Vol. 49
JANUARY 1973
No. 1
THE
Pan-Pacific Entomologist
PARSONS—The Lagriidae of California (Coleoptera) ___ 1
TURNER AND CHILLCOTT—Four new species of the Symphoromyia
pachyceras complex from California (Diptera: Rhagionidae) _ 5
GILBERT—Foraging behavior of Trigona fulviventris in Costa Rica
(Hymenoptera: Apidae) _ 21
PAETZEL—Behavior of the male Trypoxylon rubrocinctum (Hymenoptera:
Sphecidae) _ 26
CHANDLER—A revision of the New World genus Formicilla (Coleoptera:
Anthicidae) _ 31
MUCHMORE—A new genus of pseudoscorpions based upon Atemnus hirsutus
(Pseudoscorpionida: Chernetidae) _ 43
EIGHME—A new Pulverro from California (Hymenoptera: Sphecidae) . 49
GORDH—Description of a new species of Trjapitzinellus parasitic on
Coniopterygidae in California (Hymenoptera: Encyrtidae; Neuroptera) 51
FOSTER AND ANTONELLI—Larval description and notes on the biology of
Anthocomus horni (Coleoptera: Melyridae) _ 56
McCREA—A new species of the flea beetle genus Chaetocnema found on
dichondra in California (Coleoptera: Chrysomelidae) _ 61
PETERMAN—Possible behavioral thermoregulation in Tanarthrus salinus and
T. inyo (Coleoptera: Anthicidae) _ 67
EMMEL, TREW, AND SHIELDS—Chromosomal variability in a Nearctic
Lycaenid butterfly, Philotes sonorensis (Lepidoptera: Lycaenidae) _ 74
CORRECTION _ 4
EDITORIAL NOTICE _ 25
SCIENTIFIC NOTES_ 42, 59, 60, 80
COLLECTION DONATED_ 48
BOOK REVIEW _ 55
ZOOLOGICAL NOMENCLATURE _ 73
PROCEEDINGS _ 81
SAN FRANCISCO, CALIFORNIA • 1973
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
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The Pan-Pacific Entomologist
Vol. 49
January 1973
No. 1
The Lagriiclae of California
(Coleoptera)
Carl T. Parsons
Manchester Depot, Vermont 05256
In 1965 I published an account of North American Statira in which
the Lagriidae of California were represented only by an unnamed
unique labelled merely “S. Cal.” Now there are before me three more
species two of which are undescribed. Much more material for study
is urgently needed from both California and Baja California, Mexico.
Key to California Statira
1. At least middle and posterior tibiae sulcate along nearly entire outer edge.
Disc of pronotum irregularly rugulose, not punctate. Third elytral interval
with more than eight setigerous punctures . 2
Tibiae on outer edge rounded to nearly flattened towards apex. Disc of
pronotum obsoletely rugulose or finely granular and sparsely punctate.
Third elytral interval with less than eight setigerous punctures . 3
2. Elytra and pronotum dark brown to piceous with some specimens having
pronota tending to dark rufous. Apical antennal segment of female equal
to next 2.5 segments. First elytral interval with 23-33 setigerous punctures,
third with 26-39, fifth with 21-28, seventh with 17-26, and ninth with
18-30 setigerous punctures . dumalis Parsons, n. sp.
Elytra usually dark piceous and feebly shining, pronotum dark rufous and
subopaque. Apical antennal segment of female equal to next 2.75 segments.
First elytral interval with 11-16 setigerous punctures, third with 14-21,
fifth with 12-20, seventh with 10-17, and ninth with 12-20 setigerous
punctures . pluripunctata Horn
3. All femora finely sparsely pubescent. Pronotal lateral margins almost always
obliterated anteriorly but becoming distinct posteriorly. Surface of
pronotum finely obsoletely rugulose. Female apical antennal segment equal
to next 2.25 segments. First elytral interval wtih 2-3 setigerous punctures,
third with 1-3, fifth with 1-3, seventh with 1-2, and ninth with 2-3
setigerous punctures . latitator Parsons n. sp.
No alternate.
Statira dumalis Parsons, new species
Female. —Uniformly dark brown above with underside and legs slightly paler.
Vertex densely finely rugulose, as on disc of pronotum. Apical antennal segment
The Pan-Pacific Entomologist 49: 1-4. January 1973
2
The Pan-Pacific Entomologist
as long as tenth, ninth, and one-half of eighth segments. Prothorax with width:
length as 1:1.05, anterior margin feebly emarginate. Scutellum very finely obsoletely
punctate and transversely alutaceous. Elytra along inner halves somewhat flattened;
striate, with closely placed punctures along striae, intervals finely granular,
moderately convex; setigerous punctures about twice as large as strial punctures.
First elytral interval with 25 setigerous punctures, second interval with one at
base, third interval with 29, fifth interval with 22, seventh interval with 17, and
ninth interval with 25 setigerous punctures. All femora finely sparsely pubescent;
all tibiae sulcate along outer edge. Length 9.8 mm, width of pro thorax 1.7 mm,
width of elytra at humeri 2.5 mm.
Paratypes.—Color may be piceous and anterior prothoracic margin truncate in
two examples. First elytral interval may have 23-33 setigerous punctures, third
interval with 26-39, fifth interval with 21-28, seventh interval with 18-26, and
ninth interval with 18-30 setigerous punctures. Length 9.5-11.2 mm, width of
prothorax 1.6-2.0 mm, width of elytra at humeri 2.4-3.1 mm.
M ale .—Unknown.
Holotype female, Imperial County, California, July 1912, J. C.
Bridwell, E. C. Van Dyke. Known only from the type and seven females
taken in southeastern California in eastern Riverside County and
Imperial County. Paratypes: two same data as holotype; one, Holtville,
23 June 1946: three, Coachella, Riverside Co., 24 May 1928, E. C.
Van Dyke; one, Laguna Lake, Imperial Co., 9-11 June 1950. Holotype
and five paratypes are in the California Academy of Sciences, one
paratype is in the Los Angeles County Museum and one is in the writer’s
collection.
This species agrees with S. subnitida in lateral pronotal line, color,
female apical antennal segment, but has many more setigerous punctures
and the pronotal surface much more rugulose and not granular. Also
S. dumalis is related to S. pluripunctata differing as follows. Disc of
pronotum more flattened than in S. pluripunctata or S. subnitida, about
as convex as in S. dejecta. Pronotal surface slightly more coarsely
rugulose than in 5. pluripunctata but distinctly more finely rugulose
than in S. hirsuta. Elytra along inner halves somewhat flattened as in
S. dejecta, less convex than in S. pluripunctata, intervals about as convex
as in S. pluripunctata.
Statira pluripunctata Horn
Statirci pluripunctata Horn, 1888, 15:29.
This species occurs in Utah, Arizona (west to Ajo Mts.), New Mexico,
western Texas, and Mexico. Also there is at Ohio State University one
typical S. pluripunctata collected 1 August 1940 in Yosemite National
Park by D. J. and J. N. Knull. Professor Knull (personal communica¬
tion) states, “We must assume that the locality is correct. We were in
Vol. 49, No. 1, January 1973
3
1 )nm
Figs. 1, 2. Lateral and dorsal views of the aedeagus of Statira latitator Parsons
n. sp.
a crowded camping; area in Yosemite and it is possible that the specimen
could have been carried by one of the automobiles.” This record needs
confirmation.
Statira latitator Parsons, new species
Male.— Appearing teneral; testaceous, head and antennae fuscous, basal 6-8
antennal segments tending to piceous. Elytra dusky laterally and posteriorly.
Vertex very narrow, as wide as width of first antennal segment, surface densely
rugulose. Apical antennal segment as long as tenth, ninth, and three-fifths of
eighth segments. Prothorax with width:length as 1:1.06; anterior margin truncate.
Surface of pro thorax very finely obsoletely rugulose; prothoracic lateral line
obliterated anteriorly becoming distinct posteriorly. Scutellum very finely alu-
taceous and punctate, laterally and posteriorly with a smoother raised margin.
Elytra striate with punctures closely placed along striae; intervals finely granular
and moderately convex; setigerous punctures about twice as large as strial punc¬
tures. First elytral interval with one setigerous puncture at base and one at apex,
third with three at apex, fifth with two at base, seventh with two at base, and
ninth with three at apex. All femora very finely and sparsely pubescent; all tibiae
rounded on outer edge. Aedeagus with unique pair of recurved dorsal spines
(Figs. 1, 2). Length 8.4 mm, width of prothorax 1.4 mm, width of elytra at
humeri 2.0 mm.
Female. —As for male except vertex as wide as tenth antennal segment; apical
antennal segment as long as tenth, ninth, and one-fourth of eighth segments;
prothorax with width:length as 1:.97; third elytral interval with one setigerous
puncture at base and one at apex, fifth with one at base, and ninth with two at
apex. Length 9.5 mm, width of prothorax 1.5 mm, width of elytra at humeri 2.3 mm.
Paratypes.—Some specimens darker; occasionally the anterior pronotal margin
feebly emarginate; rarely lateral pronotal line extends very finely to anterior
angles; first elytral interval with 0-1 setigerous puncture at base and 1-3 at
4
The Pan-Pacific Entomologist
apex, third with 0-2 at base, 1-3 at apex, fifth with 1-2 at base, 0-1 near apex,
seventh with 1—2 at base, and ninth with 2-3 at apex; length 7.5-9.5 mm.
Holotype male and allotype, Deep Canyon, Riverside County, Cali¬
fornia collected at light 13 June 1963 E. I. Schlinger. Paratypes: 49
from type locality in 1963, 13 June (36), 16 May (2), 30 May (1),
20 June (1), 22 June (8), and 8 August (1) ; also 24 May 1964, M. E.
Irwin (1) ; and one 10 April 1961, 1.5 miles north of Puertocito, on the
gulf coast of Baja California, Mexico, A. G. Smith. Holotype and
allotype in the California Academy of Sciences, paratypes in the
Entomological Museum, University of California at Riverside, except
for one pair in the writer’s collection and a male in the California
Academy of Sciences.
This species has the appearance of S. opacicollis which differs in
having an entire lateral pronotal line, much longer apical antennal seg¬
ment, more numerous setigerous elytral punctures, and different
aedeagus. The prothorax, elytra, and elytral intervals are about as
convex as in S. pluripunctata.
Literature Cited
Horn, G. 1888. Miscellaneous Coleopterous Studies. Trans. Amer. Entomol. Soc.,
15: 29.
Parsons, C. T. 1965. A Key to North American Statira (Coleoptera: Lagriidae).
Psyche, 72: 241-254.
CORRECTION
Emmel, T. C. and J. F. Emmel. 1971. A new subspecies of Papilio indrci from
central Nevada (Lepidoptera: Papilionidae). Pan-Pac. Entomol., 47: 220-223.
On page 221, lines 3 through 11, the data for the types should read as follows:
Holotype male, Jett Canyon, 6,600 feet elevation, east side of Toiyabe
Range, Nye County, Nevada, 3 August 1967, John F. Emmel.
Paratypes. —11 males and 9 females. Data as follows (all localities east side
of Toiyabe Range, Nye County, Nevada) : 9 males, 5 females, Jett Canyon, 6,500-
6,800 feet elevation, 3 August 1967, John F. Emmel, Oakley Shields, and Scott Ellis;
2 males, 2 females, Summit Canyon, 7,000-7,200 feet elevation, 30 June 1968, John
F. Emmel and Oakley Shields; 2 females, reared from ova laid by a female collected
in Summit Canyon, 7,000 feet elevation, on 30 June 1968, and reared by Chris
Henne on Tauschia parishii C. and R. at Pearblossom, California, emerged 30 May
1969, and 29 May 1969.— Thomas C. Emmel and John F. Emmel, University of
Florida, Gainesville 32601.
Vol. 49, No. 1, January 1973
5
Four New Species of the Symphoromyia pachyceras
Complex from California 1
(Diptera: Rhagionidae)
William J. Turner and J. G. Chillcott 2
Department of Entomology, Washington State University, Pullman
The Nearctic species of biting snipe flies of the genus Symphoromyia,
last revised by Aldrich (1915), remain poorly known. On morphological
bases, including the structure of the male genitalia, the species can be
divided into several distinct groups or complexes (Turner, 1971). One
group, the pachyceras complex, was found to contain several new
entities; four of these, all from northern California and closely related
to Symphoromyia pachyceras Williston, are described herein. Other
described species contained within this group include: S. inquisitor
Aldrich, S. plumbea Aldrich and S. comata Bigot. Since the original
description by Williston (1886), however, the name Symphoromyia
pachyceras has been incorrectly applied to nearly all the species now
included in the complex.
These four new species are being described to make the names avail¬
able for several biological studies by J. B. Hoy and J. R. Anderson.
While the late J. G. Chillcott had begun work on the pachyceras complex,
his untimely death prevented the results of his initial efforts from
reaching publication. His influence on this present work cannot be
overemphasized. My appreciation goes to Drs. James B. Hoy and
Herbert J. Teskey for making Chillcott’s working notes and files avail¬
able for use during this study.
Symphoromyia pachyceras Complex Diagnosis
Color essentially black, surface often dusted with gray or brown pollen, third
antennal segment convex or straight beneath arista, usually rounded above as
much as beneath in lateral view, parafacial area sparsely to densely haired in both
sexes, halter dark (yellow only in S. inquisitor) . Males with eyes narrowly separated
forward of ocellar prominence, abdominal segments I-IV often subparallel, seg¬
ments V and following becoming noticeably constricted, segment IV with short,
stiff hairs posterolaterally, abdominal tergite IX with laterally compressed process
medially on posterior margin, dististylus elongate, slightly elbowed or curved, often
with a ventral lobe. Females dull and densely gray pollinose on head and body with
each abdominal tergite bearing dark median spot or, in some species, abdomen
1 Scientific paper 3810, College of Agriculture, Washington State University. Work was conducted
under Project 9043.
2 Entomology Research Institute, Canada Department of Agriculture, Ottawa, Ontario, Canada.
Deceased, 1967.
The Pan-Pacific Entomologist 49: 5-20. January 1973
6
The Pan-Pacific Entomologist
subshiny, only feebly pollinose and without markings, minimum distance between
eyes from one-third to two-fifths head width.
Terminology used for morphological structures and surfaces essen¬
tially those of Grimshaw (1905), Crampton (1942) and Turner (1971).
Symphoromyia cervivora Turner and Chillcott, new species
Males of this large species are recognized by long postocular hair
dorsally which is about one-half length of hair on ocellar prominence;
humeral and laterotergal hair mostly black, rarely white; wing mem¬
brane nearly clear and hyaline; dististylus with broad, quadrangular
basiventral process and short apical arm. Females have black palpi,
laterally restricted parafacial hair, abdominal tergites each with a
median brown spot surrounded by dark gray or bluish gray pollinosity,
mesoscutal vittae brown, faint.
Holotype male. —Length: 7.6 mm. Wing length: 5.9 mm. Head and its
appendages, thorax, legs and abdomen black; femur I reddish yellow narrowly at
apex, femora II, III similar but less so; halter knob dark brown, stem lighter. Sur¬
face subshiny, minutely rugulose at high magnification (45 X), lightly dusted with
gray or bluish gray pollen, most dense on antennal scape, behind eyes, on meso-
scutum forward and at sides, on proepimeron, proepisternum, meron, metepisternum,
metepimeron, coxae and abdominal sternites, less dense and noticeably more shiny
on remainder of head, thoracic pleura, femora and tibiae; abdominal tergites
sparsely brown pollinose. Hair mostly black, but golden or white on peristomal
sclerites, laterotergite forward, abdominal segments I, II, and on tergite III medially;
tergites I, II with black hair along posterior margin; mixed black and white hair
present on anterior pronotum and coxa III. Head: Eyes separated above by dis¬
tance slightly less than width of median ocellus; interocellar distance subequal;
antennal scape elongate, swollen medially beneath, tapering forward, less so
basally, height three-fourths length, greater than height of segment III, hair dense,
length beneath subequal to segmental length, longer above; segment III straight
beneath arista, forming right angle with dorsal margin about base of arista; palpus
elongate, segment III subcylindrical, blunt apically, barely arcuate postmedially,
length less than twice length of segment I, bearing dorsal subapical callus, hair
dense; labrum shorter than palpus; postocular hair dorsally one-half length of
hair on ocellar prominence; parafacial area densely haired, length of hair subequal
to that of antennal segment I and to suberect hair of ocellar prominence. Thorax:
Mesoscutal vittae black, broad, prominently outlined by wide, dense, bluish gray
pollinose lines, confluent on postscutum; median vitta bisected by thin, brownish
gray pollinose line; lateral vitta complete medially, incomplete forward. Wing
membrane nearly clear, gray hyaline, only very faintly infuscated brown about
veins apically, stigma dark brown; veins dark brown towards apex, yellow basally;
costal vein with row of black, erect hairs basally beneath. Tibia III with long,
suberect hair on basal one-half of posteroventral surface, length subequal to tibial
thickness, shorter, less erect hairs present on other surfaces. Abdomen: From
above, segments I-IY subparallel laterally, segments V and following greatly com¬
pressed; on sternite IV with broad, rounded posterolateral expansion bearing stiff
Vol. 49, No. 1, January 1973
7
0.3mm
Figs. 1-4. Male IX tergites, dorsal and left side, lateral views: Fig. 1,
Symphoromyia cervivorcr, Fig. 2, S. truncata ; Fig. 3, S. nana\ Fig. 4, S. pachyceras
Williston.
The Pan-Pacific Entomologist
hairs; tergite IX apically on posterior margin with laterally compressed process
appearing rounded in lateral view (Fig. 1) ; dististylus bearing quadrangular
basiventral lobe and short distal arm (Fig. 6) ; cercus elongate, narrow and
attenuate, apically acute.
Female. —Length: 6.7-9.2 mm. Wing length: 5.9-6.6 mm. Black color ob¬
scured by dense pollinosity; femur I narrowly at apex and humerus yellowish red,
femur II, III similar hut less so; halter knob dark brown to black, stem lighter.
Surface typically dull, opaque with almost dense gray or bluish gray pollen overall,
becoming less dense on femora and tibiae and often on clypeus, thoracic pleura,
and abdominal tergites; upper frons area, head behind and scutellum often lightly
tinted with brown pollen, scutal vittae and abdominal terga similar but darker;
tergites II and behind each marked with dark brown along front and rear margins
and medially by a separate, more or less distinct spot surrounded by dense, pale
gray, bluish gray, or dark gray pollen, replaced laterally by dense, gray pollen;
surface often subshiny or dull. Hair mostly white but appearing black on antenna,
parafacialia, head above and behind, palpus apically, mesoscutum except for
humerus and immediately behind, scutellum, femur I, often on mesoanepisternum
posterodorsally, abdominal tergites II-IV and sometimes on tergite V medially and
along rear margin, completely black on segments VI and following; hair mixed
black and white on mesocoxite A. Head : Minimum distance between eyes at
midfrons about two-fifths head width (MFW/HW ratio: 0.39-0.42) ; antennal
segment I either elongate and swollen medially, tapering at ends or shorter, sub¬
truncate forward, tapering slightly to base, height greatest subapically or medially
but varying from nearly three-fourths to almost nine-tenths length, hair sparse,
longest ventrally, subequal to segmental height, shorter above; palpal segment II
highly convex dorsally, less so beneath, height greatest postmedially, tapering to
base, length about twice length of subcylindrical segment I, with short, digitate
apical process; parafacial area with hair virtually restricted to lateral portion, bare
medially or with only one or two hairs; labrum shorter than clypeal height.
Thorax : Mesoscutal vittae pale grayish brown, faint, tapering to rear, separated
by gray or bluish gray pollinose lines; middle vitta becoming dark gray forward,
bisected by narrow, brown pollinose line; lateral vitta incomplete forward, inter¬
rupted or not medially, either bifurcate or appearing as elongate stripe and lateral
spot on postscutum. Wing membrane nearly clear, gray hyaline, stigma infuscated
brown; veins dark brown apically becoming yellow at base; costal vein beneath
at base with mixed golden and black hairs or predominantly yellow haired.
Abdomen : Disticercus elongate, subtruncate or rounded apically, attenuate toward
base.
Holotype male , Hopland Field Station (4 mi. E. Hopland), Uni¬
versity of California, Mendocino County, California, elevation
2,880 feet, 2 May 1969 (W. J. Turner), from male swarm, ca. 1145
P.D.T. CAS type no. 11250. Allotype female, same locality and date
as type.
Paratypes, 188 males, 126 females. CALIFORNIA: 43 $, 52 $, same locality
as type (all W. J. Turner collector unless otherwise indicated), “from dry ice
Malaise trap,” 4 $ 14 April 1968, 4 $ 20 April 1968, $ 4 May 1968, 15 $ , 2 $
17 May 1968, $ 18 May 1968, $, 20 $ 1 June 1968, 16 2 2 June 1968, $ 10
9
Vol. 49, No. 1, January 1973
0.3 mm
Fig. 5. Male IX tergite, dorsal and left side, lateral view: Symphoromyia
inconspicua. Figs. 6-10. Male disticerci: Fig. 6, S. cervivora; Fig. 7, S. truncata;
Fig. 8, S. inconspicua ; Fig. 9, S. nana; Fig. 10, S. pachyceras Williston.
10
The Pan-Pacific Entomologist
June 1968, 4 $ 2 May 1969, $ 2 May 1969, $ 1 June 1969; “from male swarm,”
$ 2 June 1965 (J. R. Anderson), 10 $ 2 May 1969, 9 $ 28 May 1969; 2 14
May 1964 “off ears of deer” (J. R. Anderson) ; $ 8 May 1970 “at light”;
2 2 9 May 1970. Alameda Co.: Berkeley, 2 27 April 1954 (W. W. Middlekauff).
Contra Costa Co.: Danville, 2 9 May 1952 (F. X. Williams) ; Lafayette, 2 mi.
S., 2 $ 18 April 1968, “from dry ice Malaise trap,” 11 $, 2 2 26 April 1968,
2 $, 12 2 1 May 1968, 8 ^,3 2 9 May 1968, $, 2 2 17 May 1968, 7 2 15
May 1969; “from male swarm,” 46 $ 15 May 1969, Walnut Creek, foot of Shell
Rdg., S 15 April 1964 (J. A. Powell) ; Walnut Creek, 3.5 mi. E., $, 2 2 2 May
1969 (M. F. Knudsen). Lake Co.: Hopland Grade, $ 3 May 1961 (S. M. Fidel).
Marin Co.: Alpine Lk., $ 8 May 1965 (G. Buckingham), 0.5 mi. N., 3 $ 26
April 1961 (R. Garcia) ; nr. Alpine Lk. (Lily Pond), “from dry ice Malaise trap,”
3 $ 3 May 1970 (W. J. Turner) ,9 2 9 May 1970 (W. J. Turner) ; “from male
swarm,” 6 $ 3 May 1970 (W. J. Turner), 14 ^ 9 May 1970 (W. J. Turner) ;
Carson Rdg., 2 26 April 1961 (J. A. Powell) ; Cypress Rdg., 2 29 March 1921
(T. Aitkin), $ 17 May 1922 (E. P. Van Duzee) ; Fairfax, $ 13 April 1919 (E. P.
Van Duzee), 2 mi. W., $ 15 April 1966 (J. A. Powell), 2 mi. S.W., nr. Meadow
Club, $ 7 April 1961 (D. C. Rentz) ; Inverness, 2 $ 14 April 1961 (C. A. Toschi),
2 30 May 1964 (C. A. Toschi) ; Mill Va., 2 2 26 April 1925 (E. P. Van Duzee),
$ 28 February 1926 (M. C. Van Duzee), $ 26 April 1947 (E. L. Kessel), $ 28
April 1950 (F. X. Williams) ; Mt. Tamalpais, $ 24 April 1950 (F. X. Williams) ;
Nicasio, 4.4 mi. E., $ 13 April 1968 (P. A. Opler) ; nr. Novato (Umdelelannyoni),
4 $ 10 May 1948 (E. L. Kessel), 12 3', 2 2 5 May 1951 (E. L. Kessel) ; Paradise
Cove, $ 27 April 1947 (E. L. Kessel) ; San Geronimo, 2 2 23 April 1913 (J. C.
Bridwell). Napa Co.: Lokoya, T 5 2 4 May 1958 (E. G. Linsley, R. F. Smith) ;
Napa, 2 $ 7 April 1939 (M. A. Cazier) ; Samuel Spg., 3 $ 29 March 1956 (E. I.
Schlinger). San Francisco Co.: San Francisco, $ (H. Edwards). San Luis
Obispo Co.: Creston, 10 mi. S., 2 1 May 1962 (C. A. Toschi) ; La Panza Cmp.,
8 2 25 April 1968 (J. A. Chemsak, J. T. Doyen, D. Veirs). San Mateo Co.:
Corte de Madera Crk., nr. Portola, $ 31 May 1958 (P. H. Arnaud). Santa Clara
Co.: Herbert Crk., 3 mi. W. New Almaden, “from dry ice Malaise trap,” 4 2
28 April 1969 (W. J. Turner) ; Los Altos, $ 17 April 1937 (E. S. Ross) ; Mt.
Hamilton, 2 15 April 1947 (G. E. Bohart), 5 mi. E., 2 16 April 1955 (D. J.
Burdick); San Antonio Va., 2 20 April 1948 (R. F. Smith). Solano Co.: Mix
Cyn., 2 $ 14 April 1955 (J. C. Downey), 2 13 April 1969 (M. K. Sears);
Vacaville, 2 6 April 1951 (R. C. Bechtel), $ 27 May 1955 (E. Mezger), $ 1
April 1963 (P. Richerson), $ 4 April 1967 (P. A. Opler). Sonoma Co.: Glen
Ellen, $ 29 April 1950 (H. B. Leech) ; Mt. St. Helena, 2 30 April 1949 (A. T.
McClay). Yolo Co.: Winters, 1 mi. W., S 29 April 1967 (P. B. Schultz), 9 mi.
W., $ 20 April 1966 (M. R. Gardner). County unknown: “California,” 2 2
(O. T. Baron).
Holotype male, allotype female and thirty-three paratypes, 27 $, 6 2,
will be deposited in the California Academy of Sciences, San Francisco.
Additional paratypes have been placed in the collections at the
California Department of Public Health, Bureau of Vector Control,
Berkeley; Canadian National Collection, Ottawa; Museum of Com¬
parative Zoology, Cambridge; University of California, Berkeley and
Vol. 49, No. 1, January 1973
11
Davis; U.S. National Museum, Washington, D.C.; and Washington
State University, Pullman.
Discussion.- —This species appears quite similar to S. pachyceras
Williston. Males of both species are large and bear long, dorsal
postocular hairs which are one-half the length of hair on the ocellar
prominence. Unlike S. pachyceras , the wing membrane in S. cervivora
is essentially hyaline, lacking any infuscation other than the stigma;
also the laterotergite is usually black haired. The shape of the dististylus
quickly separates males of this species from those of S. pachyceras
(Fig. 10). Ventrally, the dististylus bears a broad, moderately deep
quadrangular-shaped basiventral lobe and short distal arm, appearing
somewhat cleaver-shaped in outline (Fig. 6). Females of S. cervivora
might be confused with S. pachyceras as parafacial hair is crowded
laterally and the palpus appears black in both. Uniquely, females of
this species are bluish gray or a dark gray pollinose on the abdomen
above and the surface appears subshiny; dark brown dorsal coloration
is limited to a single median spot and narrow, posterior margination.
In S. pachyceras, for comparison, the surface is primarily dull gray or
brownish gray pollinose and often the medial brown area nearly en¬
compasses the entire tergite. In S. cervivora the mesoscutal vittae are
usually gray but faint, hardly ever as distinct or dark as seen in the
other species.
Examination of Williston’s S. pachyceras type series indicates that
the two female cotypes are actually S. cervivora. Abdominal pollinosity
of both females is dark gray without any tinge of brown; mesoscutal
vittae appear dark brown in one female and less distinct, pale gray in
the other but definitely conspecific with S. cervivora.
The single male cotype specimen is most probably S. pachyceras.
Positive identification is impossible as the genitalia have been removed
and evidently lost at some time in the past. Even without the genitalia,
several other morphological features suggest that this specimen is
S. pachyceras and not S. cervivora with which it might be confused.
The wing membrane of the type appears almost clear as one would
expect of S. cervivora and not infuscated which is characteristic of S.
pachyceras. However, the stigma is now quite pale and suggests that
the original duskiness of the wing may have faded somewhat. Williston
(1886) noted of his male type that the wings had “a strong brownish
yellow tinge” and Aldrich (1915), who later examined this specimen,
also indicated the strong wing infuscation. The first antennal segment
of S. cervivora males is typically swollen beneath medially, its height
in lateral view, greater than the height of the third segment. The
12
The Pan-Pacific Entomologist
pedicle of S. pachyceras appears swollen submedially tapering to the
apex, its height subequal to or slightly greater than that of the third
segment. This type of antennal structure is also seen in the Williston
cotype male.
Each Williston specimen bears a label reading “Baron.” Essig (1931:
p. 552) notes that 0. T. Baron, “collected Coleoptera, Lepidoptera and
other insects extensively in Mendocino and Fresno counties.” Although
the precise localities remain unknown, it is quite likely that the S.
pachyceras type material collected by Baron was taken at least in
Mendocino Co., and perhaps even near Hopland, the type locality for
S. cervivora. It is also not inconceivable that Baron captured a mixed
series of individuals representing two closely related species, S. pachy¬
ceras and S. cervivora , as females of several species may be present at
any one time in some areas. From the limited Baron material and no
further data, Williston associated males of one species with the female
pair of the other assuming them to be conspecific.
This species was referred to by Chillcott (in lilt.) as species “P” for
use by Hoy (1966) in his manuscript.
Adult seasonal activity. —Collecting dates extend from 28 February
until 2 June but this species appears most abundant from late April until
mid May.
Geographical distribution. —Collection records suggest that this
species is found primarily at margins of oak woodland areas of the Bay
Area and adjacent counties from Mendocino County in the north to
Santa Clara County in the south. At several localities, S. cervivora is
sympatric with most other species of the pachyceras group occurring
in the northern Coast Range. Several individuals from San Luis
Obispo County in the central Coast Ranges appear conspecific with
more northern populations and will be treated as such here. These pop¬
ulations may represent the southern most limit of this species.
Symphoromyia inconspicua Turner and Chillcott, new species
Males of S. inconspicua are small, faintly dusted with brownish or
bluish gray pollen; metatibia bears long suberect hair on both postero-
and anteroventral surfaces; humerus black haired; dististylus elongate
and slightly elbowed but lacking definite ventral process; ninth tergite
broadly rounded in lateral view. Females have yellow palpi and mostly
black hair well distributed over parafacial area and beneath first
antennal segment.
Holotype male. —Length: 7.0 mm. Wing length: 5.6 mm. Head and its
appendages, thorax, legs and abdomen black; femur I narrowly reddish brown
Vol. 49, No. 1 , January 1973
13
apically, femora II, III similar but less so, halter knob dark brown, stem lighter.
Surface gray, bluish and brownish gray pollinose, most dense on clypeus,
parafacialia, behind eyes, mesoscutum forward and at sides, on meron, mete-
pisternum, metepimeron, and on coxa III, becoming less pollinose, definitely
subshiny on remainder of head, antenna, thoracic pleura except as noted, on coxae
II, III, femora, tibiae, and abdominal sternites; scutellum and abdominal tergites
lightly dusted with brownish gray pollen, surface subshiny, more dense laterally.
Hair essentially black; white hair present on proepimeron, proepisternum, latero-
tergite forward, abdominal segments I—III, mixed white and black on coxa III
and somewhat on segment III behind. Head : Features of head and its appendages
as described for S. cervivorci except: antennal segment I elongate, swollen sub-
medially beneath, tapering forward, less so basally, height less than two-thirds
length, greater than height of segment III which is roughly kidney-shaped, straight
beneath arista, palpal segment II bearing weak anterodorsal callus behind attenuate
apex; postocular hair dorsally erect but short, scarcely visible from front. Thorax :
Mesothoracic vittae broad, faint, almost dull black, nearly confluent on postscutum,
separated by narrow, bluish gray pollinose lines; median vitta tapering forward,
bisected by thin, brownish gray pollinose line; lateral vitta incomplete forward,
uninterrupted medially. Wing membrane definitely infuscated brown, almost
concolorous with stigma, darkest near stigma, apically and about veins, nearly
hyaline toward base; costal vein beneath base black haired. Tibia III with long,
suberect hair on basal one-third to one-half of both postero- and anteroventral
surfaces, length greater than tibial thickness. Abdomen : Shape as described
for S. cervivora. Tergite IX apically on posterior margin with laterally compressed
process broadly rounded in lateral view (Fig. 5) ; dististylus elbowed, elongate
and narrow, somewhat flattened and polished on posterior surface, smoothly
rounded beneath, lacking definite ventral lobe (Fig. 8) ; cercus elongate, narrow
and attenuate, apically blunt. Sternite VIII and hypopygium with hair much
longer than height of basistylus.
Female.— Length: 4.7-5.8 mm. Wing length: 4.8-6.7 mm. Black except for
femora II, III, and tibiae which are dark brown, nearly black; halter knob similar
but becoming yellowish brown on stem; palpus orange yellow; femora I—III
yellowed narrowly at apices. Surface dull, rugulose under high magnification
(45x), nearly opaque with gray pollen, at most feebly shiny over most of head,
its appendages and on body except: upper frons area usually brownish gray
pollinose, becoming more gray behind head, on scutellum and abdominal sternites;
abdominal tergites each marked with indistinct, cupreous-tinged brown pollinose
spot on disc and along rear margin, otherwise with subshiny, dark gray or gray
pollen becoming brownish gray laterally; mesoscutal vittae dark brown. Hair pre¬
dominantly white but appearing black on palpus beneath and at apex, head above,
mesoscutum except humerus and behind, scutellum, tibiae, femora apically, on
abdominal tergites becoming white only laterally on tergites II-IV and on sternites,
mostly black or with very few white hairs on parafacial area and antennal segment
I beneath. Head: Minimum distance between eyes at midfrons more than one-third
but less than two-fifths head width; antennal segment I short, height greatest
subapically, subequal to length, apex almost truncate, tapering slightly to base,
hair sparse, shorter than segmental height, shortest above; segment III convex
beneath arista, more rounded ventrally than above when viewed from side, height
greater than height of segment I; palpal segment II greatly convex above, hardly
14
The Pan-Pacific Entomologist
so ventrally, length about twice length of segment I, attenuate or narrowly appended
apically, hair sparse, short, absent dorsomedially; parafacialia sparsely but evenly
haired; labrum shorter than clypeal height. Thorax-. Mesoscutal vittae wide,
brown, more or less distinct, tapering to rear, separated by wide, bluish gray lines;
middle vitta bisected by faint, light brown, pollinose line; lateral vitta incomplete
forward, interrupted medially or nearly so. Wing membrane nearly clear, gray
hyaline; veins dark brown apically, becoming light brown or yellowish brown at
base; costal vein at base yellow haired at most with few black hairs. Abdomen:
Disticercus elongate, shape subfusiform, rounded or subtruncate at apex, tapering
toward base.
