC Int. Sym., Athens,
Greece, Nov. 1982, A. A. Balkema, Rotterdam.

PAN-PACIFIC ENTOMOLOGIST
63(3), 1987, pp. 292-300

The Status of Efferia similis (Williston), with Descriptions of Three
New Nearactic Efferia Species in the Albibarbis Group

(Dip ter a: Asilidae)

Gregory S. Forbes

Box 3AF, Department of Biology, New Mexico State University, Las Cruces,
New Mexico 88003.

Abstract. —The nomenclatural history of Efferia similis (Williston) and its
relationship to E. tagax (Williston) are discussed. Examination of the syntype of
similis discloses it actually to be tagax. Similis (Will.) is therefore placed as a
synonym of tagax, and the species currently known as similis is renamed neosimilis.
Two related Efferia species of the Albibarbis group (incognita from Texas and New
Mexico and sonorensis from Mexico) are described.

Efferia similis and E. tagax were described from Arizona by Williston (1885). The
identity of E. similis (based on an extremely brief description) and its taxonomic
relationship with E. tagax have long been a source of confusion and
misidentification. The confusion has been compounded by the presence in west
Texas and New Mexico of a third closely related Efferia species described below.
Strangely, Williston compared similis not with tagax, which had nearly identical wing
venation, but with a specimen near E. staminea (Will.), which was also described in
the same paper. Hine (1919), noting that similis had not been seen since its
description, justifiably considered it a synonym of tagax, but his associated discussion
of tagax suggested that his material contained more than one species. Wilcox and
Martin (1965) also treated similis as a synonym of tagax. Wilcox (1966) showed,
however, that a second species, closely related to tagax, does in fact occur in southern
Arizona. Assuming this entity to be Williston’s similis, Wilcox described or
redescribed both sexes of it and tagax, describing for the first time the relatively
uncommon "'similis'' male.

Although E. similis was described from one female (not a male as indicated by
Wilcox, 1966), Kansas University has two female “syntypes” which bear a printed
label “Arizona/C.U. Lot 35” and a handwritten orange label "Erax similis Will."
Neither of these specimens is similis auct.; one is E. triton (Osten Sacken), which on
the basis of wing venation could not have been Williston’s “type.” The other,
although the ovipositor is broken, appears to be E. tagax: the femora, trochanters,
and scutellum are reddish, and the black bands on tergites 3-6 are triangular and are
narrowed successively with each segment. This specimen agrees with Williston’s
description, including the 24 mm length (with the broken ovipositor).

E. tagax was described from a male, with a brief comparative description of two
“rubbed” females appended. The location of these “syntypes” is unknown, but the
detailed male description is recognizable. Based on the superiority of the E. tagax
description, E. similis (Will.) is placed as a synonym of E. tagax (Will.), and E.

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similis sensu Wilcox (1966) must be named. As Wilcox presumed he was redescribing
Williston’s similis (based on Williston’s type), his account does not constitute a
formal description, and this species is named below.

The three species described here, along with E. armata (Mine), E. bicolor
(Bellardi), E. grandis (Mine), and E. tagax form a distinct subgroup of the Albibarbis
group in the southwestern United States and Mexico. These species are
characterized by large size, black, bulbous, male genitalia, modified hind tibiae in
males (except in bicolor and grandis ), and shrub perching behavior, which appears to
be nearly obligate in bicolor and grandis. E. leucocoma (Williston) may also belong
here but its habits are unknown. There are a number of described and undescribed
neotropical members of this subgroup, particularly in Mexico.

Measurements in the following descriptions follow the procedures of Bullington
and Lavigne (1984), with “body length” being exclusive of the epandrium or
ovipositor.

Efferia neosimilis, New Species (Fig. 5)

(This description is in part based on Wilcox’s (1966) redescription of E. “similis, ”
in which the male hypopygium and hind tibia and female ovipositor are figured.)

Male .—Body length 18.0-25.0 mm (holotype 23.0 mm); head ground color black
to red-black, thinly white tomentose, tomentum often very thin white on gibbosity,
vertex, antennae, and ocellar tubercle, tomentum dense white between gibbosity
and compound eye margin; proboscis shining black; antenna (including base) and
area adjacent to ocellar tubercle black to red-black; hairs on scape and pedicel
mostly white, sometimes mixed black/white dorsally; ocellar tubercle black with two
black bristles and 3-5 black or mixed black/white hairs; mystax white (some bristles
yellow-white) with black hairs (7 in holotype) in upper 1/3 of gibbosity, variable black
bristles and hairs occur along lower lateral margins of gibbosity to oral margin; beard
white; palpi black with white basal hairs and black apical bristles; upper occipital
bristles white to partly black (6 and 7 black on each side in holotype).

Mesonotum black, brownish-gray tomentose, usually with a broad black stripe,
divided by gray tomentum, on either side of meson (these stripes reddish in some
specimens); humeral callosity red-brown, hairs white anteriorly, black posteriorly;
anterior mesonotal hairs black, shorter than scape, posterior hairs black on upper
slope and as long as pedicel, shorter white on lower slope; scutellum thin gray
tomentose, hairs on disk mostly black, sometimes white basolaterally, 6-10 black
marginal bristles (10 in holotype); pleura gray-white tomentose, hairs mixed black
and white; halteres yellow-brown.

Coxae and trochanters grayish white tomentose, femora black (sometimes very
dark red-black) reddish apically, hairs mostly white, bristles black; tibiae reddish
brown, nearly black at apices, bristles black, apical portion of hind tibia with
rounded dorsal and ventral swellings (Wilcox, 1966, Fig. 44), hairs long white
ventrally on basal 2/3 of hind tibia, short dense black on ventral swelling, short dense
white or orange on dorsal swelling; tarsi red-brown, bristles and hairs black, hairs
black, hairs short dense orange and black on first tarsomere of hindleg, claws black,
red basally, empodia and pulvilli light reddish brown.

Wings hyaline, light brown apically and along veins, veins brown, subcostal cell
dark brown adjacent to dilated costa.

Tergites 2-5 broadly black dorsally, laterally edged with brown tomentum.

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PAN-PACIFIC ENTOMOLOGIST

Figures 1-4. 1 Male terminalia of E. incognita. 2 Male terminalia of E. sonorensis. 3-6. Apex of
basistylus of (3) E. incognita, (4) E. sonorensis, (5)E. neosimilis, (6)E. tagax. 7 Hind tibia of E. incognita.
8 Hind tibia of E. sonorensis. Terms: ab = apex of basistylus, bst = basistylus, dst = dististylus,
ep = epandrium, sp = subapical process of epandrium.

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295

gray-white tomentose apically, dark brown at apical edges; venter, tergites 2-5
laterally, and 6-7 entirely gray-white tomentose, hairs of tergites white, longer
dorsolaterally on 2-3, short black dorsally on 2-5 and sometimes 6; erect white hairs
on sternites 1-5, short white on 6, longer white at apex of 7, sternite 8 black with
apical black hairs.

Terminalia black, hairs black, some hairs reddish dorsoapically on epandria,
ventral fringe with dense black hairs; inner margin of basistylus red-brown;
epandrium 3.2 to 3.7 mm long (3.7 mm in holotype), with a ventroapical “tooth”;
basistylus forked apically with shorter upper lobe (Fig. 5).

Female. —Body length 16.5-24.0 mm (allotype 19.0 mm); allotype with 6 black
scutellar bristles; tergite 7 black, dorsal hairs short black; sternite 7 with erect mixed
black and white hairs; ovipositor black, 3.9-4.5 mm long (4.0 mm in allotype); black
hairs and bristles in mystax reduced, most bristles yellow-white; lateral gray
tomentum on tergites 3-6 more extensive, narrowing dorsal black bands in some
specimens; subcostal cell hyaline to light brown in middle; otherwise similar to male.

Etymology. —Use of the name neosimilis in part conserves the familiar name
similis (sensu Wilcox and subsequent authors).

Diagnosis. —Male: The combination of broad black bands on the tergites,
subapical epandrial process, shape of the basistylus, and rounded hind tibial
swellings will identify E. neosimilis. Female: The very short ovipositor will readily
separate E. neosimilis from related species except for extremely small specimens of
E. bicolor. See Table 1.

Habitat. — E. neosimilis occurs in southeastern Arizona, extreme southwestern
New Mexico, and undoubtedly northern Sonora and Chihuahua, from the upper
elevation limits of saguaro-ocotillo well into oak-pine habitat. It is sympatric with E.
tagax at the base of the Santa Rita (and possibly Santa Catalina) Mts., and tends to
replace tagax at higher elevations. Both species occur on shrubs or other elevated
perches (frequently ocotillo); neosimilis perches on acacias, grass (i.e. Sorghum)
stems, manzanita, and oaks. In Pinery Canyon, Chiricahua Mts., neosimilis was
found in late afternoon on low rocks, in grassy clearings among oaks and junipers.

Holotype male and allotype. —Ruby Rd. at Summit Motorway, 4 mi. W of Pena
Blanca Lake (1424 m), Santa Cruz Co., Arizona, 31 July 1979 (G. S. Forbes).

Paratypes. —same data: 1 male, 2 females; 7 July 1985, 2 males, 10 females;
ARIZONA: Graham Co.: Stockton Pass (1648 m), 18 July 1970,1 male (S. Draper,
J. Bigelow, O. Francke, M. Cazier) (CAS). Cochise Co.: top of Huachuca Mts., 10
Aug. 1940, 1 female (E. S. Ross) (CAS); Bernardino, 23 July 1966, 3 males, 5
females (J. Davidson, M. Cazier) (1 male, 2 females from CAS); Lower Pinery
Canyon, Chiricahua Mts., 23 June 1985, 4 males, 2 females (GSF); Chiricahua Mts.,
Cave creek, 6.5 mi. W Portal, 11 July 1981, 1 female (H. Hespenheide); Peloncillo
Mts., Clanton Draw Rd. 3.1 mi. WNM line, 5 July 1985,2females (GSF); Pima Co.:
lower Madera Canyon, 200 m NE of Proctor Ranch Rd. Jet, 6-7 July 1985, 4 males,
10 females (GSF) (sympatric with E. tagax); Box Canyon Rd., 1 mi. NE Santa Rita
Ranch, 7 July 1985, 1 female (GSF) (with E. tagax). Santa Cruz Co.: Washington
Mts., 15 July 1920, 1 male (J. A. Kusche) (CAS); NEW MEXICO: Hidalgo Co.:
Clanton Draw, Peloncillo Mts., 9.5 mi. W Rt. 338 (1635 m), 5 July 1985, 2 males
(GSF). OTHER ARIZ. LOCALES: Cochise Co.: Sunnyside Cyn., Huachuca Mts.;
W. Entrance, Ft. Huachuca; Miller Cyn., Huachuca Mts.; Pima Co.: Sabino Cyn.,
Santa Catalina Mts.; N. of Gibbon Mtn. (1420 m), S. Catalina Mts.; Box Cyn., Santa

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PAN-PACIFIC ENTOMOLOGIST

Rita Mts.; Florida Cyn., S. Rita Mts.; Santa Cruz Co.: 1 mi. SW Pena Blanca Lake;
Ruby. See also Wilcox (1966).

Holotype, allotype, and paratypes labelled CAS will be returned to the California
Academy of Sciences.

Specimens reported as E. “tagax” from the Davis Mts., Texas, (Bromley, 1934)
and New Mexico are here recognized as a distinct species in the Albibarbis group.

Efferia incognita, New Species (Figs. 1, 3, 7)

Male .—Body length 19.0-23.0 mm (holotype 21.5 mm); head color andtomentum
differ from E. neosimilis as follows: head ground color black, tomentum variably thin
white or brown-white on antennae, ocellar tubercle, and vertex; antennal base,
scape, and pedicel black, apex of scape and pedicel usually reddish-brown;
flagellomere and stylus black; hairs on scape and pedicel predominantly black dorsally,
white ventrally; mystax white, with numerous black hairs on upper 1/3 of gibbosity
and a variable row of black hairs and bristles along lower margins of gibbosity to oral
margin; ocellar tubercle with two black bristles and 3-9 predominantly black hairs;
4-8 black upper occipital bristles on each side (4 and 5 in holotype).

Mesonotum brownish-gray tomentose, dorsum usually with two broad lateral
brown stripes and one narrow central brown stripe, delineated by areas of
grayish-white tomentum, mesonotal hairs and bristles black, anterior hairs subequal
in length to scape, posterior hairs as long as antenna; humeral callosity reddish, hairs
white anteriorly, black posteriorly; pleura gray-brown tomentose (thinly so on
anepisternum), hairs and bristles black (a few may be white or orange-red); postalar
callosity reddish-brown, gray tomentose; haltere brownish-white; scutellum light
brown tomentose, hairs on disk black, sometimes white basolaterally, margin of
scutellum with 6-11 black bristles (8 on holotype) and with black submarginal
bristlelike hairs, 273 - 4/5 as long as bristles.

Femora black, reddish at apex, spines black, hairs mostly white, long white
ventrally on fore femora; tibiae reddish-brown, darker at apices, with rounded
swellings distally on hind tibiae (Fig. 7); tibial bristles mostly black, hairs variable:
long white dorsally, often short dense black on ventral swelling, hairs on remainder
of tibia mostly white; short, dense, orange pile may be present on tibiae and
tarsomeres (especially first tarsomere) on fore and hind legs; tarsi dark red-brown,
tarsal spines black, claws black, reddish basally; empodia and pulvilli yellowish
white.

Wings as in E. neosimilis.

Tergites 1-5 laterally and 6-7 entirely silver-white tomentose, tergites 2-5 dorsally
dull black, the black margined laterally with brown tomentum; the posterior 1/4 to 1/3
of tergites white tomentose, brown tomentose on extreme posterior margin; tergite 2
often entirely black dorsally; hairs white laterally, mostly black dorsally; sternites
1-7 gray-white tomentose with erect white hairs; sternite 8 black with long black
apical hairs.

Terminalia black, hairs black (sometimes reddish white on apex of epandrium),
dense black on ventral margin of basistyli; epandria lacking ventral subapical
process; inner margin of basistyli reddish-brown; basistyli with a subapical rounded
lobe and dorsal triangular flange (Fig. 3). Epandrial length 3.4-4.0 mm (3.7 mm in
holotype).

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297

Female. —Body length 17.5-22.0 mm (allotype 20.5 mm); tergite 7 dark
brown-black dorsally, sometimes brown tomentose laterally; ovipositor black,
5.2-6,3 mm long (allotype 5.7 mm); lateral gray tomentum on tergites 2-5 may be
extensive, narrowing the dorsal black bands; sternite 7 black-haired; black bristles
and hairs often much reduced in mystax; allotype with 9 black scutellar bristles; hind
tibiae not swollen, dense orange pile on fore and hindlegs may be absent or much
reduced; subcostal cell variable from nearly hyaline to brown in middle, but not as
dark as in male; otherwise similar to male.

Etymology. —From the Latin for “unknown” or “hidden,” with reference to its
long confusion with E. tagax.

Habitat. — E. incognita occurs in foothill and montane habitats in northern
Chihuahua, Mexico, west Texas, and New Mexico. A disjunct population occurs in
Union Co., NM. A record from Grant Co., NM is near the range of E. neosimilis. E.
incognita is unrecorded from the Guadalupe Mts. of NM/Texas but probably occurs
there. Specimens labelled E. bicolor or E. tagax from Santa Fe, NM, are probably
incognita. The species typically occurs in oak-juniper woodland, perching on low
rocks, fallen branches, grass stems, and occasionally shrubs, although shrubs are
apparently not a preferred perching site.

Diagnosis. —Male: The shape of the apical basistylus is distinctive. The broad
bands on the tergites and rounded hind tibial swellings (Fig. 7) are shared with E.
neosimilis, but that species has the toothed processes on the epandria. E. tagax has
redder femora and weakly enlarged hind tibiae. Female: Very similar to E.
neosimilis, but with a longer ovipositor. E. tagax has generally narrower tergal bands
and redder femora. See Table 1.

Holotype male and allotype. —Davis Mts., S. side Rt. 166, 6.3 mi. SW jet. with Rt.
118 (1790 m), Jeff Davis Co., Texas, 27 June 1985 (G. S. Forbes).

Paratypes. —Same data as holotype and allotype: 13 males, 12 females; TEXAS:
Brewster Co.: Road Summit, Green Gulch, Chisos Mts., 8 June 1985, 2 males (D.
Lightfoot); Chisos Basin Campground, 29 June 1985, 1 male (D. Lightfoot); Green
Gulch, 5-6 mi. S Panther Jet. Rd., 29 June 1985, 7 males, 1 female (GSF); Chisos
Basin, 14 August 1976,1 female (L. Bezark collection). Jeff Davis Co.: Rt. 118,19.7
mi. S Kent, 2 July 1985, 1 female (GSF); Rt. 166, 6 mi. SW jet. Rt. 118 (1790 m),
19-20 June 1977, 3 males, 1 female (E. M. Fisher); Limpia Crossing, Davis Mts., 28
June 1985, 1 female (D. Lightfoot). NEW MEXICO: Grant Co.: Mimbres, 25 July
1986, Imale (E. and W. MacKay). Lincoln Co.: Nogal, 21 July 1952,1 female (H. H.
andL. D. Beamer, W. LaBerge, C. Liang) (KU); 5 mi. N. Angus (2136 m), Highway
37, 7 August 1965,1 male, 1 female (H. B. Leech) (CAS); NM214,1.7 mi. N Rt. 70
(NE of Ruidoso) (1760 m), 9 July 1986, 2 males, 2 females (GSF). Union Co.:
Clayton Lake, 12 mi. N Clayton (1545 m), 23 July 1965, 2 males (G. W. Beyers)
(KU). MEXICO: Chihuahua: Rancho Experimental La Campana (80 km N of
Chihuahua City), 27 June 1981,1 male.

The holotype and allotype are deposited in the California Academy of Sciences.
One male and female paratype are deposited in the following: U.S. National
Museum, American Museum of Natural History, and San Diego Natural History
Museum. The Bezark, CAS, Fisher, and Kansas University paratypes are returned.

Eric M. Fisher has kindly provided specimens from western Mexico of a third
undescribed Efferia of the Albibarbis group.

298

PAN-PACIFIC ENTOMOLOGIST

Efferia sonorensis, New Species (Figs. 2, 4, 8)

Male .—Body length 20.5 to 24.5 mm (holotype 22.0 mm); head with these
differentiating characters: ground color reddish black, scape and pedicel of antenna
black, red apically; flagellomere red basally, reddish black apically; style reddish;
hairs on antennae white, a few may be black dorsally; 4 to 8 (5 and 6 in holotype) of
occipital bristles on each side are black or mixed black and yellow; ocellar tubercle
with 2 long black bristles and 4-7 mixed black and white hairs; mystax white, with
stout yellow-white bristles above an on oral margin; 5-10 black bristles present on
upper 1/3 of gibbosity centrally, variable black hairs and bristles along lower margin
of gibbosity to oral margin.

Mesonotum red, mostly white tomentose with brown tomentum above near
central stripe; meson with a broad brown-black longitudinal stripe, which is often
subdivided centrally by areas of gray-white tomentum; humeral callus reddish, hairs
white anteriorly, black posteriorly; postalar callus reddish; anterior mesonotal hairs
black, subequal in length to scape, posterior hairs black, some as long as scutellar
bristles; scutellum black, white tomentose, reddish basolaterally, with 4-7 marginal
bristles (4 on holotype) and black or white submarginal hairs (some as long as
bristles), discal hairs mixed black and white, 1/4 to 1/3 as long as bristles.

Pleura reddish, thin grayish white tomentose, anepisternum tending to be black
anteriorly; hairs fine, mostly white, with individual hairs black or yellow, usually
black on posterior anepisternum; stem and knob of haltere reddish-brown.

Wings as in the previous species.

Tergites black or reddish-black, 1-5 laterally and 6-7 entirely grayish white
tomentose, tergites 2-5 with dull shining brownish-black areas, margined with brown
tomentum dorsally, tergite 2 often entirely black dorsally, posterior 1/4 to 1/3 of
dorsum of tergites 3-5 white tomentose, becoming reddish apically, tergite 6 often
with a small, elongate dorsal brown spot; hairs on tergites white laterally, mostly
black on dorsal bands; sterna grayish-white tomentose with erect white hairs on
sternites 1-4, shorter and sparser on 5-7, 8 with long apical white hairs.

Terminalia variable from black to reddish-brown, especially dorsally (epandria
dark red-brown on holotype); epandria with a curved, apically directed subapical
process (Fig. 2), length 3.7-4.2 mm (3.7 mm in holotype); basistylus black or
red-brown, light red-brown on inner margin; genitalic hairs mostly black, some
reddish-white apically on epandria; basal hairs of basistylus black or white, ventral
fringe dense black; apex of basistylus in lateral view is forked, with a broad, apically
truncated upper lobe (Fig. 4).

Femora light reddish-brown dorsally, black ventrally, red-brown at apices, spines
black on middle and hind femora, hairs mostly white, some black, especially
dorsoapically; tibiae light reddish-brown basally, dark red-brown at apices, most
hairs white, some black hairs dorsoapically; hind tibia with acute dorsal and slightly
rounded ventral swellings (Fig. 8); fore and hind tibiae with short, dense,
yellow-white to orange pile dorsally; tarsi dark red-brown, spines black, hairs mixed
white and black, claws black, empodia light red-brown; tarsomeres (especially the
first) on fore and hindlegs may have dense ventral red-orange pile.

Female .—Body length 19.5 to 25.0 mm (allotype 20.5 mm); tergite 7 dully shining
brown-black dorsally with black hairs, the hairs usually white laterally (longer near
apex) and mixed black and white on sternite 7; ovipositor 5.9 to 6.8 mm long (5.9 mm

Table 1. Comparison of Nearctic shrub-perching Efferia of the Albibarbis group (females). Lengths in mm.

neosimilis

incognita

tagax

sonorensis

armata

bicolor

grandis

Body length

mean

21.2

20.0

23.1

21.8

25.0

20.8

23.8

range

16.5-24.0

18.0-22.0

21.0-28.0

19.5-25.0

21.5-29.0

14.0-27.0

20.0-27.5

n

7

7

8

5

13

15

13

Ovipositor length

mean

4.2

5.6

5.8

6.3

5.3

5.4

6.2

range

3.9-4.5

5.2-6.3

5.3-6.2

5.9-6.8

4.4-6.1

3.8-6.2

5.6-6.8

n

16

9

18

5

13

15

13

Mystax

White w/blk
brist

White w/blk
brist

White w/blk
brist

Mostly white

Yellow to
yellow-wht

White w/blk
brist

Mostly black

Scutellar hairs

Black

Black

Black

Black

Mostly white

White

Black

Femur color

Black to
red-black

Black

Red

Red to orange

Black

Black

Black

Characters of tergites

Tergites 2-4
broadly black

Tergites 2-4
broadly black

Bands often

narrow,

triangular

Variable

Tergite 6
usually broadly
black

Banding weak,
gray-brown

Tergites 6-7
entirely black

VOLUME 63, NUMBER 3

300

PAN-PACIFIC ENTOMOLOGIST

on allotype), usually reddish-black; subcostal cell hyaline basally but often dark
brown in middle; females also show a tendency to narrowed black banding on the
abdomen and to fewer black bristles in the mystax (especially below laterally).

Diagnosis. —Male; The truncated upper lobe of the apical basistylus will separate
E. sonorensis from related species. E. tagax, although having a reddish thorax and
femora, lacks the subapical epandrial process. The acute ventral swelling of the hind
tibia is much more pronounced in E. armata. E. incognita and E. neosimilis have
much blacker femora, thoracies, and tergites. Female: the orange tone of the femora
and thorax separate E. sonorensis from most related Albibarbis group species.
Characters that consistently separate females of sonorensis and tagax were not
found. The pleura in tagax tend to be more densely white tomentose and the femora a
darker red.

Habitat. —Sonoran desert scrub and thorn scrub in Sonora and Sinaloa. Fisher
(pers. comm.) suggests this is also a shrub perching species. Two of the CIS males
were collected at blacklight.

Holotype male and allotype. —17 mi. N of El Caballo (S of Hermosillo), Sonora,
Mexico, 20 May 1962, (E. Michelbacher) (California Insect Survey collection).

Paratypes. —Same data as above, 4 males, 2 females. MEXICO; Sonora: 23 mi. S
Navajoa, 8 Sept. 1965, 1 female (A. H. Gillogly) (CIS); 7 km NW Navajoa, 7 July
1963, 1 female (E. M. Fisher); 39 mi. N Guaymas, 1 Aug. 1971, 2 males (E. M.
Fisher); Guaymas, Estero Soldado, 8 Aug. 1969,1 male (L. T. Findley) (CIS); 9 mi.
SE Vicam, 22 July 1979,1 male, 1 female (E. M. Fisher). Sinaloa: Rio Piaxtla, 1 mi.
W Mex. 15, 29-30 June 1962 (at blacklight), 2 males (E. Sleeper, R. Anderson, A.
Hardy, R. Somerby) (CIS).

Holotype and allotype are deposited in the California Academy of Sciences.
The remaining paratypes are returned to Eric M. Fisher and the California Insect
Survey.

Acknowledgments

I am grateful to the following individuals for advice and/or loans of specimens;
P. H. Arnaud, Jr. and N. D. Penny (California Academy of Sciences); R. W. Brooks
(Kansas Univ.); M. A. Cazier and F. F. Hasbrouck (Arizona State University);
E. M. Fisher (Calif. Dept, of Food and Agriculture, Sacramento); D. K. Faulkner
(San Diego Natural History Museum); W. L. Murphy (USDA: Beltsville); K. Olson
(University of Arizona); R. T. Schuh (American Museum of Natural History); and
D. C. Lightfoot and J. R. Zimmerman (New Mexico State University).

Literature Cited

Bromley, S. W. 1934. The robber flies of Texas (Diptera: Asilidae). Ann. Entomol. Soc. Amer.
27:74-113.

Bullington, S. W. and R. J. Lavigne. 1984. Description and habitat of Efferia kondratieffi sp. nov. with
notes on Efferia aestuans (L.) (Diptera: Asilidae). Ann. Entomol. Soc. Amer. 77:404—413.

Hine, J. S. 1919. Robberflies of the genus Erax. Ann. Entomol. Soc. Amer. 12:103-157.

Martin, C. H. and J. Wilcox, /n Alan Stone et a/. 1965. Acatalogof Diptera of North America. Asilidae.

United States Dept, of Agriculture, Agriculture Handbook no. 276, pp. 360-401.

Wilcox, J. 1966. Efferia Coquillett in America north of Mexico (Diptera: Asilidae). Proc. Calif. Acad. Sci.
34:85-234.

Williston, S. W. 1885. On the North American Asilidae (part II). Trans. Amer. Entomol. Soc. 12:53-76.

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DOYEN, J. T.—New and little known Tenebrionidae from Central America and Mexico, with

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HAJEK, A. E. and D. L. DAHLSTEN—The exotic aphids (Homoptera: Drepanosiphidae) on

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LANHAM, U. N.—A new species of Andrena at the Micrandrena-Scaphandrena boundary

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PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 301-318

New and Little Known Tenebrionidae from Central America and
Mexico, with Remarks on their Classification (Coleoptera)

John T. Doyen

Department of Entomological Sciences, University of California, Berkeley,
California 94720

Abstract. —The tribe Misolampini is restricted to the Old World. New World
species are transferred to Coelometopini. Details are presented of morphological
similarities between Hegemona and Saziches (Neotropical) and
Promorphostenophanes (Oriental). New genera described; Cnephalura,
Bothynocephalus; new species: Cnephalura umbrata; Bothynocephalus cristatus]
Isaminas breedlovei, I. reticuloides, /. sullivani; Saziches giesberti. New synonymy;
Pteroglymmius erotyhides GQh\Qn = Isaminas erotyhides (Gebien).

The taxa treated here comprise a group of flightless, arboreal Tenebrionidae
restricted to Central and South America. The genera Saziches and Isaminas were
described by Champion (1886), who placed them, along with Oxidates and
Hegemona, in Misolampini. As discussed by Doyen et al (in press), Misolampini
presently construed consists of various geographically restricted groups of flightless
Coelometopini. The South and Central American taxa may form a valid clade, but
with few exceptions do not show close relationships to the Misolampini of Africa or
tropical Asia. For this reason, Misolampini should properly be applied only to
Misolampus and its relatives, if at all. The new world genera are more conveniently
considered as members of Coelometopini. Reconstitution of New World
Coelometopini will be considered at greater length by Doyen (in prep.). The purpose
of the present contribution is to describe several new genera and species which
significantly broaden the range of variation of the New World fauna of these beetles.

Isaminas Champion

Isaminas Champion 1886; 266.

Pteroglymmius Gebien 1928; 223.

Type species: Isaminas gibbipennis Champion, Gebien (1942-44) designation.

Champion (1886) described the salient external features of this genus. It may be
added that the antennae bear compound sensoriae on the apical five antennal
segments and that the internal female reproductive tract and glands are typical of
Coelometopini. The female tract consists of the vagina, without a bursa copulatrix,
and a single, long diverticulum, glandular except on the abruptly enlarged, apical
spermatheca. The first lobe of the coxite of the ovipositor is very elongate,
comprising about 3/4 of the length of the ovipositor tube. The gland reservoirs are

301

302

PAN-PACIFIC ENTOMOLOGIST

large and annulate, with the secretory tissue draining through a pair of enlarged
ampullae at the exit ducts. All of these features show that Isaminas is derived from a
relatively apotypic group of Coelometopini. In the New World, this group includes
most of the tribe, with the exception of the genera Camaria and its relatives,
Taphrosoma, Mylaris ( = Nyctobates; Spilman, 1973), and Hapsida which are
plesiomorphic in one or more features (Tschinkel and Doyen, 1980). Isaminas is very
similar to Oxidates Champion, differing in the configuration of the metasternum. In
Isaminas the mesocoxal and metacoxal cavities are nearly contiguous, with the
intervening strip of metasternum narrower than the metepipleuron. In Oxidates the
strip of intervening metasternum is much wider than the metepipleuron. Isaminas is
superficially similar to Saziches Champion, but in the latter the elytral punctures of
each series are connected by a fine, longitudinal furrow. In Isaminas the punctures
are isolated. There are major internal differences between these two as well,
discussed under Saziches.

I have not examined the type of Pteroglymmius Gebien (1928), but his detailed
description of the peculiar elytral sculpturing clearly identifies his species erotyloides
as synonymous with specimens at hand. Gebien separated Pteroglymmius from
Isaminas on the basis of the scutellum (punctiform in Pteroglymmius; distinct in
Isaminas). In the material examined, including cotypes of I. gibbipennis Champion
and brevicollis Champion, the scutellum is punctiform, scarcely entering the elytral
disk. In all other features erotyloides is extremely similar to other species of
Isaminas.