Holotype male, Hopland Field Station (4 mi. E. Hopland), Uni¬
versity of California, Mendocino County, California, elevation
2,350 feet, 21 May 1969 (W. J. Turner), from male swarm, 1055 P.D.T.
CAS type no. 11251. Allotype female, same data as type except 28 May
1969, elevation 2,880 feet, and from dry ice Malaise trap.
Paratypes, 77 males, 187 females. CALIFORNIA: 64 $, 152 2, same locality
as type (all W. J. Turner collector unless otherwise indicted), “from dry ice
Malaise trap,” 2 $ 6 May 1964 (J. B. Hoy), 5 $ (13-14) May 1964 (J. R.
Anderson), 3 2 15 May 1964 (J. R. Anderson), $ 19 June 1964 (J. B. Hoy),
2 20 April 1966 (J. R. Anderson), 2 18 May 1968, 42 2 1 June 1968, 2 $,
24 2 2 June 1968, $, 14 2 10 June 1968, 3 2 21 June 1968, 2 2 20 April 1969,
2 $ 21 May 1969, 4 $ 28 May 1969, 3 2 H June 1969; “from male swarm,”
7 $ 14 May 1969, 25 $ 21 May 1969, 23 $, 4 2 28 May 1969; “from deer,”
29 2 21 May 1964 (J. B. Hoy), 14 2 28 May 1964 (J. B. Hoy) ; “off small doe,”
2 2 11 June 1964 (J. B. Hoy), 2 12 June 1964 (J. B. Hoy). Lake Co.: No.
frk. Cache Crk., Hwy. 20, 2 17 May 1961 (D. R. Miller). Marin Co.: nr. Alpine
Lk. (Lily Pond), “from dry ice Malaise trap,” 2 21 April 1969 (W. J. Turner).
Mendocino Co.: Booneville, 4^1 June 1957 (J. A. Powell). Monterey Co.:
Arroyo Seco Cmp., 2 23 May 1958 (D. D. Linsdale), 2 24 May 1958 (D. D.
Linsdale), “from dry ice Malaise trap,” 4 2 2 June 1970 (W. J. Turner) ; Bryson,
2 20 May 1920 (E. P. Van Duzee). Napa Co.: Angwin, 1 mi. N., 7 $ 9 May
1964 (H. B. Leech) ; Lokoya, 15 2 4 May 1958 (E. G. Linsley). Sonoma Co.:
Armstrong St. Park., 2 20 May 1958 (A. C. Smith) ; Plantation, 4 mi. W., $ 8
May 1958 (D. J. Burdick). Yolo Co.: Rumsey, 5 mi. W., 10 2 29 April 1969
(M. F. Knudsen) ; 9 mi. W., Hwy. 16, $ 13 May 1962 (M. E. Irwin).
Holotype male, allotype female and four paratypes, 2 $, 2 2 , will
be deposited in the California Academy of Sciences, San Francisco.
Additional paratypes have been placed in the collections at the California
Department of Public Health, Bureau of Vector Control, Berkeley;
Canadian National Collection, Ottawa; Cornell University, Ithaca;
University of California, Berkeley and Davis; U.S. National Museum,
Washington, D.C.; and Washington State University, Pullman.
Discussion. —Males of S. inconspicua are black with lightly dusted
brownish or bluish gray pollen, wing membrane is lightly infuscated and
hair vestiture is mostly black, especially on the head, palpus, humerus,
Vol. 49, No. 1, January 1973
15
thoracic pleura and fourth abdominal tergite. Males may be confused
with the smaller, sympatric S. nana. This species, however, may be
identified by the slightly elbowed dististylus which appears elongate
and narrow, but most importantly, flattened and somewhat polished
on the posterior surface above an elongate carina; noticeably absent is
a definite ventral lobe. The dististylus of S. nana appears similar but
with a shallow process beneath. Also the metatibia of S. inconspicua
bears long, suberect hairs on both the postero- and anteroventral sur¬
faces; length of this hair greatly exceeds tibial thickness. In other
species, suberect hair is usually present only on a single surface of the
tibia. Females typically have orange palpi and black hair present on the
basal antennal segment and well distributed over the parafacial area.
Color of the parafacial hair should identify this species from similar
appearing females having yellowed palpi, particularly S. nana. The
palpus of some individuals is somewhat grayish along the ventral sur¬
face but, in general, the palpus remains wholly orange.
Females of this species were designated species “N” by Chillcott
(in litt.) for use by Hoy (1966).
Adult seasonal activity. —Collection dates range from 20 April to
29 June with most records occurring from mid May until early June.
Geographic distribution. —Known from the type locality and from
several other localities in nearby Napa, Lake, Marin, Sonoma and
Yolo counties. The range of this species extends south at least into
Monterey Co.
Symphoromyia nana Turner and Chillcott, new species
Males of this species are small and sparsely dusted with brownish gray
pollen; postocular hair is short and scarcely visible from front; humerus
white haired; ninth abdominal tergite small, rounded apically in lateral
view; dististylus slightly elbowed with small, shallow medioventral lobe.
In females the second palpal segment is orange, palpus and evenly
haired parafacial area with mostly white hair.
Holotype male.— Length: 5.2 mm. Wing length: 4.7 mm. Head, antenna,
thorax, abdomen and tarsomeres black; humerus, palpal segment II, halter knob,
femora, tibiae dark brown, almost black; femora I—III narrowly at apex yellow.
Surface dull, almost opaque gray or bluish gray pollinose on parafacialia, clypeus,
head behind eyes, antenna, and scutum at sides, becoming less dense and feebly
shiny on thoracic pleural area, abdominal sternites, coxae, femora and tibiae;
brown or grayish brown, almost dense pollen present on scutum between vittae,
less dense on scutellum, abdomen above and mesokatepisternum ventrally, last with
cupreous stripe medially. Hair predominantly black, but white on peristomal
sclerites, head behind, humerus, on thoracic pleural area except for few black
16
The Pan-Pacific Entomologist
hairs on mesoanepisternum at rear and on mesokatepisternum below, also white
on coxa III, abdominal tergite and on disc of sternites I-IV, mixed white with
black on coxa I, femora II, III, abdominal tergites II-IV. Head: Eyes separated
above by distance subequal to width of median ocellus; interocellar distance sub¬
equal; antennal segment I elongate, moderately swollen beneath, tapered slightly
towards ends, height three-fifths length, less than height of segment III, hair almost
dense, length above subequal to segmental length, shorter beneath; segment III
roughly kidney-shaped, straight beneath arista, equally rounded above and below,
height nearly three-fifths length of segment I; palpus elongate, narrow, subcylin-
drical, bearing subapical dorsal callus, segment II slightly incrassate basally, sub-
cylindrical to acute apex, length over twice length of segment I; labrum shorter
than palpus; postocular hair dorsally erect but short, scarcely visible from front;
hair of ocellar prominence long, curved forward, length subequal to hair of antennal
segment I beneath; parafacialia evenly, almost densely haired, length subequal to
hair of antennal segment I. Thorax: Mesoscutal vittae evident only as wide, black
stripes separated by wide, brownish gray pollinose lines; middle vitta bisected by
thin brown pollinose line; lateral vitta wider, interrupted medially, delimited
laterally and forward by bluish gray pollen. Wing membrane lightly infuscated
brown, darkest behind stigma, apically and about veins, stigma barely darker;
veins dark brown apically becoming lighter brown toward base; costal vein with
black hair beneath. Tibia III with long, suberect hair on basal one-half of postero-
ventral surface, length greater than tibial thickness. Abdomen: Shape as described
for S. cervivora. Tergite IX with laterally compressed apical process which is small
and rounded in lateral view (Fig. 3) ; dististylus elongate, narrow, curved dorsad,
posterior surface flattened, bearing small ventral lobe (Fig. 9) ; cercus nearly
triangular shaped, widest basally, attenuate and blunt apically.
Female. —Length: 4.3-5.3 mm. Wing length: 4.5-4.8 mm. Color and pol-
linosity as described for S. inconspicua. Hair predominantly white as in S.
inconspicua but appearing black on palpus at apex, antennal segment I above and
on femur III; parafacial area and antennal segment I white haired. Head: As
described for S. inconspicua except antennal segment III almost equally rounded
above and beneath when viewed laterally. Thorax: Mesoscutal vittae, wing mem¬
brane and veins as described for S. inconspicua ; costal vein at base with yellow
and black hair. Abdomen: As described for S. inconspicua female.
Holotype male , Hopland Field Station (4 mi. E. Hopland) Uni¬
versity of California, Mendocino County, California, elevation 880
feet, 21 June 1968 (W. J. Turner), from dry ice Malaise trap. CAS type
no. 11248. Allotype female, same data as type.
Paratypes, 36 males, 165 females. CALIFORNIA: 30 $, 159 $, same locality
as type (all W. J. Turner collector unless otherwise indicated), “from dry ice
Malaise trap,” $ 26 May 1966 (J. B. Hoy), $ 31 May 1966 (J. B. Hoy), 2 $
1 June 1966 (J. B. Hoy), $ 2 June 1966 (J. B. Hoy), $ 4 June 1966 (J. B. Hoy),
2 $ 10 June 1966 (J. B. Hoy), $ 12 June 1966 (J. B. Hoy), 5 $ 16 May 1968,
3 2 17 May 1968, 11 $, 45 2 1 June 1968, ^,24 2 2 June 1968, 3 ^,32 2
10 June 1968, 4 $, 10 2 21 June 1968, 2 21 May 1969, 2 $ 28 May 1969, 2 14
June 1969; “from black tail deer,” 2 2 21 May 1964 (J. B. Hoy), 2 2 June 1964
(J. B. Hoy), 3 2 16 May 1965 (J. B. Hoy), 2 20 May 1965 (J. B. Hoy), 8 2
Vol. 49, No. 1, January 1973
17
26 May 1965 (J. B. Hoy), 2 27 May 1965 (J. B. Hoy), 2 $ 30 May 1965 (J. B.
Hoy), 2 1 June 1965 (J. B. Hoy), $ 2 June 1965 (J. R. Anderson), 5 $ 3 June
1965 (J. R. Anderson), 2 $ 5 June 1965 (J. R. Anderson), 8 2 23 June 1965
(J. R. Anderson), 2 7 July 1965 (J. R. Anderson). Marin Co.: Alpine Lk.,
$ 19 May 1969 (M. F. Knudsen). Mendocino Co.: Booneville, $ 1 June 1957
(J. A. Powell). San Luis Obispo Co.: Pozo, “at light,” $ 3 May 1962 (J. A.
Powell), $ 28 April 1962 (C. A. Toschi). Santa Clara Co.: Monte Vista, 2
15 June 1956; Stevens Crk., nr. San Jose, 2 6 July 1954; Wrights, 2 2 12 June
1956. Santa Cruz Co.: Aptos Crk., 2 13 May 1961 (C. N. Slobodchikoff).
Trinity Co.: Zenia, 2 $ 20 June 1935 (M. A. Cazier). Yolo Co.: Winters, 9
mi. W., 2 21 April 1962 (H. V. Daly).
Holotype male, allotype female and four paratypes, 2 $, 2 2, will
be deposited in the California Academy of Sciences, San Francisco.
Additional paratypes have been placed in the collections at the American
Museum of Natural History, New York; California Department of
Health, Bureau of Vector Control, Berkeley; Canadian National Collec¬
tion, Ottawa; University of California, Berkeley and Davis; U.S.
National Museum, Washington, D.C.; and Washington State University,
Pullman.
Taxonomic discussion. — Symphoromyia nana is one of the smallest
of Symphoromyia species. Males are also unique in that their dististyli
are narrow and elbowed with a very shallow medioventral lobe and the
ninth tergite is small, rounded apically in lateral view. Females of
S. nana are quite similar to S. inconspicua females. Both species have
orange palpi and evenly haired parafacial area but these surfaces appear
white haired in S. nana and black in S. inconspicua. Each abdominal
tergite in S. nana appears dark brown medially and almost shiny, usually
surrounded by gray but often brownish gray pollen. The abdomen of
S. inconspicua appears more densely gray pollinose and the median
spot on each tergite is reduced in size and appears not as dark nor shiny.
Hoy (1966) referred to this species as “0” based on tentative
recognition of the species by Chillcott (in litt .).
Adult seasonal activity. —Collection records extend from 21 April
until 7 July. This species is most abundant from mid May to mid June.
Geographical distribution. —This species is recorded from Trinity
County to San Luis Obispo County along the Coast Ranges in associa¬
tion with oak woodland habitats.
Symphoromyia truncata Turner and Chillcott, new species
Males of this species are small; head and thorax somewhat densely
dusted with gray to brownish gray pollen; postocular hair short, barely
visible from front; ninth abdominal tergite flairing apically, dorsal
18
The Pan-Pacific Entomologist
angle acute; dististylus elongate, narrow and curved but not elbowed,
lacking ventral lobe. Females may be recognized by black palpi;
parafacial area sparsely but uniformly black haired; abdomen dark
brown, almost shining, only lightly dusted with brownish gray pollen
which tends to gray laterally.
Holotype male. —Length: 6.5 mm. Wing length: 5.8 mm. Head and its
appendages, thorax, legs and abdomen black; femur I, II narrowly reddish yellow
apically, femur III similar but less so; halter knob dark brown, stem lighter.
Surface appears minutely rugulose at high power (45 X) weakly subshining, gray
or brownish gray pollinose on head, antennal segment I, scutum on non-vittate
area, thoracic pleura, coxae, abdominal sterna and tergites at sides; frons, ocellar
prominence, scutellum and abdominal tergites lightly dusted with brown pollen,
becoming quite sparse on tergites. Hair essentially black but noticeably white
on humerus and behind, thoracic pleura except rear of mesoanepisternum, on
laterotergite at sides, coxa III, and abdominal segments I—III and disc of IV;
mixed yellow or white and black hair present on femora II, III, and on abdominal
tergites IV posterolaterally, and V; sternite IV with white, stiff hairs on postero¬
lateral lobe. Head : Eyes separated above by distance subequal to width of medial
ocellus; interocellar distance subequal; antennal segment I swollen medially
beneath, tapering forward, less so basally, height three-fourths length, greater than
height of segment III, hair dense, length beneath subequal to segmental length,
longer above; segment III kidney-shaped, convex beneath arista, more rounded
below than dorsally, height one-half length of segment I; palpus elongate, segment
II tapered from slightly incrassate base, almost carrot-shaped, scarcely arcuate
medially, length just twice length of segment I, bearing small, anterodorsal callus,
hair dense; labrum shorter than palpus; postocular hair dorsally erect but short,
scarcely visible from front, hair on ocellar prominence long, suberect, subequal
to hair of antennal segment I, parafacial area almost densely haired, of similar
length. Thorax : Mesoscutal vittae present as wide, black median stripe and dark
grayish brown, pollinose, lateral stripes separated by medium width gray lines;
middle vitta bisected by thin, brownish gray pollinose line; lateral vitta incomplete
forward, uninterrupted medially, tapered to rear. Wing membrane lightly infuscated
brown, darkest behind stigma and apically, nearly hyaline toward base; costal vein
basally beneath with mixed golden and hlaclc hair. Tibia III with long, suberect
hair on basal one-half of posteroventral surface, length greater than tibial thick¬
ness. Abdomen : Form as given for S. cervivora. Tergite IX with laterally com¬
pressed, somewhat flaired apical process, nearly triangular in lateral view (Fig. 2) ;
dististylus elongate, narrow throughout and curved, lacking ventral lobe (Fig. 7) ;
cercus elongate, flat, attenuate from wide base to acute apex.
Female. —Length: 4.9-6.1 mm. Wing length: 4.3-5.3 mm. Color essentially
black but obscured by dense or almost dense gray or bluish gray pollen; femur I
yellow narrowly at apex, femora II, III similar but less so; halter dark brown,
stem brownish yellow. Surface dull, opaque, at most feebly subshiny and gray or
bluish gray pollinose on parafacialia, upper frons area, head behind, and on
scutellum, becoming less dense, subshiny on clypeus, antennal segment I, non-
vittate areas of mesoscutum, thoracic pleural area, head behind and sometimes on
mesokatepisternum ventrally; upper frons area often tinged with brown pollinosity,
Vol. 49, No. 1, January 1973
19
becoming darker, more dense on mesoscutal vittae; abdominal segments subshining,
moderately dusted with gray pollen having cupreous tinge above, more dense and
brownish gray laterally, and beneath, tergite I slightly more densely pollinose than
following terga. Hair mostly white but usually black on antenna, parafacialia,
head above and behind, on mesoscutum, scutellum, mesoanepisternum behind,
medially on abdominal tergites II and following, white haired only laterally on
tergites II, III and along posterior margin of tergite II; hair mixed black and
white on femur I; palpus essentially white haired with black hairs apically. Head :
Minimum distance between eyes at midfrons between one-third and two-fifths
head width, rarely greater (MFW/HW: 0.35-0.42); antennal segment I short,
subtruncate forward, tapering slightly to base, height greatest postmedially, ranging
from nearly two-thirds of to subequal to length, hair sparse, longest less than
segmental height, shortest above; segment III convex beneath arista, subequally
rounded above and beneath, height greater than height of segment I; palpal seg¬
ment II convex above, barely convex or flat beneath, narrowly appended apically,
length about twice length of segment I; parafacialia sparsely but evenly haired;
labrum just shorter than clypeal height. Thorax: Mesoscutal vittae wide, indis¬
tinct, dark gray or brown to black, separated by wide, bluish gray pollinose lines;
middle vitta widest, indistinct to rear, bisected by faint, narrow, brown pollinose
line; lateral vitta interrupted medially or nearly so, often appearing as two elongate
spots. Wing membrane clear, gray hyaline, stigma faintly infuscated brown;
veins dark brown apically becoming yellow at base; costal vein at base beneath
with mixed yellow and black hair. Abdomen : Disticercus elongate, rounded
apically, tapering toward base.
Holotype male , Hopland Field Station (4 mi. E. Hopland), Uni¬
versity of California, Mendocino County, California, elevation
1,280 feet, 14 May 1969 (W. J. Turner), from male swarm, 1343
P.D.T. CAS type no. 11249. Allotype female, same data as type except,
17 May 1968, elevation 880 feet, and from dry ice Malaise trap.
Paratypes, 106 males, 162 females. CALIFORNIA: 90 $, 150 2, same locality
as type (all W. J. Turner collector unless otherwise indicated), “from dry ice
Malaise trap,” 2 15 May 1964 (J. R. Anderson), $ 30 May 1966 (J. B. Hoy),
$ 10 June 1966 (J. B. Hoy), 2 $ 11 June 1966 (J. R. Anderson), 2 2 20 April
1968, 2 2 4 May 1968, 2 $, 7 2 17 May 1958, 2 $, 2 2 18 May 1968, 5 $,
33 2 1 June 1968, $, 10 2 2 June 1968, 10 2 10 June 1968, 2 2 21 June 1968,
4 $ 21 May 1969, 3 $ 22 May 1969, 4 2 28 May 1969, 2 7 June 1969, 9 2 11
June 1969 (J. R. Anderson), 2 2 14 June 1969; “from male swarm,” 11 $ 12
May 1968, 11 $ 21 May 1968, 39 $ 14 May 1969, 34 $ 21 May 1969, 2 $ 28
May 1969; “from black-tail deer,” 2 29 April 1964 (J. B. Hoy), 7 2 6 May 1964
(J. B. Hoy), 3 2 21 May 1964 (J. B. Hoy), 4 2 9 May 1965 (J. B. Hoy), 2 2
11 May 1965 (J. B. Hoy), 7 2 12 May 1965 (J. B. Hoy), 6 2 16 May 1965 (J. B.
'Hoy), 3 2 23 May 1965 (J. B. Hoy), 7 2 26 May 1965 (J. B. Hoy), 3 2 27
May 1965 (J. B. Hoy), 3 2 28 May 1965 (J. B. Hoy), 5 2 2 June 1965 (J. B.
Hoy), 6 2 23 June 1965 (J. B. Hoy), 2 29 June 1965 (J. B. Hoy), 2 10 July
1966 (J. B. Hoy) ; 2 15 May 1964 (J. R. Anderson), $ 30 May 1966 (J. B. Hoy),
2 $ 11 June 1966 (J. R. Anderson). Marin Co.: Alpine Lk., 3 $,3 2 19 May
1969 (M. F. Knudsen) ; nr. Alpine Lk. (Lily Pond), “from male swarm,” 6 $
20
The Pan-Pacific Entomologist
17 May 1969 (D. D. Munroe) ; Carson Rdg., 3 2 26 April 1961 (J. A. Powell).
Napa Co.: Middletown, 10 mi. S., $ 27 April 1958 (T. R. Haig) ; Pope Va.,
$ 24 May 1969 (M. K. Sears) ; Samuel Spr., 2 $ 13 May 1956 (C. M. Gardella).
Santa Clara Co.: Wrights, “collected biting,” $ 12 June 1956. Santa Cruz
Co.: Felton, Sta. Cruz Mts., 2 2 (15-19) May 1907 (J. C. Bradley). Sonoma
Co.: Mt. St. Helena, 2 15 May 1951 (E. I. Schlinger) ; Plantation, 2 22 May
1954 (A. Collins). Trinity Co.: Fawn Ldg., 2 28 May 1955 (E. I. Schlinger).
Yolo Co.: Rumsey, 5 mi. W., 3 $ 29 April 1969 (M. F. Knudsen).
Holotype male, allotype female and four paratypes, 2 $ , 2 2, will
be deposited in the California Academy of Sciences, San Francisco.
Additional paratypes have been deposited in the collections at the
California Department of Public Health, Bureau of Vector Control,
Berkeley; California State Department of Agriculture, Sacramento;
Cornell University, Ithaca; Canadian National Collection, Ottawa;
University of California, Berkeley and Davis; U.S. National Museum,
Washington, D.C.; and Washington State University, Pullman.
Chillcott (in litt.) and Hoy (1966) recognized S. truncata as species
“C”
Adult seasonal activity. —Collection records range from 20 April
until 29 June. This species most common from mid May to mid June.
Geographical distribution. —This species is found from Trinity
County south along the northern Coast Ranges to Santa Cruz County.
Throughout its distribution, S. truncata seems associated with oak
woodland habitats and occurs there sympatrically with several other
pachyceras group species.
Literature Cited
Aldrich, J. M. 1915. The dipterous genus Symphoromyia in North America.
Proc. U.S. Nat. Mus., 49: 113-142.
Crampton, G. C. 1942. Guide to the insects of Connecticut. Part VI. The
Diptera or true flies of Connecticut. First fascicle. The external mor¬
phology of the Diptera. Bull. Conn. Geol. Natur. Hist. Sur., No. 64,
pp. 10-175, 17 figs.
Essig, E. O. 1931. A History of Entomology. The Macmillan Co., New York.
1029 pp.
Grimshaw, P. H. 1905. On the terminology of the leg bristles of Diptera.
Entomol. Mon. Mag., 16: 173-176.
Hoy, J. B. 1966. The behavior of members of the genus Symphoromyia attacking
deer in Northern California (Diptera: Rhagionidae). Univ. Kansas,
unpublished Ph.D. dissertation. 162 p., 17 tabs., 31 figs.
Turner, W. J. 1971. Biosystematics of the California species of Symphoromyia
(Diptera, Rhagionidae). Univ. California, unpublished Ph.D. disserta¬
tion. 412 p., 110 figs.
Williston, S. W. 1886. Dipterological notes and descriptions. Trans. Amer.
Entomol. Soc., 13: 287-307.
Vol. 49, No. 1 , January 1973
21
Foraging Behavior of Trigona fulviventris in Costa Rica
(Hymenoptera: Apidae)
William M. Gilbert 1
Department of Zoology, University of California, Davis 95616
A colony of the stingless bee Trigona fulviventris fulviventris Guerin
was observed during the dry season between 8 and 12 March 1969, at
The Tropical Science Center field station, about four km southwest of
Rincon de Osa, Puntarenas Province, Costa Rica. Observations cen¬
tered around a nest of T. fulviventris at the hollow base of a large,
buttressed tree one m from the bank of the Rio Agua Buena. The
entrance tube projected between two buttresses, about 20 cm above the
ground surface.
The diurnal activity cycle of T. fulviventris was determined by sam¬
pling the numbers of bees leaving and entering the nest on 10 March.
Six one minute counts, three of bees leaving and three of bees entering
the nest entrance tube, were made at hourly intervals from 0552 until
1750 (sunrise, 0540, sunset, 1741), with one series of counts being
made on the half hour around 1730. The bees were also observed at
the hive before sunrise on 11 March. Other aspects of T. fulviventris
behavior, as well as the behavior of other stingless bees, were observed
and recorded at the nest site and elsewhere in the study area. Addi¬
tionally, eight cardboard feeding stations were placed on trees at
varying distances from 5 to 100 m and in various directions from the
study nest. These stations were baited with 50% honey solution and
checked for bees at regular intervals.
Results. —The diurnal activity cycle of T. fulviventris was recorded
as departure and return rates per minute from the study nest (Fig. 1).
The first observed activity was at 0523 on 11 March (sunrise 0540)
when bees, observed by flashlight, were clustered at the lip of the mud
nest entrance. Individual bees would leave the cluster and fly toward
the flashlight beam at that time, but natural departure from the nest
did not appear to have begun. Bees could be seen in the dim light at
0535 to be leaving the nest in great numbers. A few were returning.
By 0550 large numbers of bees were both leaving and returning to the
nest.
The greatest activity period of T. fulviventris was in the morning
hours from the onset of activity at about 0530 until about 1000, during
1 Present address—1996 Catalpa Way, Hayward, California 94545.
The Pan-Pacific Entomologist 49: 21-25. January 1973
22
The Pan-Pacific Entomologist
Fig. 1. Diurnal activity pattern of Trigona fulviventris leaving and returning
to a nest on the Osa Peninsula, Puntarenas Province, Costa Rica, on 10 March 1969.
which time departures fell from about 80/min to about 20/min (Fig.
1). The lower activity level was maintained until after 1500, when
activity began to rise to a second peak of about 40 departures/min at
1700. Beyond this time departure rate decreased rapidly, and arrival
rate increased, until by 1750 (sunset, 1741) all departure and arrival
activity had ceased.
Pollen carrying activity was greatest in the early morning hours,
when an estimated 50% of the returnees carried pollen. Only an oc¬
casional bee returned with pollen in the afternoon.
The flight of T. fulviventris was very rapid and direct in leaving
and returning to the nest tube. At other times, however, individuals
could be found hovering low over the ground, occasionally alighting,
usually around tree bases or stumps. Bees were also observed to hover
near feeders at times, while not alighting on the feeding boards. Hover¬
ing of T. fulviventris was noticed at all times of day, and was usually
the only activity noticed in bees of this species found away from the
study nest. No stingless bee species other than T. fulviventris was ob¬
served hovering in such a manner, although at least three other species
were commonly seen in the study area.
Trigona fulviventris failed to forage at any of the several feeding
stations established around the study nest from 8 to 10 March. In
Vol. 49, No. 1, January 1973
23
separate observations made on 7 March, however, in an area approxi¬
mately 300 m upstream from the nest site, T. fulviventris foraged in
small numbers at three feeding stations similar to the ones used in the
study. As many as three individuals were seen at one time on a feeder.
It was not known whether or not these individuals originated from the
observed nest.
All observations of T. fulviventris were made during fair weather
under non-overcast skies.
Discussion. —The absence of T. fulviventris at honey-baited feeding
stations at varying distances from the study nest is surprising consider¬
ing that the species visits many different kinds of flowers (Schwarz,
1948, p. 333) and makes honey. The situation seems all the more
strange because some feeding stations treated on 7 March similarly to
those around the study nest, but placed in a different part of the forest,
attracted a small number of T. fulviventris within a few hours. Possibly
individuals of T. fulviventris from a given nest do not establish at food
sources near to that nest. This possibility is supported by Michener
(1946, p. 181) who states “At old Panama T. fulviventris was never
seen visiting flowers of herbaceous plants growing near its nest although
these same plants were visited regularly by T. jaty and T. nigra pauper a”
Observations on 11 March indicated that T. fulviventris began to
leave the nest naturally under light intensities so low that departures
could not be detected without a flashlight. How the bees could navigate
at these times, when visual cues were apparently unavailable, and be¬
fore scent trails could have been laid for that day, is unknown.
Early flight activity appears to be characteristic of other Trigona
species. Schlising (1970) for example, studied morning foraging of
bees on four species of plants in three different areas in Costa Rica
during the dry season. At each site Trigona bees, foraging in large
numbers on two plant species, were the earliest bees to arrive after it
was light. Only bees of the genus Ptiloglossa (Colletidae), which often
foraged before light, were earlier. Schwarz (1948, p. 83) cites in¬
stances of a Trigona species of the subgenus Partamona being active
before sunrise, although he states that this activity might have resulted
from disturbance of the nest. Peckolt (1894) found that in Brazil the
stingless bees Trigona jatay, T. bipunctata , and T. mosquito (all re¬
ported as Melipona) showed first foraging activity respectively at about
0700, 0700, and between 0530 and 0600 during good weather. In rainy
weather, however, T. mosquito did not appear until about 0800, and
this species and T. jatay were not seen to initiate foraging at all during
thunderstorms.
24
The Pan-Pacific Entomologist
By contrast von Ihering (1903) reported that T. schrottkyi Friese,
occurring in Brazil, did not open its nest entrance until between 0800
and 1000, even in fair weather. Additionally Lutz (1931) cites eight
first flights for T. ( Plebeia ) mosquito variety domiciliorum Schwarz
on Barro Colorado Island, Canal Zone, all of which occurred relatively
late in the morning. Lutz concluded that opening of the nest and
initiation of foraging activity occur in response to both light conditions
and to a circadian clock mechanism. The observations of Lutz were
made in November and December; near the end of the rainy season in
the canal zone. This condition perhaps explains the apparent contra¬
diction between his observations for T. mosquito and those of Peckolt,
who worked in the dry season in Brazil.
The activity pattern recorded in the present study for T. fulviventris
(Peak activity in the mornings and evenings; moderate activity at
midday) is similar to activity patterns observed by Peckolt (1894)
for two stingless bee species which he identified as T. mosquito and
T. bipunctata. However, foraging activity reportedly lasted only until
about 1500 or 1600, normally, for those bees; not until nearly dusk
as was the situation with T. fulviventris in the present study.
A different foraging pattern is reported by Peckolt for T. jatay, and
Ducke (1901) for T. fuscipennis Friese (later considered to be T.
cupira F. Smith) and for T. ( Plebeia ) goeldiana Friese. These species
were observed to do the majority of their foraging from early morning
until about noon, to cease flight activity during the heat of midafter¬
noon, and to once again initiate foraging toward evening. Peckolt
observed the latter flight period of T. jatay to last from about 1500
until about 1800.
The nest of T. fulviventris fulviventris in the present study seems
similar in structure and location to those described for T. fulviventris
fulviventris by Michener (1946, p. 189). There was no association
with a termite nest, as has been reported for the subspecies guianae.
Similar to the experience of Michener (1946) in studying the species
in Panama, T. fulviventris in the present study were very gentle and
not easily excitable, continuing apparently normal activity during close
and prolonged observation at the nest site. Threat of attack was not
observed.
Summary. —Observations were made at and around a nest of Trigona
fulviventris from 8 through 12 March 1969, on the Osa Peninsula,
Puntarenas Province, Costa Rica. Foraging activity of T. fulviventris
began before light, declined in intensity throughout the morning hours
to a stable level maintained through midday, and then increased to a
Vol. 49, No. 1, January 1973
25
second peak before dark. Foraging ended abruptly after the evening
activity peak. The foraging pattern of T. fulviventris in the present
study is similar to that of some other Trigona species, but still other
Trigona species terminate foraging completely during midday. Early
foraging activity appears to be characteristic of many species of Trigona ,
including T. fulviventris, but this behavior apparently is altered by
inclement weather in some species, and at least one species, T. schrottkyi,
characteristically opens its nest tube late in the morning, even in fair
weather.
Trigona fulviventris failed to forage at feeders baited with honey
solution and placed at varying distances and directions from the nest.
Failure in the species to forage near to the nest is also reported by
Schwarz (1948). A unique “hovering” flight, not seen in other Trigona
species in the study area, was observed for T. fulviventris.
Acknowledgements. —Critical evaluation of this manuscript by Drs.
Robbin Thorp, Norman Gary, and William Hamilton III is gratefully
acknowledged. I am also indebted to Dr. Alvaro Wille, Universidad
de Costa Rica, for identification of specimens. The present study was
conducted while attending the Organization of Tropical Studies funda¬
mentals course in Costa Rica.
Literature Cited
Ducke, A. 1901. Beobachtungen uber Blutenbesuch Ershcenungzeit. u.s.w. der
bei Paravorkommenden Bienen. Z. Syst. Hymenopt. Dipt., 1: 25-32,
49-67.
Ihering, H. von. 1903. Biologie der stach ellosen Honigbienen Brasiliens. Zool.
Jahrb., Abt. Syst. Oekol. Geogr. Tiere, 19: 179-287.
Lutz, F. E. 1931. Light as a factor in controlling the start of daily activity of a
wren and stingless bees. Amer. Mus. Novitates, No. 468: 1-9.
Michener, C. D. 1946. Notes on the habits of some Panamanian stingless bees
(Hymenoptera, Apidae). J. N. Y. Entomol. Soc., 54: 179-261.
Peckolt, T. 1894. Uber brasilianische Bienen. Die Natur (Halle), 43: 87-91.
Schlising, R. A. 1970. Sequence and timing of bee foraging in flowers of
Ipomoea and Aniseila (Convolvulaceae). Ecology, 51: 1061-1067.
Schwarz, H. F. 1948. Stingless bees (Meliponidae) of the Western Hemisphere.
Bull. Amer. Mus. Natur. Hist., 90: 1-546.
EDITORIAL NOTICE
The editorship of the Pan-Pacific Entomologist will change hands beginning
with the July 1973 issue (Vol. 49, No. 3). New manuscripts should be submitted
to Dr. John Doyen, Division of Entomology, University of California, Berkeley,
California 94720.
26
The Pan-Pacific Entomologist
Behavior of the Male Trypoxylon rubrocinctum
(Hymenoptera: Sphecidae)
Margaret Mary Paetzel
5031 Donaldson Road, Grants Pass, Oregon 97526
The following observations on male Trypoxylon behavior were made
five miles north of Grants Pass, Oregon, at 1,150 ft. elevation. The
nests were in 6.4 mm holes bored in several old posts set up on the
north side of a building. Notes were taken during approximately thirty
hours of observation over a period of two years; July and August 1968
and 1971. Seven nests were studied, two in 1968, five in 1971. In each
case the males were actively helping the females. Identification of
specimens was made by Mr. Robert Every of Oregon State University,
Corvallis, Oregon.
That, in some species, the male Trypoxylon guards the nest and
occasionally helps the female has been known since the original work
of the Peckhams in 1898. However, knowledge of the behavior of the
male, other than his guard duty, is fragmentary, and the object of my
studies was to determine the actions of the male from the beginning of
a nest, through provisioning and partition building, to final closure.
This was accomplished by the use of a magnifying glass wired to the
post four inches from the boring under study. Observation of activity
within the nest was facilitated by the use of a 4X hand lens and a
flashlight held to throw a beam of light into the interior. While work¬
ing, both inside and outside of the nest, the wasps were remarkably
tolerant of these arrangements, and after a short time of adjustment,
proceeded with their work in a normal manner.
Beginning and Provisioning of the Nest .—In contrast to the males
studied by the Peckhams (1895) I found three males helping the females
clean out the borings before storage of prey. Both wasps worked at the
back of the tunnel and seemed to be smoothing the walls with their
mandibles, working together from side to side. Occasionally one or the
other would come to the entrance and drop a load of fine sawdust out¬
side. After an hour or so of this preparation the female would fly off
to hunt while the male remained in the nest. With only one exception,
the males constantly guarded the doorway while their mates were absent
(Fig. 1). Also, with this same exception, the males guarded the nest
entrance at night. In three nests, during early provisioning when there
was much space in the boring, the female remained in the nest with
the male all night.
The Pan-Pacific Entomologist 49: 26-30. January 1973
Vol. 49, No. 1, January 1973
27
Fig. 1 . Male Trypoxylon guarding his nest. Fig. 2. A cuckoo wasp breaking
into nest of Trypoxylon. This nest was tightly sealed with hard clay, hut note the
large hole the parasite has already made. Fig. 3. Pair of Trypoxylon wasps mating
at nest entrance.
All males studied exhibited a challenging attitude upon the approach
of the female with prey, but after identifying her, their actions varied
somewhat while allowing her to enter the nest. Some backed down the
tunnel when there was room to do so, while others came outside and
rode in on the female’s back. One individual came out, slipped beneath
the female as she entered, seized the spider, placed it in the cell, doubled
himself in half and returned to the entrance facing outward.
Two pairs had a very distinctive ritual of recognition on the return
of the female. Before allowing her to enter, the male grasped each of
her antennae in his mandibles and stroked it with a cleaning motion,
then with much head bobbing on the part of both, he permitted her to
enter. The other pairs performed their greetings more after the manner
of the wasps mentioned by Rau (1928)—merely touching antennae and
nodding heads briefly at the entrance.
From the beginning the males appeared motivated to participate in
storing the prey, but in four of the nests the female was sometimes
reluctant to drop the spider and often a wrestling contest ensued. This
contending for the prey started at the entrance and continued the length
of the tunnel. However, as storing of the nests progressed, and under
28
The Pan-Pacific Entomologist
pressure of work, all but one of the females eventually allowed the
males to participate, often dropping the prey inside the door and
leaving the male to arrange it as he pleased.
Three females, even at the onset of provisioning, allowed the males
to take the prey at the entrance, two willingly giving up the spider when
the male reached outside and took it from them. These were the three
pairs that exhibited the most cooperation throughout, and whose males
appeared the most skilled in helping their mates. But in all these nests
there were periods when the female reversed her cooperative behavior,
and for a time refused to allow the male to handle the prey. This
seemed to occur after she had been out hunting longer than usual.
Then as she began bringing in prey at more frequent intervals (15 to
20 minutes) she again relinquished the prey to the male.
All the males who helped their mates store prey worked in the same
manner. At first, when there was plenty of room, they merely placed the
spider at the back of the boring and shoved it lightly into place with
the head, turned around, (usually inside) and began cleaning face and
antennae. Later when the provisions were closer to the entrance, after
a few ramming motions with the head, they would turn around outside,
back in and shove and push the bulky prey into place with the feet or
tip of abdomen—some even hooking their legs over the outside edge
of the boring to obtain greater leverage.
The Partition .—After a sufficient number of spiders (from 5 to 15)
had been put away and an egg laid, usually on the last (Krombein,
1956), the female brought in mud and made a partition to seal off the
cell. All my observations agree with Hartman (1905) that the mud
partitions and outer barriers are started from the bottom up. In three
of the nests the males appeared to be helping the female in trowelling
the mud for this inner wall. Both wasps stood together, rotating in
unison, as they worked from side to side spreading the wet clay. The
wings of both vibrated rapidly and each emitted a high pitched buzzing
sound while working. The males who did not participate in this activity
merely moved to the front of the boring and allowed the female to place
the mud unhindered. One male seemed particularly stubborn about
moving aside, and his female had difficulty building her partitions.
Parasites .—The wasps had few parasites, but one or two females of
the brilliant metallic green cuckoo wasps (Chrysididae) were always
hanging about inspecting empty borings, and warily eyeing the males
guarding their doorways. Medler (1967) suggests that the presence
of Chrysis and others in a nest may be due to inadequate protection
by the male Trypoxylon. However, on several occasions I observed a
Vol. 49, No. 1, January 1973
29
prowling Chrysis wasp attempt to enter a boring when the male was
back in the tunnel, only to come out immediately, to all appearances
greatly agitated, pursued by the alert guard. If there was much activity
about a nest, or if the guard was sitting at the entrance where he could
be seen the parasites never attempted to enter, but they seemed to take
a great deal of interest in the closed borings. One afternoon I watched
one of these wasps gnawing at the closed and hardened mud door of a
Trypoxylon nest until it had succeeded in cutting through into the
vestibular cell where it immediately began working at the inner parti¬
tion (Fig. 2). Systematically, in the days to follow, it worked at one
nest after another until I put adhesive tape over all the completed
borings to protect them from being ruined.
The Last Cell .—While the female was building her cells deep within
the nest I was unable to observe egg laying, but as the last cell was
provisioned both wasps were unable to completely enter the boring,
and they were close enough to the entrance to be seen easily. On four
occasions, involving two nests, 1 was able to see the mating and the
actions of both wasps as the female deposited an egg on the last spider
brought in. After arranging the prey, the wasps took up positions fac¬
ing each other, one inside, the other outside the nest. After much head
bobbing, antennae waving and mandible clashing, the wasp inside came
out and they mated while clinging to the post an inch or so from the
entrance to the nest (Fig. 3). At once the female backed into the nest,
head, front feet and thorax outside, and remained perfectly still. From
time to time there was a slight quivering and tensing of the antennae.
Three times the male waited, rigidly alert, watching the female. One
male, when the female’s antennae began to vibrate, became quite agitated
and reared up on four legs, holding the bent first legs up under his
head. So absorbed was he that I was able to approach within inches
and make a number of pencil sketches of his unusual attitude.
Closure of the Nest .—All the Trypoxylon pairs observed closing nests
(five out of seven) made an inner door, leaving a vestibular cell be¬
tween the last occupied cell and the outer barrier. Two of the males
left after the final cell was provisioned, and before the entrance was
sealed, but three of the more attentive stayed with the females until the
closing, though they were forced to rest on the post beside the entrance.
One persistent male spent the last night resting in the vestibular cell,
and halfway out of the nest when darkness came before the female
finished the closure.
None of the males who remained until the nest was finished attempted
to help the female with building of the mud barrier, though they had
30
The Pan-Pacific Entomologist
seemed to be assisting with the inner partitions. While not actively
participating, these males appeared intensely interested when the female
began making the vestibular cell wall, and hovered protectively over
her as she worked. The attentive one mentioned above stood with a
leg over the female’s back as he appeared to be trying to watch the
work in progress.
In general, it seemed that the more attentive and helpful the male, the
longer he stayed with the nest. The males from nests where there was
little cooperation between the pair were the ones who left early, often
before the female began making the inner door. The one male of all
the nests studied who stayed until the last mud pellet was spread on
the final barrier, and who even came back several times with the female
to inspect the closed nest, was the one who had been allowed by the
female to share most in the work.
The last nest was provisioned and sealed on 13 August, and from
that time I saw no more females although the males continued to fly
about until 28 August. Every night several males rested in the empty
borings until that date, when the weather turned unusually cold, then
they disappeared from their old haunts, nor were they to be found on
the asters or Michaelmas daisies of the garden.
Literature Cited
Hartman, C. 1905. Observations on the habits of some solitary wasps of Texas.
Univ. Tex. Bull. No. 65, Sci. Series No. 7. 72 p.
Krombejn, K. V. 1956. Proc. Ent. Soc. Wash. 58. No. 3: 155-156.
Medler, J. T. 1967. Biology of Trypoxylon in trap nests in Wisconsin. Amer.
Midland Natur., 78: 344-358.
Peckham, G. W. and E. G. Peckham. 1895. Notes on the habits of Trypoxylon
rubrocinctum and Trypoxylon cilbopilosum. Psyche, 7: 303-306.
1898. On the instincts and habits of solitary wasps. Wis. Geol. Natur. Hist.
Surv. Bull. No. 2. 245 p.
Rau, P. 1928. Field studies in the behavior of the non-social wasps. Trans.
Acad. Sci. St. Louis, 25: 489 p.
Vol. 49, No. 1 , January 1973
31
A Revision of the New World Genus Formicilla 1
(Coleoptera: Anthicidae)
Donald S. Chandler
University of Arizona, Tucson 85721
The genus Formicilla , designated for F. munda LeConte, is now
distinct due to the removal of most of the neotropical species which
had accumulated in the genus to the leporinus- and scitulus- groups of
Acanthinus (Werner 1960, 1970). Beal (1950) had defined the genus
for the United States. However, due to the removal by Werner of two
species included in Beal’s paper and the anatomical similarities of the
remaining United States species, the need for more study of the rela¬
tionships within the genus was indicated.
Formicilla is here defined as those New World Anthicidae with
mesosternum expanded laterally; a tuft of long curved setae on inner
margin of mesepisterna reaching humeral angle; pronotum deeply
constricted in basal half; surface polished, with little pubescence; long,
tactile setae; right mandible with single to triple tooth in middle of
cutting edge; last segment of maxillary palpi securiform; simple tegmen
of aedeagus.
The species of Formicilla are here separated into two groups with
the bruchi- group containing a single species, F. bruchi Pic, and the
munda-group containing the remaining. The bruchi-gxowp is distinct
in having a deep depression on the dorsum of the pronotum and in
having the gonopore armature separated into two lateral sclerites and
a biarmed sclerite. The munda- group is characterized by having a
shallow depression or no depression on the dorsum of the pronotum
and the gonopore armature consisting of a biarmed sclerite.
Both external and genitalic characters were used in defining species.
The tegmina of the various species are very similar, but differences in
the gonopore armature were noted and can be used for identification.
The gonopore armature is best seen when the genitalia are withdrawn
and cleared in KOH and then examined with a compound microscope.
For best viewing the internal sac should be teased free of the phallobase
so that it can be viewed from all angles. The internal sac in all species
has an area of small, anteriorly projecting spines completely encircling
it. The aedeagus is bilaterally symmetrical.
The populations of Formicilla munda in the United States vary in
color, which resulted in the species F. gilensis and F. evanescens being
1 University of Arizona Agricultural Experiment Station Journal Series ff: 2024.
The Pan-Pacific Entomologist 49: 31-42. January 1973
32
The Pan-Pacific Entomologist
designated by Casey. The main difference was that F. munda and
F. evanescens were relatively light in color while the intervening F.
gilensis was darker. More extensive collecting has revealed that light
populations occur in Texas, along the lower Colorado River, the San
Joaquin Valley of California and in Nayarit, Mexico. Dark populations
occur in the Sacramento Valley of California, from Arizona to Sinaloa,
Mexico and in Morelos, Mexico. Areas of intergradation of color were
found in all cases. Examination of the gonopore armature of specimens
from all populations revealed they are similar and distinct from all
other species. The anatomical similarities and the devaluation of color
led me to conclude that the names gilensis and evanescens should be
treated as synonyms of F. munda.
The holotypes are to be deposited in the California Academy of
Sciences.
I would like to thank Dr. Floyd Werner for his help and encourage¬
ment in the preparation of this paper.
Key to the Species of Formicilla
1. Dorsum of pronotum with deep depression at area of constriction (Fig. la) ;
pubescence elevated at 30 degree angle or greater; gonopore armature with
lateral sclerites separated from biarmed sclerite. Northeastern Argentina.
{bruchi- group) _ bruchi Pic
Dorsum of pronotum with shallow or no depression at area of constriction
(Fig. lb) ; pubescence variable; gonopore annature consisting of biarmed
sclerite {munda- group) _ 2
2. Pubescence on flanks of elytra approximately two-thirds as long as tactile
setae, elevated at 30 degree angle. Westcentral Mexico _
_ longipilosa Chandler n. sp.
Pubescence on flanks of elytra appressed to elevated at 30 degree angle, but
always distinctly less than half the length of tactile setae _ 3
3. Pubescence appressed, short; elytral postbasal transverse impression indis¬
tinct (Fig. 2b). Venezuela to Argentina _ grandiceps Pic
Pubescence raised from surface at 10 to 30 degree angle; elytral postbasal
transverse impression distinct in lateral view (Fig. 2a) _ 4
4. Elytral markings in form of spots, posterior marking small; gonopore arma¬
ture thick in lateral view and with broad projections (Fig. 6). Central
to Northcentral Mexico . vitrea Chandler n. sp.
Elytral markings in form of complete bands across elytra; gonopore armature
narrow in lateral view (Figs. 5, 8) _ 5
5. Eyes large, round except truncate at base of mandibles (Fig. 3a) ; gonopore
armature in lateral view with no projections visible. Michoacan, Mexico
_ alta Chandler n. sp.
Eyes smaller, round to oval (Fig. 3b) ; gonopore armature with distinct
projections in lateral view. Central Mexico to California and Texas
_ munda LeConte
Vol. 49, No. 1 , January 1973
33
Fig. 1. Lateral view pronotum of: a. F. bruchi ; b. F. munda. Fig. 2. Lateral
view elytra: a. F. munda; b. F. grandiceps. Fig. 3. Eye of: a. F. alta; b. F. munda.
Fig. 4. Lateral view F. munda. Figs. 5-6. Genitalia: a. ventral view, b. lateral
view; gonopore armature: c. ventral view, d. lateral view. Fig. 5. F. munda.
Fig. 6. F. vitrea.
34
The Pan-Pacific Entomologist
Formicilla bruchi Pic
(Figs, la, 10)
Formicilla Bruchi [sic] Pic 1904, 329 (Type loc.: Prov. Buenos Aires, Argentina) ;
1911, 22. Bruch 1917, 463; 1928, 193.
Formicilla bruchi, Werner 1965, 3, 11.
Polished and sparsely punctate, head and pronotum castaneous with appendages
of similar color, elytra fuscorufous with luteous marking on area of postbasal
transverse impression of each elytron, sometimes with lighter area around scutellum
but not reaching elytral markings.
Male.— Llavalol, Argentina: 2.61 mm long, head rounded at base, width behind
eyes 0.44 mm, width at antennal insertions 0.28 mm, length 0.60 mm, eyes oval,
broadest dorsally 0.10 X 0.13 mm, antennae moderately pubescent, segments II—IV
narrow, VI-XI gradually increasing in width; head very sparsely pubescent, with
few tactile setae.
Pronotum polished, sparsely pubescent with few tactile setae, pubescence
appressed, pronotum deeply constricted in basal half, with saddle-like depression
on dorsum, pronotal length 0.69 mm, width at base 0.30 mm, width at constriction
0.19 mm, distance from base to constriction 0.20, greatest width 0.42 mm at 0.47
mm from base.
Elytra polished with distinct pubescence at 30 degree angle or greater, length
0.08 mm, tactile setae length 0.16 mm, length of elytra 1.32 mm, width at humeri
0.60 mm, greatest width 0.77 mm at 0.51 mm from base, marking 0.20 mm from
base, 0.11 mm long, elytral depression indistinct in lateral view.
Mesepisternal setae reaching humeral angle, length 0.14 mm, profemur swollen,
length 0.41 mm, width 0.13 mm; protibia slender, length 0.43 mm; mesofemur
swollen in distal two-thirds, length 0.47 mm, width 0.12 mm; mesotibia length
0.42 mm, metafemur swollen in distal one-half, length 0.61 mm, width 0.12 mm,
metatibia length 0.58 mm.
Gonopore armature with a biarmed sclerite with very short stem, an additional
sclerite projects anteriorly from distal portion of each arm, reaching mouth of
primary gonopore. In lateral view biarmed sclerite curved, lateral sclerite straight.
Female.— Similar to male.
Distribution. —Northeastern Argentina in Buenos Aires Province;
one doubtful record from Chile in the Bowditch Collection (Harvard).
ARGENTINA: Buenos Aires: Llavalol, 1906, R. Thaxter; vie. La Plata, 2
January 1963, F. G. Werner; Prov. Buenos Aires, 26 August 1917, C. Bruch.
CHILE: Chi., F. C. Bowditch Collection.
Formicilla grandiceps Pic
(Figs. 2b, 7)
Formicilla grandiceps Pic 1913, 130 (Type-loc.: Chaco du Paraguay). Bruch 1928,
192-193.
Formicilla Garciai [sic] Pic 1948, 15-16 (Type-loc.: Buenos Ayres, Argentina).
NEW SYNONYMY.
Formicilla garciai, Werner 1965, 4, 8.
Vol. 49, No. 1, January 1973
35
Length 1.80-2.00 mm, polished, sparsely punctate, head and pronotum castaneous,
elytra slightly darker with two luteous markings on each elytron, both markings
sometimes reaching sides but only anterior markings may meet at elytral suture,
anterior marking in elytral depression.
Male. —Salobra, Brazil: length 1.84 mm, head subtruncate at base, length of
head 0.42 mm, width behind eyes 0.33 mm, width at antennal bases 0.17 mm,
eyes oval, broadest dorsally 0.07 X 0.08 mm, antennae moderately pubescent,
antennal segments II-IV slender, V-XI slowly increasing in width; head very
sparsely pubescent and with few tactile setae.
Pronotum constricted in basal half, very shallow depression on dorsum, sparsely
setate and pubescent, pubescence appressed, length of pronotum 0.45 mm, width
of constriction 0.15 mm, distance from base to constriction 0.10 mm, greatest
width 0.28 mm at 0.29 mm from base, width at base 0.21 mm.
Elytra polished with numerous tactile setae and with pubescence decumbent,
length of pubescence 0.04 mm, length of tactile setae 0.12 mm, length of elytra
0.97 mm, width at humeri 0.42 mm, greatest width 0.58 mm at 0.46 mm from
base, elytral depression indistinct.
Metepisternal setae reaching humeral angle, 0.05 mm long, profemur swollen,
length 0.33 mm, width 0.12 mm, protibia slightly dilated to apex; 0.24 mm long,
mesofemur swollen in distal two-thirds, 0.34 mm long, width 0.08 mm; mesotibia
slender, 0.29 mm long, metafemur swollen in distal one-half, length 0.33 mm,
width 0.07 mm, metatibia 0.39 mm long.
Gonopore armature with tips broadened, in lateral view curved and appearing
spoon-shaped.
Female. —Similar to male, but longer.
This species is most similar to F. longipilosa because of the shallow
pronotal depression and the indistinct elytral depression.
Distribution.— From northern Argentina and southcentral Brazil
through the higher lands of the Amazon drainage to Venezuela.
ARGENTINA: Misiones: Iguazu, 20 January/13 March 1945, Hayward, Willink,
Golbach. BOLIVIA: Sta. Cruz : El Cidral, ca. 100 km. NNW Sta. Cruz, 450 m.,
UV It. trap, January 1962, R. Golbach. BRAZIL: Mato Grosso: Salobra, January
1955, P. Pereira; Xingu National Park, Jacare, November 1961, M. Alvarenga, W.
Bokermann; Cuyaba. VENEZUELA: Miranda : Agua Fria, El Jacillo, 14 July
1964, 1500777, C. J. Rosales, P. J. Salinas. Portuguesa: Guanare, 10/13 September
1957, Borys Malkin.
Formicilla longipilosa Chandler new species
(Fig. 9)
Polished, sparsely punctate, head and pronotum testaceous, elytra same color
with humeri and two markings on each elytron brown, markings oval, not reaching
flanks of elytron, posterior markings joining at elytral suture, appendages similar
in color to pronotum.
Holotype male. —Length 2.12 mm, head evenly rounded when viewed in plane
from base to apex, width behind eyes 0.34 mm, width at antennal insertions 0.19
mm, length 0.49 mm, eyes rounded, truncate toward base of antennae, broadest
dorsally 0.11 X 0.12 mm. Antennal segments moderately pubescent, pubescence
36
The Pan-Pacific Entomologist
Vol. 49, No. 1, January 1973
37
increasing distally, segments II-V narrow, YI-XI gradually increasing in width,
head very sparsely pubescent and with few tactile setae.
Pronotum polished, with both sparse pubescence and tactile setae, deeply con¬
stricted in basal half, dorsum almost level, length 0.57 mm, width at base 0.25 mm,
width at constriction 0.22 mm, length from constriction to base 0.16 mm, greatest
width 0.33 mm at 0.38 mm from base.
Elytra polished with distinct pubescence and numerous tactile setae, pubescence
at 30 degree angle from elytra, longest along flanks 0.09 mm, tactile setae length
0.12 mm, row of distinct punctures adjacent to and paralleling elytral suture,
indistinct elytral depression, length of elytra 1.12 mm, width at humeri 0.45 mm,
greatest width 0.60 mm at 0.41 mm from base, first marking 0.30 mm from base,
0.21 mm long, second marking 0.70 mm from base and is 0.19 mm long.
Mesepisternal setae not reaching humeral angle, length 0.10 mm, profemur
swollen, length 0.30 mm, width 0.08 mm, tibia slender, length 0.33 mm; mesofemur
swollen in distal two-thirds, length 0.41 mm, width 0.07 mm, mesotibia length 0.34
metafemur swollen in distal one-half, length 0.52 mm, width 0.08 mm, meta-
tibia straight, length 0.04 mm.
Gonopore armature narrow with indistinct projections, no widened areas, in
lateral view, a narrow triangle.
Female.- —Similar to male, but with mesepisternal setae reaching humeral angle,
head sub truncate. Females in type series are slightly shorter than the male,
1.98-2.07 mm.
This species is most similar to F. grandiceps by its indistinct elytral
depression and very shallow depression on the dorsum of the pronotum.
However, the gonopore armature is most similar to that of F. alta, not
broadened anteriorly as in F. grandiceps.
Holotype male , Chilpancingo, Guerrero, Mexico, D. M. Delong,
1941, at light. Paratypes: 3 females, eutopotypical.
Formicilla munda LeConte
(Figs, lb, 2a, 3b, 4, 5, 11)
Formicilla munda LeConte 1851, 152-153 (Type loc.: on lower part of Colorado
River). Casey 1895, 646. Pic 1911, 22. Beal 1950, 126.
Anthicus [ Formicomus , Formicilla] mundus, LeConte 1852, 95.
Formicilla evanescens Casey 1895, 646-647 (Type loc.: Austin, Texas). Pic 1911,
22. Beal 1950, 128-129 ( Formicilla scitula evanescens in part). NEW
SYNONYMY.
Formicilla gilensis Casey 1895, 647 (Type loc.: Tucson, Arizona). Pic 1911, 22.
Beal 1950, 125-126. NEW SYNONYMY.
<e-
Figs. 7-9. Genitalia: a. ventral view, b. lateral view; gonopore armature: c.
ventral view, d. lateral view. Fig. 7. F. grandiceps. Fig. 8. F. alta. Fig. 9. F.
longipilosa. Fig. 10. F. bruchi genitalia: a. ventral view, b. lateral view; gonopore
armature: c. lateral view; internal sac: d. ventral view.
38
The Pan-Pacific Entomologist
Formicilla munda munda, Beal 1950, 126-127. NOT: Formicilla munda gracilipes,
Beal 1950, 127-128.
Color of head and pronotum varying from luteous to fulvous, head sometimes
darker than pronotum, elytral markings always luteous with elytral basement color
varying from fuscopiceous to brown, light markings vary in width depending on
darkness of specimen, markings wider in lighter specimens and narrower in darker
specimens, markings of lighter specimens may join on elytral flanks, first marldng
continuous across elytra while second marking continuous only in lighter speci¬
mens. Elytral depression distinct, shape of base of head varies from subtruncate
to rounded and pubescence on pronotum and elytra varies from appressed to elevated
at 30 degree angle, length 1.80-2.40 mm.
Male. —Hermosillo, Mexico: 2.31 mm long; base of head subtruncate, sparsely
pubescent and setate, pubescence appressed, length of head 0.52 mm, width behind
eyes 0.41 mm, width at antennal bases 0.22 mm, eyes oval, broadest dorsally 0.11 X
0.12 mm, antennal segments II-VI slender, VII-XI gradually increasing in width,
antennae moderately pubescent, pubescence increasing in distal segments.
Pronotum deeply constricted in basal half, dorsum varies from slightly concave
to leve], sparsely setate and pubescent, pubescence depressed to elevated, length
of pronotum 0.55 mm, width at constriction 0.25 mm, length from base to con¬
striction 0.15 mm, widest anteriorly 0.42 mm at 0.39 mm from base, width at base
0.32 mm.
Elytra polished with pubescence varying from depressed to elevated at 30 degree
angle, length of pubescence 0.06 mm, tactile setae length 0.12 mm, length of elytra
1.26 mm, width at humeri 0.51 mm, maximum width 0.69 mm at 0.48 mm from
base, distance from base to first marking 0.13 mm, length of marking 0.13 mm,
distance from base to posterior marking 0.65 mm, length 0.19 mm.
Mesepisternal setae reaching humeral angle, length 0.14 mm, profemur swollen,
length 0.38 mm, width 0.11 mm, protibia 0.37 mm long; mesofemur swollen in
distal two-thirds, length 0.40 mm, width 0.09 mm; mesotibia 0.41 mm long, meta¬
femur swollen in distal one-half, length 0.56 mm, width 0.11 mm, metatibia 0.56
mm long.
Gonopore armature biarmed with median projection on each arm reaching to
primary gonopore; in lateral view projections very distinct.
Female. —Similar to male.
This species is most similar to F. alta because of the distinct elytral
depression and the gonopore armature having a median projection,
although the gonopore armature is slender and not so developed as
in F. alta.
Division into subspecies has been considered for F. munda. However,
I feel this classification would not be realistic because the different
colored populations are separated into several disjunct populations
(Fig. 11).
Distribution. —This species ranges from Morelos, Mexico, through
the lower elevations of Sonora and Arizona into the Central Valley of
California and east to Arkansas.
Vol. 49, No. 1, January 1973
39
UNITED STATES: Arizona: Apache Co.: White Mtns. Cochise Co.: Willcox.
Coconino Co.: Sedona. Gila Co.: Globe, Wheatfields-nr. Globe, Roosevelt Lake.
Graham Co.: Safford, Thatcher. Maricopa Co.: Buckeye, Gillespie Dam-Gila
Bend, Phoenix, Wickenburg. Pima Co.: Brown Cyn.-Baboquivari Mtns., Green
Valley, Picture Rock Pass and Ariz-Son. Desert Museum-Tucson Mtns., Robles
Ranch, Sabino Cyn.-Sta. Catalina Mtns., Tucson, 8 mi. N. Vail. Pinal Co.:
Coolidge, Superior. Santa Cruz Co.: Madera Cyn., Nogales. Yavapai Co.: Bloody
Basin, Cottonwood. Yuma Co.: Dateland, Ehrenberg, Laguna Dam, Palomas, Roll,
Yuma. Arkansas: Howard Co.: 5 mi. N. Nashville. Pike Co. California: Imperial
Co.: “Brawleys,” “Fr. Yuma,” Imperial, E. shore Salton Sea. Madera Co.: Visalia.
Merced Co.: 4 mi. S. Dos Palos, Merced. Riverside Co.: Cathedral City. San
Bernardino Co.: 4 mi. S. Colton. San Diego Co.: La Mesa. Tehama Co.: Red
Bluff, 20 mi. NW. Red Bluff, 1 mi. SE. Vina. Colorado: El Paso Co.: Colorado
Springs. Louisiana: Bossier Co.: Bossier Parish. New Mexico: Dona Ana Co.:
Las Cruces, Mesilla Park. Oklahoma: Texas Co.: Texahoma. Texas: Brazos Co.:
College Station-Texas A&M. Brown Co. Comal Co.: New Braunfels. Dallas Co.:
Dallas. Grayson Co.: Denison. Hidalgo Co.: Brownsville. Llano Co.: Llano.
Nueces Co.: Corpus Christi. Patricio Co.: Sinton. Potter Co.: 14 mi. E. Amarillo.
Terrell Co.: Sanderson. Victoria Co. MEXICO: Baja California: La Paz, 4 mi. N.
Todos Santos. Morelos: 5 mi. S. Cuernavaca. Nayarit: Acaponeta, Ixtlan del Rio.
Sinaloa: 6 mi. S. Culiacan, Los Mochis, Mazatlan, 26 mi. N. Pericos. Sonora:
Alamos, Desemboque, 5 mi. N. Guaymas, Hermosillo, 40 mi. N. Hermosillo,
Navajoa, Nogales.
Formicilla alta Chandler new species
(Figs. 3a, 8)
Polished, sparsely pubescent, head and pronotum flavous to tawny, elytra fulvous
to tawny with two castaneous markings on each elytron, both of which may or
may not meet at elytral suture or extend to sides, humeri darkened, elytral depres¬
sion distinct.
Holotype male. —Length 1.98 mm, head rounded at base, pubescence sparse
and appressed, few tactile setae, length of head 0.46 mm, width behind eyes 0.33
mm, width at antennal bases 0.18 mm, eyes rounded and truncate close to mandibles
0.08 X 0.09 mm, antennae moderately pubescent, pubescence increasing distally,
segments II-IV of equal width, VII-XI gradually increasing in width.
Pronotum polished, sparse pubescence appressed, few tactile setae; deeply con¬
stricted in basal half with dorsum flat, lengh 0.48 mm, width at base 0.26 mm, width
at constriction 0.20 mm, length from base to constriction 0.14 mm, greatest width
0.32 mm at 0.33 mm from base.
Elytra polished with pubescence varying from depressed to elevated at 30 degree
angle, length pubescence 0.06 mm, length tactile setae 0.12 mm, length elytra 1.04
mm, width at humeri 0.50 mm, greatest width 0.58 mm at 0.58 mm from base, first
marking 0.31 mm from base and 0.27 mm long, second marking 0.72 mm from
base and 0.14 mm long.
Mesepisternal setae reaching humeral angle, length 0.15 mm, profemur swollen,
length 0.34 mm, width 0.11 mm, protibia 0.31 mm long; mesofemur swollen in
distal two-thirds, length 0.38 mm, width 0.10 mm, mesotibia length 0.34 mm;
metafemur swollen in distal half, length 0.45 mm, width 0.08 mm, metatibia length
0.43 mm.
40
The Pan-Pacific Entomologist
Gonopore armature slender and curved in lateral view with short, stubby
projections.
Female. —Not known.
The shape of the gonopore armature closely resembles that of F.
longipilosa, but the flat pronotal dorsum and distinct elytral depression
indicate close relationship to F. vitrea.
Holotype male , Mil Cumbres, 30 km. W. Hidalgo, Michoacan,
Mexico, (Rt. 4, Km. 244), 7,600 ft., in grass on edge of road, 29 June
1948, F. G. Werner. Paratype: 1 male, eutopotypical.
Formicilla vitrea Chandler new species
(Fig. 6)
Polished, sparsely pubescent, head and pronotum castaneous, fuscopiceous elytra
with two luteous markings on each elytron, first marking in elytral depression, not
reaching distal edge of elytra but meeting at elytral suture, second marking smaller,
sometimes meeting at elytral suture, elytral depression distinct, length varies from
1.85-2.28 mm.
Vol. 49, No. 1, January 1973
41
Holotype male. —Length 2.17 mm, head subtruncate at base, pubescence sparse
and appressed, few tactile setae, length of head 0.45 mm, width at antennal bases
0.20 mm, width behind eyes 0.39 mm, eyes oval, broadest dorsally 0.07 X 0.08 mm,
antennae moderately pubescent and setate, pubescence increasing distally, seg¬
ments II-V of equal width, VI-XI barely increasing in width.
Pronotum polished, sparse appressed pubescence, few tactile setae; deeply
constricted in basal half with dorsum flat, length of pronotum 0.54 mm, width at
base 0.28 mm, width at constriction 0.24 mm, length from base to constriction 0.14
mm, widest point 0.39 mm at 0.38 mm from base.
Elytra polished with pubescence depressed to elevated at 30 degree angle, length
pubescence 0.05 mm, length tactile setae 0.12 mm, length elytra 1.18 mm, width
at humeri 0.45 mm, greatest width 0.07 mm at 0.49 mm from base, first marking
0.14 mm from base and 0.08 mm in length, second marking 0.63 mm from base
and 0.14 mm in length.
Mesepisternal setae reaching humeri, length 0.14 mm, profemur swollen, length
0.40 mm, width 0.13 mm, protibia slender, length 0.36 mm; mesofemur swollen
in distal two-thirds, length 0.41 mm, width 0.09 mm, mesotibia 0.39 mm long;
metafemur swollen in distal half, length 0.51 mm, width 0.11 mm, metatibia
length 0.50 mm.
Gonopore armature thick in lateral view with large, blunt projections, very
short basal arm.
Female. —Similar to male.
The gonopore armature is very distinct and quite different from
that of the other species. The flat pronotal dorsum indicates affinities
to F. alta.
Holotype male , 20 km. W. Lagos, Jalisco, Mexico, dry agricultural
area (running on damp ground among grass stems next to ditch), 6,130
ft., 2 August 1948, F. G. Werner.
Paratypes. —13 males, 7 females, eutopotypical; 1 male, 49 mi. SE. Chihuahua,
Chihuahua, MEX., 4,000 ft., light, San Pedro River, 20 July 1952, RB & BK
Selander; 1 female, 16 km. W. Jalostotitlan, Jalisco, MEX., 5,920 ft., (Rt. 170),
(Dry grazing area, under human dung), 1 July 1948, W. Nutting.
Literature Cited
Beal, R. S. Jr. 1950. Systematic notes on the genus Formicilla in the United
States and Mexico. Pan-Pac. Entomol., 26: 122-130.
Bruch, C. 1917. Nuevas capturas de insectos mirmecofilos. Physis, Rev. Assoc.
Argent. Cienc. Natur., 3: 458-466.
1928. Suplemento al catalogo sistematico IV, (Addenda, corrigenda, y lista
de especies). Physis, Rev. Assoc. Argent. Cienc. Natur., 9: 186-204.
Casey, T. L. 1895. Coleopterological notices, VI. Ann. N.Y. Acad. Sci., 8: 435-
838.
LeConte, J. L. 1851. Descriptions of new species of Coleoptera from California.
Ann. Lyc. Natur. Hist. N.Y., 5: 125-184.