Key to the Species oi Isaminas

1. Elytron with distinct epipleural carina; epipleuron continuing to apex of

elytra or nearly so . 2

Elytron with epipleural carina visible only from humerus to metacoxa,

obsolete posteriorly.Champion

2(1). Elytra with more or less uniform series of punctures; sequential punctures
in series sometimes contiguous or coalesced, but punctures in adjacent

series never coalesced. 3

Elytra with punctures in adjacent series coalesced to form transverse

furrows behind humeri and sometimes centrally or on declivity. 5

3(2). Pronotum broadest before middle; lateral margins shallowly crenulate,

nearly straight in posterior half . gibbipennis Champion

Pronotum broadest behind middle; lateral margins evenly arcuate . 4

4(3). Elytra with punctures relatively small, separated by much more than

puncture diameter both longitudinally and transversely, except on

declivity. breedlovei, new species

Elytra with punctures relatively large, subcontiguous; adjacent punctures
in series sometimes coalesced, forming elongate foveae .... reticuloides,
new species

5(2). Each elytron with punctures in series four through eight coalesced just
behind humerus and four through seven on declivity, forming depressed
areas covered with bright yeUow secretion in life; punctation otherwise

uniform. sullivani, new species

Each elytron with irregular transverse depressed bands about one third
and two thirds posterad, uniting punctures in rows two or three to eight;

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303

pairs or triplets of adjacent punctures irregularly coalesced over remainder
of surface, especially on declivity . erotyhides Gebien

Isaminas breedlovei, new species

(Fig. 1)

Frons and vertex with exceedingly shallow punctures about half eye facet in
diameter, separated by one to several puncture diameters; punctures slightly
coarser, deeper, and denser on genae and epistomum, finer and denser along rim.
Pronotal disk sculptured like frons centrally, becoming more shallowly and finely
punctate laterally; with very shallow, median longitudinal depression and distinct
fovea slightly behind middle at medial third on each side. Anterior angles obtuse,
angulate but rounded at apex; lateral margins more strongly arcuate in anterior
third, weakly so to base, sometimes very weakly undulate in basal third. Prosternal
process with longitudinal, parallel depressions along lateral quarters, becoming
deeper posteriorly, producing trilobed apex. Elytra with strial punctures four to
eight times eye facet in diameter, separated longitudinally by one to three puncture
diameters; striae separated by two to four puncture diameters; sutural stria with
finest punctures, becoming gradually coarser laterally and elongate in striae seven
through nine, with punctures three to four times longer than wide in epipleural stria;
epipleural carina complete to anterior margin of sternite five, continuing as rounded
prominence to apex.

Measurements: median pronotal length (PL), 2.5-2.7 mm; greatest pronotal
width (PW), 3.9-4.4 mm; elytral length (EL), 6.7-7.6 mm; greatest elytral width
(EW), 6.2-7.1mm.

Isaminas breedlovei is similar to reticuloides, new species, in thoracic shape,
configuration of prosternal process and in the extremely gibbous elytra, which are
abruptly elevated behind the prothorax as steeply as on the declivity. In breedlovei
the elytral punctures are separated by at least twice their diameter. In reticuloides the
punctures are subcontiguous.

Holotype female (California Academy of Sciences) from Mexico, Chiapas, 32 km
N Ocozocoautla, onrd. to Malpaso,762m, 6-X-1974, D. E. and J. A. Breelove. Two
paratypes, Mexico, Chiapas, 13 km N Berriozabel, 975 m, 29-V-1973, D.E.
Breedlove.

The holotype is larger than the paratypes and has more finely punctate elytra, but
is similar in all other features.

Isaminas reticuloides, new species

Frons and vertex impunctate; genae and epistomum with exceedingly fine, shallow
punctures along rim. Pronotal disk with sparse, exceedingly fine, shallow punctures
centrally, impunctate marginally; with extremely shallow, scarcely perceptible
median longitudinal depression and distinct fovea just before posterior margin,
two-thirds toward lateral margin. Anterior angles slightly obtuse, broadly rounded;
lateral margins arcuate in anterior three fourths, then parallel to hind angles.
Prosternal process with longitudinal depressions in lateral quarters, diverging
posteriorly and declivous, producing trilobed apex. Elytra with strial puncture
diameter one-half to one times length of dorsal eye lobe; smaller and subcontiguous
in sutural stria, coarser and contiguous or sometimes coalesced in striae two to seven.

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PAN-PACIFIC ENTOMOLOGIST

Figure 1. Isaminas breedlovei, new species, holotype

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305

finer in striae eight to nine and seldom coalesced; epipleural carina distinct to sternite
five, continuing as rounded prominence to apex.

Measurements: PL, 2.7 mm; PW, 4.1 mm; EL, 7.3 mm; EW, 6.5 mm.

Isaminas reticuloides is similar to breedlovei, new species in its extremely inflated
body and trilobed posternal process. See remarks under the latter species. The
reticulate sculpturing of reticuloides is similar to that of erotyhides (Gebien), but in
the latter punctures from adjacent series are coalesced, forming transverse
depressions about one-third and two-thirds posteriad on the disk. In reticuloides
punctures from adjacent series are never coalesced. In eroty hides the lateral
pronotal margins are crenulate; in reticuloides there is no trace of crenulation.

Holotype (sex undetermined; California Academy of Sciences) from Mexico,
Chiapas, north slope Cerro Bola, N. Cerro Tres Picos, 1524-2134 m. 5-V-1972.
D. E. Breedlove.

Isaminas gibbipennis Champion

Isaminas gibbipennis Champion, 1886: 267.

Cranium punctate dorsally; punctures exceedingly fine posteriorly, becoming
about one-half to one times eye facet in diameter, separated by one to several
puncture diameters on epistomum and genae; punctures slightly denser along
epistomal rim. Pronotal disk with exceedingly fine, shallow punctures centrally,
becoming impunctate marginally; with distinct medial longitudinal depression and
distinct fovea just behind middle at medial third on each side. Anterior angles
broadly rounded; lateral margins arcuate in anterior half, then nearly straight,
weakly crenulate to posterior angles. Prosternal process with longitudinal, shallow
depressions along lateral quarters, these slightly divergent posteriorly but not
deepened; apex only weakly trilobed, obtusely angulate. Elytral disk with strial
punctures one-half to one times length of dorsal eye lobe in diameter, separated by
about one-half to one times puncture diameter in each series; smallest anteriorly, on
declivity and near epipleuron; distinctly elongated in rows seven to nine, especially
anteriorly; epipleural carina complete to elytral apex.

Measurements (of paralectotype): PL, 7.4 mm; PW, 4.3 mm; EL, 6.8 mm; EW,
6.0 mm.

Isaminas gibbipennis is similar to brevicollis Champion in sculpturing and body
configuration. In gibbipennis the epipleural carina is distinct to the elytral apex and
the prosternal process has the apex obtusely angulate. In brevicollis the epipleural
carina is present only as far posterad as the metacoxae and the prosternal process is
acutely angulate.

A lectotype is hereby designated from the original series of cotypes in the British
Museum (Natural History). Type locality, Nicaragua, Dept. Chontales. Additional
paralectotypes are from Costa Rica (no further information). One paralectotype is
located in the Essig Museum of Entomology, University of California, Berkeley.

Isaminas brevicollis Champion

Isaminas brevicollis Champion, 1886: 267

Cranium punctate dorsally, punctures about one-half to one times eye facet in
diameter, separated by about one to two puncture diameters, becoming finer, denser
along epistomal rim. Pronotal disk with punctures about one-fourth to one-half eye

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PAN-PACIFIC ENTOMOLOGIST

facet in diameter centrally, becoming impunctate near margins; with distinct medial
longitudinal depression and distinct fovea at middle at medial third on each side.
Anterior angles broadly rounded; lateral margins arcuate except straight just before
base. Prosternal process with shallow, longitudinal, parallel depressions along
lateral quarters, becoming obsolete behind coxae; apex acutely angulate. Elytral
disk with strial punctures about one-third to one times length of dorsal eye lobe in
diameter, separated by about one to two puncture diameters in each series; smallest
anteriorly, on declivity and near epipleuron; slightly elongate in rows seven to nine;
epipleural carina distinct just behind humerus, becoming obsolete posteriorly, and
disappearing at about metacoxa.

Measurements (specimen from Chiapas): PL, 2.5 mm; PW, 4.0 mm; EL, 6.7 mm;
EW, 6.0 mm.

Isaminas brevicollis is similar in sculpturing and body shape to /. gibbipennis
Champion, differing as described under the latter.

A lectotype is hereby designated from the original series of cotypes in the British
Museum (Natural History). Type locality, Guatemala, Dept. Alta Vera Paz,
Senahu. Additional paralectotypes are from Dept. Baja Verapaz, Sinanha.
Additional record, Mexico, Chiapas, Tuxtla Gutierrez, 23-VI-1973, G. Ekis.

Isaminas sullivani, new species

(Fig. 2)

Frons and vertex with punctures about one-fourth to three-fourths times eye facet
in diameter, separated by about one puncture diameter, becoming slightly smaller
and denser along epistomal rim. Pronotal disk with fine, exceedingly shallow, almost
obsolete punctures centrally, impunctate marginally; with distinct median
longitudinal depression and distinct fovea at middle at median third on each side.
Anterior angles broadly rounded; lateral margins strongly, evenly arcuate in anterior
one-third, then nearly straight, slightly convergent and crenulate to posterior angles.
Prosternal process with longitudinal depressions along lateral quarters, diverging
slightly and becoming deeper posteriorly, producing trilobed apex. Elytra with strial
puncture diameter about one-half to one times length of dorsal eye lobe, separated
longitudinally by about one-fourth to one-half puncture diameter; punctures slightly
smaller in sutural stria, anteriorly and on declivity; punctures in striae four through
eight coalesced immediately behind humerus and in striae four through six on
declivity, forming irregular transverse depressions, in life filled with bright yellow
exudate; epipleural carina complete to elytral apex or nearly so.

Measurements: PL, 1.5-2.4 mm; PW, 2.9-4.5 mm; EL, 4.9-6.4 mm; EW,
4.4-7.2 mm.

Isaminas sullivani is most similar to erotyhides (Gebien). In sullivani the
transverse elytral depressions are located immediately behind the humeri and the
declivity. The elytral punctation is quite regular aside from the depressions. In
eroty hides the depressions are located about one-third and two-thirds posteriad, and
coalesced pairs or triplets of punctures are scattered irregularly over the elytra,
especially laterally.

Holotype (sex undetermined; California Academy of Sciences) from Costa Rica,
Cartago Prov. Turrialba, 14/19-V-1974, E. Giesbert. Paratypes: 3, same date as
holotype; 1, Cartago Prov., C.A.T.I.E., 3 km SE Turrialba, 13/16-V-1985, J.
Doyen; 5, same locality, 29/30-V-1985, J. Doyen; 1, same locality, 28-V-1985, C.

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307

Figure 2. Isaminas sullivani, new species.

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PAN-PACIFIC ENTOMOLOGIST

Sevilla; 1, same locality, 8-1-1984, R. L. Penrose, F. T. Hovore, P. H. Sullivan; 9,
Limon Prov., 22 km S. Siquirres, 9/12-1-1984, R. L. Penrose, F. T. Hovore, P. H.
Sullivan.

At the C.A.T.I.E. facility near Turrialba most of the beetles were knocked onto
beating sheets from tangles of dead vines in dense forest understory. They are active
nocturnally, crawling slowly along vines and twigs.

Isaminas erotyloides (Gebien)

Pteroglymmius erotyloides Gebien, 1928: 224.

Frons and vertex with punctures about as large as eye facets, usually separated by
less than one puncture diameter, densest near epistomal suture, becoming finer,
sparser along epistomal rim. Pronotal disk with very shallow punctures about twice
eye facet diameter medially, becoming impunctate in lateral quarters; with shallow
longitudinal median depression, becoming foveate slightly before middle; distinct
fovea slightly behind middle at median third on each side. Anterior angles broadly
rounded; lateral margins evenly arcuate, crenulate, slightly depressed just inside
Carina, this depression filled with bright yellow exudate in life. Prosternal process
with longitudinal depressions posteriorly convergent, meeting well before acutely
angulate apex. Elytra with strial puncture diameter about one-half to slightly larger
than length of dorsal eye lobe; punctures most regular in sutural stria, subcontiguous
anteriorly, becoming contiguous or occasionally coalesced on declivity; punctures in
striae two through nine subcontiguous to contiguous, occasionally coalesced into
pairs or triplets; those in three through nine occasionally coalesced with punctures in
adjacent striae, especially laterally and posteriorly, producing a coarsely rugose
texture; adjacent punctures in striae two or three through eight always coalesced
about one-third posterad and two-thirds posterad, forming irregular transverse
depressions, in life filled with bright yellow exudate; epipleural carina complete to
elytral apex or nearly so.

Measurements: PL, 2.5-2.9 mm; PW, 4.1-4.7 mm; EL, 6.8-7.8 mm; EW, 5.7-6.7
mm.

Isaminas erotyloides is similar to sullivani in bearing transverse, depressed regions
on the elytra. It differes as described under the latter.

Holotype (sex undetermined; deposition not stated) from Honduras, Dept.
Cortes, San Pedro Sala [sic], no additional data. Additional material examined:
Honduras (no additional data) (7); Santa Barbara Departimiento, Lago Yojoa,
7/21-VII-1978, S. Dubon (2); La Paz Departimiento, La Paz, 21-VII-1978, B.
Herrera (2); Comayagua Departimiento, Siguatepeque, 22-VI-1978 (1).

Saziches Champion

Saziches Champion, 1886: 261.

Type species: Saziches subcaudatus Champion, 1886, by monotypy.

Champion described the important external features of Saziches, relating it to
Isaminus, Oxidates and the South American Sphaerotus. The last three are derived
members of Coelometopini, as discussed above, but Saziches has distinctive
ovipositor, internal female reproductive tract, and defensive glands which show that
it is closely related to Hegemona Champion. In both these genera the ovipositor is

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309

Strongly modified as a stout, sclerotized, laterally compressed, blade-like structure
(Fig. 3). Reduced gonostyles are visible apically, and faint transverse lines may
correspond to divisions of the gonocoxite or between coxite and paraproct. In other
regards the features of this ovipositor cannot be homologized with those of typical
tenebrionids. Presumably the ovipositor is forced into some substrate, but
oviposition and larvae are unknown.

The internal female reproductive tract consists of a slender vagina, opening into an
enlarged bursa copulatix, with a single diverticulum (spermathecal accessory gland)
attached to the bursa. The gland is of nearly uniform diameter throughout, and
glandular except at the extreme base. In Saziches the gland attaches dorsolaterally
on the body of the bursa (Fig. 4). In Hegemona it attaches dorsolaterally on the neck,
which is involuted (Fig. 5). The relatively thick walled bursa of Hegemona has
numerous longitudinal pleats which maintain their shape, while that of Saziches is
fragile and irregularly saccate when cleared.

The defensive reservoirs in both genera are large elongate sacs with distinct medial
lobes basally. Oblique folds are apparent in the reservoir walls, especially in
Hegemona, but the helical folds of derived Coelometopini are absent. There is
considerable common volume between the reservoirs, which receive the secretion
from a few basal collecting ducts.

Except for the highly derived ovipositor, most of these features are plesiomorphic.
A bursa copulatrix is retained in many of the Old World coelometopines which are
included in Cnodalonini in checklists, and also in My laris and Taphrosoma in the
New World. In these taxa, however, the apex of the spermathecal accessory gland,
though not enlarged, is nonglandular, whereas in Hegemona and Saziches it is
glandular to the apex. Defensive glands similar to those of Hegemona and Saziches
are found in Camaria, Hapsida and Talanus in the new world, and in many old world
“Cnodalonini” (see Tschinkel and Doyen, 1980, Fig. 13). In Catapiestus and
Strongyliini the reservoirs are non-annulate, but much smaller.

In external appearance Hegemona is extremely similar to Promorphostenophanes
Kaszab (1960). The holotype of P. atavus vietnamicus Kaszab (1980) (BMNH) has
the ovipositor visible. Without dissection it is apparent that it is modified in exactly
the same manner as in Hegemona and Saziches. It seems certain that these taxa, and
probably Morphostenophanes Pic form a natural clade.

In the Gebien catalog (1942-44) Morphostenophanes and Saziches are included
under Misolampini ( = Coelometopini, in part), while Hegemona appears under
Helopini. None of the important features of the defensive glands or female tracts of
any of these beetles suggest Helopini. In addition, all these genera have
characteristic compound sensoriae on the apical antennal segments, while Helopini
have only simple, hair-like sensilla. While cladistic placement is somewhat
problematic, this clade appears to contain the most primitive members of the
Coelometopine lineage, lacking defensive reservoir annulation, retaining a primary
bursa copulatrix, and lacking the spermathecal specialization of the distal accessory
gland. The ovipositor also lacks the reflexed paraprocts of nearly all members of the
coelometopine lineage (oblique in Menephilus), but this may represent a secondary
specialization, related to the blade-like structure of the coxites. Talanus has a
superficially similar blade-like ovipositor, but the coxite structure differs in detail,
and the reflexed paraproct is retained, indicating separate derivation. Subdivision of
Coelometopini would be premature until a much greater proportion of the world

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PAN-PACIFIC ENTOMOLOGIST

anus _ \

Figure 3. Ovipositor, Saziches giesberti.

fauna is examined in detail. At present it is prudent to recognize these genera as a
distinctive clade within Coelometopini.

Saziches subcaudatus Champion

Saziches subcaudatus Champion, 1886: 262.

Cranium with extremely shallow, almost obsolete punctures about one-fourth to
one-half eye facet in diameter, separated by one to several puncture diameters,
becoming slightly finer, denser on epistomal rim. Pronotal disk with sparse punctures
much finer than eye facets. Elytra with strial punctures about one-fourth to one-half
dorsal eye lobe in diameter, finer anteriorly, in sutural stria and on dechvity, coarsest
on humped part of disk; stria indicated by very fine line in sutural series, lines
becoming gradually coarser laterally and obsolescent near apex; punctures in eighth
stria connected by depressed furrow with incised line in bottom; interstriae two
through nine subequal, flat; epipleural stria disappearing about halfway posteriad
along sternite five; elytral apices forming small, rounded prominences.

Measurements (of paralectotype): PL, 2.4 mm; PW, 3.6 mm; EL, 8.0 mm; EW,
5.9 mm.

Saziches subcaudatus differs from S. giesberti as described under the latter.

A lectotype is hereby designated from the original series of cotypes in the British
Museum (Natural History). Type locality: Guatemala, Dept. Alta Vera Paz, San
Juan. Additional paralectotypes are from Dept. Alta Vera Paz, Senahu. One
paralectotype is located in the Essig Museum of Entomology, University of
California, Berkeley.

Saziches giesberti, new species
(Fig. 6)

Cranium with shallow, setigerous punctures about one-fourth to one-half eye facet
in diameter, separated by one to several puncture diameters, becoming coarser.

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311

Figures 4-5. Internal female reproductive tracts. 4. Saziches giesberti. 5. Hegemona filibuster
Champion.

denser on epistomum, then finer on epistomal rim. Pronotal disk with very shallow
setigerous punctures about one to two times eye facet in diameter, separated by one
to several puncture diameters. Elytra with striae indicated by finely incised lines;
punctures minute, scarcely demarked from strial lines; striae one and two meeting
anteriorly; striae one and nine, three and four fusing posteriorly; remainder fusing
irregularly; interstriae convex, shining; first interstria narrowest, subparallel, with
scattered, irregular depressions; second interstria widest, about 2.5 times as broad in
middle as on declivity, and bearing scattered, irregular punctures and depressions;
interstriae three through nine subequal, slightly broader in middle and traversed by
five or six transverse depre sions, producing undulating surface; irregular
depressions just behind humeri and on declivity with dull surface, in life containing
yellow-orange exudate; epipleural stria continuing almost to elytral apex; elytral
apices not produced.

Measurements: PL, 1.8-2.1 mm; PW, 2.7-3.5 mm; EL, 5.9-7.3 mm; EW,
4.4-5.9 mm.

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PAN-PACIFIC ENTOMOLOGIST

Figure 6. Saziches giesberti, new species, holotype

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313

Saziches giesberti differs from subcaudatus Champion in its extremely fine elytral
punctures, the impressed strial lines, and the subhumeral and declival depressions.
In subcaudatus the strial punctures are very coarse, the strial lines barely defined
except laterally, and the subhumeral and declival depressions absent.

Holotype (California Academy of Sciences) from Costa Rica, Cartago Prov.
Turrialba, 14/17-V-1974. E. Giesbert. Paratypes, Turrialba, 31-V-1951, O.L.
Cartwright (2); CATIE, 3 km SE Turrialba, 600 m, 13/16-V-1985, J. T. Doyen (3).

At the C.A.T.I.E. facility Saziches giesberti was collected onto beating sheets
from tangles of dead vines in dense forest understory. The beetles were taken in
company with Isaminas sullivani, which they resemble in bearing patches of yellow
exudate subhumerally and on the elytral declivity. Adults are probably active
nocturnally.

Cnephalura, new genus

Epistomum nearly straight medially, shallowly emarginate at lateral epistomal
sutures; epistomal suture obsolete medially, faint laterally; epistomal membrane
concealed. Eyes shallowly emarginated by epistomal canthus; dorsal lobe slightly
larger than ventral, bordered medially by shallow groove, deepening and extending
posteroventrally to mid-lateral postgena, delimiting distinct postocular lobe; eye
bounded posteriorly by shallow groove. Antenna with third segment about twice
length of fourth; segments four through eight becoming gradually shorter, broader;
segments nine through 11 forming weak club and bearing compound sensoriae
apically. Mentum trapezoidal, anterior central portion forming elevated tuberosity.

Pronotum subquadrate, moderately convex; lateral carina complete, very
narrowly upturned; prosternal process about as wide as procoxa, declivous behind
coxae, apex broadly rounded. Elytra moderately convex, evenly ovate; epipleuron
narrowing very little from humerous to elytral apex; scutellum about twice as broad
as long. Mesosternal fossa obtusely concave, lateral margins scarcely elevated;
metasternum between coxae almost as long as mesocoxal diameter; intercoxal
process three-fourths as wide as metacoxa, truncate. Femora and tibiae nearly
cylindrical; femora reaching slightly beyond head and approximately to abdominal
apex; tibiae with apices pilose; tarsi with plantar surfaces of all but apical segment
pilose. Abdominal defensive reservoirs large, saccate, with annular foldings and
considerable common volume; aedeagus inverted, simple, fusiform; median lobe
sessile. Female reproductive tract unknown.

Cnephalura is similar to Oxidates in body form. In Cnephalura the epistomal
suture is obsolete to absent, at least medially, whereas in Oxidates it is distinct. In
Cnephalura the supraocular groove extends posteroventrally behind the eye,
delimiting a distinct postocular lobe. In Oxidates the supraocular groove is restricted
to the dorsolateral margin of the eye. In addition, the male of Cnephalura umbrata
has a sharp tooth three-fourths of the distance toward the apex of the femur (see
below). This secondary sexual feature is unknown in Oxidates.

Type species: Cnephalura umbrata, new species, here designated.

Cnephalura umbrata, new species
(Fig. 7)

Male.—Cranium with deep, setigerous punctures about one to two times eye facet
in diameter, separated by about one puncture diameter, becoming shallower and

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PAN-PACIFIC ENTOMOLOGIST

Figure 7. Cnephalura umbrata, new species, holotype.

finer on epistomal margin; setae pale, procumbent. Pronotal disk with deep,
setigerous punctures about twice eye facet in diameter, contiguous to separated by
about one puncture diameter, forming irregularly rugosopunctate surface; setae
recumbent. Anterior angles obtusely rounded; lateral margins nearly evenly

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315

arcuate; posterior angles right angled; posterior border weakly bisinuous;
hypomeron sculpted as disk; prosternum scabrous; prosternal process with marginal
groove along lateral borders. Elytra with regular series of strial punctures about 1.5
times diameter of eye facet, separated by about two puncture diameters on disk,
decreasing to one diameter on declivity; interstriae irregularly set with rounded
shining tuberosities one to three times punctures in diameter, each bearing several
minute setae and one to several very fine punctures. Abdominal sternites with
punctures a little larger than eye facets, separated by about one to two puncture
diameters, becoming denser, finer on apex of sternite five. Legs rugosely punctate;
femora bearing sharp, short tooth at about distal three-fourths.

Measurements: PL, 2.5-2.7 mm; PW, 2.8-3.0 mm; EL, 5.7-5.8 mm; EW,
3.9-4.0 mm.

Female.—U nknown.

Holotype male (California Academy of Sciences) from Mexico, Chiapas,
Municipio la Trinitaria, Dos Lagos, 14 km E paved road at Lagos de Colores, 1219M,
29-XI-1976. D. E. and J. A. Breedlove. Paratype, Chiapas, Municipio de Cintalapa,
4-6 km NW La Cienega, 40 km NW Las Cruces, 1400 m, XII-1-1980. D. E. and J. A.
Breedlove.

The combination of coarsely punctate pronotum and head with coarsely, densely
tuberculate elytra distinguishes Cnephalura umhrata from all other Central
American Coelometopini.

Bothynocephalus, new genus

Epistomum straight just before eyes, then evenly arcuate between antennal
sockets; lateral epistomal sutures faint, median suture absent; epistomal membrane
concealed. Eye with dorsal and ventral lobes subequal, shallowly emarginated by
epistomal canthus; dorsal lobe bordered anteriorly and medially by a shallow groove
terminating posteriorly in a deep cavity adjacent to the posterodorsal corner of the
eye (Fig. 8); eye bordered posteriorly by shallow groove. Frons forming prominent,
abruptly declivous, curving brow between postocular cavities. Antenna with third
segment about 1.7 times longer than fourth; segments five through 10 becoming
gradually shorter, broader; 11 almost twice as long as broad, asymmetrically
rounded apically; compound sensoriae distributed on mesal and lateral angles of
segments eight through 10, on apical half of 11. Mentum trapezoidal; central third
forming prominent, elevated tuberosity.

Pronotum globular, lateral carina obsolescent in posterior third, anterior border
not margined; posterior border broadly margined, deeply grooved just before
margin; prosternal process slightly wider than coxa, abruptly declivous immediately
behind coxae. Elytra strongly convex, evenly ovate; epipleural margin ventrolateral
in aspect; scutellum triangular, moderate in size, about twice as broad as long.
Mesosternal fossa very shallow, lateral margins scarcely elevated; metasternum
between coxae about as long as mesocoxa; intercoxal process nearly as broad as
coxa, truncate with broadly rounded corners. Femora and tibiae subcylindrical;
femora reaching slightly beyond head and to fourth abdominal sternite; tibiae with
apex pilose; tarsi with plantar surfaces of all but apical segment pilose, apical
segment more sparsely setose. Abdominal defensive reservoirs large, saccate, with
annular foldings and considerable common volume; collecting ducts emptying
through single large ampulla on each reservoir. Ovipositor coelometopoid, with

316

PAN-PACIFIC ENTOMOLOGIST

Figure 8. Bothynocephalus, cranium, lateral aspect.

paraprocts rotated, basal lobe of coxite extremely elongate; internal tract without
bursa copulatrix; spermathecal accessory gland about as long as ovipositor,
terminating in large, saccate spermatheca. Male reproductive tract unknown.

Bothy nocephalus is similar to Oxidates and Cnephalura in body form, but has a
relatively smaller, more globular pronotum, and a greater constriction between the
prothorax and the hindbody, which superficially gives it the appearance of members
of the tenebrionid tribe Triorophini. In the characteristics of its defensive glands,
ovipositor and internal female reproductive tract, Bothy nocephalus is a typical,
highly derived member of the Coelometopini.

Type species: Bothynocephalus cristatus, new species, here designated.

Bothy nocephalus cristatus, new species
(Fig, 9)

Female.—Cranium with very fine, minutely setigerous punctures, very much
smaller than eye facet diameter, separated by about two to five puncture diameters;
cranial surface very finely shagreened, except for polished declivity below brow
ridge. Pronotal disk with anteromedial punctation as on cranium; punctures
becoming slightly coarser, shallower in anterolateral corners, gradually coarser
posteriorly to slightly larger than eye facets along posterior margin; large punctures
more or less coalesced along lateral carina, forming irregular marginal groove; carina
rounded anteriorly, weakly explanate and upturned near middle, becoming
obsolescent posteriorly; anterior corners broadly rounded, without definite angles;
lateral margins strongly arcuate; posterior angles obtuse; posterior margin evenly
arcuate. Hypomeron with shallow punctures about 1.5 times eye facet in diameter,
separated by about one to three puncture diameters. Prosternal process with deep
marginal groove along lateral and posterior borders. Elytra with regular series of
deep strial punctures about 1.5 to two times eye facet in diameter, separated by about
one to three puncture diameters; sutural stria of one or two punctures; striae one and
nine, two and seven, three and six, four and five joining posteriorly; one and two, five
and six joining anteriorly. Interstriae weakly sulcate, finely shagreened. Epipleuron
obsolete anteriorly, becoming visible as fine ridge at about anterior margin of fifth
sternite and extending almost to elytral apex. Abdominal sternites finely
shagreened, with fine, obsolescent punctures barely visible on last three sternites.

VOLUME 63, NUMBER 4

317

Femora with punctures about one-half to one times as large as eye facets, separated
by about one puncture diameter to subcontiguous. Tibiae more finely punctate.

Measurements: PL, 2.6-3.3 mm; PW, 3.0-3.6 mm; EW, 4.2-5.2 mm; EL,
6.5-8.2 mm.

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PAN-PACIFIC ENTOMOLOGIST

Male.—Unknown.

Holotype female (California Academy of Sciences) from Honduras, Comayagua
Departimiento, Siguatepeque, 8-VI-1976. J. V. Mankins. Paratype (abdomen
missing, probably female), same locality 26-X-1975. J. V. Mankins.

Bothynocephalus cristatus differs from all other new world Tenebrionidae in its
prominently browed frons and deep postocular cavities.

Acknowledgments

Some of the specimens used in this study were loaned by M. J. D. Brendell and
M. E. Bacchus, British Museum (Natural History); D. Kavanaugh, California
Academy of Sciences; T. J. Spillman, Systematic Entomology Laboratory,
Agricultural Research Service, Washington, D.C., P. H. Sullivan, private
collection, and D. Cota, C.A.T.I.E., Turrialba, Costa Rica. Christina L. Jordan
rendered the illustrations of beetles.

Literature Cited

Champion, G. C. 1886. Biologia Centrali-Americana, Insecta, Coleoptera (Tenebrionidae), vol. 4, pt.
1:137-204.

Doyen, J. T., Matthews, E. G., and Lawrence, J. F. 1987. Classification and annotated checklist of the
Australian genera of Tenebrionidae (Coleoptera). Invert. Taxonomy (in press).

Gebien, H. 1928. Uber einge Gruppen amerikanischer Tenebrioniden (Col. Heter.). 2. Teil. Stett.
Entomol. Zeit., 89:167-234.

Gebien, H. 1942-44. Katalog der Tenebrioniden, Teil III. Mitt. Miinch Entomol. GeselL,
32-34:746-899 (repaged).

Kaszab, Z. 1960. Neue oreintahschenMisolampinen (Coleoptera, Tenebrionidae). Ann. Hist.-Nat. Mus.
Nat. Hung., 52:265-294.

Kaszab, Z. 1980. AngabenzurKenntnis der Tenebrioniden Nordvietnams (Coleoptera). Ann. Hist.-Nat.
Mus. Nat. Hung., 72:169-221.

Spilman, T. J. 1973. Nomenclatural problems in six genera of Tenebrionidae (Coleoptera). Proc.
Entomol. Soc. Wash., 75:39-44.

Tschinkel, W. R. and Doyen, J. T. 1980. Comparative anatomy of the defensive glands, ovipositors and
female genital tubes of tenebrionid beetles (Coleoptera). Int. Jour. Insect Morphol. and Embryol.,
9:321-368.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp.319-323

The Exotic Aphids (Homoptera: Drepanosiphidae)
on Ornamental Birch in Northern California

Ann E. Hajek^ and Donald L. Dahlsten

Division of Biological Control, Department of Entomology, University of
California Berkeley, Berkeley, California 94720.

Abstract. —Five introduced species of aphids occur on ornamental birch trees in
northern California. A key to these aphids is provided along with species diagnoses
and biological notes.

In northern California, five species of aphids of the Family Drepanosiphidae,
Tribe Phyllaphidina (sensu Heie 1980) feed on introduced trees of the genus Betula,
the birches. The most common birch in northern California is the ornamental
European white birch, Betula pendula Roth. The downy birch, Betula pubescens
Ehrhart, has also been planted but much less frequently (Hajek 1986). These host
trees and their aphid fauna are endemic to Europe where a guild of eleven birch
aphid species occur sympatrically (Fleie 1972). These introduced species of birch
aphids have only recently been studied in California (Grushkowitz 1976; Hajek
1984), although they frequently create an urban pest problem due to production of
copious amounts of honeydew. Dates of introduction for these species are not known
although one species, Callipterinella minutissima (Stroyan), was only recently
recorded from North America (Hajek 1985).

Apterous and alate viviparae of all five species can often be distinguished with the
naked eye. The only English language keys presently available for identification of
these birch aphid species include the entire European fauna of the aphid family
including these species (Stroyan 1977; Heie 1982). For ease in future studies of this
birch aphid fauna in California, a key to apterous and alate viviparae of these five
aphid species is presented.

Key to Birch Aphids in Northern California

1 Apterous (wingless) viviparous females. 2

Alate (winged) viviparous females. 5

2(1) Dorsum with blackish banding. 3

Dorsum not conspicuously blackish banded. If bands present, they are
pale and indistinct. 4

3(2) Dark banding only on abdominal tergite VIII. Very small, plump, oval.

greenish aphids (body length 0.9-1.4 mm). Short, 5-segmented
antennae.

Callipterinella minutissima (Stroyan)

Tresent address: Department of Entomology, Cornell University, Ithaca, N.Y. 14853-0999.

319

320

PAN-PACIFIC ENTOMOLOGIST

Dorsum with regular blackish banding on all abdominal segments. Color
variable from green to red. Body length 1.6-2.5 mm. Antennae
6-jointed.

Callipterinella calliptera (Hartig)

4(2) Cauda broadly triangular (Fig. 1). Small (1.5-2.0 mm), rather flat, oval
aphids. Frons with six capitate hairs placed on 4 tubercles.

Apices of antennae dark.

Betulaphis brevipilosa Bdrner

Cauda slightly to distinctly constricted (Fig. 2). Delicately built aphids
with long, thin legs. Body length 1.9-2.7 mm. Frons lacking capitate hairs
on tubercles. Antennae pale with dark apices of segments.

Calaphis flava Mordvilko

5(1) Large aphids (body length >3 mm). Pale green, thorax with brown
coloration. Abdomen sometimes with short, dark cross bars on tergite IV
and/or V. Sometimes covered with flocculent white or bluish-white wax.

Euceraphis betulae (Koch)
Body length < 3 mm. 6

6(5) Cauda broadly triangular (Fig. 1). Dorsum with very short, almost
invisible hairs; margins of posterior segments glabrous. Abdomen usually
with dark spot on tergites III-V (-VI) and dark marginal markings.

Betulaphis brevipilosa Bdrner

Cauda slightly constricted or knobbed (Fig. 2). Dorsum with distinct,
visible hairs. 7

7(6) Frons concave (Fig. 3). Siphunculi with a thickened flared apex (Fig. 5).

Calaphis flav a Mordvilko

Frons almost straight (Fig. 4). Siphunculi without flared apex (Fig. 6). 8

8(7) Processus terminalis about 1.75 or more times as long as basal part of
antennal segment VI (Fig. 7). Abdomen sometimes with dark, dorsal
cross [bars]. Body length 1.5-2.4 mm.

Callipterinella calliptera (Hartig)

Processus terminalis about 1.25-1.50 times as long as basal part of
antennal segment VI (Fig. 8). Abdomen without dorsal cross bars. Body
length 1.5-1.9 mm.

Callipterinella minutissima (Stroyan)

Treatments of Species

Betulaphis brevipilosa Bdrner

Apterous viviparous female. —Small (body length 1.5-2.0 mm), rather flat oval,
grass- or lime-green aphids. Abdominal cuticle usually wrinkled. Abdomen with
very short hairs except for marginal hairs on segments (IV-) V-VIII. Frons with 6
capitate hairs on four tubercles. Tarsi and apices of antennae dark. Siphunculi low,
truncate. Cauda broadly triangular.

Alate viviparous female. —Abdomen usually grass-green or yellowish with dark
spot on tergites III-V (-VI) and dark marginal sclerites. Hairs on frons fine and
acute. Siphunculi sometimes dark.

Eggs. —^Black, ellipsoid and without distinguishing markings. Length: x = 0.516 mm
(s^ = 0.007, n = 20); Width: x = 0.258 mm (s- = 0.005, n = 20).

VOLUME 63, NUMBER 4

321

9

Figure 1. Betulaphis brevipilosa. 2, 3, 5. Calaphis flava. 4, 6, 7. Callipterinella calliptera. 8.
Callipterinellaminutissima. 9. Euceraphis betulae. 1,2. cauda. 3,4. head. 5,6. siphunculus. 7,8. processus
terminalis. 9. antennal segment III.

Biology .—Common on B. pendula. Avoids new leaves in spring. Dense
aggregations are frequent. Extremely sessile aphid. Not ant tended.

Calaphis flava Mordvilko

Apterous viviparous female .—Pale green or yellowish, delicately built aphids with
long, thin legs. Body length 1.9-2.7 mm. Hairs on dorsum long and capitate. Frons

322

PAN-PACIFIC ENTOMOLOGIST

concave. Antennae a little longer than body. Apices of antennal segments dark. Legs
pale with knees, apices of tibiae and tarsi dark. Pale siphunculi low and truncate with
thickened, flared apices. Cauda slightly constricted.

Alate viviparous female. —Pale green or yellowish. Hairs on dorsum short and
blunt. Wing veins strong and black bordered.

Biology. —Uncommon. Populations occur predominantly on B. pubescens.
Developing leaves are preferred in spring. Not ant tended.

Callipterinella calliptera (Hartig)

Apterous viviparous female. —Green to red with blackish dorsal cross bars on all
abdominal segments. Body length 1.6-2.5 mm. Eyes red. Antennae pale with apices
of segments dark. Siphunculi and legs dark. Cauda slightly constricted, knobbed.

Alate viviparous female. —Abdomen sometimes with dark dorsal cross bars.
Processus terminalis 1.75 or more times the length of the basal part of antennal
segment VI.

Eggs. —Black, taper toward one end, meso-dorsal suture quite evident. Length:
X = 0.524 mm (s- = 0.006, n = 20); Width: x = 0.243 mm (s- = 0.005, n = 20).

Biology. —Common, predominantly found on B. pendula. In spring, developing
leaves preferred. Throughout the year, found in leaves tied together by insects, e.g.,
spiders or Lepidoptera larvae. Often found in aggregations of many individuals
which are ant tended. Fairly mobile aphid, running when disturbed.

Callipterinella minutissima (Stroyan)

Apterous viviparous female. —Very small (body length 0.9-1.4 mm), plump, oval
aphids. Greenish with one dark dorsal band on abdominal tergite VIII. Vertex with a
large dark sclerite. Short, 5-segmented antennae. Apical antennal segments usually
dark. Siphunculi shorter than wide. Cauda slightly constricted, knobbed. White tuft
of wax present below cauda.

Alate viviparous female. —Abdomen greenish. Apices of antennae, tibiae and
whole tarsi dark. Antennae 6-segmented. Processus terminalis 1.25-1.50 times as
long as basal part of antennal segment VI.

Biology. —Localized populations present inside of female catkins throughout the
summer. On leaves during early spring and fall, preferring leaves spun together by
insects. Usually aggregated when found on leaves. Not ant tended.

Euceraphis betulae (Koch)

Viviparous female. —All viviparous females alate. Large aphids (3.0-3.6 mm
long), elongate, pale green. Body sometimes clothed in flocculent white or
bluish-white wax. Head with dark dorsal spot or longitudinal stripe. Prothorax with
two brown spots, meso- and metathorax brown. Abdomen sometimes with short,
dark cross bars on segments IV-V or IV only. Antennae as long as body. Basal parts
of antennal segments III and IV dark in some specimens from spring and autumn.
Antennal segment III with 14-26 characteristic narrow transverse oval rhinaria
almost in a line (Fig. 9). Long legs sometimes with darkened tarsi and distal ends of
tibiae. Siphunculi about 0.03 times body length, truncate, sometimes dark. Cauda
knobbed.

Eggs. —Black, ellipsoid, and without distinguishing markings. Length: x = 0.802
mm (s- = 0.008, n = 20); Width: x = 0.394 mm (s^ = 0.007, n = 20).

VOLUME 63, NUMBER 4

323

Biology .—Common on B. pendula. Prefers new, developing leaves in spring and
senescent leaves in early autumn. Forms loose aggregations usually of few
individuals. Highly mobile aphid which runs or drops from leaves readily. Not ant
tended.

Acknowledgments

We thank K. S. Hagen for encouraging the writing of this key and for reviewing an
earlier draft of this manuscript. We also thank J. K. Liebherr for his comments on
this manuscript. H. L. G. Stroyan, formerly of the Ministry of Agriculture, Fisheries
and Food, England, and the late D. Hille Ris Lambers, Center for Aphid Research,
The Netherlands, identified aphid specimens.

Literature Cited

Grushkowitz, B. W. 1976. The population dynamics of two species of aphids, Callipterinella calliptera
(Hartig) and Euceraphis punctipennis (Zetterstedt), influencing ornamental birch trees in San
Leandro, California. M.A. thesis, San Jose State Univ., 151 pp.

Elajek, A. E. 1984. The population ecology of three species of aphids occurring on Betulapendula Roth in
northern California. Ph.D. diss., Univ. of Calif., Berkeley, 219 pp.

Hajek, A. E. 1985. Callipterinella minutissima, an aphid living in birch catkins. J. Natl. Hist. 19:623-626.
Hajek, A. E. 1986. Aphid host preference used to detect a previously unrecognized birch in California.
Env. Ent. 15:771-774.

Heie, O. E. 1972. Bladlus pa birk i Danmark (Horn., Aphidoidea). Ent. Meddr. 40:81-105.

Heie, O. E. 1980. The Aphidoidea (Hemiptera) of Fennoscandia and Denmark. 1. General Part. The
Families Mindaridae, Hormaphididae, Thelaxidae, Anoeciidae and Pemphigidae. Fauna
Entomologica Scandinavica, Vol. 9. Scand. Sci. Press, Klampenborg, Denmark, 236 pp.

Heie, O. E. 1982. The Aphidoidea (Hemiptera) of Fennoscandia and Denmark. IT The Family
Drepanosiphidae. Fauna Entomologica Scandinavica, Vol. 9. Scand. Sci. Press, Klampenborg,
Denmark, 176 pp.

Stroyan, H. L. G. 1977. Homoptera, Aphidoidea (Part): Chaitophoridae and Callaphididae. Handbooks
for Ident. of Brit. Ins. 114(a). Royal Ent. Soc., London, 130 pp.

Scientific Note

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 324

Heimbra opaca (Ashmead) (Hymenoptera: Eurytomidae) Discovered in Idaho; A
New State Record for the Subfamily Heimbrinae^

During the years 1981-1985, four specimens of Heimbra opaca (Ashmead) were
collected at three widely separated localities in Idaho. All three sites are quite typical
northern Great Basin grasslands with scattered shrubs. This is the first report of H.
opaca, and thus the subfamily Heimbrinae, in Idaho. This species was previously
known from California, Arizona, New Mexico, Utah, Colorado, Kansas and
Montana (Burks 1979, in Krombein et al., Cat. Hymen. Am. N. of Mex., Vol. 1, p.
846). The specimens of this distinctive species were identified by the authors using
key criteria (Burks 1971, Trans. Am. Ent. Soc. 97:1-89).

Specimens of H. opaca examined; Idaho: 1 $ Nez Perce Co., Hellsgate State Park,
4 mi. S. Lewiston, 8 Sept. 1981, T. D. Miller, collector; 19 same locality, 20 July to
15 Aug. 1985, Moericke trap, T. D. Miller, collector; 19 Clark Co., Blue Dome, 6
July 1982, J. B. Johnson, collector; 19 Elmore Co., 14.7 mi, NE Anderson Ranch
Dam, 3 Aug. 1982, sweeping grass, J.B. Johnson, collector. Specimens were
deposited in the William F. Barr Entomological Museum, University of Idaho.

James B. Johnson and Terry D. Miller, Department of Plant, Soil and
Entomological Sciences, University of Idaho, Moscow, Idaho 83843.

^Published with the approval of the director of the Idaho Agricultural Experimental Station as Research
Paper No. 86743.

324

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 325-327

A New Species of Andrena at the Micrandrena-Scaphandrena
Boundary (Hymenoptera: Apoidea)

U. N. Lanham

University of Colorado Museum, Campus Box 218, Boulder, Colorado 80309.

Abstract. —A new species of Andrena, A. (Micrandrena) robinsoni is described
from Colorado. Two related species, trapezoidea Viereck and primulifrons Casad
are transferred from the subgenus Scaphandrena where they were placed by Ribble
to the subgenus Micrandrea.

The two subgenera Micrandrena and Scaphandrena are closely related, and both
have been monographed by Ribble (1968, 1974). Scaphandrena has been given
special attention on account of the extensive introgressive hybridization by two or
three of its species in the Rocky Mountain region, the trihybrid complex ''scurra
Viereck x capricornis Casad and Cockerell x arabis Robertson” of Ribble (1973,
1974) and further described by me (1981,1984).

In my experience in Pennsylvania and the Rocky Mountain region, Scaphandrena
is an oligolege of small-flowered crucifers. Species of Micrandrena are frequently
collected on the same hosts as pollen-gathering females of Scaphandrena, but
Micrandrena as a group has a wide range of pollen sources. Ribble thought that the
primulifrons group, originally assigned to Micrandrena was a crucifer oligolege, and
finally suggested that on the basis of morphology and nomenclatorial convenience it
should be transferred to Scaphandrena, as it now appears in the Catalogue of
Hymenoptera in America North of Mexico (Krombein et al, 1979). Ribble states that
within Scaphandrena the closest relative of the sibling species primulifrons Casad
and trapezoidea Viereck (the primulifrons group) is capricornis.

In collecting specimens of the hybrid complex for the University of Colorado
Museum there accumulated a small lot of a species of the primulifrons group which I
provisionally considered to be trapezoidea Viereck. Recently Frank Parker and
Terry Griswold of the USDA Bee Systematics and Biology Laboratory at Logan,
Utah, lent me specimens of trapezoidea from Texas determined by Ribble. The
Colorado specimens turned out to be a third species of the primulifrons group, which
is described below.

Andrena (Micrandrena) robinsoni, New Species

This Colorado species has the strongly punctate abdominal terga characteristic of
the primulifrons group, but can be distinguished at a glance from the females of the
other two species, the rare trapezoidea and common primulifrons Casad, whose
distribution lies to the south and southwest, by a shiny, very sparsely punctate
median area of about one third the length of the 2nd abdominal tergite, with both
anterior and posterior margins becoming closely and strongly punctate. The general

325

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PAN-PACIFIC ENTOMOLOGIST

appearance of the bee is dominated by the three white abdominal hair bands and the
contrasting bare, shiny anterior third of the abdomen. Rohinsoni is also more robust,
8 mm in length instead of 7.

Female .—Length 8 mm. Integument black to brown black with obscure metallic
reflections of blue and green, except for the obscure amber apices of the tergites and
the bright reddish brown flagellum of the antennae.

Head with brownish white hairs, labral fringe golden-amber; measurements of the
facial quadrangle (distance between middle of eyes and vertex and bottom of
clypeus) with w:l ratio of 3.6:4.5; clypeus with upper half markedly flattened, dull
and strongly reticulate, strongly contrasting with the shiny and polished lower half
which is coarsely and irregularly punctate; facial fovea seen from front dark,
golden-tomentose seen obliquely, upper end occupying less than half space between
it and ocellus, widening centrally then narrowing below to about 1/2 width of space
between it and antennal insertion, ending just below level of insertion; upper ocelli
1/2 ocellus width from vertex; antenna with segment 3 not quite as long as 4 -h 5;
process of labrum 2-3 times as wide at base as at tip, which is truncate, sometimes
slightly emarginate; labrum shallowly concave, ending with a narrow raised rim;
galea reaching as far as apex of 4th segment of maxillary palpus.

Thorax covered with long, slender grayish-white hairs dorsally, so sparse that the
moderately reticulate, semi-shining, closely (2-3 p-w apart) and strongly punctate
integument of notum is not obscured; hair on pleura thickened, whiter; scutum with
integument like that of notum, scutellum and triangle of propodeum
strong-reticulate, triangle with weak longitudinal wrinkles above, rest of propodeum
strongly reticulate but semi-shining and obscurely punctate; pronotum with lateral
lines extending from pit at base of pronotal lobe diagonally across side of pronotum,
fading out at about 1/3 distance to midline; corbiculum without anterior fringe,
posterior half with sparse long simple hairs; legs with integument black to
brown-black, tibial spurs white-translucent, middle basitarsis broadened, lateral
margins curved outward, hind leg with tibia strongly widened apically, the basitarsis
about half its width at point of attachment, face of tibial scopa of short and simple
hair, the profile (not hair-length) of posterior fringe of scopa at midpoint of tibia a
little less than half width of tibia at apex, trochanteral floccus imperfect, with basal
hairs short and not recurved; tooth of hind tarsal claw with length about 1/4 distance
from inner base of tooth to tip of claw; forewing 6 mm, veins amber, membrane
clear, wingtips slightly infused with amber, pterostigma rather slender, less than I 1/2
times width of prestigma, first transverse cubital vein ending 3-4 vein-widths distant
from pterostigma, basal nervure meeting or falling slightly short of nervulus.

Abdomen with three strong, appressed hair bands on apices of tergites 2-4, that of
second complete or interrupted medially by about i/4th the width of the tergite, hairs
white, as are the sparse fine decumbent hairs between the bands, caudal fimbria
brownish orange; integument of 1st three tergites non-reticulate, polished, 4th
weakly reticulate, 1st with wide-scattered weak punctures, the 2nd with anterior and
posterior margins closely and coarsely punctate, with punctures 1-3 p-w apart,
becoming increasingly sparse toward mid-third of third of tergite, where the
punctures are coarse but very widely scattered, 3rd and 4th tergites entirely closely
strong-punctate.

Holotype female. —COLORADO, 10 mi SW Loveland, 5500', 26 May 1980, fls.
Physaria belli, P. Robinson.

VOLUME 63, NUMBER 4

327

Paratypes female. —2, same data as holotype; 2 with same data except U. Lanham;
1, COLORADO Weld Co., 6 mi ENE Nunn, 8 June 1981, fls. Rorippa sinuata, U.
Lanham; 1, COLORADO, Weld Co., Briggsdale, 19 May 1979, fls. Descurainia, U.
Lanham.

An additional female, COLORADO, Baca Co., 31 mi SSW Springfield, 4 May
1977, U. Lanham and P. Byron.

A byproduct of this study was the discovery of a character which distinguishes
females of the primulifrons group from capricornis and from the females of the
several other species of Scaphandrena available to me. The visible labrum in the
primulifrons group is shallowly concave in profile, while that of capricornis and the
sundry species of Scaphandrena is evenly convex. It seems to me that the three
species of the primulifrons group do not pose the danger of sinking Scaphandrena in
Micrandrena as feared by Ribble, and should be returned to Micrandrena. Also the
females of six species of the piperi group of Micrandrena, thought by Ribble (1968) to
intergrade with Scaphandrena, seen by me, have the labium similarly concave.

Literature Cited

Krombein, K. V. et al. 1979. Catalogue of Hymenoptera in America North of Mexico. Smithsonian
Institution Press, Washington, D.C. 3 vols.

Lanham, U. N. 1981. Some Colorado Andrena of presumed hybrid origin, with special reference to the
tarsal claws. J. Kansas Entomol. Soc. 54:537-546.

-. 1984. The hybrid swarm of Andrena in western North America: a possible source for the

evolutionary origin of a new species. J. Kansas Entomol. Soc. 57:197-208.

Ribble, D. W. 1968. Revisions of two subgenera of Andrena: Micrandrena Ashmead and Derandrena,
new subgenus. Bui. Univ. Nebraska State Museum 8:237-394.

-. 1973. Hybridization between three species of andrenid bees. Ann. Ent. Soc. America.

66:1281-1286.

-. 1974. A revision of the bees of the genus Andrena of the Western Hemisphere subgenus

Scaphandrena. Trans. American Ent. Soc. 100:101-189.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 328

Publication Received

Cladistic Analysis of North American Platynini and Revision of the Agonum
extensicolle Species Group (Coleoptera: Carabidae). By James Kenneth Liebherr.
University of California Publications in Entomology, vol. 106, x +198 pp., 70 figs.,
36 tables, 7 appendixes. Issue date on review card given as “August 1986.” Received
by PCES at CAS 11 August 1986. Published and distributed by University of
California Press, 2120 Berkeley Way, Berkeley, California 94720, U.S.A.,
telephone (415) 642-4562. Price $16.95 paperbound. ISBN 0-520-09958-3.

The introduction states in part:

“John L. LeConte wrote of the Agonum extensicolle species group in 1854, saying
that it is “a group of extreme complexity, containing winged species of metallic
colors, bluish or greenish, with the base of the antennae, the feet, and occasionally
the thorax rufous or pale piceous.” LeConte recognized that the species within the
Agonum extensicolle group are difficult to diagnose. Several of the species are widely
distributed geographically, and the accompanying variability in biometric features
has caused past authors to describe and name many invalid taxa. Six of the seven
species are found in the American Southwest and in adjacent Mexico, one of the last
areas in North America for which the carabid fauna remains poorly understood. This
study circumscribes the species using qualitative morphological characters,
electrophoretic data analyzed at the population level, and discriminant function
analysis based on biometric measurements.

The A. extensicolle group is distinct within the genus, supporting recognition of it
as a monophyletic lineage. Relative confidence that a taxon is monophyletic is a
prerequisite for certain aspects of systematic study. The study explores the
phylogenetic affinities of the species in the A. extensicolle group based on genetic
data obtained from starch-gel electrophoresis and on internal and external
morphological characters. The hypothesis of phylogenetic relationships is then
analyzed in light of the present-day distributions of the species. This analysis
provides insight into the speciation events and mechanisms that have brought about
the present day diversity of the A. extensicolle group.”

—Paul H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San
Francisco, California 94118.

328

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 329-332

A New Species of Paruroctonus from Coastal California

(Scorpiones: Vaejovidae)

Stanley C. Williams

Department of Biology, San Francisco State University, San Francisco, California
94132.

Abstract. —A new species of scorpion from coastal California is described and
named Paruroctonus maritimus Williams. Its nearest relatives appear to be
Paruroctonus boreus (Girard) and Paruroctonus silvestrii (Borelli).

A few years ago a small series of Paruroctonus was collected under cardboard and
other surface debris by Roy Johnson, along railroad tracks at Sea Side, Monterey
County, California. At the time, this sample posed certain problems of
interpretation, although it appeared to be Paruroctonus boreus (Girard). Since the
collection site was well outside of the known distribution of P. boreus, restricted in
area, and ecologically disturbed, the identification of this species was tentative.
Reinvestigation of these specimens indicates they are not P. boreus, but an
undescribed new species that appears to inhabit the coastal sand dune community.
This new species is here described and named. The measurements cited are as
defined by Williams (1980).

Paruroctonus maritimus Williams, New Species
(Fig. 1, Table 1)

Diagnosis. —Members of subgenus Paruroctonus. Total length up to 50 mm. base
color of exoskeleton pale yellow with contrasting dusky-black marbling dorsally on
carapace and mesosoma, dark marbling not extending to posterior margin of
mesosomal terga, metasoma with ventral and ventrolateral keels outlined in dusky
pigment; frontal margin of carapace convex; pectine teeth 24-27 in males, 18-20 in
females; metasoma with ventral keels smooth to obsolete on I, smooth on II-IV,
serrate on V; metasoma with ventrolateral keels smooth to granular on I-II, smooth
on III, smooth to crenulate on IV; chela with supernumerary denticles 6 on fixed, 7
on movable finger; chela with primary row denticles divided into 6 subrows on fixed
finger, 7 subrows on movable finger.

Related to Paruroctonus boreus (Girard) and Paruroctonus silvestrii (Borelli).
Distinguished from P. silvestrii by pigment pattern of mesosomal terga not extending
to posterior margin of terga, metasoma with 4 pairs of ventral macrosetae on
segment II. Differs from P. boreus by less distinct proximal gap between fingers of
chela in males; slightly fewer pectine teeth; median ocelli more forward on carapace;
ratio of carapace length to frontal margin distance less than 2.0; metasoma of males
less elongate, metasomal segment IV length to width ratio less than 1.9.

Holotype. —Male. Coloration: Base color of cuticle pale yellow, carapace with
black marbling, frontal margin outlined in dusky-black; mesosomal terga with

329

330

PAN-PACIFIC ENTOMOLOGIST

Figure 1. Paruroctonus maritimus Williams, holotype, dorsal and ventral views.

underlying dusky black markings, these not extending to posterior margin of terga,
dusky markings mostly limited to anterior 1/4 of tergum 7; walking legs with
inconspicuous, localized, dusky-black markings prolaterally; brachium, humerus,
and chela with inconspicuous underlying dusky-black markings; fingers of chela
similar to palm in color; pectines white, mesosomal sterna lacking dark markings;
metasoma with ventral and ventrolateral keels outlined in dusky pigment on
segments II-V. Prosoma: Carapace frontal margin slightly convex, with 2-3 pairs of
macrosetae; lateral ocelli 3 per group, median ocelli on smooth, raised ocular
tubercule; sternum elongate pentagonal, 4 pairs of sternal macrosetae, median
posterior depression, deep, broad. Mesosoma: Terga finely granular, terga 5-7 with
subtle obsolescent median keel, tergum 7 with 2 pairs of granular lateral keels;
genital opercula triangular, 10-11 pairs of genital macrosetae, distinct genital
papillae; pectine basal sternum with deep median notch on anterior margin; comb
with three marginal lamellae, middle lamellae with angular basal piece and 19
subcircular sclerites, fulcra triangular, 4-6 ventral macrosetae per fulcrum; stigma
elongate, 3.5 times longer than wide; sterna 2-6 smooth, agranular, sternum 7 with 1
pair of granular submedian keels over one-half of sternum. Metasoma: Dorsal keels
granular on I-V; dorsolateral keels granular on I-V; lateral keels granular on I,

VOLUME 63, NUMBER 4

331

Table 1. Measurements (mm) of Paruroctonus maritimus Williams, new species, holotype (male) and allotype. Ab¬
breviations as follows: 1 = length, w = width, d = depth, fmd = frontal margin distance, ditd = distal internal tricho-
bothrium distance, p-row = primary denticle row of chela, ff = fixed finger, mf = movable finger.

holotype (male)

allotype

Total length

41.0

48.0

Carapace, (1/w at median eyes)

4.6/3.6

5.8/4.5

Diad (width/fmd)

0.95/2.1

1.1/2.7

Metasoma, length

19.0

21.3

Segment I (1/w/d)

2.5/2/7/2.2

2.9/3.2/2.5

Segment II (1/w/d)

3.0/2.6/2.1

3.3/3.1/2.5

Segment III (1/w/d)

3.3/2.5/2.1

3.6/3.0/2.4

Segment IV (1/w/d)

4.2/2.4/2.0

4.5/2.8/2.4

Segment V (1/w/d)

6.0/2.1/1.8

7.0/2.6/2.3

Telson, length

5.4

6.5

Vesicle (1/w/d)

3.3/2.0/1.6

7.0/2.6/2.3

Aculeus (1)

2.1

2.5

Pedipalp, Humerus (1/w)

3.8/1.3

4.8/1.7

Brachium (1/w)

4.2/1.8

5.1/2.3

Chela (1)

7.2

8.6

Palm (1/w/d)

4.1/2.2/2.9

4.8/2.4/3.2

Movable finger (1/base)

4.2/1.05

5.7/1.4

Fixed finger (1/ditd)

3.1/2.7

3.8/3.2

Supernumerary denticles (ff/mf)

6/7

6/7

Fixed finger p-row denticles

5-7-7-9-13-9

6-8-8-10-11-17

Movable finger p-row denticles

1-6-10-10-12-15-7

1-6-9-10-13-15-11

Pectine teeth (left/right)

24/26

18/18

Stigma 3 (1/w)

0.35/0.1

0.4/0.1

granular on posterior 1/4 of II, obsolescent except for 3 posterior granules on III,
absent on IV, granular on anterior half of V; ventral lateral keels smooth to granular
on I-II, smooth on III, smooth to crenulate on IV, serrate on V; ventral keels smooth
to obsolete on I, smooth on II-IV, serrate on V. Telson: Vesicle smooth and
lustrous, subtle subaculear tubercule flanked laterally by 1 pair of long reddish
macrosetae. Pedipalps: Chela with swollen palms, keels of palm roughly granular;
subtle scallop between fingers proximally when fingers closed; supernumerary
denticles 6 on fixed finger, 7 on movable finger; primary row denticles divided into 6
subrows on fixed finger, 7 on movable finger by distinctly enlarged denticles; palm
with well-developed ventral prolateral and ventral retrolateral granular keels.
Chelicerae: Ventral margin of movable finger with 2-3 subtle crenulations, fixed
finger lacking apparent denticles or crenulations; ventral surface of movable finger
not conspicuously hirsute, with about 6 long ventral macrosetae.