1852. Synopsis of the Anthicites of the United States. Proc. Acad. Natur. Sci.
Philadelphia, 6: 91-104.
42
The Pan-Pacific Entomologist
Pic, M. 1904. Anthicides nouveaux de la Republique Argentine recueillis par M.
Carlos Bruch. Rev. Mus. La Plata, 11: 329-331.
1913. Nouveaux anthicides exotiques. Echange, 29: 130-132.
1948. Notes et nouveautes diverses. Diversites Entomologiques, 4: 12-16.
Werner, F. G. 1960. A redefinition of the genus Acanthinus as an important
element in the Anthicid fauna of the neotropical region. Verh. XI Int.
Kongr. Ent., Wien, I: 109-112.
1965. A key to the described species of New World Anthicidae of the genus
Acanthinus La Ferte, with new synonymy. Pap. Avulsos Zool. Sao Paulo.
18: 1-15.
1970. A revision of Acanthinus (Coleoptera: Anthicidae) IX. The Leporinus,
Scitulus, Cuyabanus, and Myrmecops-Groups. Ann. Entomol. Soc. Amer.,
63: 718-731.
SCIENTIFIC NOTE
A Note on Dacnochilus fresnoensis ( Coleoptera :Staphylinidae). —Hugh
B. Leech described Dacnochilus fresnoensis in 1939 (Can. Entomol., 71: 261) on
the basis of a male collected at Fresno, California 5 June, 1931 and a female from
the same locality 23 June, 1930, both by Roy S. Wagner. The species has not been
mentioned in the literature since and to my knowledge until now no other specimen
has been found.
Only two other species are known in this genus, the habits of the members of which
are unknown. The closely related Acalophaena is represented by several species
in Latin America one of which, A. hasalis Lynch-Arribalzaga, was reported in 1884
by its describer from termite nests (Bol. Acad. Nac. Cienc. Cordoba, 7: 270).
I have received from Derham Giuliani a single male of Dacnochilus fresnoensis
collected by him at “Riverside, California on August 16, 1971 on log with ants
and termites.” In reply to my request for more information about this rare find
Giuliani wrote:
“Concerning the specimen of Dacnochilus fresnoensis, I collected it at Dr. Fisher’s
house when I was in Riverside last August. You were away on a trip at the time.
He could show you the log and perhaps recall some details I’ve forgotten. I helped
him move the log his wife wanted put at a spot in the garden. It contained large
numbers of ants and termites. Debris with ants and termites fell off here and there
during the move and it was near this that I spotted the staph walking about. I
looked on the log and back at the spot it had been moved from but saw no more.
Unfortunately I did not collect any of the ants or termites even though I felt
certain that the staph had been among them.”
Conversations with Dr. T. W. Fisher (U.C.R.) revealed that the log was one
of three alder logs which had been brought to his home about two years before
from Lytle Creek, San Bernardino County, California about thirty miles to the
northwest of the collecting site. Visits to his place, on a chaparral-covered hill
at Sunnymead, Riverside County, California, were made on 31 April and 17 June,
1972, a year of very little rainfall. No ants or termites were found on or under
the three logs, two of which were still in their original position. No more Dac¬
nochilus were found. —Ian Moore, Division of Biological Control, University of
California, Riverside 92502.
Vol. 49, No. 1, January 1973
43
A New Genus of Pseudoscorpions Based upon
Atemnus hirsutus 1
(Pseudoscorpionida: Chernetidae)
William B. Muchmore
Department of Biology, University of Rochester, Rochester, N.Y. 14627
The species Atemnus hirsutus was described by N. Banks in 1914 on
the basis of a single individual “found 10 feet from ocean” at Laguna
Beach, California. Evidently, no further specimens have been discovered,
and the species has remained an enigma ever since. Beier (1932) and
Roewer (1937) overlooked it completely in their worldwide reviews
of the pseudoscorpions, and Hoff (1958, p. 22) only noted that “From
Banks’ figure, the species probably belongs to an undescribed genus in
the Lamprochernetinae.” Through the courtesy of Dr. H. Levi, I have
borrowed the mounted holotype from the Museum of Comparative
Zoology, Harvard University. I have remounted it and reexamined it
in detail, and find that it belongs to an undescribed genus of the family
Chernetidae.
Mucrochernes Muchmore, new genus
Diagnosis. —-(Based upon female only.) Surfaces generally smooth, but carapace
granulate laterally and posteriorly, and palpal segments each with a few granules.
Setae generally long, thin and tapering to fine point, many with single, small,
sharp spinule at about the middle of the shaft. Carapace a little longer than
broad; with two transverse furrows and two distinct eye spots; covered with about
200 vestitural setae. Tergites and sternites, except 11th, divided; with many setae
arranged in one, two or three irregular rows; sternites 4, 5 and 6 with numerous
small slit sensilla; pleural and interscutal membranes densely papillate; anterior
genital operculum of female with about 45 acuminate setae broadly scattered over
surface. Spermathecae of female apparently consisting of two delicate, lateral
tubes of moderate length and barely enlarged at ends. Genital features of male
unknown. Cheliceral palm with five setae, all long and simple; flagellum of four
setae, all finely denticulate along anterior margins; galea with five branches
emerging from common base, each branch with 2 to 4 rami. Palps fairly heavy;
chelal hand (female) very broad at base; fingers with accessory teeth; venom
apparatus well developed only in movable finger, vestigial in fixed finger;
trichobothrium st near middle of movable finger, closer to t than to sb; ist at
same level as est, distal to middle of fixed finger, esb, eb, isb and ib closely
grouped at base of finger. Legs monotarsate, moderately slender; tibia IV with
tactile seta far distad (also on III) ; tarsus IV with tactile seta just distad of middle
(also on III) and prominent slit sensillum close to base (also on I, II, III) ; sub¬
terminal tarsal setae curved, acuminate; claws simple.
1 This work was supported in part by a grant (GB 17964) from the National Science Foundation.
The Pan-Pacific Entomologist 49: 43-48. January 1973
44
The Pan-Pacific Entomologist
Type species.— Atemnus hirsutus Banks, 1914.
Distribution. —Known only from Laguna Beach, California.
Remarks. —This genus is a member of the family Chernetidae by
virtue of the monotarsate legs, the occurrence of a well developed venom
apparatus in the movable chelal finger only, the accessory teeth on the
chelal fingers, and the presence of a prominent, longitudinally oriented,
slit sensillum basally on each pedal tarsus.
The relationship of Mucrochernes to other genera in the family is
obscure because of the unique nature of the vestitural setae. Some other
genera, such as Dasychernes Chamberlin, have large numbers of setae,
but no other pseudoscorpion has been reported to have setae similar in
shape to those of Mucrochernes . In fact, all other known chernetids
have at least some setae which are denticulate or clavate at the tips,
while those of Mucrochernes are, without exception, simple at the tips.
On the basis of this and other features, the present genus does not fall
naturally into any of the subfamilies of Chernetidae as presently formu¬
lated. It is possible that reevaluation of these groups on the basis of a
broader constellation of characters will reveal close relatives of
Mucrochernes.
Mucrochernes hirsutus (Banks), new combination
(Figs. 1-6)
Atemnus hirsutus Banks, 1914, p. 203.
Genus? hirsutus, Hoff, 1958, p. 22.
Material.— Holotype female (WM 2456.01001), “found 10 ft. from
ocean. No trees near,” at Laguna Beach, Orange County, California
(no date or collector indicated on slide, though Banks states that the
specimen was sent by Prof. Hilton).
Because the original description given by Banks is inaccurate in
some details and generally not sufficient to define the species, the
following more complete description is presented.
Female.— All parts rather lightly sclerotized and light yellowish. Carapace
slightly longer than broad, with two transverse furrows and two large, distinct
eye spots; surface nearly smooth medially, but weakly granulate on sides and
posteriorly; covered with more than 200 setae. Setae on carapace, as well as on
remainder of animal, different from those of other pseudoscorpions—all long, thin
and completely simple at tips, but often with single, small, sharp spinule about
halfway along length of shaft (Fig. 1).
Abdomen twice as long as broad; tergites and sternites, except 11th, divided,
surfaces very finely granulate; pleural membranes and interscutal surfaces densely
papillate. Tergal chaetotaxy 26:29:28:39:51:51:57:60:54:53:43:2; setae in single
marginal row on tergites 1 to 3, but in irregular double or triple rows on tergites
Vol. 49, No. 1, January 1973 45
Fig. 1-6. Mucrochernes liirsutus (Banks), holotype female. Fig. 1. Vestitural
setae with lateral spinules. Fig. 2. Spermathecae. Fig. 3. Tip of movable finger of
chelicera, showing complex galea. Fig. 4. Dorsal view of right palp (chela in
dorsolateral aspect and with crack in hand). Fig. 5. Lateral view of left chela
(hand broken). Fig. 6. Tibia and tarsus of leg IV, showing tactile setae and tarsal
sensillum.
4 to 11. Sternal chaetotaxy 43: (1) 22 (2) : (3) 14(3) :64:68:50:49:48:47:36:2; the
setae in double or triple rows on sternites 5 to 11. Sternites 4, 5 and 6 with
numerous small, slit sensilla scattered over surfaces (each sternal half with about
40 on 4, about 85 on 5, and about 75 on 6), other sternites with few such sense
organs (each half with about 12 on 3, only 2 on 7, 1 on 8, etc.). Anterior genital
46
The Pan-Pacific Entomologist
operculum with a broad field of 43 small setae; posterior operculum with irregular,
marginal row of 22 setae of similar size. Setae on spiracular plates simple.
Spermathecae difficult to observe but appear to consist of two thinwalled lateral
tubes, barely enlarged at ends (Fig. 2). Coxal area generally typical; all coxae
bearing numerous, apparently simple setae.
Chelicera generally typical of the family, about one-third as long as carapace.
Palm with five setae, all relatively long and simple. Fixed finger with three tiny,
subterminal denticles and five teeth of decreasing size along the margin. Movable
finger (Fig. 3) with distinct subapical lobe and one medium sized tooth just
proximad of this. Galea composed of five branches arising from common base,
each branch with two or four rami. Flagellum of four setae, all finely denticulate
along anterior margins. Serrula exterior with 19 blades; serrula interior of six
dentate lobes and basal velum.
Palp essentially as illustrated by Banks (1914, p. 203), though in the copy of the
Journal available to me, the lateral outline of the tibia is incomplete. Remounted
palp (chelal hands slightly broken by original, unsupported cover glass) as in
Fig. 4. Surfaces smooth, except for a few small granules on flexor surfaces of
trochanter, femur, and tibia; covered with numerous setae, mostly simple but
some with spinules on shafts. Trichobothria of chela as in Fig. 5. Chelal fingers
slightly gaping; fixed finger with 63 low, strongly cusped, marginal teeth, and
five internal and eight external accessory teeth; movable finger with 59 similar
marginal teeth, and five internal and seven external accessory teeth. Venom
apparatus well developed in movable finger, with nodus ramosus just proximad
of trichobothrium t; fixed finger without specialized venedens, but with small,
poorly preserved venom duct. Trochanter 1.7, femur 3.1, tibia 2.65, and chela
about 5.2 times as long as broad; hand 1.25 times as long as deep; movable finger
1.17 times as long as hand.
Legs typical in form, rather slender; leg IV with entire femur 4.25, tibia 5.1
and tarsus 4.35 times as long as deep. Tactile setae on tibia 0.93 and on tarsus
0.62 the length of segment from proximal end (Fig. 6). Large, prominent slit
sensillum at proximal fifth of outer margin of segment. Subterminal tarsal setae
curved, simple.
Male.— Unknown.
Measurements (mm).—Body length 3.06. Carapace 0.99 long, posterior breadth
0.89. Chelicera 0.36 long by 0.185 broad; movable finger 0.295 long. Palpal
trochanter 0.54 by 0.32; femur 0.985 by 0.32; tibia 0.925 by 0.35; chela (without
pedicel) 1.725 by 0.33; hand (without pedicel) 0.79 by 0.63; movable finger 0.925
long. Leg I: basifemur 0.25 by 0.17; telofemur 0.48 by 0.15; tibia 0.41 by 0.12;
tarsus 0.43 by 0.095. Leg IV: entire femur 0.85 long; basifemur 0.30 by 0.20;
telofemur 0.63 by 0.20; tibia 0.665 by 0.13; tarsus 0.52 by 0.12.
Remarks. —There are two inaccuracies in Banks’ description of the
species which deserve some comment.
1) As mentioned above, and as observed by Banks, all setae on
the animal are simple at the tips. However, probably because of in¬
sufficient magnification, Banks failed to notice the small spinules on
the shafts of many of the setae. It is impossible to determine how many
setae actually possess this feature because the spinule is usually obscured
Vol. 49, No. 1, January 1973
47
by the shaft of the seta unless the seta is positioned so that the spinule
extends out to one side.
2) It is difficult to understand why Banks recorded “mandible . . .
with a short stylet,” when the cheliceral galea is, in fact, large and
multiply branched. It is conceivable, however, that one of the chelicerae,
which seems to have been treated differently from the other, might have
appeared to have a simple galea when viewed in a certain position
under low magnification.
It is of special interest to note the occurrence of a large, prominent,
slit sensillum basally on the outer surface of each pedal tarsus. The
slit is oriented along the long axis of the segment and opens across the
surface of a rounded elevation. In lateral view, as the legs are usually
oriented, the organ is seen as a distinct, curved bump on the tarsal
surface. This structure apparently was first noted by Chamberlin
(1924), who called it a “domedike ‘sense-spot’later (1938, etc.)
he referred to it as a “sense dome.” Oddly, it is not mentioned at all
in Chamberlin’s large monograph “The arachnid order Chelonethida”
(1931). Vachon (1945) observed that he had found this “organe
tarsal” in representatives of all three subfamilies of the Chernetidae
but in no other family, and he suggested that it is a good diagnostic
character of the family Chernetidae. Unaccountably, then, Vachon did
not include this character in his revised diagnosis of the Chernetidae
(1951, p. 171), though he did mention it again two paragraphs beyond.
My own experience agrees with that of Vachon and I concur that these
slit sensilla are unique to the Chernetidae, the diagnostic features of
which can be listed as follows: all legs monotarsate; venom apparatus
well developed in movable finger of chela, less well developed or absent
in fixed finger; fingers of chela usually with accessory teeth external and
internal to the marginal row; tarsus of each leg with a prominent slit
sensillum on the outer surface prox'mad of the middle of the segment.
It is to be hoped that the foregoing will stimulate enough interest in
this remarkable pseudoscorpion to result in the collection of additional
specimens of the same or related species.
Literature Cited
Banks, N. 1914. A new pseucloscorpion from California. J. Entomol. Zool.,
Pomona Coll., 6: 203.
Beier, M. 1932. Pseudoscorpionidea. II. Subord. Cheliferinea. Teirreich, 58:
1-294.
Chamberlin, J. C. 1924. Hesperochernes laurae, a new species of false scorpion
from California inhabiting the nest of Vespa. Pan-Pac. Entomol. 1:
89-92.
48
The Pan-Pacific Entomologist
1931. The arachnid order Chelonethida. Stanford Univ. Publ. Biol. Sci. 7,
no. 1: 1-284.
1938. New and little-known false scorpions from the Pacific and elsewhere.
Ann. Mag. Nat. Hist., ser. 11, 2: 259-285.
Hoff, C. C. 1958. List of the pseudoscorpions of North America north of Mexico.
Amer. Mus. Novitates, 1875: 1-50.
Roewer, C. Fr. 1937. Chelonethi order Pseudoskorpione. In Bronns, H. G.,
Klassen und Ordnungen des Tierreichs, Leipzig, vol. 5, div. 4, book 6,
fasc. 2, pp. 161-320.
Vachon, M. 1945. Chernetes. Mem. Mus. Nat. Hist. Natur. Paris, Ser. A, Zool.,
19: 187-197.
1951. Les Pseudoscorpions de Madagascar. I. Remarques sur la famille des
Chernetidae J. C. Chamberlin, 1931, a propos de la description d’une
nouvelle espece: Metagoniochernes Milloti. Mem. Inst. Rech. Sci.
Madagascar, ser. A, Biol. Anim., 5: 159-172.
COLLECTION DONATED
Chris Parrish Scorpion Collection Donated to California Academy of
Sciences. —Mr. Chris Parrish of La Mesa, California, has recently donated his
collection of scorpions and other arthropods to the Department of Entomology,
California Academy of Sciences. This collection, consisting of 3,784 specimens
(plus a collection of diplocentrid scorpions out on loan for study) is important
and valuable since it contains series of scorpions both from the mainland of Baja
California, Mexico and the islands of the Gulf of California. The majority of the
Mexican collections were made on expeditions organized by Dr. George E. Lindsay,
Director of the California Academy of Sciences.
The Parrish collection contains 2,241 scorpions (plus the diplocentrid scorpions)
including seven holotypes and two allotypes described by Dr. Herbert L. Stahnke
(holotype female, Bioculus aguajensis; holotype female, allotype male, B. belvederi;
holotype female, B. danzantiensis ; holotype female, B. lindsayi; holotype female,
B. luteus ; holotype female, B. parraensis; holotype male, allotype female, B.
parrishi ), 1,399 specimens from Mexico, and 833 from the western United States.
Most of these specimens were collected by non-ultra-violet detection methods. They
will complement the extensive series of scorpions which Dr. Stanley C. Williams,
who is monographing the scorpions of Baja California, has collected and donated
to the Academy. The collection also contains 36 Uropygi and Amblypygi, 42
Solpugida, 61 Chilopoda, 50 Diplopoda, 128 Opiliones, 74 Chelonethida, 599
Araneida, 1 Onychophora, 276 Acarina, and 266 Insecta. —Paul H. Arnaud, Jr.,
California Academy of Sciences, San Francisco, 94118.
Vol. 49, No. 1, January 1973
49
A New Pulverro from California
(Hymenoptera: Sphecidae)
Lloyd E. Eighme
Pacific Union College, Angwin, California 94508
A study of the genus Pulverro has disclosed some interesting new
material especially from the North Coast Ranges of California. A new
species, Pulverro monticola Eighme was described from specimens col¬
lected in the Salmon-Trinity Mts. in 1968. Subsequent collecting trips
in the same area have resulted in specimens of another new species
described in this paper.
Specimens of the new species were first collected by the author in
August 1969 in the Deer Creek Basin at 5,300 feet elevation in the
Salmon-Trinity Mts. of Trinity County, California. The following sum¬
mer, June 1970, one of my students, Terry Griswold, collected specimens
of the same species near Healdsburg, Sonoma County, and in July 1971,
another student, Earl Aagaard, collected more specimens of the same
species in Young’s Valley at 4,600 feet elevation in Siskiyou County.
Dr. R. M. Bohart (Dept, of Entomology, Univ. of Calif., Davis) dis¬
covered eight specimens in the U. C. Davis collection from Shasta County
and recognized them as a new species, but graciously loaned them to
me as a part of the series to be described in this paper.
The holotype of this new species will be deposited in the collection at
the California Academy of Sciences. Paratypes will be placed in collec¬
tions at the University of California at Davis, the U.S. National Museum,
and the Philadelphia Academy of Sciences.
Pulverro californica Eighme, new species
Male.— Length 3 mm. Color, black; foretibia and tarsi, apex of fore femur, mid
and hind tarsi and base of mid and hind tibiae golden brown; labrum, palpi and
mandibles ivory except apices and tip of ventral tooth of mandibles red; apices of
antennal segments brown underneath. Wings clear hyaline, stigma and veins dark
brown.
Head subquadrate when viewed from above, narrower in occipital width than
ocular width. Mandibles bidentate with distinctly pointed midventral tooth placed
at right angle to ventral margin. Clypeus with distinctly raised medial subquadrate
platform as broad as the distance between the antennal sockets. Ventral margin
of clypeus with sharply angled median truncate lobe. Clypeus and front adjacent
to antennal sockets densely clothed with flattened silvery pubescence. Remainder
of front and vertex finely granular with few scattered fine hairs. Supraorbital
fovea a minute round puncture at apical margin of eye. Antenna filiform with
first segment of flagellum greater in diameter than second segment.
The Pan-Pacific Entomologist 49: 49-50. January 1973
50
The Pan-Pacific Entomologist
Thorax shining, finely punctate. Scutum projected forward above pronotum,
smoothly rounded anteriorly and laterally with notauli prominent anteriorly for
almost half length of scutum. Lateral furrow between scutum and scutellum deep
and lightly foveolate. Propodeum granulate on sides, dorsal face coarsely reticulate,
posterior face flattened with median furrow and dorsolateral margins outlined by
sharp carina.
Abdomen shining, minutely rugulose. Tergites 2-5 with prominent transverse
groove at middle, appearing as secondary division of somite. Posterior border of
tergites 2-5 depressed and marked by narrow row of irregular punctures.
Last tergite coarsely punctured and armed with short stiff bristles. Second
sternite with transverse arcuate groove anteriorly, third sternite with straight
transverse groove. Hypopygium (last sternite) spatulate, apical margin rounded,
sharply serrate with tuft of fine pubescence underneath, silvery hairs less than
one-half as long as hypopygium.
Female. —Similar to male except: Central clypeal disc roundly tumid, extending
upward between antennal sockets and projecting anteriorly into broad subtruncate
flange. Clypeal disc smooth, shining, with large scattered punctures. Labrum
not extending beyond clypeal flange. Mandibles bidentate but lack midventral
tooth. Supraorbital fovea linear, as long as diameter of lateral ocellus. Frons with
faint median carina terminating in shallow pit about % distance from clypeus to
median ocellus. Pygidial plate trigonal, margined by sharp carina, apex narrowly
truncate, shining, with few large punctures.
Holotype male , (CAS), Deer Creek, 5,300 feet, Trinity County,
California, 7 August 1969 (L. Eighme). Paratypes, 7 males, 5 females
from same locality as holotype. Additional specimens: 16 males and
4 females from Siskiyou County, 5,000 feet, California; 3 males and
5 females from Shasta County, California; 4 males and 1 female from
Healdsburg, Sonoma County, California.
This species is similar to P. chumashano Pate which is found in
coastal southern California, but strikingly different from P. monticola
Eighme which is found in some of the same areas as the new species.
Pulverro californica differs from P. chumashano in the males in that
the midventral mandibular tooth is smaller and more pointed, the last
sternite is evenly rounded instead of broadly truncate, and sparsely
hirsute instead of densely hirsute, the integument is generally black
instead of dark mahogany. The females of P. californica differ from
those of P. chumashano in that the labrum does not extend beyond the
clypeal flange in P. californica whereas it does in P. chumashano.
Literature Cited
Eighme, L. E. 1968. A New Species of Pulverro from California. Pan-Pac.
Entomol., 44(4) : 261-264.
Vol. 49, No. 1, January 1973
51
Description of a New Species of Trjapitzinellus Parasitic on
Coniopterygidae in California 1 ’ 2
(Hymenoptera: Encyrtidae; Neuroptera)
Gordon Gordh
Department of Entomology, University of California, Riverside, 92507
Slides in the collection of the Citrus Experiment Station, University
of California, Riverside, indicate the presence of a new species of
encyrtid from California as suspected by P. H. Timberlake in 1928.
Subsequent collections by Paul DeBach and Harold Compere in south¬
ern California during the 1930’s and 1940’s revealed more individuals
similar to Timberlake’s material. Intensive collecting by J. A. McMurtry
during the 1960’s associated the parasite with species of Conwentzia
and Parasemidalis (Coniopterygidae). Owing to the taxonomic status
of the Bothriothoricini (= Mir ini Ashmead), no attempt was made by
Timberlake, Compere, or DeBach to describe the species. Recently,
Viggiani (1967) described a genus of encyrtid parasitizing Coniopterygi¬
dae in southern Italy which is congeneric with the material collected
by DeBach, Compere, and Timberlake. Research by R. Malone on
synecological aspects of Trjapitzinellus microrphanos , sp. nov., neces¬
sitates description of this species.
Trjapitzinellus microrphanos Gordh, new species
Female. —Body length 2.3 mm. Head, mesosoma, gaster blackish brown; malar
space, face with mauvish refringence; antennal scape reddish brown, remaining
segments lighter; wings hyaline, without infuscation; coxae, femora, basal half of
tibiae concolorous with body; trochanters, apical portion of tibiae, tarsi tan.
Head hypognathous, in dorsal aspect more than twice as wide as median length;
anterior margin transverse; posterior margin concave between compound eye;
frontovertex (at narrowest) about % head width; ocelli forming acute triangle,
lateral ocelli touching inner margin of compound eyes; median ocellus in center
of frontovertex lateral ocelli less than diameter of one ocellus from posterior
margin. Frontal aspect of head (Figs. 4, 5) triangular, compound eye height
slightly greater than malar space; toruli at ventrolateral margin of clypeus, sepa¬
rated by 4 times diameter of single torulus; scrobe indistinct, interantennal ridge
not conspicuous; inner margin of compound eye with conspicuous line of setae
extending along inner margin of subocular suture, ending beneath torulus. Antenna
11-segmented (Fig. 2) ; scape with ventral flange; pedicel twice longer than first
1 1 am grateful to D. P. Annecke, Plant Protection Institute, Pretoria, Republic of South Africa,
for comparing the material with his collection of African Encyrtidae and V. Trjapitzin, Soviet
Academy of Science, Leningrad, for comparison with the Palearctic material at his disposal; Harold
Compere kindly provided access to his extensive collection of Encyrtidae. Professor C. D. Michener,
University of Kansas, Lawrence, reviewed the manuscript and suggested improvements.
2 Support of NSF GB-17829 to Paul DeBach is gratefully acknowledged.
The Pan-Pacific Entomologist 49: 51-55. January 1973
52
The Pan-Pacific Entomologist
flagellomere; flagellomeres uniformly setose with few carinae, becoming progres¬
sively larger distally; club compact, setose, with curved sensory margin dorsad.
Head lightly shagreened, compound eye with uniform covering of short setae;
mandibles tridentate; maxillary palp 4-segmented (Fig. 7), labial palp 3-segmented
(Fig. 6).
Mesosoma (Fig. 8) twice as long as wide; dorsal surface flattened. Mesoscutum,
scutellum, axilla lightly shagreened; tegula strongly shagreened, setose; propodeum
with few coarse striations; mesoscutum with uniform vestiture of pale thin setae;
scutellum with longer, more irregularly dispersed setae; axilla, metanotum,
propodeum asetose. Legs not modified; middle tibial spur (Fig. 3) % length of
basitarsus and subequal length. Forewing (Fig. 9) about 2% times longer than
greatest width, lacking infuscation; submarginal, marginal vein limits not evident,
forming single straight vein, with marginal vein distinct distally; costal cell well-
developed, setose; postmarginal vein at least as long as stigmal vein, terminus
indistinct; speculum present, but lateral margins not straight or distinct; hind¬
wing (Fig. 10) with submarginal, marginal veins subequal, submarginal vein
arched forward; entire wing surface distal to imaginary line perpendicular to
union of marginal and submarginal veins covered with setae of uniform length.
Gaster shorter than mesosoma, rounded at apex, ovipositor and gonostylus not
visible from above; ovipositor and hind tibia subequal in length.
Male. —Body length 2.0 mm. Color as female except antenna light brown with
reddish refringence. Head shape as female; ventral margin of antennal torulus
above clypeal margin by height of one torulus. Antenna (Fig. 1) 9-segmented;
six flagellomeres subequal in size, twice as long as wide. Aedeagus exterted sub¬
equal in length to hindbasitarsus and second tarsomere.
Biological associations. —Fleschner and Ricker (1953) indicate
four genera of coniopterygids are found in southern California, Para-
semidalis , Conwentzia, Coniopteryx , and Malacomyza. Parasemidalis
flaviceps Banks and Conwentzia nigrans Carpenter are most abundant
in citrus groves. Preliminary studies by McMurtry indicate T. mi-
crorphanos parasitizes Conwentzia sp. and Parasemidalis sp. during
the larval stage before the cocoon is spun. The parasite pupates within
the prepupal skin of its host.
Comparative comments.— In his generic description, Viggiani
(1967) relied upon the observations of Trjapitzin in establishing af¬
finities between Trjapitzinellus and related bothriothoracines. Trjapitzin
supposed the genus to be related to Aphidencyrtus Ashmead, Aphiden-
Fig. 1. Inner aspect left antenna $ T. microrphanos (X45). Fig. 2. Inner
aspect left antenna $ T. microrphanos (X 45). Figs. 3-10. 2 T. microrphanos
(X40): Fig. 3. Right tibiotarsus, Fig. 4. Head anterior aspect, Fig. 5. Head
lateral aspect, Fig. 6. Left labial palpus, Fig. 7. Left maxillary palpus, Fig. 8.
Dorsal view of meso- and metasoma, Fig. 9. Right forewing, and Fig. 10. Right
hindwing.
Vol. 49, No. 1, January 1973
53
54
The Pan-Pacific Entomologist
cyrtoides Ishii, Baeonusia Girault, Geniaspidius Masi, Mercetencyrtus
Trjapitzin and Ooencyrtus Ashmead. Viggiani added that on the basis
of body shape Trjapitzinellus is intermediate between Ooencyrtus and
Psyllaephagus. Comparison of T. microrphanos with Compere’s collec¬
tion of encyrtids confirms Viggiani’s suspicions. However, the sys¬
tematic position of Trjapitzinellus is still unsatisfactory and revision
of the Bothriothoracini seems essential before relationships are estab¬
lished. Perhaps the most distinctive feature of T. semidaliphagus
Viggiani and T. microrphanos is the dorsal curved sensory margin of
the female’s antennal club.
Biologically, T. semidaliphagus has been recovered from Semidalis
aleurodiformis Steph. in southern Italy only.
Both sexes of T. microrphanos may be distinguished from T. semi¬
daliphagus on the basis of the following characters: (1) the distal end
of the marginal vein of the forewing is broken in T. microrphanos , but
continuous in T. semidaliphagus ; (2) the forewing of T. microrphanos
bears setal tracts absent from T. semidaliphagus ; and (3) T. semi¬
daliphagus lacks the conspicuous line of setae extending along the
inner margin of the compound eye and subocular suture. The second
segment of the female maxillary palpus of T. microrphanos is glabrous
while the corresponding segment of T. semidaliphagus bears setae.
Material examined.—California: San Diego Co., 13 9,8 £, Pauma Valley,
26 March 1963, leg. J. A. McMurtry ex. coniopterygid from citrus; 11 9,3 £,
El Toro, 20 July 1964, J. A. McMurtry from coniopterygid on citrus. Orange Co.:
17 9,7 £, Santa Ana Canyon, 9 July 1963, from coniopterygid on citrus, J. A.
McMurtry. Santa Barbara Co.: 1 9,2 £, 6 February 1964, Carpinteria, from
coniopterygid on Avocado, L. Bailard; 5 9,3 £, Santa Barbara, 25 July 1964,
from coniopterygid on citrus, J. A. McMurtry. Ventura Co.: 5 9,3 £, Ladera
Ranch, Fillmore, 16 May 1963 and 2 August 1963, from coniopterygid on citrus,
J. A. McMurtry. San Bernardino Co.: 4 9,2 £, Redlands, 24 July 1963, from
coniopterygid on citrus, J. A. McMurtry. Riverside Co.: 4 9,1 £, Riverside,
10 May 1963, from coniopterygid on citrus, J. A. McMurtry.
Holotype female and allotype , Ladera Ranch, Fillmore, Ventura
County, California 2 August 1963, Coniopterygidae on citrus, J. A.
McMurtry, Collector, both card-point mounted.
Paratypes. —3 9 and 3 £, Santa Ana Canyon; 1 9, Riverside; 7 9 and 4 £,
Pauma Valley; 1 9, Redlands; 1 9, Santa Barbara; 1 £, Fillmore; all card-
point mounted.
Disposition of types. — (Names in parentheses indicate individual to whom
material was sent.) Holotype, Allotype, 1 9 and 1 £ Paratypes, Snow Ento¬
mological Museum, University of Kansas, Lawrence, Kansas (G. W. Byers).
Paratypes have been assigned to the following institutions: 2 9 and 1 £, U.S.
Vol. 49, No. 1, January 1973
55
National Natural History Museum (B. D. Burks) ; 1 2 and 1 $, Plant Protection
Research Institute, Pretoria, Republic of South Africa (D. P. Annecke) ; 1 2
and 1 $, British Museum (Natural History), London, England (G. A. Kerrich) ;
1 2 and 1 $ , Zoological Institute, Soviet Academy of Sciences, Leningrad, USSR
(V. A. Trjapitzin) ; 1 2 and 1 $, Department of Entomology, University of
California, Riverside, California.
Literature Cited
Fleschner, C. A. and D. W. Ricker. 1953. Food habits of coniopterygids on
citrus in southern California. Econ. Entomol., 46: 458-461.
VlGGiANi, G. 1967. Richerche sugli Hymenoptera Chalcidoidea XIII. Sugli ento-
moparassiti della Semidalis aleurodiformis Steph. (Neur. Coniopterygi-
dae), con descrizione di un nuovo genere di Encyrtidae. Boll. Lab.
Ent. Agr. Portici, 25: 164M.75.
BOOK REVIEW
Principles of Insect Chemosterilization. Edited by Germain C. La Brecque
and Carrol N. Smith. Appleton-Century-Crofts, New York. 354 p., 13 figs.,
32 tables. 1968. $16.00.
This is the first comprehensive survey of chemosterilization for population control
in insects. Nine contributors, eight from the U. S. Department of Agriculture
and one from the U. S. Public Health Service cooperated in producing the seven
chapters.
In Chapter 1, A. W. Lindquist introduces the topic by evaluating insects as
pests, reviewing the current methods of control, and introducing the sterile-male
concept. Chapter 2 by E. F. Knipling presents the theoretical background for
sterilizing and releasing laboratory reared insects among natural populations and
for direct field sterilization of portions of natural populations. Germain C. La
Brecque discusses in Chapter 3 the laboratory procedures for screening chemo-
sterilants, testing their quantitative effects by different methods of application,
and determining field potential from detailed studies of the effects of a sterilant
in or on an insect. In Chapter 4, Leo E. La Chance, David T. North and Waldemar
Klassen discuss the basis of chemosterility at the cellular and cytogenetic levels.
Ralph B. Turner presents a summary of the physical and chemical properties of
some representative groups of chemosterilants and their biochemistry in Chapter 5.
Field studies and evaluations of chemosterilants on specific insects (e.g., House
fly, Screwworm, Cabbage looper, Boll weevil, Mosquitoes) and the use of chemo¬
sterilants as a tool in basic biological studies of insect reproductive behavior and
population structure and dynamics are presented in Chapter 6 by D. E. Weidhaas.
In Chapter 7 Wayland J. Hayes, Jr. reviews information on aspects of mammalian
toxicity of chemosterilants for use in insect population control.
Each chapter is followed by its own set of references and a terminal index to
the volume is provided. This book is an invaluable review which deserves a place
on the bookshelves of all those interested in possible alternatives to pesticides as
methods of controlling insect populations.— Robbin W. Thorp, University of Cal¬
ifornia, Davis, 95616.