Allotype. —Female. Similar to holotype in color and structure except as follows:
Longer in total length; pectines much smaller, shorter, with fewer teeth; no genital
papillae; median ocelli slightly smaller; metasomal segments slightly less elongate

332

PAN-PACIFIC ENTOMOLOGIST

(ratio of metasomal length to width 6.7); chela slightly more elongate (ratio of chela
length to width 3.6).

Topoparatype variation. —Similar to holotype and allotype except: Total length
21.3-50.0 mm; pectine tooth counts (per comb) 24-27 in juvenile males (mode 26),
18-20 in females (mode 19); juveniles with base color of cuticle whitish, dusky-black
marbling more contrasting, more extensive; cheliceral denticles on ventral margins
of movable finger more developed, 4-5 crenular denticles; ventral margin of fixed
finger with 2 small granular denticles; adults with cheliceral denticles more worn and
subtle on movable finger, not apparent on fixed finger.

Type data. —Holotype (male), allotype, California: Monterey County, Seaside, 7
Apr. 1985, Coll. Roy Johnson. Depository: California Academy of Sciences,
Entomology Type No. 15791. Named Paruroctonus maritimus in reference to its
coastal habitat.

Topoparatypes studied. —California: Monterey County, Seaside, 3, 5, 7 Apr.,
1985, Coll. Roy Johnson, 21 females, 3 juvenile males.

Remarks. —This species is known only from coastal habitats of central California.
In the type locality it was found under surface debris on dry, fine coastal dune sand.
Field collections suggest that its preferred habitats may be coastal sand dune
communities in this region. It was curious that of the 26 specimens collected all were
mature females except one mature male and 3 juvenile males.

Acknowledgments

I am particularly indebted to Roy Johnson for collecting the specimens used in this
study. I also gratefully acknowledge the following colleagues for materially assisting
this study: Vincent F. Lee and Jack T. Tomlinson critically read this manuscript; Jett
S. Chinn assisted with the illustrations; Paul H. Arnaud, Jr. provided research
facilities at the California Academy of Sciences. The West Point Academy of Arts
and Sciences partially supported this project.

Literature Cited

Williams, S. C. 1980. Scorpions of Baja California, Mexico, and adjacent islands. Occas. Pap. Calif.

Acad. Sci. 135:1-127.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 333

The Southern Green Stink Bug, Nezara viridula Linnaeus
(Heteroptera: Pentatomidae); New Location

M. P. Hoffmann, L. T. Wilson and F. G. Zalom^

Department of Entomology, University of California, Davis, California 95616.

The southern green stink bug, Nezara viridula Linnaeus was collected from a 30.6
hectare commercial field of processing tomatoes located five kilometers north of
Davis, Yolo Co., California on 12 September 1986. N. viridula occurs across much of
the southeastern United States and as far west as Texas. Menezes, Herzog, Teare
and Sprenkel (1985. Phenological events affecting southern green stink bug on
soybean, Proc. Crop Sci. Soc. Florida, 44:227-231). This species has not previously
been reported in California.

The abundance of N. viridula in this tomato field was determined by recording the
number of nymphs and adults found in 60 plots each 2m in length. In each plot the
tomato plants were cut off at ground level and shaken onto a drop cloth. The number
of N. viridula found on the drop cloth and upper 2.5-5.0 cm of soil was recorded. The
mean number of N. viridula per meter of tomato bed was 5.97. Approximately 32,28
and 40 percent of the population were small-medium nymphs (1st to 3rd instar), large
nymphs (4th and 5th instars) and adults, respectively. One hatched N. viridula egg
mass was found near a group of first and second instars. None of the eggs appeared to
have been parasitized. On September 29 we collected 50 adult N. viridula and caged
them in our laboratory. As of October 30 no parasites had emerged from these
individuals.

The southern green stink bug has a very wide host range including: fruit and
ornamental trees, field crops, vegetables, and weeds. Todd and Herzog (1980,
Sampling phytophagous Pentatomidae on soybean, pp. 438-478 in Kogan and
Herzog ed. Sampling methods in soybean entomology, Springer-Verlag, N.Y. 587
pp.). If successfully established and in the absence of satisfactory natural enemies
this stink bug could have a significant economic impact on numerous crops in
California.

Personnel from the California Department of Food and Agriculture and personnel
from surrounding County Agricultural Commissioners offices are conducting a
survey to determine the distribution of N. viridula in California.

We thank Ray Gill, George Buxton and Alan Hardy (CDFA) for confirming our
identification of N. viridula.

^U.C. Integrated Pest Management-Implementation Group, University of California, Davis, CA.

333

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 334

Publication Received

The Butterflies of North America. A Natural History and Field Guide. By James
A. Scott. Stanford University Press, Stanford, California 94305. xiii + 583 pp., 71
figs., 64 col. pis., distribution maps. ISBN 0-8047-1205-0. Publication date given as
“September 26, 1986”; review copy received by PCES at CAS on 12 September
1986. Price $49.50 hard cover.

The book jacket states:

“The only field guide to cover all North American butterfly species, this
monumental work is also a complete natural history, fully describing the biological
and ecological world of butterflies in general. It is without question the most
important book on butterflies in several decades, and the most complete treatment of
a major butterfly fauna ever published.

The book is written at several levels of detail, most of it accessible to anyone, and
employs the minimum of technical terms necessary for ensuring scientific accuracy.
Extensive introductory material—a book in itself—stresses butterfly biology and
ecology: structure, flight, metamorphosis, hibernation, physiology, roosting,
migration, mating, egg laying, intelligence, social behavior, larval and adult foods,
enemies, mimicry, variation, evolution, habitats, distribution and conservation. The
main text is arranged in phylogenetic sequence, and characteristics or behavior
common to all members of a family, subfamily, or tribe are discussed at those levels.
The skippers, a large group often excluded, are treated in full.”

—Paul H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San
Francisco, California, 94118.

334

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 335-337

A New Species and Records of the Genus Toxorhina, Subgenus
Eutoxorhina From the South Pacific (Tipulidae, Diptera)

C. Dennis Hynes

Department of Biological Sciences, California Polytechnic State University, San
Luis Obispo, California 93407

Abstract .—Alexander (1934) described Toxorhina (Eutoxorhina) simplex from a
badly broken specimen which consisted of only the thorax, one wing, and the first
two abdominal segments. Two more specimens of the subgenus were found in 1963,
in the same geographical location (Fiji). These specimens correspond precisely with
the original description given by Alexander and, having studied the holotype, I have
little doubt that they represent the same species. I here add further notes on the
description of this species. I have examined another specimen from New Caledonia,
which is different from the specimens from Fiji, and I describe it here as a new
species. All specimens are being returned to the Bishop Museum.

Toxorhina (Eutoxorhina) simplex. Alexander 1934.

Two Females: length excluding rostrum 6.8-6.9 mm; wing 4.9-5.1 mm.; rostrum,
5.7-5.8 mm. The following are details other than those given by Alexander in the
original description. Rostrum light yellow-brown, darker at base and near tip.
Antennae dark brown, verticils at tip 4-5 times longer than the segments. Frons and
vertex light brown, pruinose; back of head and neck region dark brown. Dorsum of
the thorax pruinose, paratergites a lighter brown separating dark brown dorsal and
pleural areas. Coxae dark brown, femur light brown, grading to darker brown at tip;
tibia brown grading to darker brown at tip; tarsi uniformly brown. Abdominal
tergites and sternites dark brown; posterior sternites lighter. Ovipositor dark brown
at base; tips of valves much lighter brown. The two specimens differ only in the
intensity of the coloration.

Female. —Fiji, Viti Levu, Colo-i-suva, 3-6.iii.63 (C. Yoshimoto).

Female. —Fiji, Viti Levu, Nadarivata 850 m, 8-13.iii.63 (C. Yoshimoto).

Toxorhina (Eutoxorhina) parasimplex New Species

Male. —Length, excluding rostrum 4.1 mm; wing 4.3 mm; rostrum 3.6 mm;
antennae 5.3 mm.

Antennae with scape and pedicel yellow, fusion segment and flagellar segments
dark brown. Rostrum brown, shorter than the remainder of the body. Area
surrounding eyes and vertex pruinose black. Pronotum and mesonotum dark brown.
Propleural plates yellow-brown, remainder of thoracic pleura dark brown.
Mesonotal praescutum pruinose gray at lateral edges. Halteres uniformly reddish
brown. Legs with coxae dark brown; trochanters and basal portion of femur
yellow-brown grading to dark brown for the remainder of the leg; the hind legs
slightly darker. Terga of abdomen brown, sterna lighter to reddish brown. Segment

335

336 PAN-PACIFIC ENTOMOLOGIST

1

Figure 1. Toxorhina (Eutoxorhina) parasimplex n. sp. (b = basistyle; i = interbase; p = phallosome;
s = sternite)

just before the hypopygium appears as a dark brown ring. Last sternal segment with
an extended process which is furcate, the arms very short and blunt. Ninth tergite
with central area bordered by five short points. Wing venation exactly as described in
subgenus, tinted with brown, more hyaline between stigmal area and tip of wing; a
whitish stripe crossing m-cu; veins uniformly dark brown. Hypopygium with
elongate basistyle reddish brown; inner surface concave giving hollowed-out
appearance. A distinct line of large setae on interior surface of basistyle extending
diagonally from the ventral anterior margin to the dorsal posterior margin.

VOLUME 63, NUMBER 4

337

Interbases sigmoid in shape. Outer dististyle a broad triangular shaped base, which
then curves sharply cephalad into a darkened, elongate, triangular blade with one
small seta at the tip. Inner dististyle sharply curved, basally broad blade with a clear
fimbriated membrane on the outer edge. Phallosome large, stoutly bifurcate at tip,
base longer than arms of fork.

Holotype. —(male) New Caledonia, Cole des Roussettes 450-550 m, 4-6.ii.63
(J. L. Gressitt, C. Yoshimoto, N. Krauss),. BPBM slide 2037.

Acknowledgments

I wish to thank Dr. Neal Evenhuis for his courtesies while at the Bishop Museum
and for the privilege of describing these specimens.

Literature Cited

Alexander, C. P. 1934. New or little-known Tipulidae from eastern Asia. (Diptera), XVIII. Philippine
Journ. Sci. 53(3):267-298.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 338

Publication Received

Spiders. Webs, Behavior, and Evolution. Edited by William A. Shear. Stanford
University Press, Stanford, California 94305. xiv + 492 pp., illus. ISBN
0-8047-1203-4 (alk. paper). Publication date given as “September 12, 1986,” with
review copy received by PCES at CAS on 22 August 1986. Price $55.00 hard cover.

Collection of papers presented at the 1981 meeting of the American
Arachnological Society at the University of Tennessee. Contributors—James Edwin
Carico (Web removal patterns in orb-weaving spiders, pp. 306-318), Jonathan
Coddington (The monophyletic origin of the orb web, pp. 319-363), Frederick A.
Coyle (The role of silk in prey capture by nonaraneomorph spiders, pp. 269-305),
William G. Eberhard (Effects of orb-web geometry on prey interception and
retention, pp. 70-100), Robert R. Jackson (Web building, predatory versatility, and
the evolution of the Salticidae, pp. 232-268), Anthony C. Janetos (Web-site
selection: Are we asking the right questions?, pp. 9-22), Yael D. Lubin (Web
building and prey capture in the Uloboridae, pp. 132-171), W. Mitch Masters,
Hubert S. Markl, and Anne J. M. Moffat (Transmission of vibration in a spider’s
web, pp. 49-69), Susan E. Riechert and Rosemary G. Gillespie (Habitat choice and
utilization in web-building spiders, pp. 23-48), William A. Shear (Introduction, pp.
1-7; The evolution of web-building behavior in spiders: A third generation of
hypotheses, pp. 364—400), Mark K. Stowe (Prey specialization in the Araneidae, pp.
101-131), William James Tietjen (Social spider webs, with special reference to the
web of Mallos gregalis, pp. 172-206), and George W. Uetz (Web building and prey
capture in communal orb weavers, pp. 207-231). Taxonomic Glossary, pp. 403-432.
Bibliography, pp. 433-472. Index, pp. 474-492.

—P. H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San
Francisco, California 94118.

338

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 339-340

Geographical Inexactitude

Roy R. Snelling

Natural History Museum of Los Angeles County, Exposition Blvd., Los Angeles,
California 90007

In reading over recent issues of the Pan-Pacific Entomologist, I was, once more,
reminded of a persistent error which many entomologists in this country consistently
commit. For example, in his paper describing new tenebrionids from western North
America, John Doyen (1983) cited specimens from “Baja California del Norte” (p.
83) and from “Baja California Norte” (p. 90). Another such paper is that of Allen
and Murvosh (1963). The authors uniformly cited mayfly distribution records under
“Baja California Norte” and from “Baja California Sur.” Their map (Fig. 12) has the
two States properly identified.

These designations in both papers are incorrect and point to a problem to which I
have alluded in the past (Snelling, 1970), one that emphasizes the general ignorance
of historical events south of our border.

Prior to 1952, the peninsula of Lower California was divided into two territories:
Baja California del Norte and Baja California del Sur. In 1952 the northern territory
became the State of Baja California (Estado de Baja California: note that the
designation “del Norte” was dropped). At the same time, the southern portion
became the Territorio del Sur de Baja California. This Territory became Estado de
Baja California Sur in 1974 (Anonymous, 1984).

Obviously, it is convenient to designate “del Norte” or “del Sur,” as is commonly
done, but the fact remains that it is also geopolitically incorrect. Within their own
country, Mexicans have no problem. The peninsula, as a whole, is “Baja
California,” or simply “Baja”; the northern State is “Estado de Baja California,”
and the southern State is “Baja Sur.” Authors north of the border tend to be less
precise.

Thus, in looking at Doyen’s paper, I wonder how I am to interpret the statement:
“Adelonia filiformis Laporte from southern Baja California . . .,” etc.? Is the
reference to the southern part of the State of Baja California or is it to the southern
part of the peninsula (in other words, to the State of Baja California Sur)? Such
imprecision in geographic citations is common, as a quick perusal of recent literature
on insects of western North America will make clear.

There exists confusion of still another sort: the failure to recognize that the entire
peninsula is not a single state. For example, Evans (1966) consistently cites records
from the entire peninsula under “Baja California” as though it were a state
equivalent to Sonora or Michoacan.

All of this confusion is certainly not limited to published papers. Most
entomologists who collect there seem not to know the situation. This is attested by
the many specimens from recent collections that I have seen labelled “Baja
California del Norte” and “Baja California del Sur.” Then, there are the specimens

339

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PAN-PACIFIC ENTOMOLOGIST

from La Paz, Todos Santos, and other southern peninsular localities that are labelled
“Baja California.”

There is a solution, a way out of this confusion, and it is based on historical
precedent. It is a solution which, perhaps, might appeal to systematists, since it is
merely an extension of the principle of nomenclatural priority.

The entire peninsular region (Estado de Baja California plus Estado de Baja
California Sur) can be referred to as Lower California; this, of course, is a
straightforward translation of Baja California. The use of this term in entomological
literature is more than 100 years old (e.g., LeConte, 1861; Saussure, 1875). It has
also been used much more recently (Bohart, 1948). The two States may then be
properly designated Baja California and Baja California Sur with little or no
confusion.

There certainly should be no reason to suppose that this is linguistic “chauvinism”
on my part. It seems to me that it should be just as acceptable to use the Enghsh, viz.
Lower California, as to use Mexico, Panama, Peru, Spain, or Germany (rather than
the indigenous languages: Mexico, Panama, Peru, Espana, or Deutschland, etc.).

Then, there is “Sierra Nevada Mountains” (Gilbert, 1982). This is not an
uncommon error of redundancy. So, we see “Sierra Ancha Mountains,” “Sierra
Madre Mountains,” etc. Since the Spanish word “sierra” = “mountains,” there truly
is no excuse for this sort of error, other than simple carelessness.

Authors, of course, must assume initial responsibility and should make every
effort to avoid such errors. Reviewers, too, are not doing their job very well when
they allow such lapsi to pass. Finally, editors must assume their share of guilt.

In short, I suggest that we devote the same attention to detail and accuracy in
dealing with geography as we give to taxonomy in our work.

Literature Cited

Allen, R. K., and C. M. Murvosh. 1963. Taxonomy and zoogeography of the mayflies (Ephemeroptera;

Insects) of Baja California. An. Entomol. Soc. Amer., 76:425-433.

Anonymous. 1981. Mapa turistico de Carreteras. Secretaria de Asentamientos Humanos y Obras
Pubhcas, Mexico City.

Anonymous. 1984. Baja California. Automobile Club of So. Calif., Los Angeles, 128 pp. (This
Guidebook clearly and properly identifies the two States in the maps on pages 5 and 40).

Bohart, R. M. 1948. Contributions toward a knowledge of the insect fauna of Lower Cafifornia. No. 9.

Elymenoptera: Eumeninae. Proc. Cahf. Acad. Sci., ser. 4,24:313-336.

Doyen, J. T. 1983. New species of Tenebrionidae from western North America (Coleoptera). Pan-Pacific
Entomol. 58:81-91.

Evans, H. E. 1966. A revision of the Mexican and Central American spider wasps of the subfamily
Pompilinae (Hymenoptera: Pompihdae). Mem. Amer. Entomol. Soc. 20:1-442.

Gilbert, A. J. 1982. A new species of Coscinoptera Lacordaire from California (Coleoptera:
Chrysomelidae). Pan-Pacific Entomol. 57:364-370.

LeConte, J. L. 1861. Notes on the coleopterous fauna of Lower Cafifornia. Proc. Acad. Nat. Sci. Phila.
1861:335-338.

Saussure, H. de. 1874. Synopsis of American wasps. Smithsonian Misc. Colin. 254:1-392.

Snelfing, R. R. 1970. The social wasps of Lower Cafifornia, Mexico (Hymenoptera: Vespidae). Contri.
Sci., L. A. Co. Mus. Nat. Hist. 197:1-20.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 341-345

Marking Technique for Larvae

Raymond R. White and Michael C. Singer

Biology S-56, City College of San Francisco, 50 Phelan Ave., San Francisco, CA
94112, USA; Department of Zoology, University of Texas, Austin, TX 78712, USA

Keywords: larval marking, marking technique, field marking, insect larvae.

Abstract .—A technique for marking insect larvae as individuals is described and
the kinds of data derivable from studies of individually coded larvae are discussed.
The relative importance of larval stages is discussed.

Introduction

Biological explorations of plants and of insects still suffer from defects of omission.
In the case of plants, the roots have been too often ignored (Cody 1986). In the case
of insects, the immature stages have in general been much less studied than the
mature stage (adult). This understandable but unfortunate bias in scientific effort
limits the ultimate value of many population dynamic studies. If key factors
controlling population dynamics act during the life stages not studied, then analyses
of dynamic data remain either inconclusive or misleading. Hypotheses depending on
population dynamic data, such as those involving population genetics, suffer
accordingly. Control programs designed without accurate knowledge of the factors
affecting immature stages may have minimal effects on target population sizes.

Not only do population numbers often depend on phenomena occurring during
the immature stages, but the larval stage is frequently where the physical size
achieved by an adult insect is determined. For holometabolous insects that are short
lived as adults (the vast majority of Lepidoptera, for example), the mature larval size
(at least of females) relates directly to reproductive capacity and therefore to
potential for population increase. The last larval stadium is where most of an insect’s
weight is gained. In the case of Euphydryas editha (Boisduval) (Lepidoptera:
Nymphalidae), for instance, 60 to 80% of the mature larva’s weight is gained then
(Weiss, White, & Murphy unpublished). Therefore study of this single stadium may
contribute disproportionately more to knowledge of the biology of this and similar
insects than would the study of other stadia or stages.

For field study of phenomena such as mortality, growth, and dispersal it is
necessary to reliably identify individuals. For adult Lepidoptera this is commonly
done by means of some variation of the magic marker technique (Ehrlich &
Davidson 1960, Brussard 1970, Scott 1975, Singer & Wedlake 1981, Gall 1985). A
number of workers have used techniques for marking larvae, but such studies
remain the exception and techniques such as multilation are still current (Weseloh

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PAN-PACIFIC ENTOMOLOGIST

1985). Here we describe a technique for marking individuals that works within a
stadium for larvae of insects.

Description of Technique

Larvae of Euphydryas editha bayensis Sternitzky were collected at the Morgan Hill
(MH) site in Santa Clara County, CA, USA, in February and March 1985 and
January through March of 1986. Each larva was given a unique mark with Testors
enamel paint (available through such outlets as Long’s Drugs and hobby shops).
Among the dozens of colors available, several worked well with these dark-colored
larvae: 1103 red, 1108 light blue, 1114 yellow, 1145 white, 1127 orange, and 1134
purple. On these larvae, 1111 dark blue and 1124 green were not easily readable.
Usable light blues and greens could be produced by mixing the darker colors with
white. For lighter colored larvae, colors such as dark blue, green and black might be
effective. Metallic colors such as silver and gold cause violent reactions: vomiting and
fleeing. Testors 1170 light tan was too runny, producing large, messy marks.

Though there are thirteen body segments in the larvae of Lepidoptera that can,
with care, be identified (Howe 1975), we found that we could reliably distinguish dots
of paint on the left and right sides of segments A) near the head, B) near the middle,
and C) near the end. Very small dots of paint were applied with a sharpened
toothpick or with an insect pin (#3) to the subdorsal and/or lateral scoli (bristles) of
the appropriate segments. The paint did not wear off, nor did it seem to affect
behavior. Marked larvae were picked up 2 to 7 days after marking and release. So far
the maximum observed duration of the marks in the field is 21 days, on 2 larvae that
had hung up to pupate. Larvae that died of unknown causes and larvae that were
stepped on were found to retain their marks identifiably. In this species the final
stadium lasts just long enough (7 to 14 days during sunny weather) to be studied.
Keeping records of individual morphological traits where they exist, along with
painted codes, can help maintain the integrity of the system. Codes, by their nature,
are inevitably misread at some frequency.

When the last larval stadium ends and the skin is shed, it usually remains with the
pupa. Careful examination of shed skin allows some codes to be distinguished from
others (different colors, anal vs. cephalad, left vs. right; potentially also ventral vs.
dorsal). Thus individual pupae can be matched to larvae whose traits have been
measured.

By assigning numbers (Fig. 1) to the marking positions, each larva could be coded
with any of fifty different numbers, without changing colors of paint. It is, however,
better for the organism, the experiment, and the investigator to impose as few spots
of foreign material as possible. Restriction of the marking scheme to a single dot of
paint would allow six unique marks per color of paint used. Restriction to two dots
would allow 21 unique marks per color, three dots would allow 40 marks per color,
four dots would allow 50. Thus, a three dot system, given six positions and six colors,
provides for unique codes to be given to 240 larvae. Obviously the system can be
extended by careful use of additional marking locations (eight positions would
provide for 444 unique marks). Two colors can be used together on the same
individual to, in effect, add another color to the system.

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343

Figure 1. System for coding insect larvae individually. Upper left, key; others, examples.

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PAN-PACIFIC ENTOMOLOGIST

Uses for Marked Larvae

Marking larvae individually makes it possible to track weight gains of individuals
through time under field or laboratory conditions. This is very important because
other sampling methods are subject to significant error. For example, we (Weiss,
White, & Murphy, unpubl.) have taken field samples of larvae every ten days to
assess field growth rates. The resulting data generally show steady growth, but
exceptions occur. Even the steady growth curves misrepresent growth to some
degree because males lose weight in preparation for pupation while females are still
gaining. Preliminary data indicate that in this species the mature larva at its
maximum will lose 20% of its weight by the time the pupa is formed (larva, 400 mg:
pupa, 320mg). Since males and females of most holometabolous insects differ
significantly in size, the problem is a common one. Larvae may be kept individually
without marking in the laboratory, but laboratory studies of heliothermic insects are
often negatively influenced by the effects of replacing solar heat with ambient heat.
Migration of larvae, perhaps surprisingly, can also bias field samples of anonymous
(previously unidentified) larvae. Checkerspot butterfly larvae may commonly move
ten meters per day when experiencing adverse local conditions. For other
experiments, groups of marked larvae can be kept together in the laboratory without
the loss of individual identification. The behavior of particular individuals can be
tracked over time. Individuals may differ also in other ways (color morphs, for
instance) that may affect their growth or survival. Such traits may be recorded if
individuals are uniquely identified.

Finding Larvae

When larval hosts of phytophagous insects are known they can be searched at the
right time of year and larvae can often be found. However, for many species this
works poorly at best, and for many other species it does not work at all or the host is
still unknown. In the case of Euphydryas editha bayensis (the bay checkerspot
butterfly), the host is so common and dense that searching the host plants is very
similar to searching the entire habitat. These dark larvae can be found by looking
into one’s shadow. The spots that remain dark when in shadow may be larvae. These
animals are essentially thermal collectors and spend a lot of time basking in areas of
the sparsest vegetative cover. It is thought that speed of digestion is a limiting factor,
requiring time spent in the sun (Porter 1982). In the case of E. editha rubicunda (the
large collinsia checkerspot), the postdiapause larvae seem to be crepuscular, feeding
at dusk and dawn (Singer, pers. comm.). My own recent experience with isopods,
earwigs, and weevils on my garden Passiftora reminds me to emphasize the potential
value of night searches. The serious investigator may have to be active at midnight
with a powerful flashlight. He may also have to lie prone on a muddy substrate (as
one of us did this January) in order to see larvae that weigh under 20mg, but that have
already doubled in weight since breaking diapause.

Summary

All too often the more difficult life stages of experimental organisms are ignored in
terms of literature discussion as well as experiment. Marking individuals of such
stages is an invaluable tool for extending knowledge of the biology of insects. The

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345

technique described here is usable on the later instars of many species of insects. So
far we have found differential dispersal of larvae on slopes of different exposure and
have determined growth rates under some field conditions (Weiss, White, &
Murphy, unpubl.). We expect to use the technique described here for further
growth, microhabitat, and dispersal studies and to estimate larval survival to
pupation.

Acknowledgments

We owe Dennis Murphy and Waste Management, Inc. thanks for use of the
Morgan Hill field site. Helpful comments on the manuscript were made by Gordon
Pratt.

Literature Cited

Brussard, P. F. 1970. Field techniques for investigations of population structure in a ‘ubiquitous’
butterfly. J. Lepid. Soc. 25:22-29.

Cody, M. L. 1986. Roots in plant ecology. Tree 1:76-78.

Ehrlich, P. R. and S. E. Davidson. 1960. Techniques for capture-recapture studies of Lepidoptera
populations. J. Lepid. Soc. 14:227-229.

Gall, L. F. 1985. Measuring the size of lepidopteran populations. J. Res. Lepid. 24:97-116.

Howe, W. H. 1975. The Butterflies of North America. Doubleday, Garden City, N.Y.

Porter, K. 1982. Basking behaviour in larvae of the butterfly, Euphydryas aurinia. Oikos 38:308-312.

Scott, J. A. 1975. Flight patterns among eleven species of diurnal Lepidoptera. Ecology 56:1367-1377.

Singer, M. C. and P. Wedlake. 1981. Capture does affect probability of recapture in a butterfly species.
Ecol. Ent. 6:215-216.

Southwood, T. R. E. 1978. Ecological Methods. Second Edition. Halsted Press, N.Y.

Weseloh, R. M. 1985. Dispersal, survival, and population abundance of gypsy moth larvae determined by
releases and mark-recapture studies. Annals Ent. Soc. Am. 78:728-735.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 346

Publication Received

California Butterflies. By John S. Garth and J. W. Tilden. Illustrations by David
Mooney and Gene M. Christman. University of California Press, xvi + 246 pp., 33
text figs., 24 colored pis. (first 20 colored). ISBN 0-520-05249-8. Price $19.95 hard
cover. $10.95 paperback. Publication date given as November 13,1986. Review copy
received at CAS by PCES on October 27, 1986. California Natural History Guides:
51 (Arthur C. Smith, general editor). The book jacket states:

“With the help of this field guide, users can learn where to find butterflies and
skippers ranging in size from the tiny Pygmy Blue, with a wing span sometimes as
small as half an inch, to the Two-tailed Swallowtail which may have a wing span of
half a foot. The volume is filled with the fascinating lore of butterflies: their
migrations (some marked individual Monarchs have flown more than 2,000 miles),
their congregating on hilltops, their intriguing caterpillar and chrysalis stages.

This guide is small enough to be carried in the field, yet complete enough to enable
users to identify the more than 230 kinds of butterflies they may encounter anywhere
in California or nearby areas. This comprehensive coverage is achieved through
generous color or black-and-white illustrations of all but a very few species, together
with a text that provides compact descriptions followed by information on size,
habitat, distribution, abundance, and larval food plant preferences.

Detailed tips are provided for observing, rearing, collecting, and preserving
butterflies. An introductory section treats their intriguing anatomy and behavior. A
checklist of scientific and common names is included, along with a key to the eleven
families of California butterflies, plus a glossary of terms. A special section recounts
the history of butterfly collecting in California, and others deal with variation and
hybridization, classification, and name changes.

For all butterfly enthusiasts, this volume will provide the key to new enjoyment
and understanding.”

—Paul H. Arnaud, Jr., California Academy of Sciences, Golden Gate Park, San
Francisco, California 94118.

346

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 347-351

A Key to Trichrysis and New Species From Sri Lanka and Africa

(Hymenoptera: Chrysididae)

R. M. Bohart

Department of Entomology, University of California, Davis, 95616

Abstract. —A key is given to 20 species of the genus Trichrysis. New species
described are eardleyi from South Africa, Nigeria and Tanzania; hexapholis from Sri
Lanka; and lomholdti from South West Africa.

In connection with a generic revision of Chrysididae by Lynn Kimsey and me, 3
species of Trichrysis appear to be undescribed. Some 22 other species of the genus
occur in the Ethiopian, Palearctic and Oriental Regions. Except for a few, rarely
collected forms of other Old World genera, Trichrysis can be recognized by the
tridentate form of tergum III. Abbreviations used in the descriptions are: F-1 etc.,
flagellomeres; TEC, transverse frontal carina; MOD, median ocellus diameter; T-I
etc., terga; S-I etc., sterna.

The following key includes 20 species known to me. Others, not now available,
are coreana (Uchida), longispina (Mocsary),purpuripyga Edney, seducta Smith, and
jwJaz Tsuneki.

Key to Trichrysis (based on females)

1. Pronotum with a complete sublateral, longitudinal carina. 2

Pronotum without a complete sublateral, longitudinal carina, such carina
obsolete medially or absent. 11

2. F-I bright green in front, pronotal median groove well developed (Palearctic

Region). 3

F-I all dark or somewhat greenish toward base, pronotal median groove
various. 4

3. Hindbasitarsus bright green on outer side; T-III lateral tooth sharp; pit row

weakly indented. excisifrons (Mocsary)

Hindbasitarsus weakly colored; T-III lateral tooth obtuse, pit row strongly
indented. secernenda (Mocsary)

4. Pronotal median groove sharp and extending more than half of dorsal

surface, T-III lateral tooth merely an angle (Palearctic Region). 5

Pronotal groove weakly indicated at most; T-III lateral tooth sharp. 6

5. Moderate-sized (4-6 mm long). cyanea (Linnaeus)

Medium large (8-11 mm long). buyssoni (Mocsary)

6. F-I less than 2.5 X as long as broad, tarsi dark. 7

F-I more than 2.5 x as long as broad, tarsi various. 8

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PAN-PACIFIC ENTOMOLOGIST

7. T-III mid tooth strong, larger than lateral teeth; forewing discoidal cell well

pigmented throughout (N. Africa). scioensis (Gribodo).

T-III mid tooth sharp but smaller than lateral teeth; forewing discoidal cell
weakly pigmented distally (Philippines). aspera Brulle

8. Median bridge of pit row sharply depressed, leaving apical tooth hooklike

(view laterally) (Philippines, Taiwan). luzonica (Mocsary)

Median bridge of pit row at most slightly depressed.

9. T-III tooth intervals markedly convex (fig. 6) (Oriental Region).

. vestigator (Smith)

T-III tooth intervals at most faintly convex.

10. T-III mid tooth stronger than lateral teeth (fig. 5); highly colored green,

purple, gold, and red; tarsi yellowish (S. India). lanka (Bingham)

T-III mid tooth not stronger than lateral teeth, not highly colored, tarsi dark
(Oriental Region). triacantha (Mocsary)

11. Pronotum with a longitudinal sublateral carina which is obsolescent

medially.

Pronotum without a longitudinal sublateral carina.

12. TFC strong, straight, nearly reaching eyes (fig. 4); scapal basin partly

polished mediad (Ethiopian Region). eardleyi Bohart

TFC developed toward middle where it forms an inverse V (fig. 3); scapal
basin various.13

13. Scapal basin completely sculptured with punctures and microridging, tarsi

reddish yellow (Palearctic Region). lacerta (Semenov)

Scapal basin punctate but polished between punctures mediad, tarsi various

14. T-III tooth intervals unusually deep and evenly incurved (fig. 3); terga
highly colored, green, blue, purple, and gold; tarsi light red (Sri

Lanka).../icxop/io/w Bohart

T-III tooth intervals shallow; terga green and purple; tarsi dark (Oriental
Region). trigona Mocsary

15. Malar space 3 or more MOD (fig. 7), T-III middle tooth long and curved
downward toward tip (lateral view), S-II spots joined medially to form a
large shieldlike spot (fig. 7); dorsum of body microsculptured between

punctures (fig. 8).

Malar space 2 MOD or less, T-III middle tooth short; S-II spots rather small,
rounded, slightly separated; body dorsum without conspicuous
microsculpture between punctures.

16. Body dorsum mostly with coarse punctures, microsculpture imparting a
frosted look (fig. 8), middle half of scapal basin microsculptured (S.

Africa). impressifrons (Mocsary)

Body dorsum moderately to finely punctate, microsculpture quite fine,
middle half of scapal basin extensively polished (fig. 7) (S.W.
Africa)./om/zoWfi Bohart

17. T-III tooth intervals strongly convex (as in fig. 6).

T-III intervals evenly concave to almost straight (fig. 9).

18. Brow with a definable TFC as a downcurved crescent (Palearctic

Region). mendicalis (Cameron)

Brow with at most slight traces of a TFC among coarse punctures (Ethiopian
Region). heliophila (Mocsary)

9

10

12

15

14

16

17

18

19

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349

3. hexapholis ?

1. hexapholis 2. hexapholis

4. eardleyi

7. lomholdti ?

5. lanka

6. vestigator $

8. impressifrons

9. bohemanni +

Figure 1, pronotum, S-II. Figure 2, face. Figures 3,5,6,8,9, T-III apex. Figure 4, face, T-III apex, S-II.
Figure 7, face, pronotum, S-II. Small case letters, a, blue-green; b, golden; c, purple.

19. Forewing discoidal cell obsolescent toward wing apex; pit row not much
impressed and pits tiny (fig. 9) , T-III punctation unusually fine and close (fig.

9), F-I a little less than 2 x as long as broad (Ethiopian Region).

. bohemanni Dahlbom

Forewing discoidal cell complete; pit row well developed; T-III punctation

moderate; F-I nearly 3 x as long as broad (Ethiopian Region).

. polinierii Guerin

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PAN-PACIFIC ENTOMOLOGIST

Trichrysis eardleyi Bohart, new species

Male holotype: Length 5.5 mm. Green, grading to blue, and dark blue to black
toward base of T-II and III, F-I and following dark, tarsi off-white, wings weakly
stained, darkest in marginal cell. Vertex, notum and terga with moderate close
punctation; scapal basin finely punctate and silvery pubescent. F-I twice as long as
broad, 1.6 x as long as II; scapal basin concave, moderately deep; TFC strong,
nearly straight, almost reaching eyes (as in fig. 4); malar space and subantennal space
each about 1 MOD; pronotum with a weak dorsomedian groove, lateral margin
sharply incurved in dorsal view, and with a carina broken toward middle; discoidal
cell of forewing complete and with upper outer vein strong toward base; propodeal
projection sharp; T-III lateral margins sinuous, pit row deep, preceded by a low
transverse swelling, apical 3 teeth small but sharp, S-II spots small and oval (as in fig.

4 )-

Female: About as in male except: length 6.5 mm, tarsi light brown, face narrowly
polished medially (fig. 4), T-III somewhat saddled, apical teeth stronger, intervals
evenly concave (fig. 4).

Holotype male. South Africa: Pafuri, Kruger National Park, Transvaal, Jan. 1984
(C. D. Eardley, National Insect Collection, Pretoria). Paratype male, Nigeria:
Umuahia, IV-9-75 (J. T. Medler, U. C. Davis); paratype female, Tanzania: East
Usambera, Amani, II-2-77 (O. Lomholdt and O. Martin, Zoological Museum,
Copenhagen).

Discussion. In many respects eardleyi is similar to scioensis (Gribodo) but there
are important differences. TFC in scioensis is developed mediad as an inverted V (as
in fig. 2) rather than nearly straight and well developed all across, the sublateral
pronotal carina is complete, and intervals between T-III teeth are slightly convex
rather than evenly concave as in eardleyi (fig. 4).

Trichrysis hexapholis Bohart, new species

Female holotype: Length 5.5 mm. Green with bright purple and gold markings as
follows: purple are ocellar area, neck, transverse pronotal spots, broad median notal
stripe, rim above tegula, large semimedian spots on T-I-II-III, basal band on T-II-III;
gold are apicolateral spots on T-I, apical band enlarged sublaterally on T-II, prepit
band on T-III extended onto middle tooth; flagellum dark, wings faintly stained, tarsi
light reddish. F-I 2.5 x as long as broad, 1.5 x as long as II; scapal basin shallow,
coarsely punctate laterally, becoming mostly polished in median one-third; brow
prominent but short, topped by an inverted broad V-hke TFC in middle one-third
(fig. 2); malar space 1.6 MOD, subantennal space 1.2 MOD; pronotum with a weak
dorsomedian groove, lateral margin sharply incurved (fig. 1), and with a carina which
is broken toward middle; discoidal cell of forewing complete; propodeal projection
sharp; T-III lateral margin weakly convex, pit row with distinct pits preceded by a
low transverse swelling and moderate saddle; apical 3 teeth long and sharp, intervals
deeply concave (fig. 3); S-II spots joining to form a sharp median triangle (fig. 1).

Male: Unknown.

Holotype female, Sri Lanka: Colombo, IV-10-68 (T. F. Halstead, California
Academy of Sciences).

Discussion. The striking dorsal pattern is similar to that of other Sri Lankan
chrysidids, particularly Trichrysis lanka (Bingham). However, that species has a

VOLUME 63, NUMBER 4

351

complete sublateral pronotal carina; T-III stouter, the teeth shorter, and the
intervals nearly straight rather than deeply concave (compare figs. 3,5).

Trichrysis lomholdti Bohart, new species

Female holotype: Length 3.8 mm. Blue becoming purple toward base of T-II and
T-III; flagellum dark; wings water clear; tarsi dark. F-13.5 x as long as broad, 1.5 x
as long as II; scapal basin moderately concave, coarsely punctate in outer
one-quarter, polished in middle half; brow rounded, TFC faint, laterally recurved,
malar space 4 MOD (fig. ?); subantennal space 1.5 MOD; midocellar area outlined
by a fine carina; punctation of vertex and notum moderate, punctures of terga finer
and separated by weak microsculpture, those of T-III saddle 1-3 puncture diameters
apart; pronotum without a dorsomedian groove, weakly incurved laterally (fig. 7)
and ecarinate; discoidal cell veins of forewing weak but pigmented; propodeal
projection short and acute; T-III lateral margin slightly convex, pit row consisting of
14 moderately impressed pits preceded by a low transverse swelling and moderate
saddle; apical 3 teeth sharp, lateral ones receding, intervals evenly but shallowly
concave; S-II spots joining to form a large, shieldlike mark (fig. 7).

Male. Unknown.

Holotype female, S. W. Africa: 110 km e. Windhoek, X-25-72, (C. L. Hogue,
National Insect Collection, Pretoria).

Discussion. The only other species of Trichrysis with a long malar space is
impressifrons (Mocsary), also from the Ethiopian Region. However, that species has
the scapal basin microridged mediad, punctures of vertex and notum more coarse
and with obvious intervening micropunctation, T-III strongly convex on the lateral
margin, prepit swelling rather sharp, pit row deeply impressed, and postpit area
longer (fig. 8). This species is named for Ole Lomholdt of the Zoological Museum in
Copenhagen for his special interest in African Hymenoptera and his overall
cooperation in our chrysidid study.

Scientific Note

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 352

Parasitism of Salticid Spiders in Oregon by Two Species of Acroceridae (Diptera)

Arboreal jumping spiders (Salticidae) were collected during June 1985 from
Union County, Oregon, by beating lower branches of Douglas-fir, Pseudotsuga
menziesii var. glauca (Beissn.) Franco, and grand fir, Abies grandis (Dough ex D.
Don) Lindh, over dropcloths (Paul and Mason 1984, USDA For. Serv. Res. Note
PNW-421:1). About 165 specimens of Metaphidippus spp. were collected and
individually reared in covered petri dishes; the spiders were fed Drosophila fruit flies
until needed in field experiments on the western spruce budworm, Choristoneura
occidentalis Freeman. Nine of the spiders (about 5%) produced dipterous
parasitoids belonging to the family Acroceridae. The host spiders spun thick
protective webbing in dishes before emergence of parasitoid larvae; larval
emergence was similar to that described by Schlinger (1952, Pan-Pac. EntomoL,
28:7). The adult parasitoids were later identified by Dr. Evert Schlinger, University
of California, Berkeley, as eight specimens of Acrocera bulla Westwood and one
specimen of Ogeodes boharti Schlinger. The spiders were identified by Wayne
Maddison, Museum of Comparative Zoology, Harvard University, Cambridge,
MA, as M. aeneolus (Curtis) (or probably M. aeneolus). The immature spiders were
difficult to identify, especially because they were damaged by parasitoid emergence.
This collection extends the known range of O. boharti into Oregon; it has previously
been recovered from Arizona and New Mexico (Schlinger, personal
communication). Related species of Ogcodes have been reared from numerous
spider-host species (Eason et al. 1967, J. Kansas, Entomol. Soc., 40:422).

R. C. Beckwith,^ R. R. Mason, and H. G. Paul, Pacific Northwest Research
Station, LaGrande, OR 97850.

Tresent address: Forestry Sciences Laboratory, 3200 Jefferson Way, Corvallis, OR 97331.

352

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 353-357

Electrophoretic Comparison of European Dendroctonus micans and
Ten North American Dendroctonus Species (Coleoptera: Scolytidae)

Molly W. Stock, Jean-Claude Gregoire and Malcolm M. Furniss

Department of Forest Resources, University of Idaho, Moscow, Idaho 83843;
Laboratoire de Biologic Animale et Cellulaire, Universite Libre de Bruxelles, 1050
Brussels, Belgium; Department of Plant, Soil, and Entomological Sciences,
University of Idaho, Moscow, Idaho 83843

Abstract. —Electrophoretic techniques were used to estimate genetic relationships
among the great European spruce bark beetle, Dendroctonus micans (Kugelann),
and 10 North American Dendroctonus species. Average heterozygosity for D.
micans was .053; the North American species ranged from .114 to .226. Cluster
analysis suggests that D. micans is more closely related to D. terebrans and D. valens
than to other species in the genus.

The great European spruce bark beetle, Dendroctonus micans (Kugelann), occurs
in spruce forests of Eurasia from the United Kingdom to Siberia. It is believed to
have originated in Siberia from a North American spruce-feeding ancestor (Wood
1963) and subsequently spread across Europe. It was discovered in England in 1982
(Bevan and King 1984, Evans et al. 1984).

Relationships among adult Dendroctonus species have been studied by Hopkins
(1909), Wood (1963, 1982), and Lanier (1981). Immatures were compared by
Thomas (1965). Except for its greater size, D. micans appears virtually identical to
the North American D. punctatus LeConte, and may be conspecific (Bright 1976).
D. micans (Bevan and King 1984), and probably D. punctatus, are similar to D.
valens LeConte, D. terebrans (Olivier), and D. rhizophagus Thomas and Bright in
that adults construct egg galleries in the lower bole and large roots and the larvae
mine communally rather than make discrete individual tunnels. However,
morphologically (Hopkins, 1909, Wood 1982) and cytogenetically (Lanier 1981) D.
micans and D. punctatus have been allied with D. rufipennis Kirby and D.
murrayanae LeConte. Some aspects of the biology and behavior of D. micans are
unusual. For example, successful attacks by solitary females are common and
inbreeding is the rule. Vouland et al. (1984) report that usually more than 90% of the
females are fertilized before they emerge.

Electrophoretic studies of 10 representative North American Dendroctonus
species revealed species clusters very similar to those developed using cytogenetic
and anatomic evidence (Bentz and Stock 1986). D. micans and D. punctatus,
however, were not available for inclusion in those studies. Preliminary
electrophoretic comparisons of D. micans from France and Great Britain have
revealed consistent differences within this species (Evans et al. 1984). More recently.

353

354

PAN-PACIFIC ENTOMOLOGIST

acquisition of live D. micans from Belgium permitted electrophoretic comparison
with the 10 Dendroctonus species studied earlier. Results of this comparison are
reported here.

Methods

Larvae were grown in the laboratory in Belgium from females introduced into
fresh Picea excelsa (P. abies) logs. Fifth instar larvae were shipped from Belgium and
received at the University of Idaho, on May 25,1985, in a layer of fresh phloem taped
tightly between two 15 x 15 cm pieces of plate glass. Pupae were noted beginning
June 4 and all had transformed to adults by June 25. On June 27 and July 5, two lots of
50 females each were frozen at —30° C.

Electrophoretic analysis of these beetles followed methods described by Higby
and Stock (1982) and Bentz and Stock (1986) for other Dendroctonus species.
Genetic diversity was estimated and compared among groups using percent
polymorphism and Nei’s (1975) average heterozygosity. A locus was considered
polymorphic when the frequency of the common aUele was less than or equal to .99.
Relationships between D. micans and other Dendroctonus species were assessed,
using BIOSYS-1 (Swofford and Selander 1981), by hierarchical cluster analysis of
Nei’s (1978) genetic distance values.

Results and Discussion

Data from 15 gene loci were obtained (Table 1). Of these, four (AAT, IDH-1,
ME, and PEP-gl) were polymorphic, and 11 (CK, EST-1, EST-2, EST-4, IDH-2,
MDH-1, MDH-2, MPI, PEP-la, PGI, and SOD) were monomorphic. These beetles
were thus much less genetically diverse (27% polymorphism) than any other
Dendroctonus species that have been studied electrbphoretically: polymorphism
over the same 15 loci in 10 other species ranged from 40% to 67%. Average
heterozygosity for the species was .053, compared to a range of .114 to .226 in the
other 10 Dendroctonus species at these 15 loci. Cluster analysis suggests that D.
micans is more closely related to D. terebrans and D. valens than to other species in
the genus (Figure 1).

Specimens of D. micans used in this study were from broods of only two females
but may still represent the species’ diversity. Because the species is isolated from
others of the genus and inhabits a unique niche in European forests, with little or no

Table 1. Allele frequencies at 15 gene loci^ in Dendroctonus micans and D. ponderosae. Data for D. ponderosae are
taken from Bentz and Stock (1986). Percent polymorphism (P) and average heterozygosity (H) are also given.

Enzyme

Locus

D. micans

D. ponderosae

AAT

A

.606

—

B

.387

.662

C

.007

.338

(N)

(91)

(334)

CK

B

1.0

.989

C

—

.011

(N)

(60)

(323)

VOLUME 63, NUMBER 4

355

Table 1. continued

Enzyme

Locus

D. micans

D. ponderosae

EST-1

A

1.0

1.0

(N)

(64)

(421)

EST-2

A

—

.019

B

—

.277

C

—

.278

D

—

.137

E

—

.234

F

—

.055

G

.991

—

H

.009

—

(N)

(73)

(399)

EST-4

A

—

.993

B

—

.007

C

1.0

—

(N)

(54)

(291)

IDH-1

B

.995

—

C

.045

1.0

(N)

(44)

(361)

IDH-2

A

—

1.0

B

1.0

—

(N)

(64)

(297)

MDH-1

B

1.0

1.0

(N)

(58)

(377)

MDH-2

A

o

—

1.0

D

C

1.0

_

(N)

(58)

(348)

ME

A

.905

1.0

B

.095

—

(N)

(67)

(478)

MPI

B

1.0

1.0

(N)

(56)

(300)

PEP-gl

A

.989

—

B

.011

.932

C

—

.068

(N)

(44)

(420)

PEP-la

A

1.0

.017

B

—

.832

C

—

.151

(N)

(58)

(235)

PGI

C

—

.009

D

.009

.978

E

.991

.014

(N)

(74)

(404)

SOD

B

1.0

1.0

(N)

(53)

(421)

P

27

40

H

.053

.114

^AAT = aspartate aminotransferase, CK = creatine kinase, EST = esterase (three loci), IDH =
isocitrate dehydrogenase (two loci), MDH = malate dehydrogenase (two loci), ME = malic enzyme,
MPI mannose phosphate isomerase, PEP-gl = peptidase glycyl-leucine, PEP-la = peptidase
leucyl-alanine, PGI = phosphoglucose isomerase, SOD = superoxide dismutase (sometimes called TO
or tetrazolium oxidase in other reports).

356

PAN-PACIFIC ENTOMOLOGIST

1.2 .9 .6 .3 0

GENETIC DISTANCE

Figure 1. Phenogram derived from electrophoretic data illustrating relationships among 11
Dendroctonus speeies. (PON = ponderosae, ADJ = adjunctus, APX = approximatus,
RUF = rufipennis, BRV = brevicomis, SIM = simplex, PSD = pseudotsugae, FRN = frontalis,
VAL = valens, TER = terebrans, MIC = micans)

competition from other insect species, and because the species is highly inbred, it is
possible that the low level of genetic diversity observed here may be typical.

Based on the hosts and earlier anatomic and cytogenetic studies, we expected that
our electrophoretic data on D. micans would correspond most closely to D.
rufipennis. We observed, however, a closer relationship to the pine-infesting D.
terebrans and D. valens with which D. micans shares gregarious larval feeding in the
phloem of living hosts.

In order to further clarify D. micans’ genetic relationship within the genus, we
hope to obtain additional broods of D. micans, as well as samples of D. punctatus and
D. murrayanae for electrophoretic comparison. Such work has been hindered to date
by the relative scarcity of D. punctatus and the difficulty of obtaining live specimens
of D. micans from distant locations.

Acknov/ledgments

We thank Christine Kelly, Morgan Stage, and Sandra Cast for technical help with
this study, and Dr. Gerald N. Lanier and Dr. Stephen L. Wood for reviewing the
manuscript.

Literature Cited

Bentz, B. J., and M. W. Stock. 1986. Genetic relationships among ten species of Dendroctonus bark
beetles (Coleoptera: Scolytidae). Ann. Entomol. Soc. Amer. 79:527-534.

Bevan, D., and C. J. King. 1984. Dendroctonus micans Kug.—A New Pest of Spruce in U.K.
Commonwealth Forestry Review, 11 pp.

Bright, D. E. 1976. The Bark Beetles of Canada and Alaska (Coleoptera: Scolytidae). Can. Dept. Ag.
Pub. no. 1576. 241 pp.

VOLUME 63, NUMBER 4

357

Evans, H. F., C. J. King, and D. Wainhouse. 1984. Dendroctonus micans in the United Kingdom. Pp.
20-34 in: Proc. EEC Seminar on Biological Control of Bark Beetles (Dendroctonus micans).
Universite Libre de Bruxelles, Brussels, Belgium, 3-4 October 1984.

Higby, P. K., and M. W. Stock. 1982. Genetic relationships between two sibling species of bark beetle
(Coleoptera: Scolytidae), Jeffrey pine beetle and mountain pine beetle, in northern California.
Ann. Entomol. Soc. Amer. 75:668-674.

Lanier, G. L. 1981. Cytotaxonomy of Dendroctonus. Pp. 33-66 in Application of Genetics and Cytology
in Insect Systematics and Evolution, M. W. Stock (ed.), Forest, Wildlife, and Range Experiment
Station publication, Univ. Idaho, Moscow.

Nei, M. 1975. Genetic variability in natural populations. Pp. 127-174 in Molecular Population Genetics
and Evolution. North Holland Research Monographs, Frontiers of Biology, vol. 40. American
Elsevier, New York.

Nei, M. 1978. Estimation of average heterozygosity and genetic distance from a small number of
individuals. Genetics 89:583-590.

Swofford, D. L., and R. B. Selander. 1981. BIOSYS-1, A Computer Program for the Analysis of Allelic
Variation in Genetics. User’s Manual. Dept. Genetics and Development, Univ. Illinois,
Urbana-Champaign. 65 pp.

Thomas, J. B. 1965. The immature stages of Scolytidae: the genus Dendroctonus Erichson. Can.
Entomol. (7:374-400.

Vouland, G., M. Giraud, and D. Schvester. 1984. La periode tenerale et I’envol chez Dendroctonus
micans Kug. (Coleoptera, Scolytidae). Pp. 68-79 in Proc. EEC Seminar on Biological Control of
Bark Beetles (Dendroctonus micans). Universite Libre de Bruxelles, Brussels, Belgium, 3-4
October 1984.

Wood, S. L. 1963. A revision of the bark beetle genus Dendroctonus Erichson (Coleoptera: Scolytidae).
Great Basin Nat. 23:1-117.

Wood, S. L. 1982. The Bark and Ambrosia Beetles of North and Central America (Coleoptera:
Scolytidae): A Taxonomic Monograph. Great Basin Nat. Mem. no. 6. 1359 pp.

Scientific Note

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 358

First Records of the German Yellowjacket Paravespula germanica (L.)
from the East San Francisco Bay (California, U.S. A.) Area

The German yellowjacket, Paravespula germanica (L.), is native to Europe,
northern Africa, and western Asia (Spradbery, 1973, Wasps, University of
Washington Press). Its geographical range has increased largely due to the activities
of man, and it is now established in Australia, New Zealand, South Africa, North
America, and South America (MacDonald etal., 1980, Bull. Entomol. Soc. Amer.,
26(4):436-442). The North American population was introduced on the east coast
and gradually spread west. The first California record was from South Lake Tahoe in
1983 (anonymous, 1985, Calif. Dept. Health Services Vector Update, 2-85:1-2).
The present report extends the confirmed range to coastal California.

On 1 August 1986, in response to a call from a homeowner, I discovered an active
colony of P. germanica in a house in Berkeley, California (elevation = 152 m). The
exterior entrance hole led to a hollow space above a false wooden ceiling, but the
location of the nest within the structure was not determined. This choice of a
man-made structure as a nesting site is common in the “North American strain” of P.
germanica (MacDonald et ai, 1980). Workers made approximately 80 sorties per
minute, a rate indicative of a thriving colony. No reproductives were observed.

In addition to workers observed in the immediate vicinity of the colony, individual
P. germanica foragers were captured at other East Bay locations. On 3 September a
worker was captured at a garbage can in Berkeley (elevation = 80 m),
approximately 1.3 km south of the colony, a distance within the outer limits of the
expected foraging range of P. germanica workers. This wasp may have come from
the aforementioned colony.

Foraging workers were also captured on 8 October on vegetation in Albany
(elevation = 4 m) and on 10 October at a garbage can in Berkeley (elevation = 60
m). The sites are separated by 3.4 km, so it is not likely that these workers came from
a single colony. Since the known colony had been eradicated in late September, the
evidence suggests that at least three colonies of P. germanica were established in
1986, and that an invasion of the San Francisco Bay area is underway.

Because of the tendency to nest in buildings, P. germanica presents a greater safety
hazard than native Paravespula (Bluthgen) species, which more commonly nest in
the ground. The mild-wintered climate of coastal California may also enable P.
germanica colonies to overwinter and requeen, as occasionally occurs locally in both
P. vulgaris (L.) and P. pensylvanica (Saussure). Overwintered P. germanica colonies
are known from Australia (Spradbery, 1973) and New Zealand (Thomas, 1960, N.
Zealand Dept. Scientific and Industrial Res. Information Ser., 27:1-4), but not
North Ameriea.

Parker Gambino, Department of Entomological Sciences, University of California,
Berkeley, California, U.S.A.

358

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp.359-362

A New Genus and Two New Species of Longhorn Beetles
(Coleoptera: Cerambycidae) From Mexico and Central America

Edmund F. Giesbert

9780 Drake Lane, Beverly Hills, CA 90210

Abstract .—A new neotropical elaphidiine genus, Tropimerus, is proposed and
characterized. Two new species are described in the genus: T. cyaneus from Chiapas,
Mexico, which is figured, and T. hovorei from El Salvador and Costa Rica.

Tropimerus Giesbert, gen. nov.

Form small, elongate. Head moderately small; eyes finely facetted; palpi
subequal, terminal segments broadly triangular; frons with transverse impression
with pit at each end giving rise to a long, erect seta; antennal tubercles feebly
elevated; antennae slender, finely punctate and pubescent, with scattered flying
hairs, length subequal to body in male, shorter in female, third segment carinate,
segments 3 to 6 with short apical spine or tooth on inner side, third segment longest,
4th segment shortest, 11th longer than 10th. Pronotum nearly glabrous,
subcylindrical or with sides slightly rounded, unarmed. Scutellum small, rounded at
apex, finely pubescent. Elytra elongate, parallel sided, narrowed at posterior third,
apices narrowly rounded to subacuminate. Underside with anterior coxal cavities
rounded externally, open behind, prosternal process arcuate; mesosternum
gradually sloping in front, apex notched behind, intermediate coxal cavities closed
externally; abdomen of male with six sternites visible, female with five. Legs
moderately short, slender; femora subclavate, apices unarmed, mesofemora with
ventral and dorsal carinae; tibiae carinate; metatarsi with first segment slightly
shorter than 2nd and 3rd combined.

Type Species.—Tropimerus cyaneus Giesbert, sp. nov.

Remarks.—Tropimerus bears a superficial resemblance to Stenosphenus
Haldeman, and Psyrassa Pascoe, but differs from both by the presence of a sixth
sclerotized abdominal sternite in the males, the distinctly carinate mesofemora, and
the attenuate form of the elytral apices.

The remarkable sexual dimorphism in the abdominal segmentation presents a
taxonomic character which may prove to be significant at a higher category of
classification. Tropimerus shares this character with members of the Callichromatini,
but it would be otherwise quite anomalous in that tribe. For the present, rather than
establishing a new tribe for this genus alone, it seems more suitable to place
Tropimerus in the Elaphidiini near Stenosphenus and Psyrassa.

359

360

PAN-PACIFIC ENTOMOLOGIST

Key to the species of Tropimerus Giesbert:

1. Head and pronotum black; elytra black with blue reflections. Antennae with 3rd

segment I 1/2 times as long as scape or 4th segment. Chiapas, Mexico.

T. cyaneus sp. nov.

— Head and pronotum orange; elytra black with feeble metallic reflections.
Antennae with 3rd segment twice as long as scape or 4th segment. El Salvador,
Costa Rica. T. hovorei sp. nov.

Tropimerus cyaneus Giesbert, new species

(Fig- 1)

Male. —Form moderately small, slender, subcylindrical. Integument piceous
black, blue metallic reflections on elytra and ventral surface; head, pronotum, and
appendages feebly aeneous; palpi, mandibles, and tarsal claws paler. Head shining,
moderately rugulose-punctate, sparsely pubescent; vertex longitudinally impressed;
occiput transversely rugulose; antennae slender, nearly as long as body, scape short,
3rd segment I 1/2 times as long as scape or 4th segment, remaining segments slightly
longer than 4th with 11th segment somewhat longer than 10th, 3rd segment carinate,
segments 3 to 5 with short apical spine, 6th segment apically dentate. Pronotum
slightly longer than broad, sides feebly rounded, transversely impressed behind
apex, with impression interrupted at middle, disk shallowly indented on each side at
middle, shallowly impressed before base; surface shining, with base and apex
micro-rugulose, disk very sparsely, finely punctate, with erect, pale hairs sparse.
Scutellum rounded at apex, clothed with fine, recumbent, pale pubescence. Elytra
about 31/2 times as long as width across humeri, parallel sided, narrowing at apical 1/3
to acuminate apices; surface shining, moderately coarsely, densely punctate, with
pubescence sparse, fine, and erect. Underside with a few coarse deep punctures on
each side of prosternum in front of procoxae; abdomen with 5th sternite widely
emarginate, 6th sternite rounded at apex; prosternum, episternal pieces, and sides of
metasternum and abdominal sternites moderately densely clothed with silky,
recumbent, pale golden pubescence; remainder of ventral surface shining, finely,
sparsely punctate, with scattered, erect, pale hairs. Legs moderately slender,
shining, with femora subclavate, mesofemora carinate; tibiae carinate. Length 7-7.5
mm.

Female. —Form similar to male. Antennae reaching about apical 1/4 of elytra;
prosternum lacking coarse punctures; abdomen with 5th sternite widely rounded at
apex, no 6th sternite evident. Length 7-8.5 mm.