56
The Pan-Pacific Entomologist
Larval Description and Notes on the Biology of
Anthocomus horni
(Coleoptera: Melyridae)
David E. Foster and Arthur L. Antonelli
Department of Entomology, University of Idaho, Moscow 83843
The taxonomy of the North American representatives of Anthocomus
has been covered by Marshall (1949). Their larvae have not been
described and only one species, Anthocomus auritus (LeConte), has
been figured (Boving and Craighead, 1930). Information concerning
Anthocomus biology has not been published. For these reasons this
information, although incomplete, is being presented. Terminology used
here is adapted from Boving and Craighead (1930).
Biological notes. —On 18 April 1971, 29 May 1971 and 15 Sep¬
tember 1972 nests of the black-billed magpie, Pica pica (Linn.), were
collected on Coyote Grade, Nez Perce County, Idaho. Among insects
found to inhabit the nests were larvae of Anthocomus horni (Fall).
The larvae of A. horni are not restricted to magpie nests. The species
has been reared from tephritid galls growing on Artemisia tridentata
Nuttal (Jordan Valley, Oregon, 2 May 1960, B. A. Foote) and from
wood of Pinus flexilis James (Craters of the Moon National Monument,
Idaho, 22 June 1965, D. S. Horning).
In northern Idaho A. horni spends the winter as a mature larva,
pupating the following spring. Adults can be found from early May
through mid July. They have been collected from the following plants:
Eriodictyon sp., Ceanothus sp., A. tridentata , Urtica Lyalli Watson.
Anthocomus horni has also been taken sweeping grass.
Mature larva. —Length 8.1 mm, maximum width 1.6 mm, head capsule width
0.9 mm.
Body elongate, slightly tapered anteriorly and posteriorly; membranous areas
mottled reddish to brown, sclerites testaceous to dark brown, surface clothed with
short to long, erect or suberect, fulvous hairs.
Head with dorsal surface broadly flattened, slightly broader than long in dorsal
view (6:5). Epicranium uniformly sclerotized; surface smooth to finely roughened,
clothed with short to long hairs, reddish brown except for very dark brown pig¬
mentation in ocular area; epicranial suture much shorter than cranial width in
dorsal view (3:10) ; stemmata separate and entire, four on each side arranged in
anterior row of three and single posterior one; anterior row lying in dorsoventral
arc that projects slightly anteriorly, bottom stemma of arc largest with diameter
of each successive stemma diminished slightly; posterior stemma slightly larger
than anteroventral one. Frons suhtriangular, nearly twice as broad as long; surface
The Pan-Pacific Entomologist 49: 56-59. January 1973
Vol. 49, No. 1, January 1973
57
Fig. 1-6. Anthocomus horni, mature larva. Fig. 1. Head capsule dorsal view;
Fig. 2. Stemmata lateral view; Fig. 3. Right mandible dorsal view; Fig. 4. Right
mandible ventral view; Fig. 5. Urogomphi dorsal view; Fig. 6. Urogomphi lateral
view.
58
The Pan-Pacific Entomologist
uniformly sclerotized, irregularly sculptured and finely roughened; front margin
truncate. Clypeus transverse, very lightly sclerotized, appearing almost membranous;
surface very finely roughened; front margin feebly arcuate or truncate. Labrum
transverse and tumescent, lightly sclerotized; surface very finely roughened,
clothed with numerous short and long hairs irregularly positioned behind the feebly
arcuate front margin. Antenna three segmented; basal membrane subequal to
combined length of three distal segments, glabrous, cream colored, appearing
inflated; basal segment slightly broader than long in dorsal view (4:3), slightly
compressed laterally, circumscribed by broad reddish brown band over most of
surface; middle segment slightly longer than basal segment in dorsal view (6:5),
constricted at base and expanded gradually distally, slightly compressed laterally,
circumscribed by a broad reddish brown band that covers most of its surface, bear¬
ing three long, stiff, cream colored hairs arising near apical margin, one hair
arising laterally on each side and one middorsally, apex of middle segment bearing
conical, glabrous appendix lateroventral to base of first segment, appendix shorter
than apical antennal segment (2:3) ; apical segment subequal in length to second
segment in dorsal view, digitiform, much longer than broad (4:1), color uniformly
reddish brown except cream colored apex which bears four stiff hairs, three equal
in length and equidistantly positioned around apical margin, fourth longer, arising
from center of apex. Mandibles longer than broad in dorsal view (5:4) ; outer
margin arcuate; inner margin tridentate, retinaculum pronounced and obliquely
rounded, middle tooth small and broadly flattened, mandibular apex arcuately
rounded; surface reddish brown at apex and along lateral margins, dorsal disc
light brown, bearing two long and three short, stiff hairs arranged in two longi¬
tudinal rows; lacinia mobilis bifurcate, arising from inner margin half way between
retinaculum and posterior angle. Maxillae and labium forming a prominent, ven-
trally protuding plate; maxilla with stipes lightly sclerotized, clothed with several
short and long, erect hairs, palpifer membranous, bearing a single long, erect hair
near inner margin, palpus with basal segment short and glabrous, middle segment
about twice as long as basal segment, dorsolateral surface clothed with several
short and long, stiff hairs, terminal segment about as long as middle segment,
conical, apex with cluster of short, very stout hairs at apex; labrum with sub-
mentum and mentum membranous, clothed with scattered short and long, erect
hairs, stipes labii with sclerotized base, bearing pair of long, erect hairs at front
margin, palpus two segmented, basal segment subcylindrical, bearing several short,
erect hairs around apex, terminal segment conical, about equal in length to basal
segment, bearing a cluster of very short, stiff hairs at apex. Gula tumescent and
glabrous, sides straight, converging slightly towards anterior end, posterior width
subequal to length.
Thorax well developed, more than one-half as long as abdomen in dorsal view;
prothorax subrectangular, tergum bearing long, straight and short, feebly arcuate
sclerite on either side of ecdysial suture; mesothorax sub triangular, slightly wider
than prothorax, tergum bearing a short, feebly arcuate sclerite on either side.
Metathorax similar to but slightly broader than mesothorax.
Legs about equal in size and shape; pretarsus appendiculate, apex acuminate,
basal tooth short with apex oblique.
Abdomen broadest at fifth segment; segments one through eight similar; segment
nine with hind margin truncate, posterior surface covered by basal plate; urogomphi
arising separately from basal plate, heavily sclerotized, nearly twice as long as
Vol. 49, No. 1, January 1973
59
epicranial suture, distance between tips less than total length including basal plate
(5:7), each urogomphus acuminate and recurved, surface reddish brown, bearing
several erect hairs around middle.
Material examined.— Seven specimens from Coyote Grade, 9 miles E Lewiston,
Nez Perce County, Idaho.
Literature Cited
Boving, A. G. and F. C. Craighead. 1930. An illustrated synopsis of the principal
larval forms of the order Coleoptera. Entomol. Amer., 11: 268-269.
Marshall, M. Y. 1949. Studies in the Malachiidae II. Entomol. Amer., 28:
126-132.
SCIENTIFIC NOTE
Ectoparasites and Nidicolous Acari of the Pocket Gopher, Thotnomys
bottae (Rodentia: Geomyidae). —Ectoparasites and nidicolous Acari were
collected from 393 Thomom-ys bottae (Eydoux and Gervais) and seven gopher nests.
All gophers were live-trapped from agricultural lands around Davis, California
from April 1970 through November 1971 with 378 gophers taken from irrigated
alfalfa fields, and the remainder from barley fields. The seven nests were taken
from alfalfa fields. Table 1 presents the numerical relationships of ectoparasites
removed from both gophers and nests. Ectoparasites were removed by brushing
and detergent bath methods, and 15% of the hosts were treated with trypsin-KOH
to recover all parasites. The recovery error as determined from the trypsin-KOH
samples was 2.6% for Geomycloecus oregonus Price and Emerson, 4.8% for
Table 1. Ectoparasitic arthropods collected off of 393 Thomomys bottae and
seven gopher nests from Davis, California from April 1970 through November 1971.
Number removed
Percent infested
Ectoparasite
gophers
nests
gophers
nests
Geomydoecus oregonus
(Trichodectidae)
140,803
98.6
Foxella ignota
(Ceratophyllidae)
4
_
0.5
_
Geomylichus sp.
(Listrophoridae)
27,623
_
76.7
_
Hirstionyssus femuralis
(Laelapidae)
10,334
368
79.9
100.0
Haemolaelaps geomys
(Laelapidae)
444
1,425
22.3
85.7
Leeuwenhoekia delosa
(Trombiculidae)
1
0.2
60
The Pan-Pacific Entomologist
Hirstionyssus femuralis Allred and 5.0% for Geomylichus sp. Haemolaelaps
geomys Strandtmann was not present in any trypsin-KOH sample.
The nidicolous Acari obtained from the seven nests were: Mesostigmata—
Macrocheles merdarius (Berlese) Macrocheles sp., Parasitus americanus (Berlese),
Parasitus sp., Hypoaspis sp., Proctolaelaps sp.; Prostigmata —Biscirus sylaticus
(Kramer) ; Astigmata —Tyrophagus longior (Gervais) ; Cryptostigmata— Epiloh-
mannia sp., Euphthiracarus sp., Zygoribatula sp.
Howard and Childs (1959: Hilgardia 29(7) :277—358) listed Geomydoecus
oregonus (as Trichodectes geomydis (Osborn)), Haemolaelaps geomys, H. femuralis
(as Hirstionyssus geomydis (Keegan)) and Trombicyla californica Ewing on T.
bottae in the southern San Joaquin Valley of California. Miller and Ward (1960:
Amer. Midi. Natur. 64(2) :382-391) listed two species of Geomydoecus, the flea
Foxella ignota (Baker) and six parasitic mites and three species of Ixodes from
T. bottae in Colorado. Allred and Beck (1966: Brigham Young Univ. Sci. Bull.
8(1) :1-123) categorically listed fleas, lice, mites and ticks from T. bottae in Utah.
The low number of fleas and absence of ticks on T. bottae in the Central Valley
of California indicate that they are either geographically or ecologically separated
from those ectoparasites.
My appreciation goes to the following individuals for identification of specimens:
R. D. Price and K. C. Emerson for Geomydoecus’, B. McDaniel and C. Coffman
for Geomylichus-, and F. M. Summers for the nidicolous Acari.— R. W. Rust,
University of California, Davis 95616.
SCIENTIFIC NOTE
Uroctonoides fractus a synonymy (Scorpionida:Chactidae).—Chamberlin
(1920. Brooklin Mus. Sci. Bull., 1: 36.) described a new genus and species of
scorpion from Quito, Ecuador. This species, described from a single male spec¬
imen, has not been reported since. In Chamberlin’s description he remarked on
the close proximity of Uroctonoides fractus to Uroctonus mordax Thorell of Cal¬
ifornia and Oregon. Werner (1934. Bronns, Klassen und Ordnungen des Tier-
reichs. 5, 4, 8, 283.) and Mello-Leitao (1945. Escorpios sul-Americanos. Arq. Mus.
Nac. Rio de Janeiro, 40: 128.) included this genus in the subfamily Uroctoninae
of the Vejovidae.
I was fortunate to be able to study the genotype of Chamberlin’s species and
discovered that it was a chactid, probably of the species Teuthraustes lojanus
(Pocock). Species placement must remain somewhat uncertain since the telson
and the last three caudal segments are missing. An unauthored label, in addition
to the original label, was found in the specimen vial which identified the species
as Teuthraustes atramentarius Simon. The inferior median keels of caudal seg¬
ments III are absent however, thus eliminating T. atramentarius as a species
choice.
I wish to express my gratitude to Dr. Herbert W. Levi of the Museum of Com¬
parative Zoology, Harvard University, Cambridge, Massachusetts, for the loan
of the type specimen. —Michael E. Soleglad, 3927 Polack Street, San Diego,
California 92110.
Vol. 49, No. 1, January 1973
61
A New Species of the Flea Beetle Genus Chaetocnema
Found on Dichondra in California
(Coleoptera: Chrysomelidae)
Richard J. McCrea
Office of Agricultural Commissioner, County of Los Angeles ,
500 W. Temple, Los Angeles, California 90012
A new species of flea beetle that is doing millions of dollars damage
to dichondra lawns in Los Angeles County is described and the biology
recorded.
Important research on Chaetocnema by Duckett (1920) in Maryland
began in 1911 and ended with his death in 1918. Gentner (1928) de¬
scribed three new species including Chaetocnema magnipunctata , which
was found in California. Poos (1955) worked on the biology of Chae¬
tocnema pulicaria , C. denticulata, and C. confinis from 1934-1936. This
work was of considerable help to me in my research.
Because of the economic importance, the flea beetle will merit further
observation and study.
Materials and Methods
Study of the life cycle involved rearing these life stages by a method
that most nearly simulated its usual habitat. These beetles were reared
on full flats of dichondra grown from seed placed in a steam treated
media. The dichondra was planted four months before the start of the
test. Three to four hundred beetles were introduced into a rearing cage
capable of holding a full flat of dichondra. Samples measuring about
two inches wide and six inches long were cut from the flat of dichondra
and put in Berlese funnels daily until the first larvae appeared. Samples
were then taken every third day and the results documented.
Eggs and pupae were reared on dichondra leaves, stems, and roots
in clear plastic cages kept moist with dampened blotters. A pinch of
sterilized soil was added for moisture control when rearing pupae.
Key to the Chaetocnema of California
1. Sides of prothorax regularly arcuate from base to apex, without oblique trun¬
cations at front angles _ 2
Sides of prothorax obliquely truncate at front angles, with post apical
angulation _ 8
2. Vertex always punctate sometimes indistinctly _ 3
Vertex absolutely impunctate except near eyes _ 5
The Pan-Pacific Entomologist 49: 61-66. January 1973
62
The Pan-Pacific Entomologist
3. Rows of elytral punctures confused at bases; length 2.0 mm _
- irregularis Leconte
Rows of elytral punctures regular _ 4
4. Punctures of head distinct; length 2.3-2.5 mm _ denticulata (Illiger)
Punctures of head indistinct; length 1.5 mm _ opacula Leconte
5. Basal marginal line of prothorax entire, not defined by punctures _ 6
Basal marginal line of prothorax defined by row of punctures, sometimes
continued to middle as a line _ 7
6. Prothorax alutaceous, with median smooth space posteriorly; scutellar
stria usually confused; color greenish bronze; tibia in part and tarsi reddish
black; length 2.0-3.5 mm _ subviridus Leconte
Prothorax not distinctly alutaceous, without smooth space; scutellar stria
regular; color golden bronze; tibia and tarsi brownish yellow; length 2.0-
2.5 mm ___ opulenta Horn
7. Prothorax indistinctly punctate; alutaceous, basal punctures fine and close;
humeri rounded, black bronzed color, antennae reddish yellow at base, five
outer segments reddish black; length 1.5 mm _ ectypa Horn
Prothorax with large, round deep punctures, surface faintly alutaceous;
humeri obtuse, elytra a little wider at base than prothorax and broadly
rounded to apex; brassy bronze with slight greenish tinge; basal four seg¬
ments of antennae, tibia and tarsi more or less reddish yellow; length 1.75-
1.95 mm_ magnipunctata Gentner
8. Head impunctate or with one puncture over eyes, prothorax nearly impunc-
tate; length 1.5-1.8 mm _ confinis Crotch
Head with few distinct punctures over eyes, prothorax distinctly punctate;
length 1.5-1.8 mm _ repens McCrea, n. sp.
Chaetocnema repens McCrea, new species
Chaetocnema magnipunctata , Jefferson, et al., 1967:9 ( nec Gentner).
Male.— Length 1.5 mm, width 0.9 mm. Broadly oval, robust, reddish black and
slightly bronzed. Antennae reddish yellow. Anterior and middle legs and posterior
tibiae and tarsi reddish yellow, posterior femora slightly reddish black. Head finely
alutaceous, distinct punctures over eyes. Prothorax anterior angles obliquely
truncate, post apical angulation distinct and arcuate posterolaterally, disc dis¬
tinctly alutaceous without basal marginal line, punctures of moderate size, closely
but not densely placed, not deeply impressed, dorsum of prothorax distinctly
punctate. Humeri rounded, umbone moderate, striae impressed on disc, more deeply
at sides, punctures relatively coarse, close and deep, but not serrate, intervals
slightly convex, wider than striae on disc, but not at sides, surface smooth, shining,
with fine interestrial punctures. Body beneath light reddish black, shining. Pro¬
sternum punctate, side pieces smooth. Abdomen sparsely indistinctly punctate.
Aedeagus with smooth slope to apex when viewed laterally.
Female. —Very similar to male. Length 1.7 mm. Width 1.1 mm. Genitalia
lobes have smooth slope to base when viewed dorsally.
Holotype male and allotype, Whittier, Los Angeles County, Cali¬
fornia, 1 September 1970, R. J. McCrea, deposited in Entomological
Collection LACM. Paratypes including ten males and ten females, ex-
Vol. 49, No. 1, January 1973
63
Fig. 1 . Aedeagus of Chaetocnema repens, side view. Fig. 2. Aedeagus of C.
confinis, side view. Fig. 3. Female genitalia of C. repens. Fig. 4. Female geni¬
talia of C. confinis (line represents 0.25 mm). Fig. 5. Slide specimen of adult
C. repens (line represents 1 mm). Fig. 6. End of hind tibia of C. repens. Fig. 7.
End of hind tibia of C. confinis. Lines on Figs. 1-3, 6, 7 represent 0.5 mm.
64
The Pan-Pacific Entomologist
amples all same data as holotype. Length 1.5—1.8 mm. Width 0.8-
1.1 mm. Paratypes are placed in following collections: Los Angeles
County Museum; California State University, Long Beach; Los Angeles
County Agriculture Commissioner’s Office, Los Angeles; and California
Department of Agriculture, Sacramento.
This new species, which is very similar to Chaetocnema confinis, can
be distinguished from it by two or three distinct punctures over the eyes
and distinct punctures on the prothorax (Fig. 9) as compared to one
or no punctures over the eyes and indistinct punctures on the prothorax
(Fig. 10) ; the aedeagus having a smooth slope to the apex (Fig. 1) as
compared to a depression near the apex of the aedeagus in C. confinis
when viewed laterally (Fig. 2) ; dorsal view of the aedeagus also differs
from C. confinis by not having a slight indentation near the base (Fig.
8) as compared to the indentation of C. confinis (Fig. 11) ; female
genitalia differ by the genitalic lobes not having sharp angles at the
base (Fig. 3) as compared to the genitalic lobes of C. confinis (Fig.
4) ; inner margin of the hind tibia is thicker in C. repens (Fig. 6) than
the hind tibia of C. confinis (Fig. 7).
C. confinis and C. repens (Fig. 5) have the sides of the prothorax
obliquely truncate at the front angles, with post apical angulations, which
separates these two species from other Chaetocnema in California.
Chaetocnema repens occurs in Los Angeles County and in other south¬
ern counties. Recently the State Department of Agriculture submitted
samples to me from Sacramento which proved to be C. repens. I am
certain that when a complete distribution of this flea beetle is determined,
the area will include much of California where dichondra is grown.
Available information was taken from state and county records but
Chaetocnema specimens were usually identified only to genus.
Life Cycle
Eggs hatch three days after deposition. Eggs are deposited in di¬
chondra sod near the surface. Eggs were deposited in clear plastic cages
in the laboratory test on dichondra roots, stems, cut dichondra leaves
and even on damp blotters. These eggs were measured by their width and
length in microns. The range of width was 225—275 microns averaging
252 microns. The range of the length was 450-500 microns averaging
480 microns.
The larval stage takes 22 to 25 days to pupation. There are four
instars as was determined by measurement of the head capsules of lar¬
vae. First instar head capsules measured 115-125 microns wide and
Vol. 49, No. 1, January 1973
65
Fig. 8. Illustration of the aedeagus of Chaetocnema repens, dorsal view. Fig. 9.
Illustration of the pronotum and part of head capsule of C. repens, dorsal view.
Fig. 10. Illustration of the pronotum and part of head capsule of C. confinis,
dorsal view. Fig. 11. Illustration of the aedeagus of C. confinis, dorsal view.
Solid black line below Figs. 8-11 represents 0.5 mm.
66
The Pan-Pacific Entomologist
125 microns long. Second instar head capsules were 137-150 microns
in width and length. Third instar head capsules measured 187-200
microns wide and 200-212 microns long. Fourth instar head capsules
measured 237-265 microns in width and length. The larval stage begins
to eat dichondra roots soon after hatching. Some larvae bore into and
enter small roots while others eat the outside root tissue leaving only the
central vascular tissue. Larvae can he found down to five inches deep.
Pupation takes place in dichondra sod usually where the last instar
larva ended up feeding on roots. Pupation takes five days to the white
adult. It takes about a day for the adult to darken to its normal dark
color.
The entire life cycle ranges from 32 to 35 days depending upon
temperature and moisture conditions of the dichondra sod. The di¬
chondra flea beetle overwinters as an adult in dichondra and becomes
an active feeder during warm winter days in southern California.
Acknowledgments
I am deeply indebted to Dr. E. L. Sleeper, Professor of Entomology
at California State University, Long Beach, for the assistance and guid¬
ance he has given me throughout the entire project. I also appreciated
the help that Dr. L. G. Gentner, Emeritus Professor of Entomology,
South Oregon Experiment Station, gave me at the start of the research.
I would like also to give credit to Ralph W. Lichty, Commissioner, and
Paul B. Engler, Chief Deputy Commissioner, Los Angeles County Agri¬
culture Commissioner’s Office, for their part in permitting my staggered
work schedule and their encouragement in this research.
Literature Cited
Duckett, A. B. 1920. Annotated list of Halticini, Univ. Md., Agr. Exp. Sta.,
Bull. 241: 111-155.
Gentner, L. G. 1928. Contributions to our knowledge of North American Hal-
ticinae. I. Notes, with descriptions of new species, Coleoptera: Chry-
somelidae, Trans. Amer. Entomol. Soc., 54: 57-67.
Jefferson, R. N., F. S. Morishita, A. S. Deal, and W. A. Humphrey. 1967.
Look for turf pests now, California Turfgrass Culture, 17 (2) : 9-11.
Poos, J. 1955. Studies of certain species of Chaetocnema, J. Econ. Entomol.,
48: 555-563.
Vol. 49, No. 1 , January 1973
67
Possible Behavioral Thermoregulation
in Tanarthrus salinus and T. inyo
(Coleoptera: Anthicidae)
Randall M. Peterman 1
Zoology Department, University of California, Davis, 95616
This paper describes several aspects of the biology of two desert-
inhabiting species of anthicid beetles, Tanarthrus ( Tanarthropsis ) inyo
Wickham and the rare Tanarthrus salinus LeConte. In particular, evi¬
dence will be presented to suggest that these insects have special adap¬
tations for avoiding heat stress.
These beetles, which bear a superficial resemblance to ants, are, on
average, 6 mm long and 2 mm wide, and 3.2 mm by 1 mm, the larger
species being Tanarthrus salinus. The larger species is dark red-orange
on the head, thorax, elytra and legs, with the abdomen being black,
including the ventral and dorsal sides of its posterior tip which protrudes
beyond the elytra by 1 mm. The hind wings are semi-transparent beige.
Tanarthrus inyo is like T. salinus in coloration and general character¬
istics except that the elytra extend to the end of the abdomen and the
rear half of the elytra are black. Both species are carnivorous scavengers,
feeding on windblown material such as dead insects.
Methods
Observations were made in parts of August and September 1969 in a
four square mile area in the middle of Carson Sink, near Fallon, Nevada.
This area is a level, dry (but seasonally flooded) mud flat, lacking any
form of plant life (except perhaps algae) and having only a small en¬
demic animal population. There are few rocks and no standing water,
and the soil surface is hard-packed. “Peeling” mud cakes in the usual
sense are not found. On a macroscopic level then, the habitat appears to
be relatively homogeneous. On the microhabitat level, however, the
ground is more heterogeneous. Small cracks ranging from 1 to 5 mm
in width are often found, and there are a few small (10 cm high) sand
accumulations scattered throughout the area. The anthicids studied
spend part of their time on the surface and part in these cracks, which
offer them their only reprieve from direct sunlight.
Temperature data were gathered with a YSI telethermometer (50°
1 Present address: Institute of Animal Resource Ecology, University of British Columbia, Vancouver
8, B. C. Canada.
The Pan-Pacific Entomologist 49: 67-73. January 1973
68
The Pan-Pacific Entomologist
Fig. 1. Air temperature profiles on a typical day, 12 September 1969. The 1-cm
mounds are where “thermoregulating” beetles perch.
C upper limit) and two number 403 thermistors. The small size (about
1 mm diameter) and short equalizing time (4 seconds) of these temper¬
ature probes permitted accurate temperature measurements of very
small areas. Behavioral observations were mainly made with binoculars
because beetles were so wary that usually no approach closer than
two meters was possible.
Thermoregulatory Behavior
In the Sink winds of 0-5 mph normally prevail from the southwest
and a thin superheated air layer less than 5 mm thick is produced just
above the surface (Fig. 1). Living in such a warm environment, with
daily summer surface temperatures ranging to above 50° C, both
Tanarthrus salinus and T. inyo appear to have evolved two ways of
avoiding heat stress. The first of these adaptations is morphological:
the long, thin legs. The high surface-to-volume ratio of the legs in
combination with their length minimizes the amount of heat which can
be conducted through them from the ground to the body. In addition,
Vol. 49, No. 1 , January 1973
69
the long legs, when stretched, lift the body up out of the superheated air
layer into the wind turbulence, as noted by the rocking of the bodies
back and forth with the wind gusts. This “stilting” behavior moves
the main body axis up from 1.5 to 4.0 mm above the surface in T. salinus
and from 0.8 to 1.4 mm in T. inyo. Such small increases in height ac¬
company relatively large decreases in surrounding air temperatures
(Fig. 1), perhaps large enough to make a significant difference in the
body temperatures of these small beetles when they approach their upper
temperature tolerances. Stilting has also been observed in scorpions
(Alexander and Ewer, 1958) and tenebrionid beetles (Hamilton, 1971)
and has been thought to be related to thermoregulation.
Another factor may be important in determining the body temperature
of these very small animals: solar radiation (Pepper and Hastings,
1952). Here the second apparent adaptation comes into play—behav¬
ioral thermoregulation. The stilting described above is invariably ac¬
companied by: (1) standing immobile on top of a small mound of dirt,
(2) positioning the body approximately parallel with the sun’s rays,
with the head toward the sun and the anterior end of the body elevated
slightly, and (3) for Tanarthrus salinus only, the beige hind wings ex¬
tending out over the black tip of the abdomen (see Fig. 2). The first
two characteristics tend to minimize heat gain by (a) moving into cooler
surrounding air and (b) decreasing the angle of incidence of sunlight
on the body. Such orientation with respect to incident solar radiation
has been noted in grasshoppers, butterflies, and other beetles and has
been shown to be an important factor regulating rates of heat gain
(Clench, 1966; Edney, 1971; Heinrich, 1972; Kevin and Shorthouse,
1970; Pepper and blastings, 1952; Watt, 1968). Other cases of behav¬
ior aiding the avoidance of heat stress have been reported for reptiles
(e.g. Hamilton and Coetzee, 1969; Norris, 1953; Willard, 1966). The
third characteristic of the observed anthicid behavior, extension of the
light-colored hind wings, essentially replaces a black surface with a more
reflective one, thereby decreasing the amount of heat absorbed by that
region. Edney (1971), Hamilton (1973) and others have shown that
light-colored insect surfaces keep underlying tissues significantly cooler
than tissues under dark surfaces. Although these results have been ob¬
tained for insects larger than the present anthicids, it is probable that
the wing-extending behavior of Tanarthrus salinus can make a small
but significant difference in body temperature when the insects are
nearing their upper lethal temperatures. Tanarthrus inyo does not ex¬
hibit wing-extending behavior because its elytra extend to the end of
the abdomen.
70
The Pan-Pacific Entomologist
0 1 2 3 4 5
i-1-1_i_i_i
MILLIMETERS
Fig. 2. Tanarthrus salinus in two stances, drawn from slides. Top figure is
normal stance; bottom figure is “thermoregulatory” stance. Note the stretched
legs and the hind wings out over the abdomen end.
Vol. 49, No. 1 , January 1973
71
BEETLE
ACTIVITY'
T. saLinus
SIGHTINGS
T.iQYJi
SIGHTINGS L
o o o
NORMAL THERMOREGULATING BEETLES.
ACTIVITY 23,24 AUGUST. TEMPERATURE
ADJUSTED TO 21 AUGUST
NORMAL
ACTIVITY
Fig. 3. Beetle activity periods and environmental temperatures on a typical
day, 21 August 1969. In the beetle activity section at the bottom, one dot represents
one sighting on that day at the indicated time.
Five observations support the hypothesis that the unusual behavior
just described actually serves a thermoregulatory function (in the sense
of avoiding a heat death, rather than maintaining a constant body tem¬
perature). The first is that normally the middle three or four hours
of the day find no beetles above ground except in the shade of man-made
objects. However, on the two hottest days experienced during the study,
a total of 9 anthicids (5 of T. salinus ) were seen on the surface in the
midday period, and all were in the thermoregulatory postures (Fig. 3).
The postures were not seen at any other time in the study for more than
a few seconds. On the two hot days, these positions were maintained
for up to 30 minutes. The beetles then disappeared when a duststorm
struck the area on both days. Second, the beetles moved from this po¬
sition only when the wind came up strongly, at which time they climbed
down from their dirt perches to the shady side, only to climb back up
when the wind died down, showing a definite favoring of this position.
Third, just prior to the observed behaviors in the hot midday, literally
dozens of both species flocked to the shade of the research facility (an
open air shelter 3 m by 3 m), a phenomenon not seen previously in the
72
The Pan-Pacific Entomologist
study. Fourth, the hind wings of T. salinus were extended over their
black-tipped abdomens only when accompanied by the other character¬
istics of the apparent thermoregulatory behavior, and vice versa. Fifth,
when exhibiting this unusual behavior, both species were much less
wary than normal. They could be approached by an observer to within
a few centimeters, suggesting that they may have been in critical thermal
states.
These five observations strongly imply that the two species have
evolved behavioral adaptations for avoiding heat death. Normally, when
the surface gets too hot, they go underground. However, on exception¬
ally warm days, the subsurface may be too hot (present data are in¬
conclusive) and they might then have to resort to extreme means to
cool off above the surface. Schmidt-Nielsen (1964), Edney (1967),
Hadley (1970), Stower and Griffiths (1966), and Pepper and Hastings
(1952) have pointed out for small desert insects that evaporative cool¬
ing can only contribute a minute amount to overall heat loss. Therefore,
the only cooling mechanisms available to such insects are behavioral
and morphological. Actual body temperature data are needed to crit¬
ically test the hypothesis of behavioral thermoregulation, but present
circumstantial evidence at least fails to negate the hypothesis.
Acknowledgments
Special thanks go to Dr. F. G. Werner for his prompt identification
of the anthicids; Drs. C. J. Walters and W. R. Cothran kindly made
comments on the manuscript, and B. Minchin rendered the figures.
Literature Cited
Alexander, A. J. and D. W. Ewer. 1958. Temperature adaptive behaviour in the
scorpion, Opisthophthalmus latimanus Koch. J. Exp. Biol., 35: 349-359.
Clench, H. K. 1966. Behavioral thermoregulation in butterflies. Ecology, 47(6) :
1021-1034.
Edney, E. B. 1967. Water balance in desert arthropods. Science, 156(2) : 1059-
1066.
1971. The body temperature of tenebrionid beetles in the Namib Desert of
Southern Africa. J. Exp. Biol., 55: 253-272.
Hadley, N. F. 1970. Micrometeorology and energy exchange in two desert
arthropods. Ecology, 51(3) : 434-444.
Hamilton, W. J. III. 1971. Competition and thermoregulatory behavior of the
Namib Desert tenebrionid beetle genus Cardiosis. Ecology, 52(5):
810-822.
1973. Life’s Color Code. McGraw-Hill, N. Y., 238 pp.
Hamilton, W. J. Ill and C. G. Coetzee. 1969. Thermoregulatory behaviour of
the vegetarian lizard Anglosaurus skoogi on the vegetationless northern
Namib Desert dunes. Sci. Pap. Namib Desert Res. Sta., 47: 95-103.
Vol. 49, No. 1, January 1973
73
Heinrich, B. 1972. Thoracic temperatures of butterflies in the field near the
equator. Comp. Biochem. Physiol., 43A: 459-467.
Kevin, P. G. and J. D. Shorthouse. 1970. Behavioral thermoregulation by high
arctic butterflies. Arctic, 23: 268-279.
Norris, K. S. 1953. The ecology of the desert iguana Dipsosaurus dorsalis. Ecol¬
ogy, 34: 265-287.
Pepper, J. H. and E. Hastings. 1952. The effects of solar radiation on grass¬
hopper temperature and activities. Ecology, 33: 96-103.
Sciimidt-Nielsen, K. 1964. Desert Animals, Physiological Problems of Heat
and Water. Oxford Univ. Press, London, 277 pp.
Stower, W. J. and J. F. Griffiths. 1966. The body temperature of the desert
locust ( Schistocerca gregaria). Entomol. Exp. Appl., 9: 127-178.
Watt, W. B. 1968. Adaptive significance of pigment polymorphisms in Colias
butterflies. I. Variation of melanin pigment in relation to thermo¬
regulation. Evolution, 22: 437-458.
Willard, D. E. 1966. The thermoecology of Cnemidophorus tigris. PhD Thesis,
Zoology Dept., University of California at Davis.
ZOOLOGICAL NOMENCLATURE: Announcement A. (n.s.)89
Required six-month’s notice is given of the possible use of plenary powers by
the International Commission on Zoological Nomenclature in connection with the
following names listed by case number.
(see Bull. Zool. Nomencl. 29, part 1, 1 May 1972)
1944. Suppression of Lyda inanis Klug, 1808 (Insecta, Hymenoptera)
1979. Type-species for Uloma Dejean, 1821 (Insecta, Coleoptera)
(see Bull. Zool. Nomencl. 29, part 3, 30 November 1972)
1989. Type-species for Anobium Fabricius, 1775, Grynobius Thomson, 1859, and
Priobium Motschulsky, 1845 (Insecta, Coleoptera)
1994. Type-species for Dicyrtoma Bourlet, 1842, and Dicyrtomina Borner, 1903;
suppression of Podura minuta O. Fabricius, 1783, and Papirius cursor Lub¬
bock, 1862; validation of Papirius fuscus Lubbock, 1873 (Insecta, Collembola)
1998. Type-species for Deuterosminthurus Borner, 1901 (Insecta, Collembola)
1999. Type-species for Eusminthurus Borner, 1900 (Insecta, Collembola)
(see Bull. Zool. Nomencl. 29, part 4, 29 December 1972)
1979. Type-species for Phaleria Latreille, 1802 (Insecta, Coleoptera)
1948. Validation of RIODINIDAE Grote, 1895 (Insecta, Lepidoptera)
2000. Suppression of Ptenura Templeton, 1842, and Podura crystallina Miiller,
1776 (Insecta, Collembola)
2004. Suppression of Dapanus Hentz, 1867 (Araneae)
Comments should be sent in duplicate, citing case number, to the Secretary,
International Commission on Zoological Nomenclature, c/o British Museum
(Natural History), Cromwell Road, London SW7 5BD, England. Those received
early enough will be published in the Bulletin of Zoological Nomenclature .—
Margaret Doyle, Scientific Assistant.
74
The Pan-Pacific Entomologist
Chromosomal Variability in a Nearctic Lycaenid Butterfly,
Philotes sonorensis
(Lepidoptera: Lycaenidae)
Thomas C. Emmel, Houston R. Trew, and Oakley Shields
Department of Zoology, University of Florida, Gainesville 32601, and
Department of Entomology, University of California, Davis 95616
Variable chromosome number has been reported in many plants and
animals. Aside from polyploidy, it is clear that chromosomal loss,
Robertsonian rearrangement including centric fusion, and supernu¬
merary chromosomes represent the common modes of change in chro¬
mosome number in groups as diverse as the flowering plants (Lewis,
1970), grasshoppers (Kayano et al., 1970; White et al., 1964), fish
(Ohno et al ., 1965), lizards (Gorman and Atkins, 1967; Gorman et al.,
1968), white-throated sparrows ( Zonotrichia ) (Thorneycroft, 1966),
and mammals (Patton, 1969; Wahrman et al., 1969). In groups with
holokinetic chromosomes (e.g., butterflies and moths), fragmentation
has been hypothesized as the chief factor leading to diversity of chro¬
mosome number (Emmel, 1972; Emmel and Trew, 1973). We report
here extraordinary intraindividual as well as intrapopulation variability
in butterfly karyotype, apparently resulting from chromosomal frag¬
mentation at spermatogenesis.
Material and Methods. —-The small lycaenid butterfly Philotes
sonorensis Felder & Felder is found in scattered colonies in lowland areas
of California, frequently in seasonally dry foothill canyons. It feeds on
Dudley a (Crassulaceae) in the larval stages and flies as an adult in
early spring. Samples of testes fixed in 3:1 absolute ethyl alcohol :
glacial acetic acid were taken in 1970 and 1971 in three populations
in Placer, El Dorado, and Stanislaus counties (43, 7, and 1 male spec¬
imens, respectively) and airmailed to the University of Florida for
analysis. The testes were stained with lacto-aceto-orcein, squashed,
observed and photographed under brightfield and phase (Emmel, 1969).
Cytological Observations and Discussion
In testes with dividing cells, meiosis appeared regular with pairing
observed for all chromosomes (no univalents). One or more bivalents
per karyotype were often composed of two smaller fragments synapsing
with one large homologue in a tripartite bivalent (Fig. 1). Interest¬
ingly, Suomalainen (1969) has previously observed tripartite bivalents
The Pan-Pacific Entomologist 49: 74-80. January 1973
Vol. 49, No. 1, January 1973
75
ft
b
Fig. 1 . Pairing of apparent fragments with homologous chromosome in Philotes
sonorensis. (a) Metaphase I, tripartite bivalent; (b) orientation of triple chromatid
on spindle in late metaphase I. Redrawn from photographs; approx. 7200X and
3600 X-
(but no chiasmata) in females of the pyralid moth Witlesia murana
Curt., indicating that fragmentation of moth chromosomes also occurs
in nature.
Chromosome numbers ranging from n — 17 to 44 were found in
populations surveyed (Fig. 2), with the modal number being 22 in
the Placer County populations, 23 in the El Dorado County population,
and 19 in the single male from the Stanislaus County population. Of
the largest sample, the 13 Placer County males with clearly observable
dividing cells, 7 (54 per cent) had multiple karyotypes. Four had n =
22 and 24 cells, and one had numerous meiotic cells ranging from n =
22 to 31. This great intraindividual variability was also found in the
other sampled populations (Table 1). Cells with the higher chromosome
number had smaller chromosomes in all cases.
Lesse (1960, 1969) found great chromosomal variability in popu¬
lations of species of the Palearctic lycaenid butterfly genera Lysandra
and Agrodiaetus, but until now there was no indication of its presence
in any Nearctic butterflies. In the few Nearactic lycaenids surveyed to
date (Maeki and Remington, 1961), the haploid number tends to be
24, with no intraindividual or intraspecies variation.
The selective significance of numerical chromosomal variability of
this type would seem to lie first in the promotion of flexibility in the
genotype’s ability to respond to variable environment conditions. Un¬
published observations we have made on the cytogenetics of other butter¬
fly species demonstrate a very low chiasma frequency (usually two
chiasmata per bivalent) in spermatogenesis and Suomalainen (1969)
has previously reported a lack of chiasmata in oogenesis in Lepidoptera.
The restriction on genetic exchange through crossing over would pro¬
mote a low recombination index (Darlington, 1939) in Lepidoptera
unless other factors were operating. It is clear that the high chromosome
number (modal number being about n = 31) in moths and butterflies
76
The Pan-Pacific Entomologist
Fig. 2. The geographic relationships of three Philotes sonorensis populations
in the state of California, showing variability in haploid number (n) of chro¬
mosomes found in males of each population. From north to south: Placer County
(3-4 mi. N.E. of Auburn along Middle Fork of American River, 21 February 1970
and 13-14 February 1971) ; El Dorado County (4 mi. N. of Placerville on California
Highway 193 at South Fork of the American River, 21 February 1970) ; Stanislaus
County (18 mi. W. of Interstate Highway 5, Del Puerto Canyon, 7 March 1970).
as a group would compensate for this trend. The observations reported
here offer evidence that fragmentation of butterfly chromosomes does
occur in nature and that the holokinetic nature of the lepidpoteran chro¬
mosome allows successful diakinesis in such cases.
The data from the present study and from Lesse (1960, 1969) also
provide evidence for the suggestion that lepidopteran species in marginal
Table 1.
Multiple Genotypes in Male Individuals of Three California Populations of Philotes sonorensis (bold-face
type indicates median chromosome number for population sample).
No. of individuals with cells having chromosome number ( n) of
Total No. Cells
Counted
Population 1
17 18
19 20 21
22
23
24
25
26 29 30 31 34
44
(Individuals)
Placer
County
1
3
10
2
2
1
1111
4
50 (13)
El Dorado
County
1 1
1 1 1
1
1
1
1 1
1
99 (4)
Stanislaus
County
1
1
47 (1)
1 See Fig. 1 legend for exact location of population. All counts (including n = 41) verified in Meiosis I (usually metaphase I) where chromosomes were still paired.
<1
<1
Vol. 49, No. 1 , January 1973
78
The Pan-Pacific Entomologist
habitats or habitats with high environmental stress, such as Philotes
in the dry foothill areas of California and Lysandra in certain areas
of Spain, may often exhibit fragmentation of their chromosomes, pro¬
moting increased genetic flexibility. Such increased variability of
supernumerary chromosomes has been found in marginal weedy pop¬
ulations of the plant Claytonia virginica L. (Lewis et al ., 1971), where
bivalents form normally and meiosis is not upset. (The cytogenetics of
this situation are not completely understood as yet.) Because of the
holokinetic nature of lepidopteran chromosomes, however, their fission
products can successfully synapse and segregate at meiosis. It would
be predicted that such dramatic reassortment of the genotype through
fragmentation should occur in other stress situations, as in rapid evo¬
lution of phenotypically polymorphic mimetic systems in Lepidoptera.
Evidence for numerical polymorphism is accumulating for the genus
Heliconius, especially H. doris (L.), which commonly participates
in mimicry complexes in South and Central America (Emmel, in
preparation).
A possible revolutionary feature of this chromosomal variability
in Philotes is the polyploidy (n = 17, 34, 51, 68) found in the two
Dudleya subgenera that P. sonorensis larvae use as hosts (Moran, 1951;
Moran and Uhl, 1952; Uhl and Moran, 1953). Hybridization between
host species is common. The small seeds of Dudleya are viable only
a short time and are not adapted for long-range dispersal; hence the
host colonies are usually quite isolated. Adult Philotes sonorensis have
been shown to have a restricted home range in an extensive mark-re¬
capture study (Keller et al., 1966; Mattoni and Seiger, 1963). It would
thus appear that the butterfly is closely linked to its foodplant and some
of the same factors that make polyploidy of selective value in Dudleya
may be preserving gains and losses of chromosomes in Philotes.
Acknowledgments
This research was supported by the National Science Foundation
(grants GB-8442 and GB-32151 to T.C.E.), the U. S. Public Health
Service through Biomedical Sciences Research Grant 71-59 from the
Division of Sponsored Research, University of Florida (T.C.E. and
H.R.T.), and an NSF graduate traineeship (O.S.). We thank F. C.
Johnson, University of Florida, and Dr. John F. Emmel, University of
California Medical School at San Francisco, for critical comments on
the manuscript. Population samples were collected by 0. Shields and
R. Wells.
Vol. 49, No. 1, January 1973
79
Literature Cited
Darlington, C. D. 1989. The evolution of genetic systems. Cambridge Univ.
Press, Cambridge, 149 pp.
Emmel, T. C. 1969. Methods for studying the chromosomes of Lepidoptera.
J. Res. Lepidoptera, 7: 23-28.
1972. Mate selection and balanced polymorphism in the tropical nymphalid
butterfly, Anartia fatima. Evolution, 26: 96-107.
Emmel, T. C. and H. R. Trew. 1973. The chromosomes of skipper butterflies
from southwestern North America (Lepidoptera, Hesperiidae). Cyto-
logia, (in press).
Gorman, G. C. and L. Atkins. 1967. The relationships of the Anolis of the
roquet species group (Sauria: Iguanidae). II. Comparative chromosome
cytology. Syst. Zool., 16: 137-143.
Gorman, G. C., R. Thomas, and L. Atkins. 1968. Intra- and interspecific
chromosome variation in the lizard Anolis cristatellus and its closest
relatives. Breviora, 293: 1-13.
Kayano, H., M. Sannomiya, and K. Nakamura. 1970. Cytogenetic studies
on natural populations of Acrida lata. I. Local variation in the frequency
of B-chromosomes. Heredity, 25: 113-122.
Keller, E. C, Jr., R. H. T. Mattoni, and M. S. B. Seiger. 1966. Preferential
return of artificially displaced butterflies. Anim. Behav., 14: 197-200.
Lesse, H. de. 1960. Speciation et Variation Chromosomique chez les Lepi-
dopteres Rhopaloceres. Ann. Sci. Natur., Zool. (Paris), 12 e serie, II:
1-223.
1969. Les Nombres de Chromosomes dans le Groupe de Lysandra coridon
(Lep. Lycaenidae). Ann. Soc. Entomol. France (N.S.), 5: 469-522.
Lewis, W. H. 1970. Chromosomal drift, a new phenomenon in plants. Science,
168: 1115-1116.
Lewis, W. H., R. L. Oliver, and T. J. Luikart. 1971. Multiple genotypes in
individuals of Claytonia virginica. Science, 172: 564-565.
Maeki, K., and C. L. Remington. 1961. Studies on the chromosomes of North
American Rhopalocera. 3. Lycaenidae, Danainae, Satyrinae, Morphinae.
J. Lepidoptera Soc., 14: 127-147.
Mattoni, R. H. T., and M. S. B. Seiger. 1963. Techniques in the study of
population structure in Philotes sonorensis. J. Res. Lepidoptera, 1:
237-244.
Moran, R. V. 1951. A revision of Dudleya (Crassulaceae). Unpubl. Ph.D.
dissertation, Univ. of Calif., Berkeley.
Moran, R. V., and C. H. Uhl. 1952. Four natural hybrids in Dudleya. Desert
Plant Life, 24: 28-42.
Otino, S., C. Stenius, E. Faisst, and M. T. Zenzes. 1965. Post-zygotic chro¬
mosomal rearrangements in rainbow trout ( Salmo irideus Gibbons).
Cytogenetics, 4: 117-129.
Patton, J. L. 1969. Chromosome evolution in the pocket mouse, Perognathus
goldmani Osgood. Evolution, 23: 645-662.
Suomalainen, E. 1969. Chromosome evolution in the Lepidoptera. In Darling¬
ton, C. C., and K. R. Lewis, eds., Chromosomes today, Volume Two,
Oliver and Boyd, Edinburgh, pp. 136-137.
80
The Pan-Pacific Entomologist
Thorneycroft, H. B. 1966. Chromosomal polymorphism in the white-throated
sparrow, Zonotrichia albicollis (Gmelin). Science, 154: 1571-1572.
Uhl, C. H., and R. V. Moran. 1953. The cytotaxonomy of Dudleya and Hasse -
ajithus. Amer. J. Bot., 40: 492-502.
Wahrman, J., R. Goitein, and E. Nevo. 1969. Mole rat Spalax: evolutionary
significance of chromosome variation. Science, 164: 82-84.
White, M. J. D., H. L. Carson, and J. Cheney. 1964. Chromosomal races in
the Australian grasshopper Moraba viatica in a zone of geographic
overlap. Evolution, 18: 417-429.
SCIENTIFIC NOTE
Notes on the Distribution and Ecology of Scaphinotus bilobus (Coleop-
tera :Carabidae) .—According to Lindroth (1961. Opusc. Entomol. Suppl., 20:
1-200), “the species is rare and its demands little known.” This species is mainly
northeastern and it is distributed south to Illinois, Missouri, Ohio, and Catslcill
Mountains, N.Y., west to Nebraska. In Canada, Lindroth records this ground beetle
from Nova Scotia, Quebec (a single locality), Ontario and Manitoba.
The species is not uncommon in southern Quebec (south of 50° N Lat.). In the
locality records, the following abbreviations were used to indicate the collections
in which the specimens are located: AL—Andre Larochelle Collection, Rigaud;
CC—Claude Chantal Collection, Quebec; CH—Cercle Harricana Collection, Amos;
JCA—Jean-Charles Aube Collection, Quebec; JPL—Jean-Paul Laplante Collection,
Sainte-Foy; LEM—Lyman Entomological Museum, Sainte-Anne-de-Bellevue; LRF
—Laboratoire de Recherches Forestieres Collection, Sainte-Foy; UM—Universite
de Montreal Collection, Montreal. The distribution of Scaphinotus bilobus Say in
Quebec is: Abitibi Co., Amos, 13 May 1962 (1, CH) ; Chapais, 11 August 1972
(2, AL) ; Chibougamau, 23 September 1968 (2, CC) and 12 August 1972 (1, AL) ;
Lac Chicobi, 21 June 1968 (2, AL) ; La Ferme, 26 May 1963 (1, UM) ; Lebel-sur-
Quevillon, 16 July 1968 (3, AL) ; Matagami, 20 June 1968 (3, AL) ; Saint-Mathieu,
1 July 1964 (1, CH) ; Senneterre, 12 July 1968 (1, AL) ; Val-d’Or, 11 July 1968
(2, AL). Champlain Co., Lac Normand, 27 August 1969 (2, CC). Charlevoix-Est
Co., Baie-Sainte-Catherine, 28 June 1969 and 1970 (10, AL; 2, CC). Charlevoix-
Ouest Co., Baie-Saint-Paul, 13 September 1969 (1, CC). Frontenac Co., Woburn,
2 September 1955 (2, JPL). Lotbiniere Co., Saint-Sylvestre, 14 July 1967 (1, JCA).
Matane Co., Metis-sur-Mer, 18 August 1889 (2, LEM). Montmorency No. 1 Co.,
Grand Lac Jacques-Cartier, 31 August 1968 (1, CC). Saguenay Co., Tadoussac,
14 June 1971 (2, AL). Saint-Maurice Co., Lac Caousacouta, 21 June 1965 (2,
JPL), 17 July 1968 (1, LRF), 22 July 1965 (1, LRF) and 23 September 1965 (4,
JPL; Lindroth, 1969. Opusc. Entomol. Suppl. 34: 945-1192). Temiscamingue Co.,
Tabaret, 5 May 1968 (1, AL).
This species is confined to shaded, rather moist places, especially coniferous and
mixed forests, on banks of lakes and rivers. During the day hiding under dead
leaves and logs, it often invades the river banks, peat-bogs and clearings in the
night in search of snails. Associated ground beetles include: Agonurn retractum
LeConte, Calathus ingratus Dejean, Pterostichus adstrictus Eschscholtz, Pterostichus
coracinus Newman, Pterostichus punctatissimus Randall and Sphaeroderus nitidi-
collis brevoorti LeConte. —Andre Larochelle, College Bourget, C. P. 1000, Rigaud,
Quebec.
Vol. 49, No. 1, January 1973
31
PACIFIC COAST ENTOMOLOGICAL SOCIETY
D. G. Denning J. A. Chemsak M. S. Wasbauer P. H. Arnaud, Jr.
President President-elect Secretary Treasurer
Proceedings
Three Hundred and Forty-Fourth Meeting
The 344th meeting was held Friday, 18 February 1972 in the Morrison Audi¬
torium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Denning presiding.
Members present (40) : F. G. Andrews, C. Armin, P. H. Arnaud, Jr., W.
Azevedo, J. W. Bass, G. S. Benham, Jr., F. L. Blanc, R. M. Bohart, T. Briggs,
R. M. Brown, G. Buckingham, Cathy Cheap, S. Clement, Karen Corwin, D. G.
Denning, A. R. Dutton, J. G. Edwards, J. F. Emmel, C. Farrell, W. E. Ferguson,
M. R. Gardner, Lauren Green, E. Grissell, K. S. Hagen, E. A. Kane, B. Keh,
R. L. Langston, H. B. Leech, R. W. Lem, A. E. Michelbacher, J. Powell, E. S.
Ross, J. Shepard, R. E. Stecker, Barbara Trowbridge, D. Ubick, M. S. Wasbauer,
S. C. Williams, D. Y. Wong, R. L. Wong.
Visitors present (22) : Fern Blanc, R. Buckerly, Caroline Caw, R. Coville,
Phyllis Emmel, Stephenie Ferguson, B. and D. Grushkowitz, L. M. Halverson,
R. Hollingsworth, Netta and Susanne Leong, Carolyn Mullinex, M. Narasaki, Reny
and Taly Slay, D. Strike, S. Szerlip, Barbara Ware, A. Wong, R. Wong, Virginia
Woo.
The minutes of the meeting held 17 December 1971 were summarized.
President Denning called for announcements. Dr. Stecker announced that the
tear sheets from the first 20 volumes of the Pan-Pacific Entomologist on Coleoptera
are out and will be on sale after the meeting. There are 75 articles and they will
be priced at $0.05 each.
Dr. Ferguson announced that the program committee would be grateful for any
suggestions on future speakers for Society meetings and especially for possible
sites and times for the annual field day and picnic.
Mr. Jon Shepard announced that Mr. Scott Ellis of Colorado will be collecting
in the Four Corners area of Colorado, Utah and Arizona this coming summer and
would like to know if anyone would be interested in buying material he collects.
Mr. Leech announced the appearance of Hatch’s latest and final volume of the
Beetles of the Pacific Northwest. This volume includes the buprestids, chrysomelids,
weevils, cerambycids and other families.
Dr. J. A. Powell announced the appearance of Volume 13 of the Bulletin of the
California Insect Survey by Bohart and Horning which treats the bembicine sand
wasps of California, and includes the genera Bembix, Bicyrtes and Microbembix,
a total of eighteen species.
Dr. Powell also commented on the butterfly, Nymphalis californica, which he
had discussed at an earlier meeting. There was a great build-up in numbers of
this species at Mt. Shasta last summer so that by the fall of the year they were
migrating and were seen in the Bay Area. At that time, Dr. Powell asked anyone
who witnessed numbers of the butterfly to contact him. He mentioned the possi¬
bility of their overwintering in this area and going through a couple of generations
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The Pan-Pacific Entomologist
here, although normally large populations do not occur here. He stated that he
had observed some overwintering under bark this season in the Berkeley Hills.
They are currently flying there in good numbers but apparently with no directional
movement. Directional movement by the species appears to be in response to
population density. He again requested anyone who witnesses large populations
of this butterfly this spring and summer, especially if flying in a pattern of
directional movement, to contact him.
Dr. Ferguson mentioned having collected caterpillars of Nymphalis californica
at Escondido last spring. They all died and the one submitted to the insect
pathology lab at Berkeley was reported killed by a bacterium never before recorded
from butterflies of that species. He noted that it would be interesting to find out if
the bacterium is a common natural agent in the biotic control of Nymphalis
calif ornica.
The following notes were presented:
A remarkable Tabanid inhabitant of an ant nest in western Washington
—natural or accidental? —A year ago this month (20 February 1971), a
graduate student, Gary Alpert, at Washington State University, Pullman, discovered
two mature larvae of Tabanus sp. in the center of a nest of the western mound¬
thatching ant, Formica obscuripes, which he was exploring in his studies on local
ants. In this portion of the nest, there were a few other denizens such as isopods
and collembolans. The mound was situated among growing vines and under trash
from trimmings of blackberries in the vicinity of Walla Walla, Washington.
On return to his laboratory, Alpert succeeded in rearing one of the larvae to the
pupal stage from which a large, dark horsefly emerged on 24 April. This, together
with the other preserved larva, was mailed to me in May for identification, but
unfortunately, the fly and pupal case were broken into small fragments in transit.
It was possible to ascertain from the bits and pieces, that the fly was a large
black, bare-eyed Tabanus male of some resemblance to, but not, Tabanus kesseli
Philip, known on the Pacific Coast as far north as southern British Columbia; the
upper eye facets are a little larger, as in T. kesseli than in related T. aegrotus 0. S.
Fragments of the pupa indicate an unusually long-spined form, perhaps habitat-
oriented.
The purpose of this cursory note is to alert others to this possible special
habitat of an obscure or perhaps unknown tabanid species. Customary breeding
in tree-holes by certain bizarre tabanids in tropical Africa and of Leucotabanus
ambiguus Stone, in frass-filled termite galleries in rotting trunks of old, living
willows, discovered recently by Dr. John Burger of Tucson, Arizona, are notable
unusual tabanid breeding sites. I am not aware, however, of reported tabanid
occupation of ant hills, at least in the New World, which stimulates speculation
on the natural cycle—oviposition, larval development, pupation, and any traversal
of the ant-occupied periphery of the nests, if this is a customary, and not an
accidental, eco-niche. In the words of the collector, “The tabanid larvae came
from the center of the nest where the ant activity was high”! Although the nest
mound was located about 15 yards above a small pond, it seems unlikely that two
mature larvae would have migrated to this special situation from the pond margin
to complete their development. —Cornelius B. Philip, California Academy of
Sciences, San Francisco.
Meigenielloides cinereus (Diptera: Tachinidae) reared from Gam-
marotettix bilobatus (Orthoptera: Gryllacrididae). —The genus Meigeniel¬
loides Townsend is one of 43 North American (north of Mexico) genera assigned
Vol. 49, No. 1 , January 1973
83
to the tachinid tribe Blondeliini. The hosts for this tribe include larvae of
Lepidoptera, Coleoptera, and tenthredinid Hymenoptera. It is of considerable
interest to report the rearing of Meigenielloides cinereus (Townsend) from the
Orthopteran family Gryllacrididae on the basis of two rearings, both of which
were made at Mill Valley, Marin County, California.
The first rearing (deposited in tire collection of the California Academy of
Sciences) was made by Thomas S. Leech from a female adult specimen of
Gammarotettix probably bilobatus (Thomas) (det. A. B. Gurney, 1957). Two
tachinid larvae emerged from the host and pupated on 29 May 1957. One adult
male of Meigenielloides cinereus issued on 14 June 1957, and a fly did not issue
from the second puparium.
The second rearing (deposited in the collection of the author) was made by
Thomas W. Davies from an adult female of Gammarotettix bilobatus (Thomas)
(det. David C. Rentz). On 20 June 1967 a tachinid larva emerged from the host
and pupated. An adult female of Meigenielloides cinereus issued on 7 July 1967.
In both rearings the pupal period lasted about two and one half weeks (17
days).— Paul H. Arnaud, Jr., California Academy of Sciences, San Francisco.
The status of Loxosceles unicolor in Tesla Canyon. —In November 1969,
the late Dr. J. W. MacSwain, of the University of California, gave a report en¬
titled, “A Naturalist Returns to Tesla Canyon.” (Pan-Pac. Entomol. 46(1) :74-75,
1970.) In his report Dr. MacSwain mentioned that on one of his 1951 collecting
trips he collected “. . . numerous L. unicolor . . . around or in the sides of the
canyon . . . .” Further, he implied that the spider still existed there on his last
collecting trip on 26 October 1969. If valid, this report would represent the
northern-most record of L. unicolor in the San Joaquin Valley.
I talked with Dr. MacSwain about the occurrence and there was no doubt that
these spiders were found in the canyon, and that he had collected them there on
several occasions. I then checked with Dr. E. I. Schlinger about the spider samples
given him by Dr. MacSwain which were collected 26 October 1969. Dr. Schlinger
said there were no Loxosceles in those samples.
I have made several collecting trips to Tesla Canyon, working the entire length
on both sides, including the places where Dr. MacSwain said he collected and on
the Lawrence Radiation Laboratory property. Although many spiders resembling
violin spiders were found I have yet to collect Loxosceles spiders in Tesla Canyon
and so far the available evidence does not support the claim made by Dr. Mac¬
Swain. The most northerly collection of L. unicolor on the west side of San
Joaquin Valley is in Jacalitos Canyon, about 5 miles south of Coalinga, Fresno
County, California (collectors: R. R. Snelling, 1967; F. Ennik, 1970). —Franklin
Ennik, California Department of Public Health, Bureau of Vector Control,
Berkeley.
The principal speaker of the evening was Mr. Michael R. Gardner, Systematic
Entomologist, California Department of Agriculture. His illustrated talk was
entitled, “Ecological and Behavioral Relationships of Millipeds in California.”
Coffee and other refreshments were served at a social hour in the entomology
rooms following the meeting.—M. S. Wasbauer, Secretary.
Three Hundred and Forty-Fifth Meeting
The 345th meeting was held Friday, 21 April 1972, in the Morrison Auditorium
of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Denning presiding.
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The Pan-Pacific Entomologist
Members present (41) : F. G. Andrews, P. H. Arnaud, Jr., J. W. Bass, G. S.
Benham, Jr., G. Brady, D. Briggs, T. Briggs, M. Browning, R. Bushnell, Karen S.
Corwin, R. Coville, H. V. Daly, T. Delgman, J. G. Edwards, Beverly Ehreth, F.
Ennik, W. E. Ferguson, A. J. Gilbert, B. W. Grushkowitz, J. Guggolz, K. S.
Hagen, E. A. Kane, B. Keh, W. H. Lange, R. L. Langston, H. B. Leech, R. Lem,
W. W. Middlekauff, R. H. Peterson, C. B. Philip, H. G. Real, G. Rotrammel,
S. L. Szerlip, K. Taylor, R. Thorp, R. Toth, D. Ubick, D. Voegtlin, K. R. Wald,
D. Y. Wong, R. L. Wong.
Visitors present (19) : J. Baier, Lily Dong, J. T. Doyen, Stephenie Ferguson,
Marcia Gilbert, Dorothy Grushkowitz, R. Hall, Lucia Hui, M. Jensen, S.
Germanovitch, J. B. Kimsey, Mrs. J. B. Kimsey, Netta Leong, Susanne Leong,
W. Minkel, R. P. Papp, Norma E. Papp, Joyce Thorp, Virginia Woo.
The minutes of the meeting held 18 February 1972 were summarized.
The following notes were presented:
Mass movement of Sympetrum corruptum (Hagen) at Pacifica, Cali¬
fornia (Odonata: Libellulidae). —Observations on the undirectional mass
movements of the dragonfly Sympetrum corruptum (Hagen) have been reported
in the Kensington-Albany-Berkeley and Mill Valley areas by Turner (1965, Pan-
Pac. Entomol. 41:66-67), and Opler (1971, Pan-Pac. Entomol. 47:223). A further
record of a flight south of these previous recorded localities is presented. On
24 September 1970, Mrs. Jose Sanchez of 452 Heathcliff Drive, Pacifica, San
Mateo County, California, first telephoned the Academy at 2:10 p.m. (P.D.T.)
to report that for the past two hours there had been what she considered
“thousands” of dragonflies in the air above where she lives. They were flying at
a height so that it was impossible to catch them without a long-handled net. At
3:50 p.m., she telephoned to say that dragonflies were also very common at the
Fairmont Elementary School. One speciemn which was hit in the schoolyard and
collected by one of her children was determined by Dr. Dennis E. Paulson as a
male (young specimen) of Sympetrum corruptum (Hagen). It is presumed that
this specimen was representative of the species involved in this flight. It is to
be regretted that additional voucher specimens are not available. Mrs. Sanchez
commented that this was the first flight of Odonata that she had observed in her
five years residence in Pacifica.— Paul H. Arnaud, Jr., California Academy of
Sciences, San Francisco.
Hjelle scorpion collection donated to California Academy of Sciences.
—Mr. John T. Hjelle, has recently donated his collection of 712 specimens of
scorpions from northern California to the Department of Entomology, California
Academy of Sciences.
This collection contains the types and specimens which formed the basis of his
publication “Scorpions of the northern California coast ranges (Arachnida:
Scorpionida) ” which was published in 1972 (Occas. Pap. Calif. Acad. Sci., No. 92,
59 pages, 52 figures). It contains 202 type specimens: the holotype, allotype,
and 162 paratypes of Vaejovis gertschi striatus Hjelle; the holotype, allotype, and
17 paratypes of Uroctonus glimmei Hjelle; and the holotype, allotype, and 17
paratypes of Uroctonus mordax pluridens Hjelle. In addition, there are 249
specimens of Paruroctonus silvestrii (Borelli), 4 Superstitionia donensis Stahnke,
255 Uroctonus mordax mordax Thorell, and 2 Uroctonus new species. This collec¬
tion will complement the extensive series of scorpions being received from Dr.
Stanley C. Williams. Because of these and other important donations, the
Vol. 49, No. 1, January 1973
85
Academy’s collection of Scorpionida is one of the world’s largest.—P. H. Arnaud,
Jr., California Academy of Sciences, Golden Gate Park, San Francisco.
The principal speaker of the evening was Dr. Jerry A. Powell, Department of
Entomology, University of California, Berkeley. His illustrated talk was entitled:
“Diverse adaptations to arid habitats by ethmiid moths.”
Coffee and other refreshments were served during a social hour in the entomology
rooms following the meeting.—M. S. Wasbauer, Secretary.
Three Hundred and Forty-Sixth Meeting
The 346th meeting was the annual field day and picnic. It was held on Saturday,
20 May 1972, at Castle Rock Park near Mt. Diablo, Contra Costa County, California.
Members present (12) : J. E. Cronin, H. V. Daly, Jr., H. V. Davis, D. G.
Denning, R. Dutton, W. E. Ferguson, J. R. Gabel, E. G. Linsley, W. W. Middle-
kauff, C. B. Philip, M. S. and Joanne S. Wasbauer.
Visitors present (13) : Marcia Cronin, Loretta Denning, Pamela Dutton, Natasha
Field, Juanita Linsley, Gladys Philip, D. W. Price, Duane Price and children,
K. Wasbauer.
The day was overcast and cool, but did not prevent picnicking on the wooden
benches provided. Insects were not in evidence abundantly but Drs. Daly and
Price brought a group of entomology students from the University of California,
Berkeley, who collected, in addition to other things, a series of the Acrocerid
fly, Eulonchus sp. from sticky-monkey flowers, Mimulus aurantiacus. Dr. Daly
was nearly successful in collecting a specimen of an ant-mimic mirid bug, probably
Dacerla inflata (Uhler). -—M. S. Wasbauer, Secretary.
Three Hundred and Forty-Seventh Meeting
The 347th meeting was held Friday, 20 October 1972, in the Morrison Audi¬
torium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Denning presiding.
Members present (42) : R. P. Allen, J. R. Anderson, F. G. Andrews, C. Armin,
P. H. Arnaud, Jr., A. D. Bacon, D. J. Burdick, R. Bushnell, R. M. Brown, S.
Clement, Karen Corwin, C. Dailey, D. G. Denning, J. G. Edwards, F. Ennik,
W. E. Ferguson, A. J. Gilbert, A. R. and L. R. Gillogly, C. Goodpasture, E.
Grissell, A. R. Hardy, S. Haskett, L. Hawkins, T. E. Hewton, Jr., E. Kane, B.
Keh, H. B. Leech, A. E. Michelbacher, R. Peterson, J. A. Powell, R. F. Schoeppner,
R. Stecker, S. Szerlip, R. L. Tassan, R. W. Thorp, B. Villegas, M. S. Wasbauer,
S. P. Welles, Jr., R. H. Whitsell, S. C. Williams, D. L. Wilson.
Visitors present (39) : H. P. Allmendinger, L. Bezoh, G. Brady, Carolyn Cain,
Linda Campbell, Carolyn Dailey, Chris Duncan, Priscilla Ewing, Janie Edwards,
Stephenie Ferguson, Marcia Gilbert, Gwen Gillogly, Mr. and Mrs. B. W.
Grushkowitz, D. Guiliani, Debbi Hains, R. M. Haradon, Jr., Wanda Hardy, A.
Koplan, J. Krug, Val Landwehr, R., D. and B. Merritt, Martha Michelbacher,
R. P. Papp, R. Pemberton, Jane Rosemarin, W. E. Savary, R. E. Somerby, Chris
Springer, Wilfred Sumner, M. A. Taylor, Joyce Thorp, Martha Usinger, J. F.
Wells, D. Whitman, J. D. Wright, W. Yu.
The minutes of the meetings held 21 April and 20 May were summarized.
The following individuals were proposed for membership in the Society.
Student membership: H. P. Allmendinger, M. Browning, D. Chandler, B. W.
Grushkowitz, W. N. Mathis, R. Papp, J. R. Sawbridge. Regular membership:
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The Pan-Pacific Entomologist
J. A. Burnett, J. L. Gillespie, R. Goertzen, R. M. Haradon, Jr., W. W. Klopp,
R. J. McCrea, J. D. Pinto, R. Schick, W. S. Sibray, A. C. Smith, R. S. Toth.
Dr. Daly gave a brief resume of how the R. L. Usinger Autobiography came
into being, and suggested that the book would be inspirational reading for young
people who were thinking of careers in the natural sciences. He suggested that
members contact their local public libraries and ask that the book be placed
on their shelves.
President Denning then introduced Mrs. R. L. Usinger who gave sincere thanks
to all involved in the production of the autobiography. She also thanked the
membership for its support and for the specially bound presentation copies given
to the immediate members of the Usinger family. She pointed out that her late
husband did not conceive of the book or put it down on tape with the idea in
mind of having it published, but rather to leave a chronicle of his life for his
family members.