Holotype male, allotype (California Academy of Sciences), and 16 paratypes (10
male, 6 female) from MEXICO, Chiapas, Sumidero Canyon, 4000 ft., on blossoms
of Croton sp., July 7-8, 1986 (E. Giesbert, J. Wappes). Additional paratypes
include; 17 males, 8 females, same locality, June 14-24,1987 (E. Giesbert); 3 males,
same locality, June 14-20,1987 (J. Wappes).

Tropimerus hovorei Giesbert, sp. nov.

Male. —Form moderately small, slender, subcylindrical. Integument piceous
black with metallic reflections on elytra; head and pro thorax orange. Head s hinin g,
moderately rugose-punctate, sparsely pubescent; vertex longitudinally impressed;
occiput transversely rugulose; antennae slender, nearly as long as body, scape short.

VOLUME 63, NUMBER 4

361

Figure 1. Tropimerus cyaneus Giesbert, new species; male.

362

PAN-PACIFIC ENTOMOLOGIST

3rd segment twice as long as scape or 4th segment, remaining segments longer than
4th, with 11th segment slightly longer than 10th, 3rd segment feebly carinate,
segments 3 and 4 with short apical spine, segments 5 and 6 apically dentate.
Pronotum longer than broad, sides somewhat rounded, transversely impressed at
sides behind apex, shallowly impressed near base; surface shining, base and apex
micro-rugulose, disk finely, very sparsely punctate, with erect, pale hairs sparse.
Scutellum rounded at apex, clothed with fine, recumbent, pale pubescence. Elytra
about 31/2 times as long as width across humeri, parallel sided, roundly tapering at
apical 1/3 to narrowly rounded apices; surface shining, moderately finely and densely
punctate, with somewhat larger punctures scattered on disk and sublineate along
suture; pubescence sparse, fine, erect. Underside with prosternum lacking coarse
punctures; abdomen with 5th sternite widely emarginate, 6th sternite narrowly
rounded at apex; disk of prosternum, episternal pieces, and margins of metasternum
and abdominal sternites moderately densely clothed with pale, silky, recumbent
pubescence. Legs moderately slender, shining, with femora subclavate, mesofemora
and tibiae carinate. Length 7.25-8.5 mm.

Female. —Form similar to male. Antennae slightly shorter, attaining apical 1/4 of
elytra; abdomen with 5th sternite widely rounded at apex, no 6th sternite evident.
Length 7.5-10 mm.

Holotype male, allotype (California Academy of Sciences), and 8 paratypes from
EL SALVADOR, Quetzaltepeque, 500m., June 19, July 5,1963 (D. Q. Cavagnaro,
M. E. Erwin). 54 additional paratypes (29 males, 25 females) from COSTA RICA,
Puntarenas prov., 6 km S. Santa Elena, on blossoms of Croton sp.. May 23,1985 (F.
Hovore), May 24-28,1987 (E. Giesbert), June 7-8,1987 (F. Hovore).

Remarks.—Tropimerus hovorei differs from T. cyaneus by the orange head and
pronotum, relatively longer 3rd antennal segment, the finer elytral punctures, as well
as by the more southeastern range.

Acknowledgments

I thank my colleagues F. Hovore and J. Wappes for providing specimens and data
from their collections, and for comradeship in the field. I would also like to thank J.
Chemsak for the loan of specimens, and helpful criticism of the manuscript.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 363-367

Lectotype Designations and Redescription of
Vejovis wupatkiensis Stahnke (Scorpiones: Vaejovidae)

Stanley C. Williams

Department of Biology, San Francisco State University, San Francisco, CA 94132

Abstract. —The syntypes of Vejovis wupatkiensis Stahnke were studied. A
lectotype, allolectotype and 6 paralectotypes are designated, and the species is
redescribed based on the lectotype. It is concluded that the nearest relatives to V.
wupatkiensis Stahnke are Serradigitus gertschi (Williams) and Serradigitus torridus
Williams and Berke. The current placement of V. wupatkiensis within the genus
Serradigitus (i.e., S. wupatkiensis (Stahnke)) is confirmed.

Introduction

In 1940, H. L. Stahnke published an abstract of his 1939 doctoral dissertation on
the scorpions of Arizona. In it, his characterization of the new species encountered
were sufficient to validate and make available 9 new species-group names and one
genus-group name. Stahnke named one of these new species Vejovis wupatkiensis
after its type locality at Wupatki National Monument. This species was distinctive
because of its elongate pedipalp fingers, elongate terminal pedipalp denticle, and
swollen basal teeth of the female pectines. Subsequently, several related new species
have been found and described. The phylogenetic relations among these taxa has
remained obscure until recently because the types of V. wupatkiensis were not
available for study. The purpose of this paper is to designate a lectotype, an
allolectotype, and 6 paralectotypes for V. wupatkiensis, to redescribe the species
based on the lectotype, and to confirm its current placement within the genus
Serradigitus Stahnke. The measurements given are as described by Williams
(1980:2-3).

Serradigitus wupatkiensis (Stahnke)

(Figure 1, Table 1)

Vejovis wupatkiensis Stahnke 1940:105. Gertsch and Allred 1965:8. Sole glad
1972:180-181.

Vaejovis wupatkiensis Stahnke. Johnson and Allred 1972:168-169. Williams 1976:2.
Serradigitus wupatkiensis (Stahnke). Stahnke 1974:130-132. Williams and Berke
1986: (in press).

Diagnosis. —Small, slender species of Serradigitus, adults to 35 mm long. Body
uniform whitish yellow to amber-brown. Frontal margin of carapace straight to
slightly emarginate; chela of pedipalps long and slender, movable finger longer than
carapace, ratio of chela length to palm length 5.6 in males, 5.0 in females. Brachium

363

364

PAN-PACIFIC ENTOMOLOGIST

width approximates palm width; chela with supernumerary denticles 6/7 on fixed and
movable fingers respectively; primary row denticles subdivided into 2-3 subrows of
sharp serrate denticles by slightly enlarged denticles, chelicerae lacking denticles on
ventral margin of fixed finger; stigma short, oval, 2-3 times longer than wide, ratio of
carapace length to stigma length 14-15. Telson with small subaculear tubercule.

Similar to Serradigitus gertschi (Williams) and Serradigitus torridus Williams and
Berke. Distinguished from S. gertschi by smooth to obsolete inferior median keels of
metasomal segment I (not crenulate); differs from S. torridus by presence of 4
prolateral macrosetae on pedipalp humerus (not 5).

Redescription based on lectotype. —Female. Coloration: Carapace, mesosoma,
metasoma, pedipalps, and walking legs uniform golden brown; pedipalp fingers
similar to palm in color; pectines and genital operculum slightly more whitish, no
other contrasting markings. Prosoma: Carapace frontal margin slightly concave,
with 3 pairs macrosetae; lateral ocelli 3 per group, median ocelli small, similar to
most cephalad lateral ocelli in size, interocular space wider than median ocellus
diameter; dorsal surface finely granular; sternum broadly pentagonal, wider than
long, 3 pairs sternal setae. Mesosoma: Terga 2-6 with subtle obsolescent median
keel, tergum 7 with short, obsolescent median keel and 2 pairs well developed serrate

VOLUME 63, NUMBER 4

365

Table 1. Measurements (mm) of Vejovis wupatkiensis. Abbreviatins are as follows: 1 = length, w = width,
d = depth, fmd = frontal margin distance, ditd = distal internal trichobothrium distance, p-row = primary denticle
row of chela.

lectotype #67
(female)

lectoallotype (#75.3)

(male)

Total length

33

25

Carapace, (1/w at median eyes)

4.3/3.2

3.1/2.4

Diad (width/fmd)

0.6/1.4

0.5/1.2

Metasoma, length

15.2

11.4

Segment I (1/w/d)

2.1/2.5/2.0

1.5/1.8/1.4

Segment II (1/w/d)

2.4/2.3/2.0

1.8/1.8/1.4

Segment III (1/w/d)

2.6/2.3/2.0

2.0/1.7/1.5

Segment IV (1/w/d)

3.5/2.3/2.0

2.6/1.7/1.4

Segment V (1/w/d)

4.6/2.2/2.0

3.5/1.6/1.4

Telson, length

5.7

3.0

Vesicle (1/w/d)

4.1/1.6/1.2

1.8/1.2/0.8

Aculeus (1)

1.6

1.2

Pedipalp, Humerus (1/w)

4.3/1.2

2.9/0.8

Brachium (1/w)

4.5/1.4

3.1/0.9

Chela (1)

7.8

5.0

Palm (1/w/d)

3.2/1.4/1.4

2.1/1.0/1.1

Movable finger (1/base)

5.3/0.8

3.3/0.5

Fixed finger (1/ditd)

4.6/2.6

2.6/1.8

Fixed finger p-row denticles

47:12,35

43:12,31

Movable finger p-row denticles

43:8,7,28

43:9,7,27

Pectine teeth (left/right)

—/16

17/17

Stigma 3 (Fw)

0.3/0.1

0.2/0.1

lateral keels; genital operculum with 3 pairs setae; pectine with anterior lamella of 4
elongate sclerites, middle lamella of long quadrilateral basal piece plus 9 subcircular
sclerites, fulcra triangular, 15 teeth per comb (including missing distal tooth), 2
proximal teeth more swollen, more elongate than others; stigmata small, oval, 3
times longer than wide; sternum 7 with 1 pair lightly developed, smooth to crenular,
lateral keels. Metasoma: Dorsal and dorsolateral keels well developed, crenulate,
terminating in elongate denticle on each posterior terminus; dorsal keels I-IV with
17-17-18-26 crenulations; ventrolateral keels with 12-17-18-26 crenulations on
segments I-IV respectively; ventral keels obsolete on I, smooth to obsolete with a
few obsolescent crenulations on II, lowly crenulate on III, crenulate on IV;
metasomal segments short, stout, segment IV 1.5 times longer than wide. Telson:
Aculeus one-third length of telson, distinct subaculear tubercule flanked with pair
macrosetae, similar to aculeus in length, vesicle with about 6 pairs macrosetae
approaching aculeus in length. Pedipalps: Chela long, slender, fixed finger longer
than carapace, supernumerary denticles 6/7 on fixed and movable fingers
respectively; primary row denticles in single, continuous, linear row, these sharp,
serrate—numbering 47/43 denticles on fixed and movable fingers respectively,
primary row denticles subtly subdivided into 2/3 subrows on fixed and movable
fingers respectively by slightly enlarged denticles; fingers each with conspicuously
elongate terminal denticle, these greater than 5 times length of first supernumerary;

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PAN-PACIFIC ENTOMOLOGIST

fixed finger with trichobothrium di dorsal to supernumerary 5, trichobothrium pi
dorsally between supernumeraries 5 and 6; palm with 1 dorsal, 2 dorsal prolateral
keels, these lowly developed, crenulate; dorsal retrolateral keels smooth to obsolete.
Chelicera: Movable finger ventral margin smooth, with setal comb; dorsal margin
with broad basal bicuspid, 2 subterminal denticles and short terminal denticle. Fixed
finger with basal bicuspid, 1 subterminal and 1 terminal denticle, no ventral margin
denticles.

Variation .—Allolectotype and 6 paralectotypes examined. These similar in
structure and coloration to lectotype but varied as follows: six females varied in total
length from 29-32 mm, single male 25 mm; pectine teeth 15-16 (mode = 16) in
females, 17 in male; females with proximal two pectine teeth swollen (one apparent
subadult with proximal first tooth swollen); male pectine teeth all similar in size;
fixed finger tends to be slightly shorter than carapace length (lectotype with fixed
finger slightly longer than carapace length).

Types .—Lectotype (designated here), female, Arizona: Coconino Co., Wupatki
National Monument, November 10, 1938, D. J. Jones (“#67, Wupatki Nat. Mnt.,
11-10-38, D. J. Jones, Stahnke Collection, Type”) [missing left movable pedipalp
finger, right chela detached, right tarsi missing on leg 4, many setae missing].
Depository: California Academy of Sciences, Entomology Type Number 15173.

Lectoallotype, male, “#75.3 Wupatki Natl. Mnt., Before 8-38, D.J. Jones,
Vejovis wupatkiensis Stahnke, Type.” Depository: California Academy of Sciences,
Entomology Type Number 15173.

Paralectotypes as follows: “#72 Wupatkia, Flagstaff, Arizona, 11-7-38,
syntype-cotype” [female]; “#73.4 Wupatkia, Flagstaff, Ariz, Oct 1938, cotype”
[female]; “#73.3, Wupatkia, Flagstaff, Arizona, Oct 1938, syntype-cotype” [small
subadult female]; “#75.11, Wupatki, Flagstaff, Arizona, 8-10-38, syntype female,”
[female]; “#326, Wupatki, Flagstaff, Arizona, 6-11-39” [female, not labeled as type,
but apparently part of syntype series]; “#325, Wupatki, Flagstaff, Arizona, 6-9-39”
[female, probably subadult]. Paralectotype Depository: California Academy of
Sciences.

Remarks .—According to his doctoral dissertation (1939), Stahnke based the
description of this species on 9 specimens. Of these, only eight were received by the
California Academy of Sciences [from H. L. Stahnke] in 1983. The missing specimen
was numbered 75.2 in the dissertation, and its location is unknown.

This species is known from northern Arizona, southern Utah, southern Nevada,
and adjacent eastern California. Little geographic variation is apparent. However, I
recognize a distinctive race from the Panamint Mountains of eastern California. It is
characterized by having pedipalp fingers more darkly pigmented than palm, pedipalp
keels outlined with contrastingly dark pigmentation, and slightly more slender
pedipalp palms.

Acknowledgments

Thanks to Wojciech J. Pulawski and Vincent F. Lee, California Academy of
Sciences, for arranging the loan of the types, to Warren E. Savary for assistance with
illustrations, and to Paul H. Arnaud, Jr., Alec M. Balmy, Vincent F. Lee, and
Warren E. Savary for reading and criticizing this manuscript.

VOLUME 63, NUMBER 4

367

Literature Cited

Gertsch, W. J. and D. M. Allred. 1965. Scorpions of the Nevada Test Site. Brigham Young Univ. Sci.
Bull. 6(4): 1-15.

Johnson, J. D. and D. M. Allred. 1972. Scorpions of Utah. Great Basin Nat. 32(3):154-170.

Soleglad, M. E. 1972. Two new scorpions of the wupatkiensis group of the genus Vejovis (Scorpionida;
Vejovidae). Wasmann J. Biol., 30(1-2):179-195.

Stahnke, H. L. 1939. The scorpions of Arizona. Iowa State College, Ames, (unpublished Ph.D.
Thesis): 185 pp.

-. 1940. The scorpions of Arizona. Iowa State College, J. Sci. 15(1):101-103. (Abstr.)

-. 1974. Revision and keys to the higher categories of Vejvidae (Scorpionida). J. Arachnol. (1973)

1(2):107-141.

Williams, S. C. 1976. The scorpion fauna of California. Bull. Soc. Vector Ecol. 3:1-4.

-. 1980. Scorpions of Baja California, Mexico, and adjacent islands. Occas* Pap. Calif. Acad. Sci.

135:1-127.

- and B. T. Berke. 1986. A new species of Serradigitus from central California (Scorpiones:

Vaejovidae). Pan-Pac. Entomol. 62(4):(in press).

Scientific Note

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, PP.36S-369

A Namatode Parasite of Erebia occulta Roos & Kimmich
(Nematoda; Lepidoptera: Satyridae)

According to the list published by Poinar (1975, Entomogenous Nematodes, Brill,
Leiden), nematode parasites have been recorded for 232 lepidopteran species, of
which only 20 are butterflies. Parasitism records for North American species include:
Pieris rapae (L.) Pieridae; Polygonia comma (Harris), P. interrogationis (Fabricius),
and Vanessa atalanta (L.), all Nymphalidae (Puttier and Thewke, 1971, Ann.
Entomol. Soc. Am., 64:1177-1178). As of November 1986, Poinar {in litt.) had no
nematode parasite records for the Satyridae. The nematode group responsible for
parasitism in butterflies is the Mermithidae, which are obligate parasites. They have
no free-living or nourishment-receiving stages outside of their hosts. Most of the
moth nematode parasites are also Mermithidae, but there are 6 records of
Steinernematidae, which behave as both facultative and obligate parasites, and one
record for Rhabditidae, which are facultative parasites.

In Alaska on the morning of 3 July, 1986 I collected a male specimen of Erebia
occulta Roos & Kimmich (Satyridae) as it was flying over a low scree slope at mile
41.5 Nome-Council Road (64°39'N, 164°20'W, 30-120 m). The specimen was
papered and subsequently relaxed for spreading. When removed from the relaxing
chamber, about 5 mm of a mermithid was found protruding from the 7th abdominal
segment of the specimen, through which it had apparently bored a hole before dying.
Using forceps, the worm was carefully extracted. When measured, it extended 10.1
cm. The butterfly’s abdomen measured 9 mm. The nematode was placed in 70%
isopropanol and sent for identification to Dr. George O. Poinar, Jr. at the University
of California, Berkeley, who determined that it was a “postparasitic juvenile and
could not be identified past family level (Mermithidae).”

Figure 1. Specimen of Erebia occulta from which 10.1 cm juvenile mermithid was extracted. Scale = cm.

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VOLUME 63, NUMBER 4

369

The butterfly from which the mermithid was extracted behaved normally in the
field, and from its somewhat damaged condition it must have been on the wing for
several days prior to capture. It is shown in Fig. 1, and its parasite in Fig. 2.

Clifford D. Ferris, Bioengineering Program, University of Wyoming, Laramie,
Wyoming 82071.

Figure 2. Juvenile mermithid extracted from male specimen of Erebia occulta.

Scientific Note

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, p. 370

Heleocoris brasiliensis De Carlo is a Ctenipocoris
(Hemiptera: Naucoridae)

While recently undertaking a character analysis of a large number of naucorid taxa
in the subfamily Laccocorinae in an attempt to clarify the generic concepts in this
group I examined two specimens of Heleocoris brasiliensis De Carlo from the Mato
Grosso region of Brazil. It was immediately apparent that this species is not
congeneric with the type-species of Heleocoris, H. tabidulus Stal from Syria, but
instead belongs to the genus Ctenipocoris Montandon. The following nomenclatural
change is thus proposed.

Ctenipocoris brasiliensis (De Carlo) New Combination

Heleocoris brasiliensis De Carlo, 1968. Rev. Soc. Entomol. Arg., 30:101

This species may be assigned to Ctenipocoris by the possession of the following
characters: eyes triangular when viewed from above, with barely developed lateral
flange, converging ventrally below and behind the folded anterior margin of the
head; lateral margins of abdominal tergites lacking numerous stout spines, bearing
instead scattered short slender pale spines intermixed with long pale setae; fore tibia
and tarsi very short and stout; middle and hind tibiae with numerous very long and
stout reddish spines, their length exceeding the diameter of the middle tibia; middle
tibia lacking a dense pad of short gold setae distally on the ventral face. Based on
comments in Usinger (1935, Rev. Entomol., 5:135) I suspect that another South
American Heleocoris species, H. spinipes Montandon, may belong to Ctenipocoris
as well, but I have not examined any specimens of this taxon. La Rivers (1969, Biol.
Soc. Nev. Occ. Pap., 20:5) indicated that none of the Neotropical species held in
Heleocoris truly belonged there but suggested no alternative generic placements.

The discovery of Ctenipocoris in the Western Hemisphere adds yet another
naucorid genus to the Neotropical fauna, and provides a link between the tropical
Naucoridae of Africa and South America, implying a vicariance pattern resulting
from the Cretaceous breakup of Gondwanaland. Based on material I have
examined, the range of Ctenipocoris now includes Brazil, Zaire, Burma, Thailand,
Vietnam, Malaysia, Singapore and Java, giving it a circumtropical distribution.

Dan A. Polhemus, Univ. of Colorado Museum, 3115 S. York St., Englewood, Colo.
80110

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PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 371-376

Two New Trichogrammatidae (Hymenoptera) From North America:
I tty sella lagunera Pinto and Viggiani (N. Gen, N. Sp.) and
Epoligosita mexicana Viggiani (N. Sp.)

John D. Pinto and Gennaro Viggiani

(J.D.P) Department of Entomology, University of California, Riverside, CA
92521; (G.V.) Institute of Agricultural Entomology, University of Naples, Portici,
Italy

Abstract.—Ittysella lagunera Pinto and Viggiani, and Epoligosita mexicana
Viggiani are described. Both are parasites of leafhopper eggs. Ittysella is assigned to
the Paracentrobiini and appears closest to Ittys Girault. Epoligosita mexicana is
closest to E. clara Hay at and Viggiani, from India.

Recent surveys for parasites of leafhoppers on grape have been conducted in
northern Mexico and the southwestern United States by D. Gonzalez, Division of
Biological Control, University of California, Riverside. Among the egg parasites
collected are two new taxa of Trichogrammatidae, Ittysella lagunera Pinto and
Viggiani (n. gen, n. sp.) and Epoligosita mexicana Viggiani (n. sp.). Both are
described below. Collections of E. mexicana represent the first records of this genus
in the New World.

Ittysella Pinto and Viggiani, n. gen.

Antenna of male and female with a single anellus, two funicle segments and a
two-segmented club; funicle II annuliform, distinctly shorter than I. Maxillary palpi
one-segmented (Fig. 1). Forewing (Fig. 5) moderately narrow, ca. 0.4 as wide as
long; marginal vein contacting anterior border of wing, contiguous with submarginal
vein at base, terminating abruptly at apex; stigmal vein well-developed, slightly
constricted at base; disc with linear vein tracks; RSi track absent. Male genitalia (Fig.
6 ) with phallobase subconical in shape and a relatively small anterodorsal aperture;
parameres distinct, unciform, apices directed laterally; volsellar digit! absent.

Type species.—Ittysella lagunera Pinto and Viggiani, n. sp.

Remarks.Itty sella belongs to the Paracentrobiini as defined by Viggiani (1971).
This tribe also includes Paracentrobia Howard, Ittys Girault and Paraittys Viggiani.
Structure of the male genitalia suggests that Ittysella is closest to Ittys. In both, the
parameres are well developed and unciform. Parameres are lacking in
Paracentrobia, and are present but of an entirely different structure in Paraittys (see
Viggiani, 1973). Ittysella differs from Ittys in lacking an RSi vein track, and from all
genera of the tribe in possessing antennae with a single anellus and a club with two
rather than three segments.

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PAN-PACIFIC ENTOMOLOGIST

Figures 1-4. Ittysella lagunera. 1. Head, ?, anteroventral view (Torreon, Coah.) (940X). 2. Thorax, ?
(Torreon, Coah.) (540X). 3. Antenna (right), 2 (Torreon, Coah.) (780X). 4. Antenna (left), S (Caborca,
Son.) (780X).

Ittysella lagunera Pinto and Viggiani, n. sp.

Length 0.40-0.45 mm. Color yellow with limited brown as follows: basal portion
of pedicel and antennal club; mandibles; transverse stripe on gena behind eye;
ventral and dorsal margins of pronotum; anterior and posterior margins of
mesepisternum; metacoxa; marginal and stigmal vein, and area below them on
fore wing; maculae on dorsolateral margins of visible urotergites, together appearing
as a longitudinal stripe; apical half of ovipositor. Legs pallid. Eyes and ocelh reddish.

Head ca. 1.15 as wide as thorax; mandibles tridentate; thorax (Fig. 2) shorter than
gaster (ca. 0.6 gaster length), midlobe of mesoscutum subpentagonal, with a pair of
setae near apical third at lateral margin and another more medial pair near posterior
margin, both pair subequal in length; mesoscutellum ca. 0.67 length of mesoscutum,
with two pair of setae, posterior pair closer together and longer than anterior pair.
Forewing (Fig. 5) with marginal vein elongate; stigmal vein short, broad, only
slightly constricted at base, ca. 0.20 length of marginal vein; marginal vein with six
relatively elongate setae dors ally; premarginal vein with one elongate seta; disc
relatively sparsely setose, with six poorly developed vein tracks, tracks most distinct

VOLUME 63, NUMBER 4

373

5 6

Figures 5-6. Ittysella lagunera. 5. Forewing, 9 (Torreon, Coah.). Male genitalia, ventral view
(Hermosillo, Son.).

at apical third of wing; RSi absent or indicated by one or two setae at most; fringe
setae elongate, longest seta about half the greatest wing width (see below);
substigmal macula obsolescent. Hind wing narrow, elongate, with only one row of
setae on disc; hamuli on apex of an acute projection of anterior margin of wing;
longest fringe seta ca. 1.7 greatest wing width (at hamuli). Legs unmodified;
tarsomeres subequal; hind tibial spur ca. 0.6 length of basitarsis.

Female .—Antenna (Fig. 3) with scape 3.60 as long as wide; pedicle almost twice as
long as wide and 0.55 the length of scape; single anellus distinct; funicle segments
short, subequal in width, wider than long, distinctly narrower than club; funicle I
widening apically, 0.6 as long as wide; funicle II annuliform, only a third as long as
wide and ca. half the length of I; club elongate ca. 3.3 as long as wide, widest at basal
fourth, with two distinct but appressed segments, first segment 1.5 as long as second,
club with 3-4 linear sensilla on each segment and relatively short inconspicuous
setae.

Ovipositor moderately elongate, 0.16 mm long, length ca. half that of gaster and
1.4 that of hind tibia, not projecting appreciably beyond apex of gaster.

Male .—Coloration similar to female but with legs more brownish primarily from
the coxae to tibiae. Antenna (Fig. 4) as in female except club shorter, ca. 2.9 as long
as wide, with 1-2 linear sensilla on each segment and with more elongate setae on
apical half. Phallobase (Fig. 6) 0.07 mm long, 0.5-0.6 as wide as long; unciform
parameres laterally directed, each with a moderately long ventral seta at base,
parameres occupying 0.25 length of entire phallobase; medio-ventral ridge
obsolescent; aedeagus subequal in length to phallobase and ca. half the length of
hind tibia; apodemes occupying 0.5-0.6 length of aedeagus.

Type information .—Holotype $ and allotype from MEXICO, Coahuila,
Torreon; 16 July 1985; D. Gonzalez, collr.; “ex. grape stem cuttings.”; deposited in
the National Museum of Natural History. Twenty paratypes (6 dd, 14 $ ?) also

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PAN-PACIFIC ENTOMOLOGIST

from Torreon deposited as follows: British Museum (Natural History) 1 6,1 $;
Canadian National Collection, Ottawa, 1 c?, 1 9; University of California,
Department of Entomology, Berkeley, 1 9 ; University of California, Department of
Entomology, Riverside (UCR), 2 6 6, 5 9 9; University of Naples, Institute of
Agricultural Entomology, Portici (UNP), 2 6 6,69^.

Holotype, allotype and ten of the paratypes are slide mounted in Canada balsam;
the remaining paratypes are point mounted. The type series emerged from grape
cuttings harboring eggs of the leafhoppers Dikrella cockerelli (Gillette) and
Erythroneura ziczac Walsh.

Records. —MEXICO. Baja California Sur\ Las Barracas, ca. 30 km E Santiago, 1
6,2 9 9, 19 and 29 April 1984, yellow pan trap, P. DeBach (UCR). Coahuila:
Parras, 1 (3,3 9 9,29 August 1985, D. Gonzalez (UCR). Torreon, 9 <3 d, 13 9 9,16
and 28 July 1985, D. Gonzalez. Sonora: Caborca, 2 3 3,2 9 9,19 August 1986, D.
Gonzalez (UCR). Hermosillo, 7 33, 10 9 9,29 and 30 July 1985 (UCR, UNP).
UNITED STATES. Arizona: Cochise Co., Chiricahua Mts., Sunny Flats
Campground, 1 9,10 July 1983, screen sweeping, A. Mayor (UCR). New Mexico:
Las Cruces, 5 3 3,10 July 1986, D. Gonzalez (UCR).

All material from Coahuila, Sonora and New Mexico emerged from collections of
grape cuttings. Leafhopper eggs of the following species were deposited in the leaves
of these collections: Dikrella cockerelli (New Mexico, Coahuila), Erythroneura
ziczac (Coahuila), and Erythroneura variabilis Beamer (Sonora).

Remarks. —All material examined is relatively uniform structurally. Specimens
from Arizona and New Mexico have a somewhat longer forewing fringe (0.50-0.56
wing width; x = 0.524 ± 0.03 S.D.; n = 6) than material from Mexico (0.43-0.50
wing width; X = 0.480 ± 0.02; n = 8).

The specific epithet is derived from La Laguna, the name commonly applied to the
type locality and environs.

Epoligosita mexicana Viggiani, n. sp.

Female. —Length: 0.47 mm. Body yellow; eyes and ocelli blackish; tips of
mandibles yellowish brown; antenna, middle and hind tibiae, meso- and
metapleurae, with some brown; forewings with basal third and substigmal area
infuscated; ovipositor honey-brownish.

Head normal for the genus, ca. one-fourth wider than high. Mandibles tridentate.
Maxillary palpi uniarticulate and labial palpi reduced. Antenna (Fig. 7) with scape
3.0 as long as wide, pedicel slightly shorter but distally ca. one-third wider; single
anellus distinct; funicle segment about as long as wide, narrower than pedicel and
club; club not clearly divided in two segments, 2.0 as long as pedicel, ca. 5.0 as long as
funicle segment and ca. 3.0 as long as wide; first segment with one linear sensillum
and distal one with 4.

Thorax ca. one-third shorter than gaster, with main dorsal characters as illustrated
for male (Fig. 8). Forewing (Fig. 9) ca. 3.0 as long as wide; disc with only a single
minute seta located mid-way between center of marginal vein and posterior margin;
longest fringe setae as long as greatest wing width. Hind wing very narrow, without
setae on disc. Legs normal; fore, middle, and hind tarsomere ratios respectively as
follows: 8:9:10; 18:13:13; 13:12:12 (hind tibial spur = 8).

Gaster conic; ovipositor occupying about half length of gaster, about as long as
hind tibia; third valvulae very short, one-sixth of the entire ovipositor.

VOLUME 63 , NUMBER 4

375

8

Figures 7-10. Epoligosita mexicana. 7. Antenna, 9.8. Thorax, 3.9. Forewing. 10. Male genitalia.

Male .—Similar to female, but antenna with longer setae on the club. Male
genitalia as in Fig. 10, 0.057 mm in length.

Type information .—Holotype 9 from MEXICO, Coahuila, Torreon; 28 August
1985; D. Gonzalez, collr.; allotype, same data as holotype, except 16 July 1985. Both
deposited in the National Museum of Natural History. Paratypes from the same
locality deposited as follows: University of California, Department of Entomology,
Riverside; University of Naples, Institute of Agricultural Entomology, Portici.

Holotype, allotype and 6 paratypes are slide mounted in Canada balsam; 3
additional paratypes are point mounted. All type specimens emerged from leaves of
grape cuttings harboring eggs of the leafhoppers Dikrella cockerelli and
Erythroneura ziczac.