The following notes were presented:
Notes on a mass flight of Nymphalis californica (Boisduval) in the
Sierra Nevada. —On 26 May 1972 while driving east on California Route 140
from Merced into Yosemite National Park, my wife and I observed scattered
flights of the California tortoiseshell butterfly, Nymphalis calif ornica (Boisduval),
along the roadside and occasionally crossing the road. At 1300 hrs., at Red Bud
Campground on the Merced River in Sierra National Forest, Nymphalis calif ornica
was abundant especially on Aesculus calif ornica, which was flowering, and along
the river itself.
As we continued along the Tioga Road, tortoiseshells became increasingly
numerous, flying sometimes in small clouds in front of our car. We observed
many specimens dead or crippled in the road, which in places was dotted with
their bodies. Steller’s jays, Cyanocitta stelleri, and robins, Turdus migratorius,
were observed feeding on these insects, picking them up off the road and flying
to nearby trees with them.
Then suddenly, about 1430 hrs., near the overlook at Yosemite Creek Basin
(at 8,400 ft. elevation) the butterflies become incredibly abundant. Stopping our
car, we walked along the Tioga Road and observed large clouds of butterflies
streaming past us on the highway. The temperature in the sun, open air, at this
time was 80°F., with light northwesterly winds. As we crossed to the side of the
highway and looked down into the basin, we could clearly see large numbers of
Nymphalis californica welling up from the basin and flying NNW about 2-3 feet
above the ground. (The tortoiseshells thus appeared to be flying approximately
parallel to and west of the crest of the Sierra.) We estimated that 10-15 butter¬
flies per second were flying past a fixed point. As they crossed the terrain below
us, they were easily seen, silhouetted against the light-colored exfoliating granite.
Scanning the ground around us, we found many small patches of perhaps two
square feet containing up to 14 butterflies, which appeared to be resting. Some
patches were shaded, while others were in open sunlight but moist. A scrub jay,
Aphelocoma coerulescens, was observed here feeding on these insects. Perhaps this
bird, which was far from its normal habitat, had come up the creek basin following
and feeding upon Nymphalis.
During the next few hours we observed smaller numbers of Nymphalis californica
through Tioga Pass (9,943 ft.) and up into the meadows of Saddlebag Lake
Canyon below Mt. Conness. On 27 May during the late morning and early after-
Vol. 49, No. 1, January 1973
87
noon we found wings of Nymphalis calif ornica lying in melt holes in the snow at
altitudes up to 11,480 ft. on the ridge east of Mt. Conness.— Richard P. Papp,
University of California, Berkeley.
The principal speaker of the evening was Dr. Howell V. Daly, University of
California, Berkeley. His illustrated talk was entitled: “Bees, Twigs and Africa.”
Coffee and other refreshments were served during a social hour in the entomol¬
ogy rooms following the meeting.—M. S. Wasbauer, Secretary.
Three Hundred and Forty-Eighth Meeting
The 348th meeting was held Friday, 17 November 1972, in the Morrison Audi¬
torium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Denning presiding.
Members present (34) : F. G. Andrews, C. Armin, P. H. Arnaud, Jr., A. D.
Bacon, J. W. Bass, G. S. Benham, Jr., F. L. Blanc, G. Brady, T. Briggs, R. M.
Brown, J. A. Chemsak, Karen S. Corwin, T. Delgman, D. G. Denning, Beverly
Ehreth, C. P. Farrell, A. R. Gillogly, Lauren Green, J. Guggolz, K. S. Hagen,
A. R. Hardy, L. Hawkins, H. B. Leech, R. P. Papp, R. Peterson, C. B. Philip,
W. W. Pitcher, E. S. Ross, S. L. Szerlip, R. L. Tassan, T. J. Toren, D. Ubick,
M. S. Wasbauer, D. L. Wilson.
Visitors present (30) : Barbara Barr, Grace Bass, T. Christensen, J. Collins,
Loretta Denning, Margo Ebeling, R. Falls, Kathy Farrell, J. T. Hjelle, H. Implom,
D. Moss, N. E. Papp, R. W. Pemberton, D. Phillip, D. Roxon, Suzanne Roxon,
W. E. Savary, M. V. Silveira, J. Sorensen, Meta Sorensen, Mary Stark, G. L.
Weller, J. F. Wells, Patti Willey, Constance Wilson, R. Wong, Virginia Woo,
W. Yee, R. Zarate.
The minutes of the meeting held 20 October 1972 were summarized.
Persons who were proposed for membership in the Society: J. K. Wangberg,
R. L. Potts.
The following notes were presented:
In Memoriam—Segundo E. Valastegui B. (15 August 1910—1 July
1972) .—Senor Segundo E. Velastegui Barriga, entomologist and commercial
collector who was born at and resided in Banos, Ecuador, died from a skull
fracture and internal injuries received from a brutal beating. While returning
from his work on 28 June 1972, at a point called Ulvilla, near Banos, he was
attacked by a young man of about 25 years of age, without apparent reason. Don
Segundo was found on the road after the attack by a friend who brought him to
a hospital, but it was not possible to save his life.
Senor Velastegui was an excellent collector and he carefully prepared specimens
for his correspondents. Visiting extremely interesting areas of the Andes, he
collected all types of insects, birds, and orchids. His collections were sold to
persons and institutions throughout the world. The Academy and I received
nearly 1,000 specimens of Tachinidae in the year prior to his death. His widow
informs me that he started working at the age of 10, but I am not certain if this
refers to natural history collecting, or perhaps other type of work.
Senor Velastegui is survived by his wife, Etelvina and seven children (Marina,
Ysabel, Marta, Danilo, Guillermina, Gloria, and Oliverio). They intend to con¬
tinue the work of their husband and father and will collect all groups of en¬
tomological specimens from “todas partes del Oriente, y del Occidente” and other
parts of Ecuador. A drawer of Tachinidae collected by Senor Velastegui is on
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The Pan-Pacific Entomologist
display in the entomology rooms.— Paul H. Arnaud, Jr., California Academy of
Sciences, San Francisco.
Troglobitic harvestmen observed in lava caves of Washington and Idaho.
—Recently much interest in lava cave biology was generated by the discovery of
highly modified invertebrates in lava caves of Japan, Idaho and Hawaii. Hereto¬
fore, only limestone caves were thought to produce the characteristic modifications
of troglobites. Lava caves were assumed to be too small and too brief in geologic
time to evolve these modifications. The new discoveries call for a reexamination
of cave evolution. Recent explanations (F. Howarth, Science, 137:325, 1972)
suggest that deep breaks in lava flows provide routes of dispersal from collapsing
caves to new caves.
During the 1972 national convention of the National Speleological Society,
Robert Lem and I were able to locate several lava troglobites in Washington and
Idaho. The Washington caves were located near Mt. St. Helens in the Cascade
Range. This area is densely forested and receives much rainfall. Larger lava
caves contained a deep interior zone of a peculiar mold or yeast known as “lava
tube slime.” This moist growth tended to form under lava breakdown and occa¬
sionally coated the walls and ceilings with deposits up to an inch thick. Numerous
cavernicolous invertebrates lived in association with this food source, including
a remarkable laniatorid phalangid. It proved to be the first new world member
of family Travuniidae, a group of relicts well known in European limestone caves.
White juveniles and pale yellow adults were collected on the slimy habitat.
Specimens of another troglobitic phalangid received from Craters of the Moon
National Monument led us to travel to central Idaho. A lava cave was located
in a prairie habitat near the town of Shoshone. It had a narrow entrance to a
single large room which was particularly rich in life. The cave was unusually
warm and had a rock-strewn floor with mold-encrusted walls and ceiling. On
entering the room we noted that camel crickets and four families of flies were
numerous in the zone of darkness. Abundant evidence of mammal occupation was
seen, including three decapitated pigeons and a half-eaten pizza near a large
burrow. Under breakdown we found troglobitic beetles ( Glacicavicola of family
Leiodidae), epigean beetles, a cavernicolous centipede and large troglobitic
phalangids. These and the Craters of the Moon phalangids proved to be in
Erebomastridae, a family not previously known in Idaho.
Many workers are now looking at material from lava caves. I hope to describe
the cavernicolous harvestmen in the near future. —Thomas S. Briggs, Galileo
High School, San Francisco.
The principal speakers of the evening were Drs. W. H. Lange and J. Anderson,
University of California, Davis and Berkeley, respectively. Their illustrated talk
was entitled “Entomology Down Under.”
Coffee and other refreshments were served during a social hour in the entomology
rooms following the meeting.—M. S. Wasbauer, Secretary.
Three Hundred and Forty-Ninth Meeting
The 349th meeting was held Friday, 15 December 1972, in the Morrison Audi¬
torium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Denning presiding.
Members present (33) : R. P. Allen, P. H. Arnaud, Jr., R. G. Blair, F. L.
Blanc, G. Brady, D. Briggs, T. Briggs, D. J. Burdick, J. A. Chemsak, T. Delgman,
Vol. 49, No. 1, January 1973
89
D. G. Denning, A. R. Dutton, J. G. Edwards, F. Ennik, J. R. Gabel, M. R.
Gardner, A. J. Gilbert, A. R. Gillogly, K. S. Hagen, L. Hawkins, H. B. Leech,
R. W. Lem, A. E. Michelbacher, D. W. Moss, Jr., R. L. Peterson, C. B. Philip,
R. W. L. Potts, R. E. Stecker, S. L. Szerlip, R. W. Thorp, M. S. Wasbauer, S. C.
Williams, R. L. Wong.
Visitors present (21) : Fern Blanc, T. Coe, M. J. Croce, J. Denk, Alice Edwards,
B. Eya, G. B. Fairchild, Mrs. G. B. Fairchild, Jan and Toni Gabel, D. Gibbs,
Marcia Gilbert, Lorraine Kitajima, C. Kitayama, Martha Michelbacher, R. W.
Pemberton, W. F. Peregren, Peggie Phillips, W. Raunder, W. E. Savary, M.
Silveira, Joyce Thorp, D. Whitman.
The minutes of the meeting held 17 November 1972 were summarized.
The following names were proposed for membership in the Society. Student
membership: Joe Denk, Linda Floyd, David W. Moss, Jr., Manual Silveira and
Douglas Whitman. Regular membership: Ronald E. Somerby and Donald R.
Dilley.
President Denning asked for introductions from the floor. Dr. Philip introduced
Dr. and Mrs. G. B. Fairchild. Dr. Fairchild, a medical entomologist with a
distinguished career at the Gorgas Memorial Laboratory in Panama is now
retired. He and Mrs. Fairchild have just returned from a trip to Australia where
they were able to do some good collecting, especially for tabanids. They plan to
make their home at Gainesville, Florida.
The following notes were presented:
Prey records for robber flies of the Lake Texoma region, Oklahoma
(Diptera: Asilidae). —During June and July 1972 while I was at the University
of Oklahoma Biological Station at Lake Texoma, I collected a number of robber
flies. Most were collected during the day, but Diogmites symmachus Loew was
occasionally attracted to lights at night. The largest robber fly taken, Microstylum
moTosum Loew, had bright green eyes while alive. The most colorful robber fly
taken was Mallophora fautrix bromleyi Curran, a mimic of the common local
bumblebee, Bombus americanorum (F.). Although all my robber fly collecting was
incidental to other studies, my principal interest was in their prey associations.
The following eight species were taken with prey by me near the Biological Station,
two miles west of Willis, Marshall County, unless otherwise noted:
1. Diogmites neoternatus (Bromley), male and female with Apis mellifera
(Apidae), workers, 5 and 7 July.
2. Diogmites symmachus Loew, 3 females with Calliopsis andreniformis F. Smith
(Andrenidae), female, 30 June; Tachytes distinctus F. Smith (Sphecidae),
female, 11 July (L. S. B. Koch) ; and Sceliphron caementarium (Drury)
(Sphecidae), female, 29 July (L. S. B. Koch).
3. Efferia nemoralis Hine, female with Hemiargus ( Echinargus) isola Reakirt
(Lycaenidae), 7 June, 1.2 mi. E. Troy, Oklahoma.
4. Triorla interrupta (Macquart), male with Phyciodes phaon Edwards (Nymphali-
dae), 30 June; and female with Orthemis ferruginea F. (Libellulidae), 22 July,
5 mi. N. Gordonville, Texas (E. E. Grissell).
5. Proctacanthus hinei Bromley, 2 females with Apis mellifera, worker, 11 July;
and Bombus fraternus (F. Smith) (Apidae), worker, 20 July.
6. Promachus bastardii (Macquart), male with Apis mellifera, worker 9 June;
and 2 females with Alysson melleus Say (Sphecidae), male, 20 June; and
Halictus ligatus Say (Halictidae), male, 14 July.
90
The Pan-Pacific Entomologist
7. Saropogon dispar Coquillett, 2 females with Sceliphron caementarium, male,
5 June; and Apis mellifera, worker, 2 July; 2 males with Trypoxylon
( Trypargilum ) texense Saussure (Sphecidae), male, 9 June; and Dialictus sp.
(Halictidae), female, 21 July, Powell, 5 mi. N.W. Willis, Oklahoma.
8. Stichopogon trifasciatus (Say), male with (Araneae: Salticidae), 5 June,
1.2 mi. E. Troy, Oklahoma.
The prey of Diogmites, Proctacanthus, Promachus and Saropogon consisted
entirely of aculeate Hymenoptera. These four robber fly genera contain species con¬
sidered serious predators of the honey bee. However, Diogmites symmachus, com¬
monly referred to as a “Texas bee-killer,” was the only species of the group not
observed with honey bees as prey. Comparative studies of prey selection by the two
sympatric species of Diogmites would be of special importance in determining
how they divide up their food requisite.
Most of the predators were as large as or larger than their prey, except for the
female of Triorla which had taken a large dragonfly. The most unusual record
was the capture of a jumping spider by the male Stichopogon. This is an un¬
common habit for the family Asilidae, but has been reported for S. trifasciatus
(Bromley, 1934, Ann. Entomol. Soc. Amer., 27:74-110).
I thank the following for confirming or correcting my determinations: Dr. J.
Wilcox (Asilidae), Dr. R. M. Bohart (Sphecidae), and Mr. C. E. Goodpasture
(Lepidoptera).— Robbin W. Thorp, University of California, Davis.
The unexpected loss of one of our student members. —In 1961 Miss Linda
May Halverson was first introduced to the subject of entomology by taking a
course in General Entomology during the fall semester at San Francisco State
College. Through this experience she developed a most absorbing interest in
insects. So much so that she began to study insects every possible opportunity
and began to make a comprehensive personal collection. By the first of 1972 Miss
Halverson had acquired a considerable knowledge of insects and had decided
that she wanted to enter a career in entomology, most specifically in the area of
medical entomology. On 18 February 1972, she was elected to membership in the
Pacific Coast Entomological Society. Meanwhile, she continued her studies in
entomology at San Francisco State College, became Curator of the college’s
insect and arachnid collection, and became a laboratory assistant in the general
entomology courses. During the summer of 1972, Miss Halverson secured a
position with the Entomology Department of the California Academy of Sciences
as a clerical assistant. With the coming of the fall 1972 semester at San Francisco
State, Miss Halverson began to prepare for graduation and began preparing her
application for graduate study in entomology at the University of California at
Berkeley. By this time Miss Halverson had shown a rare gift for the study of
insects, and a personal interest not commonly encountered. She also had proven
herself as a truly exceptional student. All of her plans and aspirations came to
a sudden halt on 25 November 1972 when she died suddenly, as the result of an
automobile accident on the San Francisco Bay Bridge. With her death we have
lost both an exceptional entomology student and a good friend.— Stanley C.
Williams, Division of Biology, San Francisco State University.
A new technique for study of tarantula burrows.— Study of the burrows
of tarantulas (Theraphosidae) has always been hard, sweaty work for it has been
necessary to follow the pathways of the burrow by digging in hard soil. Frequently
Vol. 49, No. 1, January 1973
91
the “path” is lost when the shovel slides earth into the hole and obscures it.
A pliable piece of wire or some similar object thrust into the burrow during
digging helps to follow the direction. Digging also obscures any of the probable
side chambers dug by the spider. On only one occasion has the author found a
side chamber made by the spider. It contained the abandoned excuvia from a
previous shedding. Past attempts to “cast” burrows by pouring plaster of Paris
into the hole have been unsatisfactory for the hardening time of the plaster varies
with the fluidity of the mixture. If the tunnel turns the least bit upward in its
meanderings, the plaster sometimes stops flowing and the extent of the burrow
is lost.
During a visit to the Southwest Research Station of the American Museum of
Natural History near Portal, Arizona, this past summer, the director, Vince Roth,
suggested I try injecting foam plastic material into the burrow and then follow
the contours when this material hardened. Insta-Foam Products, Inc., Addison,
Illinois manufactures an aerosol eurethane foam for “insulating, repairing, packag¬
ing, sound dampening, sealing, caulking, display building and many other uses.”
It is packaged in a double tank unit called a “Froth-Pale” which comes with two
dispenser nozzles with hoses. The two liquids are mixed by the nozzle after ex¬
trusion from the pressurized cans. It hardens into a yellowish foam solid within
60 to 90 seconds. This is sufficient time for the liquids and gas to penetrate all
parts of the burrow. The burrow can be carefully excavated by following the
pathway of the urethane foam which has completely filled all the crevices and
side chambers of the burrow.
My first attempt was successful and four burrows were filled with the foam
plastic and subsequently dug out. The soil surrounding the casting was carefully
scraped away using a soft stick. It was then washed under running water and
carefully scrubbed with a stiff bristle hairbrush. The resulting cast shows all
the twists, turns and side chambers of the tunnel and even revealed that two of
the burrows had two entrances or an “escape tunnel” connected to the main
excavation.
One of the castings contained a discarded excuvia of the former occupant. All
burrows “cast” were unoccupied at the time for I had previously washed the
spiders out for transplanting experiments. The foam would undoubtedly be fatal
to the spider remaining in the burrow. The manufacturer assured me that washing
the arachnid out of the burrow prior to “casting” would in no way effect the
performance of the foaming plastic.
Because the rainy season in California comes during the winter, there is a
minor drawback in “casting” burrows at this time of the year. The Insta-Foam
has a critical temperature of 60° to 70° F. Below that temperature it does not
harden nor flow out of the cans properly. If the day is sunny and warm, there is
no difficulty in using the “twin pak,” but on overcast days it may be necessary
to use an in-car heater to assure that the two cans in the pak are at the proper
temperature. Complete instructions for use are on the side of the package, and
experience alone teaches the operator when to shut off the pressure to avoid
wasting the foam in “mushroom heads” which form above the burrow entrances
as excess plastic extrudes itself from the burrow opening. — J. Russel Gabel,
Division of Biology, California State University at San Francisco.
The December meeting is traditionally the business meeting of the year and
President Denning asked for the reports of the standing committees.
92
The Pan-Pacific Entomologist
In the treasurer’s report, Dr. Arnaud gave the total income for the Society in
1972 as $13,581.00. This was derived from dues and subscriptions, reprints and
miscellaneous page charges, sales of memoirs, advertising income, interest on
savings accounts and stock dividends. The total expenditures for the year were
$14,194.00 for the following items: publication costs of the Pan-Pacific En¬
tomologist, publication costs of the R. L. Usinger autobiography, reprints, postage
and other miscellaneous expenses. The overall decrease in fund balances was
thus $613.00. Dr. Arnaud expressed appreciation to Mr. H. V. Davis who audited
the books for the Society again this year. He also thanked Mr. Davis for his
help in maintaining the tax-exempt status of the Society with the Internal
Revenue Service.
The report of the publication committee was presented by Dr. Chemsalc in
Dr. Daly’s absence and covered the appearance of R. L. Usinger’s autobiography
in the memoirs series of the Society.
Mr. Leech presented the report of the historical committee for 1972. He
addressed himself to the use of the Society’s biographical forms which are sent
to each new member. Among his observations are the following: of 287 past
and present members who have filled out these forms, first letters of the surnames
include all letters of the alphabet except Q, X and Y. Data from these forms in¬
dicate that the Society as a group is a serious environmental factor. According
to the statements on the sizes of private collections, our group is responsible for
catching or at least acquiring 3,079,000 insects. The following persons have listed
their holdings as 100,000 specimens or more: C. P. Alexander, 100,000 Diptera;
D. D. Jensen, 100,000 Homoptera; G. H. Nelson, 100,000 Coleoptera; C. W.
O’Brien, 100,000 Coleoptera; J. T. Polhemus, 100,000 Heteroptera; J. Schuh,
100,000 Coleoptera; J. S. Buckett, 150,000 Lepidoptera; J. G. Franclemont,
350,000 Lepidoptera. The following reported 50,000 or more in their collections:
F. R. Cole, Diptera; H. R. Dodge, Diptera; C. H. Martin, Diptera; Ian Moore,
Coleoptera; J. R. Powers, Coleoptera; R. L. Usinger, Heteroptera; E. L. Sleeper,
Coleoptera; P. S. Bartholomew, Coleoptera; S. C. Williams, Scorpionida; J. G.
Edwards, Coleoptera. The total figure does not include the hundreds of thousands
of specimens collected by such members as R. M. Bohart and E. S. Ross and
later donated to various institutions.
Dr. Williams read the report of the membership committee which stated that
26 regular members and 25 student members were elected to Society membership
in 1972, a total of 51 new members for the year.
President Denning announced his appointments to the publication committee
for 1973 through 1975: Dr. J. G. Edwards and Dr. C. B. Philip.
Dr. R. Stecker, Chairman of the nominating committee, presented the slate of
candidates for office in the Society for 1973: President, John A. Chemsak;
President-elect, Cornelius B. Philip; Secretary, Michael R. Gardner; Treasurer,
Paul H. Arnaud, Jr. There were no nominations from the floor. The candidates
were elected to office in the Society for 1973 by unanimous vote.
The principal speaker of the evening was the outgoing President of the Society,
Dr. D. G. Denning. His presidential address was entitled, “A resume of the order
Trichoptera, with emphasis on western North America.”
Coffee and other refreshments were served during a social hour held in the
entomology rooms following the meeting.—M. S. Wasbauer, Secretary.
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ENTOMOLOGICAL EXCHANGES AND SALES
Space is available to members and non-members who wish to advertise entomo¬
logical exchanges, sales, announcements, etc. We can offer an ad in four issues
for $1.50 per line. Ads will be arranged in order of their receipt. Contact the
Advertising Manager, W. H. Lange, Department of Entomology, University of Cali¬
fornia, Davis, California 95616.
Current developments in the sugarbeet industry in the United States are pin¬
pointed in ADVANCES IN SUGARBEET PRODUCTION: PRINCIPLES AND
PRACTICES , just published by the Iowa State University Press. The control of
insects, plant diseases, nematodes and weeds are discussed in addition to other
phases of production. Available at $12.50 from bookstores or from the Iowa
State University Press.
FOR SALE. Orders solicited for Pacific Northwest terrestrial and aquatic insects.
Queries answered promptly. S. G. Jewett, Jr., Route 1, Box 339, West Linn,
Oregon 97068.
FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca,
Department of Biology, University of Arizona, Tucson, 85721.
FOR SALE. Over 900 species of butterflies from the Malaysian and Indonesian
Jungles, very rare moth (14" wing span), Scorpion, giant sized Indonesian flying
lizard, centipedes, stick insects, all types of rhinoceros beetles and rare beetles
in the families Scarabaeidae, Lucanidae, Mordellidae, Cerambycidae, etc. Write
Sim Yam Seng, 21, Joon Hiang Road, Singapore, 19, Singapore.
THE MOTHS OF AMERICA
NORTH OF MEXICO
THE MOTHS OF AMERICA, one of the most significant publications in the history of
American entomology, will contain full-color photographs of all 10,000-)- species of North
American moths. The color photography has been done with excruciating care and attention
to detail, and must be seen to be believed.
THE MOTHS OF AMERICA, written by authorities in each group treated, constitutes a
thorough review of the North American moths, with descriptions of new genera and species
where appropriate.
In addition to the color photographs of adults, illustrations of genitalia or other structures
are included where necessary to assure accurate identification. The comprehensive text gives
a full taxonomic treatment, including synonymy, plus information on distribution, foodplants,
biology, and other aspects of each species.
THE MOTHS OF AMERICA will be published in over 100 parts, over a period of about 20
years. Each part is individually priced, according to its size, number of color plates, and
number of copies printed. Three to five parts are expected each year.
Special pre-publication/subscription prices, about 20% less than the regular list prices,
apply to subscriptions for the entire work and to orders received prior to the publication of
specific parts. (NOTE: The only way to get the parts already published at the reduced
prices is to subscribe to the entire series.)
Please start your subscription now, or order the parts you desire, before the cost be¬
comes prohibitive.
ALREADY PUBLISHED:
Fascicle 13, Part la, Pyralidae: Scopariinae & Nymphulinae, by Dr. Eugene G. Munroe.
1972. 134 p. $20.60 ($16.50 to subscribers).
Fascicle 13, Part lb, Pyralidae: Odontiinae & Glaphyriinae, by Dr. Eugene G. Munroe.
1972. 116 p. $20.60 ($16.50 to subscribers).
Fascicle 20, Part 2a, Bombycoidea: Saturniidae, in part (Citheroniinae and Hemileucinae, in
part), by Dr. Douglas C. Ferguson. 1972. 153 p., 11 color plates, 19 figs. $39.00
($32.50 to subscribers).
Fascicle 20, Part 2b, Bombycoidea: Saturniidae, in part (Hemileucinae, in part, and Satur-
niinae), by Dr. Douglas C. Ferguson. 1972. xxi 121 p., 11 color plates, 11 figs.
$39.00 ($32.50 to subscribers).
Fascicle 21, Sphingoidea, by Dr. Ronald W. Hodges. 1971. xii -f- 158 p., 16 plates (14 in
color), 8 halftones, 19 figs. $24.00 ($19.60 to subscribers).
IN PREPARATION:
Fascicle 20, Part 1, Bombycoidea (excluding Saturniidae) (i.e., Lasiocampidae, Mimalloni-
dae, Apetelodidae, and Bombycidae), by Dr. J. G. Franclemont.
Fascicle 13, Part 1c, Pyralidae: Evergestiinae, by Dr. Eugene G. Munroe (including 14
color plates for all 3 parts of Fascicle 13. 1).
Fascicle 6, Part 2, Gelecbioidea: Oecophoridae, by Dr. Ronald W. Hodges.
Fascicle 22, Part 2, Lymantriidae, by Dr, Douglas C. Ferguson.
Also in preparation: Notodontidae, Cosmopterygidae, Walshiidae, and Aegeriidae.
THE MOTHS OF AMERICA NORTH OF MEXICO belongs on the shelf of every serious
lepidopterist and in every significant library. Please make sure that your local library sub¬
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A fully descriptive brochure, with sample color plate and order form, is available on
request.
THE MOTHS OF AMERICA NORTH OF MEXICO is distributed exclusively in North
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THE PAN-PACIFIC ENTOMOLOGIST
Information for Contributors
Papers on the systematic and biological phases of entomology are favored, including
articles up to ten printed pages on insect taxonomy, morphology, behavior, life history,
and distribution. Excess pagination must be approved and may be charged to the author.
Papers are published in approximately the order that they are received. Immediate publi¬
cation may be arranged after acceptance by paying publication costs. Papers of less than
a printed page may be published as space is available, in Scientific Notes.
Manuscripts for publication, proof, and all editorial matters should be addressed to the
Editor, Robbin W. Thorp, Department of Entomology, University of California, Davis,
California 95616.
Typing. —All parts of manuscripts must be typed on bond paper with double-spacing and
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First page. —The page preceding the text of the manuscript should include (1) the
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Names and descriptions of organisms. —The first mention of a plant or animal should
include the full scientific name with the author of a zoological name not abbreviated.
Do not abbreviate generic names. Descriptions of taxa should be in telegraphic style.
References. —All citations in text, e.g., Essig (1926) or (Essig, 1958), should be listed
alphabetically under Literature Cited in the following format:
Essig, E. 0. 1926. A butterfly migration. Pan-Pac. Entomol.,
2 : 211 - 212 .
1958. Insects and mites of western North America. Rev.
ed. The Macmillan Co., New York. 1050 pp.
Abbreviations for titles of journals should follow the list of Biological Abstracts, 1966,
47(21) : 8585-8601.
Tables. —Tables are expensive and should be kept to a minimum. Each table should be
prepared as a line drawing or typed on a separate page with heading at top and foot¬
notes below. Number tables with Arabic numerals. Number footnotes consecutively
for each table. Use only horizontal rules.
Illustrations. —No extra charge is made for the line drawings or halftones. Submit only
photographs on glossy paper and original drawings (no photographs of drawings).
Authors must plan their illustrations for reduction to the dimensions of the printed
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Page charges. —All regular papers of one to ten printed pages are charged at the rate of
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All articles are accepted for publication only on the basis of scientific merit
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PACIFIC COAST ENTOMOLOGICAL SOCIETY
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MEMOIR SERIES.
Volume 1 . The Sucking Lice by G. F. Ferris. A 320-page book
which summarizes the knowledge of the Anoplura of the world.
Published October 1951. Price $6.00. (Plus 350 postage.)*
Volume 2. The Spider Mite Family Tetranychidae by A. Earl
Pritchard and Edward W. Baker. This worldwide treatment deals
with the systematics, identification, and economics of the “Red
Spiders 5 ' and includes descriptions of 33 new species. 472 pages.
Published July 1955. Price $10.00. (Plus 500 postage.) "
Volume 3. Revisionary Studies in the Nearctic Decticinae by David
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13 new species and subspecies. Published July 1968. Price $4.00.
(Plus 250 postage.)*
Volume 4. Autobiography of an Entomologist by Robert L.
Usinger. A 343 page book relating the life story of one of the
outstanding entomologists of our time. Published August 1972.
Price $15.00. (Plus 400 postage.) *
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Vol. 49
APRIL 1973
No. 2
THE
Pan-Pacific Entomologist
SMITH—The sawfly tribe Lycaotini in South America (Hymenoptera:
Tenthredinidae) . 93
FRANKIE—Feeding habits and seasonal history of Ernobius conicola in
cones of Monterey cypress with notes on cohabiting insects (Coleop-
tera: Anobiidae) . 102
BENTZIEN—Biology of the spider Diguetia imperiosa (Araneida: Di-
guetidae) . 110
TORGERSEN—A new species of Spinolochus from North America (Hyme¬
noptera: Ichneumonidae) . 124
HARDY—A new species of Phobetus (Coleoptera: Scarabaeidae) . 127
DENNING—New species of Trichoptera . 182
ALCOCK AND RYAN—The behavior of Microbembex nigrifrons (Hyme¬
noptera: Sphecidae) . 144
GOODPASTURE—A new subspecies of Plebejus acmon (Lepidoptera:
Lycaenidae) . 149
ALLEN—New Ephemerellidae from Madagascar and Afghanistan (Ephem-
eroptera) ... 160
SOLEGLAD—A redescription of the scorpion Vejovis flavus (Scorpionida:
Vejovidae) . 165
DAILEY AND SPRENGER—Unisexual generation of Andricus atrimentus
(Hymenoptera: Cynipidae) . 171
DAILEY AND CAMPBELL—A new species of Diplolepis from California
(Hymenoptera: Cynipidae) . 174
ARNAUD AND WALE—Thomas Wrentmore Cook (1884-1962) . 177
SCIENTIFIC NOTES . 126, 182
BOOK REVIEWS . 101, 123, 143, 176, 184
RECENT LITERATURE .. 148, 173
EDITORIAL NOTICE . 184
SAN FRANCISCO, CALIFORNIA • 1973
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
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The Pan-Pacific Entomologist
Vol. 49
April 1973
No. 2
The Sawfly Tribe Lycaotini in South America
(Hymenoptera: Tenthredinidae)
David R. Smith 1
Systematic Entomology Laboratory, Agiicultural Research Service, USDA
The tribe Lycaotini is represented by two genera in South America
and is found in Ecuador, Bolivia, southern Brazil, and Chile. These
genera are related to, but distinct from, Lycaota Konow and Blenno-
generis MacGillivray, the two lycaotine genera found in western North
America. This distribution represents another example of the occur¬
rence of closely related forms in North and South America. Other
bipolar taxa of the Tenthredinidae are Pristiphora Latreille in North
America and southern Brazil, Periclista Konow in North America and
Chile and southern Argentina, and the Fenusini with the genera Pro-
fenusa MacGillivray and Bidigitus Smith in North America closely
related to Notofenusa Benson and Brasinusa Malaise in Chile, Argentina,
and southern Brazil. The northernmost species of Lycaotini in South
America is Synaptoneura vopida n. sp. in Ecuador. No other repre¬
sentatives of the tribe are found to the north until Lycaota bouquetetisis
Smith is encountered in southern California.
Benson (1966) gave a key to eight world genera of Lycaotini. He
included only the genus Trichotaxonus Rohwer (= Kuschelia Malaise)
from South America, neglecting to mention Synaptoneura Konow.
Smith (1969) defined the tribe and treated the Nearctic species. This
tribe is in the subfamily Blennocampinae and is distinguished by the
venation of the forewing as follows: Veins M and Rs + M meet Sc + R
at the same point; veins M and Im-cu parallel; and vein 2A and 3A
complete, fused to 1A for short distance at center or slightly separated
from 1A but connected to 1A by a very short crossvein. Vein 2A and
3A is sometimes separated from 1A in the species Trichotaxonus coquim-
bensis (Spinola) but this is a variable character and this vein is usually
fused to 1A. Specimens which have vein 2A and 3A separated may be
1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560.
The Pan-Pacific Entomologist 49: 93-101. April 1973
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The Pan-Pacific Entomologist
confused with the Allantinae, but the Allantinae have a long, oblique
anal crossvein and vein 2A and 3A is widely separated from 1A.
Another genus of Blennocampinae, Metaneura Malaise, has a small
basal anal cell in the forewing formed by the basal stub of vein 2A and
3A curving up and meeting 1A. This genus, however, is more closely
related to other genera of Blennocampinae in which the basal stub of
2A and 3A is curved up but not meeting 1A; therefore, it is not included
in the Lycaotini.
The kind cooperation of the following has made this review of South
American Lycaotini possible: Dr. J. Oehlke, Institut fur Pflanzen-
schutzforschung, Eberswalde, Germany; Dr. Per Inge Persson, Natur-
historiska Riksmuseum, Stockholm, Sweden; Dr. B. Pisarski, Polska
Akademia Nauk, Instytut Zoologiczny, Warsaw, Poland; Dr. M. Zunino,
Museo ed Istituto di Zoologia Sistematica, Universita di Torino, Torino,
Italy; and Dr. Paul H. Arnaud, Jr., Department of Entomology, Cal¬
ifornia Academy of Sciences, San Francisco.
Key to the Genera of Lycaotini
Tarsal claws bifid, with basal lobe; malar space distinct _ Synaptoneura Konow
Tarsal claws trifid, without basal lobe; malar space linear_
- Trichotaxonus Rohwer
Synaptoneura Konow
Synaptoneura Konow, 1908, p. 160. Type-species: Synaptoneura boliviensis
Konow. Monotypic.