One female, UNITED STATES, Arizona, Cochise Co., 25 August 1982, J.
LaSalle, collr., screen sweeping, probably belongs to the new species.

Remarks .—The new species Epoligosita mexicana runs to the group of Epoligosita
Girault with the funicle segment quadrate or wider than long (couplet 1 in Hayat and
Viggiani, 1981), in which E. biclavata (Girault and Dodd), E. nudipennis (Kryger)
and E. clara Hayat and Viggiani are included. Of these species the closest seems to be
E. clara, from which E. mexicana may be separated by having the scape 3.0 as long as
wide (in E. clara 3.5 as long as wide), the funicle segment as long as wide (in E. clara a
little wider than long), the club twice as long as length of pedicel (shorter in E. clara ),

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PAN-PACIFIC ENTOMOLOGIST

ovipositor as long as half length of gaster (in E. clara more than half length of gaster),
and the forewings with the basal third and substigmal area infuscated (in E. clara only
basal third of forewings infuscated).

Acknowledgments

Figures 5-6 were prepared by Patricia Mote. Robert Velton was responsible for
the SEM photographs and specimen curation. Individuals participating in the
leafhopper parasite surveys include J. Ellington, D. Gonzalez, F. Gonzalez, L.
Guerra Sobrevilla, M. Moratorio, A. Tijerina, and W. White.

Literature Cited

Hayat, M. and G. Viggiani. 1981. The genus Epoligosita from India, with descriptions of two new species
(Hymenoptera: Trichogrammatidae). Boll. Lab. Ent. Agr. Portici, 38:119-123.

Viggiani, G. 1971. Ricerche sugli Hymenoptera Chalcidoidea XXVIII. Studio morflologico comparativo
dell’armatura genitale esterna maschile dei Trichogrammatidae. Boll. Lab. Ent. Agr. Portici,
29:181-222.

Viggiani, G. 1973. Ricerche sugli Hymenoptera Chalcidoidea. XXXV. Nuovi tricogrammatidi dTsraele
{Paraittys latipennis, n. gen., n. sp. e Oligosita gerlingi, n. sp.), con reperti su varie specie. Boll.
Lab. Ent. Agr. Portici, 30:152-157.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 377-380

Four Species of Scuttle Fly (Diptera: Phoridae)

From Dominican Amber

R. H. L. Disney

Field Studies Council Research Fellow, University Museum of Zoology,
Cambridge, CB2 3EJ, U.K.

Abstract .—Four species of Phoridae, Dohrniphora poinari n. sp., Megaselia
amberae n. sp., M. bernsteinae n. sp. and M. dominicana n. sp. are described from
Dominican amber dated 30 m.y. B.P. ± 10m.y. (Oligocene, or possibly late Eocene
or early Miocene). Morphologically these species resemble living species.

Dr. George O. Poinar, Jr., of the University of California, Berkeley, asked me to
examine four specimens of Phoridae preserved in Dominican amber dated 30 m.y.
B.P. ± 10 m.y. They are most probably Oligocene but are possibly late Eocene or
early Miocene. All four specimens prove to be undescribed females. In dealing with
the present-day fauna one would not describe species on the basis of females alone,
unless they were highly distinctive. In the case of fossils more than 10 m.y. B .P. in age
this precaution can be safely ignored; although there remains a low probability of a
male of one of these species being discovered and, because of sexual dimorphism,
treated as a distinct species in error.

The four species are described below:

Dohrniphora poinari sp. nov.

A medium sized species with wing length about 1.2-1.4 mm. Scutum yellowish
with contrasting dark, brownish, scutellum which is yellowish at margins.
Abdominal tergites dark with median yellowish band, which expands anteriorly on
each tergite.

Legs yellowish to brownish yellow. Hind-tibia with a single dorsal hair palisade
and no pre-apical bristles. Mid-tibia apparently without dorsal hair palisade but with
a pair of bristles in basal quarter and a short anterior pre-apical bristle. Fore-tibia
with five near-dorsal bristles, with 2-5 being short.

Wings with costal index about 0.6. Haltere knob yellowish. Third antennal
segment pale brownish, the two-segmented palps yellow with short apical bristles.
Proboscis not elongated, labella apparent and a little pointed anteriorly. Frons with
chaetotaxy much as in the present-day cosmopolitan Dohrniphora cornuta (Bigot)
but with no apparent anterolaterals, although on the most visible (left) side there is
what appears to be a basal scar in the appropriate position.

Thoracic chaetotaxy with a pair of pre-scutellars, intra-alars and pre-alar bristles.
A humeral bristles and three notopleurals present. Fine hairs present on
mesopleuron. Scutellum with an anterior pair of hairs and a posterior pair of bristles.

Holotype female. D-7-51.

Affinities. While a number of species of fossil Dohrniphora have been described
from Baltic amber (e.g. Brues, 1939), these have all been subsequently transferred to

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the closely related genus Diplonevra Lioy (Borgmeier, 1968). The present-day
Neotropical species are dealt with by Borgmeier (1960, 1961), Borgmeier & Prado
(1975), Prado (1976) and Disney (1983a). These keys and descriptions deal primarily
with the males. However the lack of bristles on the hind tibia and short proboscis will
distinguish D. poinari from the majority of present day species. The long costal
index, contrasting colors of the scutum and scutellum and single pair of scutellar
bristles will distinguish it from the rest.

Genus Megaselia Rondani 1856

Megaselia is the largest genus of Phoridae in the world today, with some 1400
described species. However most species remain undescribed and estimates (Disney,
1983b) suggest that the true total lies between 5,000 and 20,000 species. The present
day Neotropical species are covered by Borgmeier (1962, 1969a, 1969b and 1971)
and Disney (1982).

The three species described below are evidently closely related and share a
number of features. These include a large Costal Index (0.5 or more), short coastal
cilia (<0.1 mm). Vein Sc confluent with Rl, Vein 3 forked, Mesopleuron bare,
haltere knob mainly yellowish and a posterior pair of bristles and anterior pair of
hairs on the scutellum. They thus belong to a sub-section of Borgmeier’s (1962)
Group VII.

Eight species of Megaselia have been described from Baltic amber and two species
from Zanzibar copal (Borgmeier, 1968). Whilst all the descriptions are inadequate
by present-day standards none agree with the three species from the Dominican
amber described below.

Megaselia amberae sp. nov.

A medium to large species (wing length 2.63 mm). Frons black. Scutum and
scutellum dusky orange yellow. Antennae orange brown. Abdominal tergites appear
to be a bit darker (but considerably obscured in specimen). The Coastal Index is
0.54-0.55. The costal ratios are 4.1:1.8:1. The costal cilia measure 0.07-0.09 mm.
There appears to be only two notopleurals, otherwise the chaetotoxy seems to be the
standard (ground plan?) for the genus. The palps are orange yellow in color with
standard bristles. The last two segments of the fore tarsus are subequal in length.
Legs yellowish with a dark apex to hind femur.

Holotype female. D-7-54.

Megaselia bernsteinae sp. nov.

A little smaller than the previous species (wing length 2.08 mm). Frons black,
scutum and scutellum yellow. Antennae brown. Abdominal tergites dark greyish
brown. Tergites 5 with posterior margin concave. Tergite 6 with anterior margin
notched in middle third.

The costal index is 0.552. Costal ratios 3.64:1.56:1. Costal cilia 0.065-0.075 mm
long. Only two notopleurals present, otherwise chaetotaxy seems to be standard.
Palps not visible in specimen. Legs yellowish except for apex of hind femur.

Holotype female. D-7-52.

Megaselia dominicana sp. nov.

A little smaller than the above species (wing length 1.60 mm). Frons brown,
scutum and scutellum yellowish brown. Antennae brown. Abdominal tergites brown

VOLUME 63, NUMBER 4

379

Figure 1. Lateral view of Dohrniphorapoinari sp. n. in amber from the Dominican Republic. Figure 2.
Dorsal view of Megaselia dominicana sp. n. in amber from the Dominican Republic.

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PAN-PACIFIC ENTOMOLOGIST

and somewhat narrow after tergite 2. Tergite 5 with concave notch in middle of hind
margin. Tergite 6 with side and rear margins forming a continuous semicircle.

Costal Index 0.497. Costal ratios 3.56:1.48:1. Costal cilia 0.50-0.051 mm long.
Three notopleurals present and rest of chaetotaxy appears to be standard. Femora
somewhat brownish, otherwise legs yellowish. Tarsal segment 5 of front leg about
1.25 X length of 4. Palps not visible in specimen.

Holotype female. D-7-53.

Discussion

The four species described above belong to the two genera which dominate
Neotropical forest Phorid faunas today. Indeed on morphological grounds all appear
modern in appearance. However our knowledge of the ground-plan characters of
both genera is still extremely slight. In the Megaselia species the long costa, short
costal cilia and vein Sc confluent with R1 are probably all plesimorphic features. All,
however, are not uncommon in the genus today. The genus Dohrniphora is almost
certainly closer to the ground-plan of the Phoridae than Megaselia, for example in
possessing a two segmented palp. D. poinari in lacking a hair palisade on the
mid-tibia and pre-apical bristles on the hind tibia stands with a minority of the species
in the genus today. The relatively short proboscis is almost certainly plesiomorphic
for the genus.

The Megaselia specimens all have balloon-like distensions of the abdominal
pleura. These are evidently post-mortem changes and not structural features.

Specimens are in the collection of G. O. Poinar, Jr., Berkeley, California and will
eventually be deposited at the American Museum of Natural History, New York.

Literature Cited

Borgmeier, T. 1960. Gefluegelte and ungefluegelte Phoriden aus der neotropischen Region, nebst
Beschreibung von sieben neuen Gattungen (Diptera, Phoridae). Studia ent. 3,257-374.
Borgmeier, T. 1961. Weitere Beitraege zur Kenntnis der neotropischen Phoriden nebst Beschreinbung
einiger Dohrniphora-Alien aus der indo-australischen Region (Diptera, Phoridae). Studia ent. 4,
1 - 112 .

Borgmeier, T. 1962. Versuch einer Uebersicht ueber die neotropischen Megaselia arten, sowie neue oder
wenig bekannte Phoriden veschiedener Gattungen (Diptera, Phoridae). Studia ent. 5,289-488.
Borgmeier, T. 1968. A Catalogue of the Phoridae of the World (Diptera, Phoridae). Studia ent. 11,1-367.
Borgmeier, T. 1969a. Bredin-Archibold-Smithsonian Biological survey of Dominica: The Phoridae of
Dominica (Diptera). Smithsonian Contrib. 23,1-69.

Borgmeier, T. 1969b. New or little-known Phorid flies, mainly of the Neotropical Region (Diptera,
Phoridae). Studia ent. 12, 33-132.

Borgmeier, T. 1971. Further studies on Phorid flies, mainly of the Neotropical Region (Diptera,
Phoridae). Studia ent. 14,1-172.

Borgmeier, T., and Prado, A. P. do 1975. New or httle-known Neotropical Phorid flies, with description
of eight new genera (Diptera, Phoridae). Studia ent. 18, 3-90.

Brues, C. T. 1939. Fossil Phoridae in Baltic amber. Bull. Mus. Comp. Zool. 85(6), 413-436.

Disney, R. H. L. 1982. A curious new species oiMegaselia from Brazil (Diptera, Phoridae). Z. ang. Zool.
68,415-418(1981).

Disney, R. H. L. 1983a. Four new species of Dohrniphora (Diptera, Phoridae) from Panama. Ent. scand.
14, 452-456.

Disney, R. H. L. 1983b. A useful new character in the giant genus Megaselia (Diptera, Phoridae) with two
new species from Britain. Z. ang. Zool. 70, 225-234.

Prado, A. P. do 1976. Records and descriptions of Phorid flies, mainly of the Neotropical Region
(Diptera, Phoridae). Studia ent. 19,561-609.

Scientific Note

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 381-382

Pteroloma nebrioides Brown in Idaho (Coleoptera: Agyrtidae)

Pteroloma nebrioides Brown was initially described from southeastern British
Columbia, and recorded from a number of sites throughout British Columbia,
western Alberta, and one site in northwest Montana. Distribution, taxonomy, and
biology, have been reported by R. Anderson and S. Peck (1985, The Insects and
Arachnids of Canada, Part 13, The Carrion Beetles of Canada and Alaska,
Coleoptera: Silphidae and Agyrtidae, 121 p). Their single United States record is
from Glacier National Park, Montana (R. Anderson, pers. comm.).

Here, I report this species from two new localities in the United States, thus
representing a significant range extension and a new state record. Distribution of this
species is now extended approximately 400 km southwest of Glacier National Park.
Specimens have been collected at: IDAHO (NEW STATE RECORD), Clearwater
County, Isabella Creek, 43 km NNE of Headquarters, T41N R7E s.31 NESW, 518
m, 21.IX.1984 (1) and 24.VII.1985 (2), P. J. Johnson; and Idaho County, Packer
Meadow, 2.4 km E. Lolo Pass, 1585 m, T38N R15E s.l5, 25.V.1986 (2) and
30. V. 1986 (2), P. J. Johnson and J. R. LaBonte. A single agyrtid larva was collected
at Isabella Creek with two adults on 24. VII. 1985 and is assumed to be this species.
Adult specimens have been deposited in the W. F. Barr Entomological Museum,
University of Idaho, and my personal collection. The larva is with A. F. Newton, Jr.,
Field Museum of Natural History, Chicago.

Isabella Creek is in the Clearwater mountains, a western extension of the
Bitterroot Range, at a site dominated by western red cedar {Thuja plicata Donn.).
Grand fir (Abies grandis (Dougl.) Forbes) and western white pine (Pinus monticola
Dougl.) dominate the steep surrounding slopes. Specimens collected were found
under moss coated cobbles on silty-sand in a highwater channel which is lateral and
parallel to the main stream channel. The site is densely shaded by red alder (Alnus
rubra Bong.), thimbleberry (Rubus parviflorus Nutt.), devil’s club (Oplopanax
horridum (Smith) Miq.), and ladyfern (Athyriumfilix-femina (L.) Roth), and is cool
and humid on warm summer days. Isabella Creek is a large second order stream with
a moderate gradient initiating in subalpine elevations, but largely flowing through
dense mesophytic mixed conifer forest on steep mountainous terrain. The water is
cold and undoubtedly assists in maintaining a cool temperature regime in the
adjacent vegetation which produces heavy, day-long shade.

Packer meadow is a natural, wet, subalpine frost pocket surrounded by subalpine
fir (Abies lasiocarpa (Hook.) Nutt.), lodgepole pine (Pinus contorta Dougl.) and
Englemann spruce (Picea englemannii Parry) forest, near the crest of the Bitterroot
Range. The meadow flora is composed largely of Camas lily (Camassia quamash
(Pursh) Greene), bistort (Polygonum bistortoides Pursh) and sedges (Carex spp.),
with occasional shrubs of bog birch (Betula glandulosa Michx.) and Labrador-tea
(Ledum glandulosum Nutt.). Winter snow packs are often deep (3-4 m), with final
snowmelt usually complete by mid-June, but soil temperatures are cold-to-the-touch
until July. A meandering stream and small island-like stands of mature subalpine fir
and Englemann spruce characterize the meadow, with these trees providing windfall

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logs and limbs under which all specimens were collected on fine to moderately
coarse, decomposed organic matter.

My thanks to J. R. LaBonte for collecting assistance; and too J. B. Johnson and
J. P. McCaffrey for reading the manuscript. This article is being published with
permission of the director of the Idaho Agricultural Experiment Station as Research
Paper no. 8772.

Paul J. Johnson, Department of Plant, Soil and Entomological Sciences, University of
Idaho, Moscow, ID 83843.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 383-388

Flume Collecting: A Rediscovered Insect Collecting Method, with
Notes on Insect Extracting Techniques

Jeffrey A. Halstead and Robert D. Haines

Department of Biology, California State University Fresno, Fresno, California
93740; present address 2110 N. Hayes, Fresno, California 93722. Tulare County
Agricultural Commissioner’s/Sealer’s Office, Visalia, California 93291

Abstract .—The use of flumes (i.e., a channel for the transportation of water) as a
collecting source is discussed. A five mile long flume near Ash Mountain, Sequoia
National Park, Tulare County, California is described, along with techniques used to
collect and extract insects from flume debris. This flume, utilizing the various
collecting techniques described, has produced an astonishing diversity and
abundance of insects. In the 1930’s and 1940’s, much of the material from this site was
cryptically labeled and thus type locations and basic locality data were misleading
and difficult to pinpoint. Notes on such labels are presented.

Flumes, as a collecting source, were heavily utilized in the 1930’s by such collectors
as Dr. E. C. Van Dyke, Mr. F. T. Scott and Mr. R. S. Wagner. Their work greatly
added to the knowledge of California species of Coleoptera, many of which were new
or rare species collected from flumes. The latter two researchers collected almost
exclusively from flumes in the Sierra Nevada foothills. The function of these flumes is
water diversion from rivers and streams for the generation of hydroelectric power
and public uses. These flumes act as a giant moving pit trap, catching insects that
either blunder in or are drawn to a water source.

Two basic techniques were used to collect these insects: (1) watching the water
surface as material approached and netting what was seen and (2) pulling out and
sorting through debris that had accumulated at the end of the flume prior to the water
being run through a powerplant or into a holding pond. The first technique was time
consuming, limited to those insects that fell into the flume that day, and yielded
mostly larger species. The second technique yielded much better results (i.e., greater
diversity and abundance), but required the use of some sort of debris collection
device.

All three of the previously mentioned collectors made use of a flume located in
Tulare County, California on the Middle Fork of the Kaweah River, initiating at
Potwisha Campground in Sequoia National Park and terminating on the hillside
south of the Ash Mountain Park Headquarters. This flume (Figures 1-4) flows
through approximately five miles of Chamise Chaparral and Foothill Woodland
plant communities at an elevation of 660 m (2200 ft.) and empties into a large (20
m X 60 m) forebay (Figure 5) whose teardrop shape and exposure to prevailing
breezes promotes clockwise surface currents. A wooden boom (10 cm x 15
cm X 3.5 m) projecting into these currents traps and accumulates the floating debris

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Figures 1-4. Figs. 1-3. Typical sections of the Ash Mountain. Kaweah Powerhouse #3 flume,
illustrating adjacent and overhanging vegetation and rugged terrain of Sequoia National Park (June). Fig.
4. The flume, denoted by arrows, curves along hillsides of Chamise Chaparral and Foothill Woodland
habitats at 660 m (2200 ft.).

(boom-trap method) (Figure 6). The researchers would scoop out the debris and
spread it over the ground for examination.

The Ash Mountain flume is unique in comparison to others in the Kaweah area and
throughout California. In addition to having a built-in debris accumulating feature
(i.e., the boom), the edge of the flume throughout most of its length is at ground
level. This greatly enhances the chance of insects falling into it—in comparison to
flumes that are supported by trestlework 1 to 7 m above the ground. The native flora
adjacent to and overhanging the flume’s edge is another factor contributing to the
great diversity and abundance of insects collected.

The early researchers labeled the material collected at the Ash Mountain flume in
a variety of ways. Because of this, a number of type localities have been difficult to
pinpoint. Examples of such labels are: near Postwisha, Sequoia National Park;
Sequoia National Park, 500-2000 ft., Potwisha (Van Dyke); Kaweah; Sequoia
National Park, 2000-3000 ft. (Scott); K.P.H.R.; Kaweah Powerhouse Reservoir;
Kaweah (Wagner).

Some examples of type material collected from this site include: Coleoptera,
Cerambycidae: Ergates pauper Linsley, Paranoplium gracile laticolle (Linsley),
Aneflomorpha California Linsley = A. parowana Casey, Neoclytus resplendens
Linsley, Leptura sequoiae (Hopping); Scarabaeidae: Pleocoma tularensis Leach,

VOLUME 63, NUMBER 4

385

Figures 5-8. Fig. 5. Overview of forebay from beside the entrance of the flume. Floating debris, having
floated down the flume, can be seen on the forebay’s surface. Fig. 6. View of forebay towards the entrance
of the flume. The wooden boom, floating on the forebay’s surface and extending from its edge, traps and
accumulates debris which has floated down the five mile flume. Fig. 7. Window screen panels are inserted
into the flume to trap and accumulate flume debris before it enters the forebay. Fig. 8.
Berlese-photoattractive trap used to extract insects from the flume debris.

Coenonychafusca McClay; Buprestidae: Polycesta tularensis Chamberlin, Polycesta
crypta Barr, Acmaeodera simulata Van Dyke; Elateridae: Euthysanius cribicollis
Van Dyke; Diptera, Acroceridae: Ocnaea sequoiae Sabrosky; Hymenoptera,
Pompilidae: Allaporus amabilis E\ 2 im = Pompilus (Aporus) smithianus Cameron.
The above information is presented so that researchers will have a clearer
understanding of how and where such material was collected.

Beginning in 1982 a renewed effort was made to utilize the Ash Mountain flume
facility by the authors, Dr. D. J. Burdick (California State University Fresno) and
Mr. W. F. Peregrin (Fresno County Agricultural Commissioner’s Office, Fresno,
California). Initial efforts using the old technique of retrieving accumulated debris
(mostly leaves) at the forebay yielded a great diversity and abundance of species.
Insects are easily observed and collected from the drying debris as they dry their
wings and/or move among the leaves. Dead insects are also found while sorting
through the debris, commonly stuck to wet leaves.

The amount of debris collected by the flume varied with the season. The greatest
accumulation of debris correlated with fall leaf drop. Two researchers during a
typical 8 to 10 hour period can sort approximately 60 to 75 square meters (200 to 250

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PAN-PACIFIC ENTOMOLOGIST

square feet) of debris, 2.5 to 5 cm (one to two inches) in depth. Because the flume is
continually accumulating debris both throughout the day and the night, the day’s
collecting was usually terminated only by the lack of visibility (i.e., sunlight). Often,
all of the debris which had accumulated behind the boom could not be sorted in a
single day. Because this unexamined debris represented material which had floated
down the flume that day (thus containing live insects) and was known to contain a
good diversity and abundance of late-day and crespuscular species, the debris was
sealed in garbage bags and transported to the laboratory for further treatment (see
discussion under Insect Extraction Techniques).

While the boom trap method was successful in 1982 and early 1983, a large
sandbar, which had accumulated over a number of years, began changing surface
currents reducing debris accumulation behind the boom. As a result, the debris was
patchily distributed over the forebay, reducing its availability. In order to collect the
debris before it entered the forebay, window screen panels were inserted into the
flume against an existing metal grating at its entrance into the forebay (Figure 7). The
debris was removed from the screens and examined about every 30 minutes. During
an 8 to 10 hour period, two researchers could collect and sort approximately 24
square meters (80 square feet) of debris. While some small (< 1 mm) insects
probably passed through, many were stuck to leaves on the screens. The small
amount of debris examined (versus the boom trap method) improved collecting
efficiency. Also, the insects collected on the screens were very active and easily
detected.

The screens gave excellent results for material-of-the-day (including activity
period information), but the accumulated debris behind the boom covered several
days and nights, and in general, held a much greater abundance and diversity
(including nocturnal species). At times, screening was not practical due to large
amounts of leaves and/or algae which blocked flow through the screens. Though the
sandbar was removed in 1985, the benefits of using screens for collecting
material-of-the-day precluded abandoning this technique to return strictly to the
boom trap method.

Another problem was encountered in the late Summer and early Fall of 1985. Due
to a lighter than normal winter snowpack in the Sierras and a lack of substantial
Spring rains, water levels in the Kaweah River dropped below diversion levels.
While there had been temporary shutdowns in prior years for flume maintenance and
sand removal, this was the most extensive dry period encountered by the authors,
lasting from August through mid November.

Insect Extraction Techniques
1) Glass-topped Sleeve Cage.

Approximately 10 to 15 cm (4 to 6 inches) of debris was placed in each of two to
three glass topped sleeve cages (0.6 x 0.6 x 1 m). An oscillating fan was used to
move air through the sleeve cage(s) to dry the debris. A white light (60 watt bulb) was
positioned on the top of each cage to attract insects. The cages were checked
periodically for two days though insect activity rapidly declined after one day. Insects
were removed with an aspirator or by hand. Cryptic species were commonly found
on the bottom of the cage or among the debris. This technique was productive though
limited by the amount of debris that could be placed in the sleeve cage(s). Also, the
debris was difficult to dry and immediately started to decay.

VOLUME 63, NUMBER 4

387

2) Enclosed Malaise Trap and Tent trap.

To increase the amount of debris that could be handled at one time, an enclosed
Malaise trap and tent trap were utilized. The enclosed Malaise trap (2 x 1 x 1.8 m)
was set up outdoors. The bagged debris and/or debris that was first examined in the
sleeve cages was placed inside the sealed trap the morning following the collecting
trip. The warmth and drying effect of the sun immediately resulted in insect activity.
Insects readily crawled up the sides of the trap, passed through an inverted funnel,
and fell into an alcohol solution in the collecting head. The alcohol solution
eliminated any damage to specimens. This technique was more productive and less
time consuming than the sleeve cages. A major drawback was that insects which did
not climb up the Malaise trap were not collected. Also, spiders occasionally spun
webs at the entrance to the collecting head, but because of the great numbers of
insects very few were eaten or deterred from entering the head.

The tent trap was devised to be used in the laboratory. This trap was a two-man
tent (2 X 1.2 x 1 m) to which an alcohol-collecting head was attached by a fine mesh
insect net. The debris was placed inside the tent on a 2.5 cm (one inch) chicken wire
screen (1.2 x 1.8 m (4 x 6 ft.)) that sat on blocks 10 cm (4 inches) above the floor of
the tent. An oscillating fan was run outside the tent’s mesh door for about 6 to 12
hours. The screen aided the drying of the debris and also burrowing insects fell from
the screen to the floor of the tent where they could be collected. Very little insect
activity occurred when the fan was on; though once off, increased immediately. This
technique was more profitable than the two previous techniques (especially because
non-climbing insects were also collected). Again, minor spider problems occurred
with the collecting head.

3) Berlese-photoattractive Trap (Figure 8).

This trap (2 x 1.2 x 1.5 m (6 X 4 X 5 ft.)) was the most productive and time
efficient insect extracting method, utilizing the collecting principles of both a large
Berlese funnel trap and a photoattractive trap. It consisted of a black plastic top,
particle board sides with fine mesh windows, two flat internal debris trays (0.9 x 1.5
m (3 X 5 ft.)) made of window screening stacked 15 cm (6 inches) apart, and clear
plastic lower sides which funnel downward into a plastic rain gutter filled with 5 to 7.5
cm (2 to 3 inches) of super saturated saltwater solution or ethylene glycol.

Upon returning home from the day’s collecting trip, about 5 to 10 cm (2 to 4 inches)
of debris was placed on each tray. A light and fan were run outside the trap’s windows
until early morning. Light seeking insects were immediately attracted to the mesh
windows and clear plastic funnel. Those which burrowed into or through the debris
eventually fell off the screens into the rain gutter. With the coming of daylight, the
black plastic top readily heated the inside of the trap, thus forcing insects downward.
The windows were occasionally fogged with a quick knockdown pyrethroid. The
debris was left in the trap for about one week by which time most insects had fallen
into the rain gutter. One last heavy fogging killed any remaining survivors. The
debris trays were removed and the inside of the trap was washed with water to
remove any clinging insects. The resulting rain gutter full of insects was then
screened and the insects preserved for future sorting.

Though much of our material is still unmounted, detailed examination of some
groups by authorities have revealed many new species, male/female associations,
range extensions, and rarely collected species (not to mention the great diversity and
abundance of the more common species). Most of this material has been deposited in

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the California Academy of Sciences, San Francisco; California Collection of
Arthropods, California Department of Food and Agriculture, Sacramento; and the
Canadian National Collection, Ottawa.

Examples of species diversity for some of the families reviewed are (# of genera, #
of species): Coleoptera, Buprestidae (11, 40), Cerambycidae (43, 61),
Chrysomelidae (47, 57); Hymenoptera, Chalcididae (12, 30), Chrysididae (9, 23),
Dryinidae (12, 23), Eumenidae (9, 18), Mutillidae (9, 21), Pompilidae (17, 41),
Sapygidae (1, 5), Sphecidae (22, 40); Diptera, Acroceridae (6, 12). Examples of
some of the rare families encountered are: Coleoptera, Amphizoidae (1, 1),
Cupedidae (1, 1); Hymenoptera, Chalcedectidae (1, 2), Cimbicidae (1, 1),
Eucharitidae (1, 1), Evaniidae (1, 1), Leucospididae (1, 1), Orussidae (1, 1),
Sierolomorphidae (1,1), Stephanidae (1,2); Neuroptera, Mantispidae (3,3).

As illustrated by this preliminary data, when the appropriate techniques are
utilized, flume collecting can be an extremely successful method; giving the collector
access to rare species that would be otherwise unavailable. A great deal of
seasonality and daily activity information can also be gathered using these
techniques. By the publication of this paper we hope to provide an understanding of
where and how the Ash Mountain flume specimens have been collected and also to
encourage the use of flumes as a collecting source.

Acknowledgments

We thank D. J. Burdick, Department of Biology, California State University
Fresno andN. J. Smith, Fresno County Agricultural Commissioner’s Office, Fresno,
California for editorial comments on this paper. We thank also Larry Bezark,
Sacramento, California for the use of his slide to make figure 7.

PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 389-393

New Distribution Records for Some Nearctic Dryinid Wasps, with a
Species List from a Flume in Tulare County, California

(Hymenoptera: Dryinidae)

Jeffrey A. Halstead and Robert D. Haines

Department of Biology, California State University Fresno, Fresno, California
93740; present address: 2110 N. Hayes, Fresno, California 93722. Tulare County
Agricultural Commissioner’s/Sealer’s Office, Visalia, California 93291

Abstract. —A species list, collection periods, and distributional notes (including 12
new California-state records) for 23 species of dryinid wasps collected from a
hydroelectric flume in Tulare County, California are presented. Additionally, new
state records for Gonatopus mimoides in California and G. cyphonotus in Minnesota
are presented. G. pallidiceps and G. portalensis are each reported from a third
locality in California. Twelve genera and 34 species of dryinid wasps are known from
California.

Recently, the world fauna of the wasp family Dryinidae was revised and
distributional information was updated for each species (Olmi, 1984). Prior to this
revision, distribution information for the Dryinidae in America north of Mexico was
available in Krombein (1979).

Presented herein are the results of collecting dryinid wasps from a hydroelectric
flume located at Kaweah Powerhouse #3, Ash Mountain, Tulare Co, California.
Included are a list of 23 species collected from this site, collection periods, the
number of specimens collected thus indicating how common or rare each species is,
and distributional comments (including 12 new California-state records and many
county records) for each species (see Table 1 and text).

The flume winds through approximately five miles of Foothill Woodland and
Chamise Chaparral habitats at an elevation of 660 m (2200 ft) (Halstead and Haines,
in prep.). A total of 62 flume collecting trips (approximately 1100 man hours of
collecting) were conducted from 1982 to 1985. Trips were made throughout the
calendar year though most were conducted between May and October. More trips
were made and man hours spent in 1982 than in other years.

Additionally, new state records of Gonotopus cyphonotus Bradley in Minnesota
and G. mimoides (Perkins) in California are presented. Also, G. pallidiceps
(Perkins) and G. portalensis Olmi are each reported from a third locality in
California.

Anteon funestum. —Widely distributed throughout North America. The flume
specimens represent a new state record for California and the westernmost record for
the species. The nearest record is approximately 1000 km (600 mi) to the southeast in
Tucson, Arizona.

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Table 1. Species Ust, collection period(s) and number of dr 3 dnid wasps collected from the Kaweah Powerhouse #3
flume, Ash Mountain, Tulare County, California in 1982-85. Species are listed alphabetically.

Scientific Name

Months

Collected

No. Specimens

Collected

Anteon funestum (Perkins)*

IV, IX

2

A. popenoei (Ashmead)*

VI

2

A. rugosiceps Kieffer

VI

2

Aphelopus albopictus Ashmead

V-VI

13

A. varicornis Bmes

III-V

6

Apterodryinus californicus (Ash.)

III-V

10

A. Perkins*

VII

1

Bocchus flavipes Kieffer

VI-VII

29

B. hainesi n. sp.^*

VI-VIII

20

Crovettia theliae (Gahan)

IV

1

Deinodryinus atriventris (Cres.)*

VI-VII

14

Dryinus halsteadi n. sp.^*

IV-VIII

10

Esagonatopus niger (Fenton)*

VII-VIII

4

Gonatopus agropyrus Fenton*

VII

2

G. herbarum (Perkins)

VI-VII, IX

10

G. mayori Olmi

VI

1

G. paraleptias (Perkins)

VI-VII, IX

15

Lonchodryinus bakeri (Kieffer)

VII-IX

7

L. fiavus Olmi*

VI

1

L. masneri (Olmi)*

VIII

2

Pseudogonatopus sjoestedti (Kief.)*

V

1

Tetrodontochelyspeculiaris Brues

VI-VII

3

T. unicus (Perkins)*

IX

1

*Denotes a new state record for California.
^Olmi, in prep.

Anteon popenoei. —Known from the central and eastern United States and
Canada. The flume specimens represent a new state record for California and the
westernmost record for the species. The nearest locality record is approximately
1700 km (1000 mi) to the northeast in Alberta, Canada or to the southeast in Texas.

Anteon rugosiceps. —The few locality records are from southeastern Canada, the
western United States, and Mexico. The flume specimens represent a new county
record, the fourth locality record for Cahfornia, and the second record for the Sierra
Nevada Mountain Range. Other California counties include Kern, Santa Clara, and
Siskiyou.

Aphelopus albopictus. —Widely distributed in North America. Recorded from
four localities in California but the flume specimens represent a new county record
and the second and southernmost record for the Sierra Nevada. The other Sierrian
record is from Somerset, El Dorado Co. Other California records (Needles,
Alameda, and Chocolate Mts.) are coastal or southern desert.

Aphelopus varicornis. —Widely distributed in North America. Several records are
noted for California but the flume specimens represent a new county record and the
fifth and southernmost record from the Sierra Nevada. Other Sierrian records
include Alta, Placer Co; Summerset, El Dorado Co; Strawberry, Alpine Co; and
Sierra City, Sierra Co. Other California counties include Monterey, Ventura,
Alpine, San Luis Obispo, and Del Norte.

Apterodryinus californicus. —Known from Los Angeles and Marsh Or. Spr. in

VOLUME 63, NUMBER 4

391

California and Prescott, Arizona. The flume specimens represent a new county
record and the third locality record for California.

Apterodryinus torvus. —Known from Arizona, Texas, and New York. The flume
specimens represent a new state record for California and the westernmost locality
record for the species. The nearest locality record is approximately 1200 km (700 mi)
to the southeast in Nogales, Arizona.

Bocchus flavipes. —Known from Plumas Co, California and Ormsby Co, Nevada.
The flume specimens represent a new county record, the second locality record for
California, and the southernmost record for the species.

Crovettia theliae. —Known from British Columbia and Ontario, Canada; New
York, Washington D.C., Michigan, Louisiana, Arizona, and California. The flume
specimen represents a new county record and the third locality record for California.
Other California localities include Glenville, Kern Co and lone, Amador Co.

Deinodryinus atriventris. —Widely distributed in North America. The flume
specimens represent a new county record and the third and northernmost locality
record for California. Other California counties include San Bernardino and San
Diego.

Esagonatopus niger .—Known from Ontario Canada, North Dakota, Iowa,
Kentucky, Pennsylvania, and Mexico. The flume specimens represent a new state
record for California. The nearest locality records are approximately 2200 km (2000
mi) to the northeast in Ames, Iowa and 4300 km (2600 mi) to the southeast in
Cuernavaca, Mexico.

Gonatopus agropyrus. —Known from southeastern Canada, Texas, Wisconsin,
Kentucky, Iowa, Georgia, New York, and Arizona. The flume specimens represent
a new state record for California. The nearest locality record is approximately 1200
km (700 mi) to the southeast in Sunnyside, Arizona.

Gonatopus cyphonotus. —Known from Florida and throughout southern Canada.
Two females from Cushing, Morrison County, Minnesota (VII-24-1983, P. S.
Simpson and J. A. Halstead, sweeping grassy vegetation near marsh) represent a
new state record for Minnesota.

Gonatopus herbarum. —Known from Arizona, Texas, and three California
localities (Summerset, El Dorado Co; San Bernardino Co; and Apple Valley, San
Bernardino Co). The flume specimens represent a new county record and the fourth
locality record for California, coming from the central region of the state in the
southern Sierra Nevada.

Gonatopus mayori. —The flume specimen represents a new country record. Olmi
(1984) incorrectly listed the type locality and distribution as El Mayor (California,
U. S. A.). A review of the holotype, deposited in the California Academy of Sciences,
San Francisco, shows the type locality to be “El Mayor, L. California.” The
distribution should therefore read, “Nearctic region: U.S.A.: Ash Mtn. (Tulare
County, California) MEXICO: El Mayor (Baja California Norte).”

Gonatopus mimoides. —Known from Utah, Arizona and Texas. Five females and
one male from 3 mi SE Madera, Madera Co, California (VII-29-1985, J. A.
Halstead, sweeping grass in pistachio orchard) represents a new state record for
California and a westward range extension of approximately 1300 km (800 mi).

Gonatopus pallidiceps. —Known from Nova Scotia and British Columbia Canada,
California, North Dakota, Kansas, Tennessee, and Florida. One female from 3 mi SE
Madera, Madera Co, California (VII-29-1985, J. A. Halstead, sweeping grass in

392

PAN-PACIFIC ENTOMOLOGIST

pistachio orchard) represents a new county record and the third locality record for
California. The two other California records are from Alameda Co.

Gonatopus paraleptias. —Known from Arizona, Wyoming, and five localities in
California. The flume specimens represent a new county record and the first record
for the Sierra Nevada. Other California counties include Riverside, Los Angeles,
Santa Clara, and Marin.

Gonatopus portalensis. —Recently described from Sagehen Creek, Nevada Co
and Omira, Lassen Co, California; Portal, Arizona; Ward, Colorado; and Kerrville,
Texas (Olmi 1984). Three females from Hopkins Well, 18 miles West of Blythe,
Riverside County, California (X-4-1984, N. J. Smith) represent a new county record,
and the southernmost and the third locality record for California.

Lonchodryinus bakeri. —Widely distributed in North America. The flume
specimens represent a new county record and the third locality record for California.
Also, they represent the first record from the Sierra Nevada. Other California
counties include Siskiyou and Del Norte.

Lonchodryinus flavus. —Recently described from throughout Canada, Michigan,
Georgia, Maryland, New Jersey, New York, and New Hampshire (Olmi 1984). The
flume specimen represents a new state record for California. The nearest locality
record is approximately 2333 km (1400 mi) to the northeast in Alberta, Canada.

Lonchodryinus masneri. —Recently described from two localities near Portal,
Arizona and Culberson Co, Texas (Olmi 1984). The flume specimens represent a
new state record for California and a northwestern range extension of approximately
1200 km (700 mi). Also, one female was collected from a Gypsy Moth Trap at
Clough’s Cave, Tulare Co, California (I-XII-1985, R. D. Haines).

Pseudo gonatopus sjoestedti. —Known from Texas, Ohio, Kentucky, North
Carolina, and New York. The flume specimens represent a new state record for
California and the westernmost locality record for the species. The nearest locality
record is approximately 1700 km (1000 mi) to the southeast in Texas.

Tetrodontochelys peculiaris. —Known from Manitoba and Ontario Canada,
Arizona, Texas, Iowa, South Carolina, and one locality in California (Stanford
University, Marin Co). The flume specimens represent a new county record and the
second locality record for California.

Tetrodontochelys unicus. —Known from throughout southern Canada,
Washington, Idaho, Arizona, Texas, North Dakota, South Dakota, and Minnesota.
The flume specimens represent a new state record for California. The nearest locality
record is approximately 1200 km (700 mi) to the southeast in Nogales, Arizona.

In summary, this data improves the distribution information for several species,
depicts their habitat and seasonality, indicates how common or rare each species is
(flume collectability), and shows that a large diversity of species occur at the Kaweah
flume location.

In all, 12 genera and 23 species of dryinid wasps were collected from the flume.
This material contains 3 genera and 12 species (including 2 new species) which were
not previously recorded from California. Additionally, G. mimoides is recorded for
the first time in California. Olmi (1984) listed 9 genera and 21 species from
California. The California fauna is now known to contain 12 genera and 34 species.
It’s astonishing to note that approximately 68 percent of the species of California’s
dryinid wasp fauna has been collected at a single locality, the Kaweah flume.

As denoted in Table 1, most adult wasp activity occurred at the flume in June and
July though various species were collected between March and October. Despite the

VOLUME 63, NUMBER 4

393

many collecting trips and man-hours of collecting, most species were rarely
collected. We feel that without the use of the great collecting ability of the flume and
our intense collecting effort, many species would have gone undetected. Also, we
feel that flumes collect a better representation of an area’s fauna, at least at the
Kaweah locality, than do other collecting techniques such as Malaise traps and
screen sweeping.

In addition to improving the knowledge of the Nearctic Dryinidae and especially
that of California, this paper illustrates that flumes can be an important collecting
source and we hope that it encourages other researchers to utilize them.

Acknowledgments

We thank M. Olmi, Universita Degli Studi Della Tuscia, Viterbo, Italy for
confirming and determining our dryinid material; D. J. Burdick, California State
University Fresno, Fresno for collecting assistance and for reviewing this manuscript
and N. J. Smith, Fresno County Agricultural Commissioner’s Office, Fresno,
California for editorial comments.

Literature Cited

Krombein, K. V. 1979. Dryinidae, pp. 860-874. In: Krombein, K. V. etal., eds.. Catalog of
Hymenoptera in American North of Mexico. Vol. I. Smith. Instit. Press. Wash., D.C. 1198 pp.
Olmi, M. 1984. A revision of the Dryinidae (Hymenoptera). Mem. Amer. Entomol. Instit. No. 37:1913

pp.

1987 SPONSORING MEMBERS
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PAN-PACIFIC ENTOMOLOGIST
63(4), 1987, pp. 395-396

A New Species of Mordellistena from North America
(Coleoptera: Mordellidae)

N. M, Downie

505 Lingle Terrace, Lafayette, IN 47901

Abstract .—A new species of Mordellistena Costa, one of the few very small ones
found in North America north of Mexico, Mordellistena stephani Downie, is
described and named.

While working over a box of mordellids for Karl Stephan, a series of specimens
that was apparently new was observed. This new species is described below.

Mordellistena stephani Downie, new species

Length to tip of anal style 1.5 mm; body shape typical of the genus; body black,
head and pronotum reddish-brown to yellowish; antennae yellow, becoming darker
distally; pronotum with the basal third or so sometimes clouded with piceous; elytra
with two transverse bars of grayish pubescence, one half way between base and
middle and the other half way between middle and apex; venter black with the last
two abdominal sternites and the style reddish yellow; legs testaceous with the
metafemora darker; pubescence, except for that on the elytral bars, fine, indistinct,
the venter and legs with longer, yellow, rather dense pubescence.

Discussion .—In this species the ridges of the hind legs are poorly developed. In
those specimens in which the ridges are most distinct, there are two ridges on the
metatibiae, two on the first metatarsal segment and one on the second. These ridges
would cause this species to key to item 17, page 70 of Liljeblad’s (1945) key to the
genus. Four species key out here, all larger than this species and differing in overall
habitus. Neither Khalaf, 1971a, 1971b, nor Ray, 1937, 1944, 1946a, 1946b, 1946c,
1947, contain material relevant to this species. In all specimens, the posterior,
transverse pubescent band is distinct, whereas the anterior one is variable in the
amount of development.

Specimens examined 14. Holotype, Latimer Co., OK., 5 miles west Red Oak,
May 1981, K. Stephan, deposited in the U.S. National Museum; paratypes: OK.,
Latimer Co., Nov. 1983; 2 specimens, OK., Latimer Co., 5 miles west Red Oak, May
1981, K. Stephan; 3 specimens, OK., Latimer Co., May 1982, K. Stephan; 4
specimens, OK., Latimer Co., May 1983, K. Stephan; 1 specimen, OK., Latimer
Co., June 1983; 2 specimens, OK., Latimer Co., July 1983; 1 specimen, GA.,
Lumpkin Co., 18 miles northwest Dahlonega, July 1982, R. H. Turnbow. The
paratypes are in the collection of the Florida Collection of Arthropods, the
California Academy of Sciences, and the collections of N. M. Downie, K. Stephan,
and R. H. Turnbow, Jr.

395

396

PAN-PACIFIC ENTOMOLOGIST

Little is known about the habits of this species. The Oklahoma specimens were
taken sweeping low vegetation in an upland forest.

References

Khalaf, K. T. 1971a. Mordellidae from Louisiana and Mississippi. Jr. Kansas Entom. Soc. 44:441-447.
Khalaf, K. T. 1971b. Five new species of Mordell. from Louisiana and Mississippi, Pan-Pacific
Entomologist 47:140-151.

Liljeblad, E. 1945. Mono, of the fam. Mordellidae of North America north of Mexico. Misc. Publ. Mus.

Zool. Univ. Michigan, 62, 229 pp., 7 pis. (Actually this was completed in 1929).

Ray, E. 1937. Studies in N. American Mordellidae, I, Canadian Ent. 68:124-129.

Ray, E. 1944. Mordelhd. beetles of West. Hemisphere. Zool. Ser. Field Mus. Nat. Hist., 29 no. 7, pp.
117-133.

Ray, E. 1946a, 1946b, 1946c, 1947. Studies in N. American Mordellidae. II, 1946, Pan-Pac. Entom.
22:41-50; III, 1946, Pan-Pac. Entom. 22:90-99; IV, 1946, Pan-Pac. Entom. 22:121-132; V, 1947,
Pan-Pac. Entom. 23:121-131.

The Pan-Pacific Entomologist
Contents for Volume 63

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

ALEXANDER, B. and J. G. ROZEN, JR.—Ovaries, ovarioles, and oocytes in parasitic bees

(Hymenoptera: Apoidea) . 155

BECKWITH, R. C., R. R. MASON and H. G. PAUL—Parasitism of salticid spiders in Oregon

by two species of Acroceridae (Diptera). 352

BJOSTAD,L.—see O’NEILL, K.M. 207

BOHART, G. E. and T. L. GRISWOLD—A revision of the dufoureine genus Micralictoides

Timberlake (Hymenoptera: Halictidae) . 178

BOHART, R. M.—A key to Trichrysis and new species from Sri Lanka and Africa

(Hymenoptera: Chrysididae) . 347

BOTTORF, R. L.—see SZCZYTKO, S. W. 65

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

saponaria Molina (Rosaceae). 60

BURNE, J. C.—A morphometric analysis of Mordellistena Costa in Southwestern United States

(Coleoptera: Mordellidae) . 224

CANE, J. H.—see PARKER, F. D. 172

CHEMSAK, J. A.—A new Mexican species of Linsleyella Chemsak (Coleoptera:

Cerambycidae) . 145

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

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

DAHLSTEN, D. L.—SEE HAJEK, A. E. 319

DALY, H. V., C. D. MICHENER, J. S. MOURE, and S. F. SAKAGAML—The relictual bee

genus Manuelia and its relation to other Xylocopinae (Hymenoptera: Apoidea). 102

DISNEY, R. H. L.—Four species of scuttle fly (Diptera: Phoridae) from Dominican amber .... 377

DOWNIE, N. M.—A new species of Mordellistena from North America (Coleoptera:

Mordellidae). 395

DOYEN, J. T.—New and little known Tenebrionidae from Central America and Mexico, with

remarks on their classification (Coleoptera). 301

EHLER, L. E.—Ecology of Rhopalomyia californica Felt at Jasper Ridge (Diptera:

Cecidomyiidae) . 237

EVANS, H. E.—Observations on the prey and nests of Podalonia occidentalis Murray

(Hymenoptera: Sphecidae). 130

FERRIS, C. D.—A nematode parasite of Erebia occulta Roos & Kimmich (Nematoda;

Lepidoptera: Satyridae) . 368

FORBES, G. S.—The status of Efferia similis (Williston), with descriptions of three new Nearctic

Efferia species in the albibarbis group (Diptera: Asilidae). 292

FRANKIE, G. W.—see PARKER, F. D. 172

FURNISS, M. M.—see STOCK, M. W. 353

GAMBINO, P.—First records of the German yellow)acket Paravespula germanica (L.) from the

east San Francisco Bay (California, U.S. A.) area. 358

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

GIESBERT, E. F.—A new genus and species in the tribe Macrotomini (Coleoptera:

Cerambycidae) from Costa Rica . 147

GIESBERT, E. F.—A new genus and two new species of longhorn beetles (Coleoptera:

Cerambycidae) from Mexico and Central America. 359

GOEDEN, R. D.—Life history of Trupanea conjuncta (Adams) on Trixus californica Kellogg in

Southern California (Diptera: Tephritidae) . 284

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

Encyrtidae) . 16

397

398

PAN-PACIFIC ENTOMOLOGIST

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

infestation in the home . 1

GREGOIRE, J. C.—see STOCK, M. W. 353

HAINES, R. D.—see HALSTEAD, J. A. 383

HAINES, R. D.—see HALSTEAD, J. A. 389

HAJEK, A. E. and D. L. DAHLSTEN—The exotic aphids (Homoptera: Drepanosiphidae) on

ornamental birch in northern California. 319

HALSTEAD, J. A .—Acanthochalcis nigricans Cameron—new distributional information,

including South America (Hymenoptera: Chalcididae) . 236

HALSTEAD, J. A.—On the rearing of Michrochridium minutum and its probably
host —Ammoplanellus (Ammoplanellus) unmatilla (Hymenoptera; Chrysididae,
Sphecidae). 256

HALSTEAD, J. A. and R. D. HAINES—Flume Collecting: A rediscovered insect collecting

method, with notes on insect extracting techniques. 383

HALSTEAD, J. A. and R. D. HAINES—New distribution records for some Nearctic dryinid
wasps, with a species list from a flume in Tulare County, California (Hymenoptera:
Dryinidae). 389

HALSTEAD, J. A. and C. G. NIWA —Rhyacionia zozana (Lepidoptera: Tortricidae), host of

Hockeria tenuicornis (Hymenoptera: Chalcididae) in Oregon. 276

HOFFMAN, M. P., L. T. WILSON, and F. G. ZALOM—The southern green stink bug, Nezara

viridula Linnaeus (Heteroptera: Pentatomidae); New location. 333

HOVORE, F. T.—A new genus and species of Cerambycidae from Costa Rica (Coleoptera) ... 151

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

(Coleoptera), study number three. 50

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

Asia (Diptera: Tipulidae) . 92

HYNES, C. D.—A new species and records of the genus Toxorhina, subgenus Eutoxorhina from

the South Paeific (Tipulidae: Diptera). 335

JOHNSON, C. D.—see LUCKOW, M. 48

JOHNSON, J. B. and T. D. MILLER —Heimbra opaca (Ashmead) (Hymenoptera:

Eurytomidae) discovered in Idaho: A new state record for the subfamily Heimbrinae .... 324

JOHNSON, P. J .—Pteroloma nebrioides Brown in Idaho (Coleoptera: Agyrtidae). 381

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

Hymenoptera) . 56

KROMBEIN, K. V.—Biosystematic studies of Ceylonese wasps, XVIII: The species of

Trachepyris Kieffer (Hymenoptera: Bethylidae: Epyrinae). 135

LANHAM, U. N.—A new species of Andrena at the Micrandrena-Scaphandrena boundary

(Hymenoptera: Apoidea) . 325

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

Desmanthus (Leguminosae) from Texas and Mexico . 48

MASON, R. R.—see BECKWITH, R. C. 352

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

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

MICHENER, C. D.—see DALY, H. V. 102

MILLER, T. D.—see JOHNSON, J. B. 324

MOURE, J. S.—see DALY, H. V. 102

NADEL, H.—Male swarms discovered in Chalcidoidea (Hymenoptera: Encyrtidae,

Pteromalidae). 242

NIWA, C. G.—see HALSTEAD, J. A. 276

O’NEILL, K. M. and L. BJOSTAD—The male mating strategy of the bee Nomia nevadensis

(Hymenoptera: Halictidae): Leg structure and mate guarding . 207

PARKER, F. D.—Nests of Callanthidium from block traps (Hymenoptera: Megachilidae). 125

VOLUME 63, NUMBER 4 399

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

PINTO, J. D. and G. BIGGIANI—Two new Trichogrammatidae (Hymenoptera) from North
America: Ittysella lagunera Pinto and Viggiani (N. Gen., N. Sp.) and Epoligosita mexicana
Viggiani (N. Sp.) . 371

POLHEMUS, D. A .—Heleocoris brasiliensis De Carlo is a Ctenipocoris (Hemiptera:

Naucoridae) . 370

POLHEMUS, D. A. and J. T. POLHEMUS—A new genus of Naucoridae (Hemiptera) from the

Philippines, with comments on zoogeography . 265

POLHEMUS, J. T.—see POLHEMUS, D. A. 265

Proceedings of the Pacific Coast Entomological Society, 1986 . 199

Publications received. 150, 328, 334, 338,346

RESH, V. H.—see McELRAVEY, E. P. 75

ROZEN, J. G., JR.—see ALEXANDER, B. 155

SAKAGAMI, S. F.—see DALY, H. V. 102

SCHMIDT, J. O.—see STARR, C. K. 37

SCHMIDT, P. J.—see STARR, C. K. 37

SCUDDER, G. G. E.—A review of the genus Nosostethus Kirkaldy (Hemiptera: Lygaeaidae:

Lygaeinae) . 270

SINGER, M. C.—see WHITE, R. R. 341

SNELLING, R. R.—A revision of the bee genus Aztecanthidium (Hymenoptera: Megachilidae) 165

SNELLING, R. R.—Geographical inexactitude . 339

SPIETH, H. T.—The Drosophila fauna of a native California forest (Diptera: Drosophilidae) .. 247

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

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

STOCK, M. W., J-C GREGOIRE and M. M. FURNISS—Electrophoretic comparison of
European Dendroctonus micans and ten North American Dendroctonus species
(Coleoptera: Scolytidae). 353

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

of western Nearctic Isoperlinae (Plecoptera: Perlodidae) . 65

THORP, R. W.—A new species of Andrena (Onagrandrena) from Utah’s San Rafael Desert

(Hymenoptera: Andrenidae) . 194

TORGERSEN, T. R.—see WICKMAN, B. E. 218

VIGGIANI, G. —see PINTO, J. D. 371

VINSON, S. B.—see PARKER, F. D. 172

WAGNER, D. L.—A new Microcalyptris species from California (Lepidoptera: Nepticulidae) . 278

WALKER, G. P.—Probing behavior of Aphis helianthi (Homoptera: Aphididae) and its

preference for Pittosporum tobira leaves of different ages . 258

WHITE, R. R. and M. C. SINGER—Marking technique for larvae. 341

WICKMAN, B. E. and T. R. TORGERSEN—Phenology of Douglas-fir tussock moth Orgyia
pseudotsugata, egg ecolosion and mortality in a thinned and unthinned stand (Lepidoptera:
Lymantriidae). 218

WILLIAMS, S. C.—A new species of Paruroctonus from coastal California (Scorpiones:

Vaejovidae) . 329

WILLIAMS, S. C.—Lectotype designations and redescription of Vejovis wupatkiensis Stahnke

(Scorpiones: Vaejovidae) . 363

WILSON, L. T.—see HOFFMAN, M. P. 333

WOOD, D. L.—see GRACE, J. K. 1

ZALOM, F. G.—see HOFFMAN, M. G. 333

The Pan-Pacific Entomologist
Index to Volume 63
(New taxa in boldface)

Acanthochalcis nigricans, 236
Acanthoscelides compressicornis, 48;

pectoralis, 49; desmanthi, 49
Acroceridae, 352
Actaletes nemyops, 52
Actaletidae, 52
Agyrtidae, 381
Ammoplanellus umatilla, 256
Andrena (Micrandrena) robinsoni, 325
Andrena (Onagrandrena) linsleyana,
194

Andrenidae, 194
Aphididae, 258
Aphis helianthi, 258
Apis dorsata, 37
Apoidea, 155, 325
Asilidae, 293

Aztecanthidium, 165; cuauhtemocum,
166; tenochtitlanicum, 166;
xochipillium, 170

Bethylidae, 135

Bothynocephalus, 315; cristatus, 316
Bruchidae, 48

Callanthidium, 125
Cecidomyiidae, 237
Cerambycidae, 43,145,147,151, 359
Chalcididae, 236, 276
Chrysididae, 56, 256, 347
Cleptes asianus, 56; canadensis, 56;

townesi, 58
Cleptoparasitosis, 1
Cnephalura, 313; umbrata, 313
Coleoptera, 43,48,50,145,147,151,
224,301,353,359,381,395
Collembola, 52
Corrigenda, 205

Cosumnoperla, 66; hypocrena, 66
Ctenipocoris brasiliensis, 370

Dasytidae, 50
Dendroctonus, 353
Desmanthus, 48

Diptera, 5, 92, 237, 247, 284, 292, 335,
352,377

Dohrniphora poinari, 377
Dolichostelis louisae, 173; costaricensis,
173; rudbeckiarum, 174
Drepanosiphidae, 319
Drosophila, 247
Drosophilidae, 247
Dryinidae, 389

Efferia neosimilis, 293; incognita, 296;

sonorensis, 298
Encyrtidae, 242
Epoligosita mexicana, 374
Erebia occulata, 368
Eurytomidae, 324

Financial statement, 203
Flume collecting, 383, 389

Gortonia, 151; linsleyi, 152

Haematopota abacis, 6;

albofasciatipennis, 6; assamensis, 7;
bealesi, 7; bicolor, 10; cilipes, 10;
cynthiae, 10; excipula, 11, gobindai,
12; howarthi, 12; pachycera, 12;
singularis, 13; splendens, 13; vimoli,
13

Halictidae, 207
Heimbra opaca, 324
Hemiptera, 265, 270, 370
Heteroptera, 333
Hippocentrodes desmotes, 5
Hockeria tenuicornis, 276
Homoptera, 258, 319
Hymenoptera, 16, 37, 56,102,125,130,
135,155,165,172,178,194, 207,

236, 242, 256, 276, 324, 325, 347,
358,371,389

Isaminas, 301; breedlovei, 303;
reticuloides, 303; gibbipennis, 305;
brevicollis, 305; sullivani, 306;
erotyloides, 308

400

VOLUME 63, NUMBER 4

401

Ittysella, 371; lagunera, 372

Lepidoptera, 218,276, 278, 368
Linsley, E. Gorton, 100
Linsleyella virgulata, 145
Lygaeaidae, 270
Lymantriidae, 218

Manuelia, 102’, gayatina, llA’,postica,
116’, gayi, 118

Megachilidae, 125,165,172
Megaselia amberae, 378; ernsteinae, 378;
dominicana, 378

Meromyzobia, 16; americana, 22;
bifasciata, 24; deserticola, 2A;flava,
26; flavincincta, 26; maculipennis, 21;
melanosoma, 28; pedicelata, 30;
texana, 33

Micralictoides, 118; chaenactidis, 181;
altadenae, 185; linsleyi, 186;
quadriceps, 186; grossus, 187;
ruficaudus, 189; mojavensis, 190;
dinoceps, 192
Microcalyptris lotella, 218
Microchridium minutum, 256
Mordellidae, 224, 395
Mordellistena, 224, 395; stephani, 395

Naucoridae, 265
Nematode parasite, 368
Nepticulidae, 278

Nesostethus, 270; bipartitus, 212;fuscus,
212; lunatus, 213; niger, 213; ornatus,
21 A; variegatus, 21A
Nezara viridula, 333
Nomia nevadensis, 207

Orgyia pseudotsugata, 218

Pachymerola, 43; vitticollis, 44;

ruficollis, 44; humeralis, A1
Parastrongylaspis, 147; linsleyi, 148
Paravespula germanica, 358
Paruroctonus maritimus, 329
Pentatomidae, 333
Perlodidae, 65

Philippinocoris, 265; usingeri, 268
Phoridae, 377
Plecoptera, 65

Podalonia occidentalis, 130
Proceedings of the Pacific Coast
Entomological Society, 1986,199
Pteroloma nebrioides, 381
Pteromalidae, 242

Quillaja saponaria, 60

Rhopalomyia californica, 231
Rhyacionia zozana, 216

Salticidae, 352
Satyridae, 368

Saziches, 308; subcaudatus, 310;

giesberti, 310
Scolytidae, 353
Scorpiones, 329, 362
Sphecidae, 130,256
Statorpruininus, 49

Styringomyia bidentata, 92; digitostylus,
93; rostrostylus, 93; vietnamensis, 94;
ysabellae, 94; dilinhi, 95 ; labuanae,

96; idioformosa, 96

Tabanidae, 5
Tenebrionidae, 301
Tephritidae, 284
Tipulidae, 92, 335
Tortricidae, 276

Toxorhina (Eutoxorhina) simplex, 335;
parasimplex, 335

Trachepyris, 135; haemorrhoidalis, 138;

indicus, lAO; spinosipes, 142
Trichogrammatidae, 371
Trichoptera, 75

Trichrysis eardleyi, 350; hexapholis, 350;
lomholdti, 351

Tropimerus, 359; cyaneus, 360; hovorei,
360

Trupanea conjuncta, 284

Vaejovidae, 329, 363
Vejovis wupatkiensis, 363

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