Description. —Antenna long, slender, first and second segments each longer
than broad, third segment slightly longer than fourth segment, segments beyond
third gradually decreasing in length (Fig. 7). Clypeus with shallow central emar-
gination; malar space distinct, nearly as wide as diameter of front ocellus; no genal
carina; eyes small, converging below but farther apart below than length of an
eye; postocellar area broader than long. No prepectus; distance between cenchri
equal to breadth of a cenchrus. Tarsal claw bifid, inner tooth slightly shorter
than outer tooth and teeth closely appressed; basal lobe distinct, acute (Fig. 5) ;
hindbasitarsus shorter than following segments combined. Forewing with vein
2A and 3A complete, fused with 1A at center. Hindwing with cell M present;
anal cell sessile; apex of radial cell close to margin of wing, with short spurious
vein. Male of S. vopida with peripheral vein in hindwing. Wings as in Fig. 1.
Fig. 1, forewing and hindwing of Trichotaxonus coquimbensis, $. Fig. 2, anal
cell of forewing of T. coquimbensis, $, variation. Fig. 3, hindwing of T. coquim-
bensis, $. Fig. 4, tarsal claw of T. coquimbensis. Fig. 5, tarsal claw of Synap¬
toneura vopida. Fig. 6, antenna of T. coquimbensis. Fig. 7, antenna of S. vopida.
Vol. 49, No. 2, April 1973
95
Fig. 8, female sheath of T. coquimbensis, a. lateral and b. dorsal. Fig. 9, female
sheath of S. boliviensis, a. lateral and b. dorsal. Fig. 10, female sheath of S.
vopida, a. lateral and b. dorsal.
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Discussion. —In the only known specimen of S. boliviensis, a female,
the antennae are missing; therefore, the above characters given for the
antenna as well as for the male are based on S. vopida.
This genus is distinguished by the broad malar space and the bifid
tarsal claws with a basal lobe.
Key to Species of Synaptoneura
Abdomen black; hindfemur and hindtibia orange; female sheath with slender,
protruding scopa (Fig. 9a, 9b) _ boliviensis Konow
Abdomen, except for apical two or three segments, orange; hindlegs black;
female sheath with short, blunt scopa (Fig. 10a, 10b) _ vopida Smith, n. sp.
Synaptoneura boliviensis Konow
Synaptoneura boliviensis Konow, 1908, p. 160. $.
Female. —Length, 7.2 mm. Antenna and head black (flagellum missing).
Thorax black. Legs black with each femur except for extreme base, outer surface
of foretibia, and hindtibia except for extreme apex, orange. Abdomen black with
very narrow white band on posterior margin of each segment. Wings uniformly
lightly infuscated; veins and stigma black. Sheath with slender protruding scopa
at apex, from above widened toward apex, in lateral view long, straight above and
rounded below (Fig. 9a, 9b). Serrulae of lancet far apart, rounded, those near
center symmetrical, those at apex asymmetrical.
Male. —Unknown.
Type. —Female, at the Institut fur Pflanzenschutzforschung, Ebers-
walde, Germany, labeled “Mapiri, Bolivia,” “Coll. Konow,” “typus”
[red label], and the handwritten name label “Synaptoneura boliviensis
Knw., Bolivia.”
Distribution. —Known only from the type locality.
Discussion.— This species is distinguished from S. vopida by the
black abdomen, partly orange legs, and the structure of the female
sheath, as illustrated.
Synaptoneura vopida Smith, new species
Female. —Average length, 7.0 mm. Antenna and head black. Thorax black,
only upper portion of metapleuron with orange spot. Legs black, apical half of
forefemur, extreme base of foretibia, and apex of midfemur orange. Abdomen
orange with basal plates, apical two or three segments, and mesal stripe on venter
black; stemites except for apical two or three segments sometimes all orange.
Wings uniformly, lightly infuscated; veins and stigma black. Antenna long,
slender, about two and one-half times width of head. Sheath with short, blunt
scopa, from above broad and slightly widened toward apex, in lateral view, straight
above and rounded below (Fig. 10a, 10b). Lancet with serrulae shallow, rounded
and far apart (Fig. 12).
Vol. 49, No. 2, April 1973
97
Male.—A verage length, 6.5 mm. Color and structure similar to that of female.
Hindwing with peripheral vein. Harpe and parapenis as in Fig. 15; penis valve
rectangular, with lateral spine (Fig. 16).
Holotype female , Lago Zurucuchu, 11 mi. W. of Cuenca, Ecuador,
16 February 1965, E. I. Schlinger and E. S. Ross. At the California
Academy of Sciences, San Francisco.
Paratypes. —Same data as for holotype (5$, 18$); Ecuador, 45
mi. S. Alausi, Chimborazo, 3,000 m, 20 February 1955, E. I. Schlinger
and E. S. Ross (12 ) ; Ecuador, 40 mi. S. Alausi, Chimborazo, 20 Feb¬
ruary 1955, E. I. Schlinger and E. S. Ross (12).
Discussion. —This species is separated from S. boliviensis by the
mostly orange abdomen, black hindlegs, and lack of a protruding scopa
of the sheath. The species name is an arbitrary combination of letters
and is to be treated as a noun.
Trichotaxonus Rohwer
Trichotaxonus Rohwer, 1910, p. 30. Type-species: Trichotaxonus reedi Rohwer.
Orig. desig.
Kuschelia Malaise, 1949, p. 21. Type-species: Netrocerus solox Enderlein. Orig.
desig. New synonymy.
Description.—A ntenna with first and second segments each longer than broad,
third segment longer than fourth segment, segments beyond third gradually de¬
creasing in length, the apical four segments not reduced (Fig. 6). Clypeus with
small circular emargination; malar space linear; no genal carina; each mandible
bidentate; eyes small, distance between eyes below longer than eye length; post-
ocellar area as long as broad. No prepectus; distance between cenchri equal to
breadth of a cenchrus. Each tarsal claw trifid, without basal lobe (Fig. 4) ; hind-
basitarsus shorter than following tarsal segments combined. Forewing with vein
2A and 3A complete, either fused to 1A at center forming suhequal basal and
distal anal cells or connected to 1A by very short anal crossvein at this point.
Hindwing with cell M present; anal cell with very short petiole or sessile; radial
cell ends slightly before apex of wing and with short spurious vein. Male of
coquimbensis with peripheral vein in hindwing. Wings as in Figs. 1-3.
Discussion. —The trifid tarsal claws will separate Trichotaxonus from
Synaptoneura. For avoidance of possible confusion of this genus with
Antholcus Konow (Allantinae), see discussion under T. coquimbensis.
Key to Species of Trichotaxonus
Long flexuous hairs on thorax, head, and legs; thorax black, abdomen, except
for apical segments and basal plates, orange; clypeus with slight central
emargination; Chile_ coquimbensis (Spinola)
Long hairs absent, vesture of short pubescence; thorax mostly red, abdomen
black; clypeus truncate; Brazil _ luteithorax (Enderlein)
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The Pan-Pacific Entomologist
Fig. 11, female lancet of Trichotaxonus coquimbensis. Fig. 12, female lancet
of Synaptoneura vopida. Fig. 13, male harpe and parapenis of T. coquimbensis.
Fig. 14, penis valve of T. coquimbensis. Fig. 15, male harpe and parapenis of
S. vopida. Fig. 16, penis valve of S. vopida.
Vol. 49, No. 2, April 1973
99
Trichotaxonus coquimbensis (Spinola), new combination
Tenthredo coquimbensis Spinola, 1851, p. 556.
Blennocampa chilensis Brethes, 1910, p. 145. $ . New synonymy.
Zarca chilensis : Brethes, 1919, p. 51 ( $ ).
Trichotaxonus reedi Rohwer, 1910, p. 30. $ (correction of sex on insert
between pages 30 and 31). New synonymy.
Netrocerus solox Enderlein, 1920, p. 372. $ $ . New synonymy.
Kuschelia solox: Malaise, 1949, p. 22; Benson, 1966, p. 75.
Female.— Length, 9.0 mm. Antenna and head black. Thorax black. Legs black
with apical half of forefemur, outer surface of foretibia, outer surface of mid¬
tibia whitish, and hindfemur entirely orange. Abdomen orange with basal plates
and apical two segments and sheath black. Wings uniformly darkly infuscated;
veins and stigma black. Head, thorax, and legs covered with long flexuous hairs,
longer than length of first antennal segment. Clypeus with shallow central emar-
gination; postocellar area about as long as broad. Vein 2A and 3A of forewing
either fused to 1A at center or separated from 1A by very short anal crossvein.
Sheath from above uniformly broad, in lateral view straight above, rounded below
(Fig. 8a, 8b). Serrulae of lance flat, with no anterior and 10 to 13 posterior
subbasal teeth (Fig. 11).
Male.— Average length, 8.2 mm. Color and structure as for that of female.
Harpe and parapenis as in Fig. 13; penis valve with lateral spine (Fig. 14).
Types. —The type of T. coquimbensis Spinola is in the Museo ed
Instituto di Zoologia Sistematica, Universita di Torino, Italy. Three
specimens are under the name coquimbensis , one of which is a female
labeled “Typus” and “Kuchelia coquimbensis Spinola, det. R. Malaise,
1955.” The other specimens are a male and female which lack labels
except for “ $ ” and “Antholcus varinervis Spinola, det. R. Malaise,
1955” on the female. The specimen labeled “Typus” is the species
treated here and it agrees well with Spinola’s description. The other
two specimens are Antholcus varinervis (Spinola) as they agree with
Spinola’s description and follow the traditional usage of the name.
Specimens which may be the types of that species have not been located.
Spinola did not state how many specimens he examined; however, in
order to fix the identity of T . coquimbensis , I am hereby designating
the specimen labeled “Typus” as lectotype.
The type of Blennocampa chilensis Brethes may be at the Museo
Argentino de Ciencias Naturales Bernardino Rivadavia, Buenos Aires.
I have not been able to examine the specimen, but it is clear from
Brethes’ descriptions of 1910 and 1919 that this is the same as T.
coquimbensis.
Rohwer’s types of T. reedi , four males, are in the U. S. National
Museum, type no. 13076, and each is labeled “Chile, E. C. Reed.”
Three specimens are the species treated here, and I am hereby desig-
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The Pan-Pacific Entomologist
nating one of them as lectotype. The other specimen is Antholcus
varinervis (Spinola).
The type of Netrocerus solox Enderlein is in the Polska Akademia
Nauk, Instytut Zoologiczny, Warsaw, a female labeled “Chile,” “Type”
[red label], and a handwritten name label “Netrocerus solox Enderlein,
2 , Dr. Enderlein det. 1918.”
Distribution. —Chile. My confirmed records: Cubillo, Cord. Curico,
24-26 January 1961; El Pangue, Coquimbo, 1,500 m., 3-5 November
1961, 4 November 1961; El Manzanito, 24 October 1948; Estero Cab¬
rera, Cord. Curico, 9 February 1961; El Coigo, Cord. Curico, February
1961; Aeon. Zapallar, 15 December 1950.
Discussion. —The black thorax, orange abdomen, and long hairs
of the head, thorax, and legs will distinguish this species from T. lutei-
thorax. It is more likely to be confused with Antholcus varinervis
(Spinola), a species in the Allantinae from Chile, because of the sim¬
ilarity of coloration and variation of the anal veins in T. coquimbensis.
Antholcus varinervis has the abdomen entirely orange, each femur and
tibia mostly orange, lacks long hairs, has a long oblique anal crossvein
in the forewing, and has the anal cell of the hindwing petiolate. In T.
coquimbensis, vein 2A and 3A may be separated from 1A but is con¬
nected to 1A by a very short crossvein.
Trichotaxonus LUTEITHORAX (Enderlein), new combination
Lycaota luteithorax Enderlein, 1920, p. 370. 2.
Female. —Length, 8.3 mm. Antenna and head black. Thorax orange with
mesoscutellum and metanotum black. Legs and abdomen black. Wings uniformly,
black infuscated; veins and stigma black. Pubescence on head and thorax much
shorter than length of first antennal segment. Clypeus truncate; labrum short
and broad, truncate; postocellar area slightly broader than long. Sheath thin
from above, in lateral view, straight above, rounded below. Vein 2A and 3A of
forewing nearly fused at center with 1A, very short crossvein present (similar
to Fig. 2).
Male.— Unknown.
Type. —At the Polska Akademia Nauk, Instytut Zoologiczny, Warsaw,
Poland, a female labeled “S. Catarina, Luderwaldt,” “Type,” [red
label], and a handwritten determination label “Lycaota luteithorax
Enderlein, 2 type, Dr. Enderlein det. 1918.”
Distribution. —Known only from the type locality in Brazil.
Discussion. —A distinctive species, T. luteithorax is separated from
T. coquimbensis by the mostly orange thorax, black abdomen, and lack
of long, flexuous hairs on the head, thorax, and legs.
Vol. 49, No. 2, April 1973
101
Literature Cited
Benson, R. B. 1966. A new genus of the Lycaotini (Blennocampinae) in
Turkey (Hymenoptera: Tenthredinidae). Proc. Roy. Entomol. Soc.
London Ser. (B) Taxon., 35: 75-77.
Brethes, J. 1910. Quelques Hymenopteres du Chili. Rev. Chilena Hist. Natur.,
14: 141-146.
1919. Tenthredines nouveaux du Chili. Rev. Chilena Hist. Natur., 23: 49-52.
Enderlein, G. 1920. Symphytologica II. Zur Kenntnis der Tenthredininen.
Sitz. Gesell. Naturf. Freunde Berlin 9: 347-374 (Nov. 1919 issue,
published Jan. 1920).
Konow, F. 1908. Neue mittel- und siidamerikanische Tenthrediniden (Hym.).
Z. Syst. Hymenopt. Dipt., 8: 144-163.
Malaise, R. 1949. The genera Waldheimia, Probleta, and other Neotropical
Tenthredinoidea (Hym.). Ark. Zook, 42A: 1-61.
Rohwer, S. A. 1910. A new genus of sawflies from Chile (Hymenoptera:
Tenthredinidae). Proc. Entomol. Soc. Wash., 12: 30.
Smith, D. R. 1969. Nearctic sawflies I. Blennocampinae: Adults and larvae
(Hymenoptera: Tenthredinidae). U. S. Dep. Agr. Tech. Bull. 1397,
179 pp.
Spinola, M. 1851. Hymenoptera. In Gay, C., Historia fisica y politica de Chile,
Zoologia, v. 6, p. 153-572.
BOOK REVIEW
Monografia da Tribo Ibidionini (Coleoptera, Cerambycinae) . By Ubirajara
R. Martins. Partes I-VI. Arquivos de Zoologia, 16 (1-6, + estampas 1-30).
Parte I (30.XI.1967), pp. 1-320, figs. 1-180. Parte II (15.VIII.1968), pp. 321-
630, figs. 181-347. Parte III (30.IX.1969), pp. 631-878, figs. 348-454. Parte
IV (5.III.1970), pp. 879-1150, figs. 455-586 (pp. 878 and 1150 are blank). Parte
V (5.VIII.1970), pp. 1151-1341, figs. 587-683. Estampas 1-30 (30.IX.1970),
120 figs, in color. Parte VI (1971), pp. 1343-1508, figs. 684-710. From the
Museu de Zoologia da Universidade de Sao Paulo, Avenida Nazare 481, Caixa
Postal 7172, Sao Paulo, S. P. Brasil. Price not stated.
The book-sized parts of this revisional study indicate the amazing extent of this
neotropical tribe (Old World genera, earlier placed in it, are now excluded), and
the high proportion of new genera show the need there has been for restudy.
North American species are of course included. The work is replete with keys for
identification, redescriptions, figures of whole beetles and dissections of mouth-
parts, genitalia, etc. There are new data on the habits of adults and the host plants
of larvae; the 120 figures in color are uniformly fine; the work contains a broad
spectrum of data, good mining ground for other than coleopterists, and Dr. Martins
should receive the thanks of all. A supplementary paper describing 17 new species
and designating lectotypes for Lacordaires species also appeared in 1971 (Arq.
Zool. Estado Sao Paulo, 21 (3) : 121-178, 25 figs.).— Hugh B. Leech, California
Academy of Sciences, San Francisco, 94118.
102
The Pan-Pacific Entomologist
Feeding Habits and Seasonal History of Ernobius conicola in
Cones of Monterey Cypress with Notes on Cohabiting Insects
(Coleoptera: Anobiidae)
Gordon W. Frankie
Department of Entomology, Texas A & M University, College Station, 77843
Most species in the family Anobiidae are poorly understood, bio¬
logically and ecologically (White 1962, pers. comm.). Members of this
beetle family are almost always characterized as infesting dying or dead
plant material (Arnett 1968, Baker 1972, Borror and DeLong 1971,
Hatch 1961). However, some species, e.g. in the genus Ernobius , are
known to occur in both living and dead plant tissue (Arnett 1968, Fisher
1919, Hatch 1961, Keen 1958, Ruckes 1957).
Ernobius conicola Fisher has been reported infesting green and dry
cones of Monterey cypress, Cupressus macrocarpa Hartwig (Fisher 1919).
Essig (1926) observed that the insect also infests dead branches of
the same host. Keen (1958) stated, “it feeds for the most part on the
blighted, dead, dry cones killed by other insects.” In 1965, I found this
insect in abundance in developing green cones of this cypress.
Since reports on the habits of this beetle are incomplete and somewhat
inconsistent, a study extending over several years was undertaken to
learn more about the relationships of Ernobius to developing, mature
and senescent cones of Monterey cypress in primarily native stands of
the host plant. Information was also collected on seasonal history, co¬
habiting insects, and feeding habits of Ernobius in those respective
cone age groups.
Distribution of host plant and beetle. —With the exception of
two uncertain host records 1 , E. conicola is known only from Monterey
cypress. The cypress is planted extensively along coastal California
and sparsely in the interior part of the state (Wagener 1939). It occurs
in the native state in two, small, coastal endemic stands at Point Lobos
and Cypress Point in Monterey County (Wolf and Wagener 1948).
Based on personal collections and museum records, it appears that
the beetle has a limited distribution within the range of the cypress.
In the two native groves, E. conicola is commonly found in cypress
cones. In all adventive localities, from San Luis Obispo County to
1 One collection was made in Monterey County, California from Myrica californica Cham. &
Schlecht, while the other specimen was taken from Pinas muricata D. Don in the same county.
However, it is not clear from the attached host labels whether the beetles were collected on the
plants or reared from them.
The Pan-Pacific Entomologist 49: 102-109. April 1973
Vol. 49, No. 2, April 1973
103
Contra Costa County, Ernobius has been collected mostly from coastal
situations where it occurs only occasionally in the cones.
Procedure. —Samples of cones, collected from Cypress Point and
Pescadero Point in Monterey County, were dissected and examined to
obtain information on the relationship of cone age to potential infes¬
tation by Ernobius. On three occasions, the samples, consisting of 200
cones from each of the available age groups, were gathered randomly
from eight trees which showed some evidence of a current infestation.
While dissecting cones, data were simultaneously gathered on seasonal
history development and the feeding behavior of the insect within each
of five recognized cone age classes.
Female strobili or cones of species in the genus Cupressus require
two years to mature. For the purposes of this investigation, the cones
were classified by age according to the following scheme:
Cone Age: 1 - Unpollinated, soft, succulent cones in first year of development.
Growth period generally begins in January and extends to De¬
cember of same year.
1.5 - Cones entering second year of development. After pollination
by wind (February-March), cones become less succulent and
noticeably harder. Period begins in January and ends in early
August of second year.
2 - Second year, mature cones which, through aging, have become
woody. Period extends from mid August of second year to about
March of third year.
3 - Third year cones 2 beginning to deteriorate from natural aging
processes and repeated insect infestations. About 85% of scales
are dry; cone cores remain resinous. Period extends from April
to December of third year.
>3 - Dry non-resinous cones which are older than three years of age.
The phenology of the developing cones (in particular, the 1 year
class) did not always proceed as described above. In some years (e.g.
1972) the appearance as well as growth of the new first year cones
was delayed as much as four months. Generally, such a paucity in
development was common to all the cypress trees in a given area.
Seasonal history development.— Larvae of E. conicola were found
throughout the year in the cypress cones (Fig. 1). Generally all instars
were well represented in each month with the exception of winter when
mid to late instar larvae predominated. Pupae were present from early
March to the middle of September (Fig. 2). The first adults flew in
mid March; adult emergence continued until the first part of October
2 Third year cones, having little to no deterioration, were not sampled due to the difficulty of
locating adequate numbers from the sample trees.
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The Pan-Pacific Entomologist
(Fig. 3). Keen (1958) mentioned that, “the principal adult emergence
occurs from late May through June and July to the first of August. 55
Based on findings from the cone dissections and adult emergence rec¬
ords, the egg-laying period is probably continuous from late March to
mid October.
Relationship of cone age to infestation. —Due to the incomplete¬
ness of the three samples, each of which was collected in a different
year, the results presented in Table 1 allowed for only limited insight
into the relationship of cone age to the level of infestation by Ernobius.
However, the nearly equal degree of infestation (about 10%) of 2
year cones observed on each sample date provided support for the
following generalizations.
Cones in the 2 and 3 year age groups sustained the highest levels
of infestation, while the 1 and >3 year groups experienced the lowest
number of infestations. The relative attractiveness of the 1.5 year
group was difficult to evaluate because of the transitional nature of
this age class.
The relatively low number of infested 1.5 year cones recorded in
January was regarded as the product of the previous year’s attraction
to the first year cones since no new infestations occur during January-
March of the second year. A cone examination from the Montara Mts.
(San Mateo Co.) added support to this observation. In January 1968,
600 cones from each of the first three age groups (i.e. 1, 1.5 and 2)
were examined for infestations of E. conicola. In the 1 year group
there were no infested cones. In the 1.5 and 2 year groups there were
five and 18 infestations respectively. After March, when the anobiid
begins to deposit the first eggs of the year, cones of the 1.5 year group
became quite attractive to ovipositing females as evidenced by the 34
infested cones recorded in April 1971 from this age group.
Feeding habits. — Ernobius conicola displayed specificity in its
selection of individual host trees. It was common to find infested trees
immediately adjacent to trees experiencing little or no evidence of
the insect. Infestation levels varied greatly among the trees sustain¬
ing attacks.
Part of this preferential selection may have been related to the spatial
arrangement of the cones (i.e. tightly vs. loosely packed in a cluster).
Although consistent for a given tree, this parameter tended to vary
within the species as a whole. In general, it seemed that Ernobius pre¬
ferred to infest trees whose cones were tightly arranged in clusters.
New and mature CONES. —In developing (1 and 1.5 year) and mature
Vol. 49, No. 2, April 1973
105
Figs. 1-3. Ernobius conicola. Fig. 1 . Larva feeding in mature cone of Cupressus
macrocarpa. Fig. 2. Pupa in mature cone of C. macrocarpa. Fig. 3. Adult.
106
The Pan-Pacific Entomologist
Table 1 . Number of Ernobius -infested cones from collections made
at Cypress and Pescadero Points in Monterey County, Calif.
Number of Infested Cones 1
Date
Cone Age 15 (yrs) :
1
1.5
2
3
>3
Jan. 1972
0
10
18
34
Apr. 1971
0
34
21
5
Aug. 1966
1
_C
16
a 200 cones dissected from each cone age class on each
b See text for description of cone age classes.
c Based on classification scheme, this age group is not
date.
represented
at this time.
(2 year) cones, first instar larvae usually entered via the basal portion
of the cone 3 . In the January 1972 sample (Table 1), 23 of 28 infested
cones were invaded through the base. Entrance into the five remaining
cones was gained through the side. After the new larvae had success¬
fully penetrated the epidermis, they began to extrude fine powdery
frass which accumulated around the small entry holes.
Upon establishment within the cone, the young larvae were generally
found feeding in the external portion of the scale tissue. If the cones
were relatively free of other invaders, then Ernobius fed locally in
one or occasionally two scales. Movement between scales seemed to
occur only when the substrate was in short supply due to a multiple
infestation by several other cone invaders.
Deteriorating cones. —As the mature cones aged or when they
became occupied by additional invaders, the feeding site of Ernobius
changed owing to a depletion of available food substrate. This occurred
most frequently during the 3 year stage and resulted in a shift of feed¬
ing activities to the fibrous cone core. It is of interest that the core
was also the site where resinous deposits persisted for the longest periods
of time, regardless of the infestation history of the cone.
In no instances were the cypress seeds fed upon by Ernobius.
In some of the cones, more than one developmental stage of the beetle
was found. Of 139 Ernobius-iniested cones examined from all age
groups (Table 1), 115 were infested with one stage, 19 with two stages,
four with three stages and one cone contained four stages. Most of the
cones containing more than one individual appeared to have been in¬
vaded independently as evidenced by the mixture of different larval
instars, pupae and/or adults that were extracted. In addition, the near¬
equal frequency of first instar larvae in newly-infested vs. previously-
3 Eggs are oviposited in secluded sites at the base of developing or mature cones and between dried
scales of senescent cones.
Vol. 49, No. 2, April 1973
107
infested cones suggests that gravid females oviposited without regard
to previous infestation history.
Cohabiting insects. —Ernobius conicola was often found associated
with two lepidopterous species, Laspeyresia cupressana (Kearfott)
(Tortricidae) and Henricus macrocarpana Walsingham (Phaloniidae)
in the cypress cones. Cohabitation of Ernobius with one or rarely both
moth species can be explained on the basis of cone age preferences,
feeding habits, and seasonal histories of Henricus and Laspeyresia.
In the case of H. macrocarpana, larvae were found infesting mostly
first year, unpollinated cones, wherein they fed almost exclusively on
the developing seeds (Frankie and Koehler 1967). New larvae entered
the soft 1 year cones during June and remained in the larval stage
until March or April of the following year. At that time the cones were
1.5 years of age. From late April through May of the second year,
adults of Henricus emerged from the cones. Simultaneously, the first
instar anobiid larvae began to invade 1.5 as well as the 2 and 3 year
cone groups. Except for this brief two month overlap period, the two
species remained relatively isolated from each other due to their re¬
spective preferences for cones of different ages. Since Henricus females
as well as the anobiid females are believed to oviposit randomly on the
cones, regardless of whether the cones are predisposed or intact, op¬
portunities for cohabitation are further diminished. An examination
of cones of the 1 and 1.5 year groups revealed that only three of 45
infested with Ernobius were concurrently invaded by Henricus.
Larvae of L. cupressana were found feeding almost exclusively in
the scale tissue of 1, 1.5 and 2 year old cones (Frankie and Koehler
1971). The larvae tunnel randomly through the scales, resulting in a
substantial amount of feeding in the interior of the cones. In contrast,
Ernobius feeds locally in the exterior of cones of these age groups. Of
85 Erno bius- infested cones examined in the above three age classes,
15 were found to contain stages of both the beetle and the moth. Periods
of coexistence between Ernobius and Laspeyresia are probably short¬
lived if cones become over-crowded with phytophagous individuals.
Under conditions of food shortage (or perhaps through water stress
in the cones brought about by multiple invasions) Laspeyresia is thought
to move to adjacent cones in search of more favorable substrate (Frankie
and Koehler 1971). Therefore, due to differences in preferred feeding
sites and the response of L. cupressana to crowded conditions, the co¬
habitation seemed to pose no problem to either of the two species.
Old infestations of either lepidopterous species and Ernobius were
easily distinguished primarily on differences in frass particle size. The
108
The Pan-Pacific Entomologist
larvae of Laspeyresia and Henricus both produce their frass in relatively
large pellets, while Ernobius excretes finely shredded frass. An addi¬
tional difference is that the larvae of both moths tend to pack their
frass in certain sections or chambers of their borings whereas the anobiid
frass merely accumulates loosely within the hollowed out cone scales.
Discussion
Early reports on E. conicola provide little insight into the relation¬
ship of this insect to its host, Monterey cypress. Field observations
and data from cone dissections reported in this paper reveal that Ernob¬
ius interacts with its host plant in several specific ways.
Numerous collections indicate that the beetle is limited primarily
to Monterey cypress in a limited portion of its range along coastal
California. This apparent restricted distribution is similar to the
pattern displayed by the scolytid beetle, Conophthorus radiatae Hop¬
kins, in its coastal confinement to central California (Schaefer 1963).
Further, Ernobius is limited to the cones of certain age groups of par¬
ticular cypress tress.
The apparent tendency of the insect to invade via the base of develop¬
ing and mature cones also suggests a very specific relationship. Using
electroantennogram recordings, a similar interaction was reported by
Asher (1970) in reference to the relatively high olfactory response of
Dioryctria abietella (Denis and Schiffermuller) to volatile compounds
escaping from the basal % of Pinus elliotii Engelmann var. elliottii cones.
However, in the case of Ernobius , penetration at the base may indicate
an adaptation to seek shelter to avoid harsh coastal climatic conditions
which often prevail in the native stands. Such a behavioral pattern
may also provide some explanation for the tendency of Ernobius to
invade trees having cones tightly packed in clusters. Closely aligned
cones would aid in protecting the newly hatched larvae.
This study may hopefully raise new questions regarding the biology
and ecology of E. conicola as well as other anobiid species, most of
which are poorly known.
Acknowledgments
I would like to thank A. E. Darling for assistance in dissecting nu¬
merous cypress cones. I am also indebted to H. R. Burke, J. A. Chemsak,
R. R. Fleet, D. R. Hamel and C. S. Koehler for reviewing and criticizing
the manuscript.
I gratefully acknowledge the cooperation of the following institutions
Vol. 49, No. 2, April 1973
109
for supplying me with collection records: California Academy of
Sciences, San Francisco; Department of Entomology, University of
California, Berkeley; Los Angeles County Museum, Los Angeles; and
Smithsonian Institution, Washington, D. C.
Literature Cited
Arnett, R. H. 1968. The beetles of the United States. Amer. Entomol. Inst.,
Ann Arbor, Mich., 1112 p.
Asher, W. C. 1970. Olfactory response of Dioryctria abietella (Lepidoptera:
Phycitidae) to slash pine cones. Ann. Entomol. Soc. Amer., 63: 474-476.
Baker, W. L. 1972. Eastern forest insects. U. S. Dep. Agr. Misc. Publ. No.
1175, 642 p.
Borror, D. J. and D. M. DeLonc. 1971. An introduction to the study of insects.
3rd Ed. Holt, Rinehart and Winston, N. Y., 812 p.
Essig, E. 0. 1926. Insects of western North America. MacMillan Co., N. Y.,
1035 p.
Fisher, W. S. 1919. Five new species of ptinid beetles. Proc. U. S. Nat. Mus.,
55: 295-299.
Frankie, G. W. and C. S. Koehler. 1967. Cypress bark moth on Monterey
cypress. Calif. Agr., 21: 6-7.
1971. Studies on the biology and seasonal history of the cypress bark moth,
Laspeyresia cupressana (Lepidoptera:Olethreutidae). Can. Entomol.,
103: 947-961.
Hatch, M. H. 1961. The beetles of the Pacific Northwest. Pt. 3. Univ. Wash.
Publ. Biol., 16: 309-327.
Keen, F. P. 1958. Cone and seed insects of western forest trees. U. S. Dep.
Agr. Tech. Bull. No. 1169. 168 p.
Ruckes, H., Jr. 1957. A synopsis of the California deathwatch beetles of the
genus Ernobius Thomson, with descriptions of two new species which
attack pine cones (Coleoptera:Anobiidae). Pan-Pac. Entomol., 33:
157-161.
Schaefer, C. H. 1963. Factors affecting the distribution of the Monterey pine
cone beetle, Conophthorus radiatae Hopk., in central California. Hil-
gardia, 34: 79-103.
Wagener, W. W. 1939. The canker of Cupressus induced by Coryneum cardinale
n. sp. J. Agr. Res., 58: 1-46.
White, R. E. 1962. The occurrence of Mesocoelopus collaris Mulsant and Rey
in Columbus, Ohio, with biological notes (Coleoptera:Anobiidae).
Coleopt. Bull., 16: 13-15.
Wolf, C. B., and W. W. Wagener. 1948. The new world cypresses. El Aliso,
1: 1-444.
110
The Pan-Pacific Entomologist
Biology of the Spider Diguetia imperiosa 1
(Araneida: Diguetidae)
Michael M. Bentzien 2
Department of Ecology and Systematics, California State University, San Francisco
Spiders of the family Diguetidae have a unique type of web and co¬
coon construction. They spin a sheet web surrounded by a tangled mesh
of silk threads, and build a retreat which serves as protection for the
spider, and in the case of females, as an enclosure for the egg sacs.
Diguetids are considered to be relatively primitive spiders and the gen¬
italia of both sexes are comparatively simple. The family was established
by Gertsch (1949) and also revised by Gertsch (1958). The nine spe¬
cies of the family all belong to the genus Diguetia and are distributed
from the southwestern United States through Mexico, and also occur in
western Argentina. Although the diguetids are often locally abundant,
few studies of their biology have been made. Cazier and Mortenson
(1962) studied the biology of Diguetia canities near Portal, Arizona;
Gerschman de Pikelin and Schiapelli (1962) described the web structure
of D. catamarquensis in Argentina, and Eberhard (1967) studied attack
and prey-wrapping behavior in D. albolineata.
In August 1968 I had the opportunity to make field studies of D.
imperiosa Gertsch and Mulaik (Fig. 1), a species distributed widely
through the southwestern United States and northern Mexico. Field
work done in the Cuatro Cienegas Basin of Coahuila, Mexico. Spiders
and retreats were brought to California for observations on behavior
and reproductive biology.
Methods and Materials
To study habitat, web structure, and behavior of Diguetia imperiosa
in the field, twenty webs were located, marked, and observed from 9 to
14 August 1968.
To determine fecundity and details of retreat construction, 45 retreats
and the spiders accompanying them were collected on 14 August and
were fixed and preserved using the methods recommended by Williams
(1968). Egg counts and dissection of retreats were done using a bin¬
ocular dissecting microscope, and measurements were taken with an
1 This work is based on research for the degree of Master of Arts completed at California State
University, San Francisco in 1970.
2 Present address: Division of Entomology, University of California, Berkeley, California 94720.
The Pan-Pacific Entomologist 49: 110-123. April 1973
Vol. 49, No. 2, April 1973
111
ocular micrometer. Maturity of female spiders was determined by ex¬
amination of the ovaries.
Ninety retreats with the accompanying spiders were collected on
15 August 1968 and were reared in the laboratory until 31 December
1969. Ten female spiders were allowed to construct webs in small bushes
simulating vegetation in the natural habitat; the remainder were con¬
fined in small plastic boxes. Laboratory temperatures varied from 20°
to 30° C and an approximate 12 hour photoperiod was maintained.
Food ( Tenebrio molitor larvae and adult Drosophila sp.) and water
were offered at weekly intervals.
Stridulation was recorded with a Sennheiser microphone and a Nagra
IIIB tape recorder (15 inches per second). A sound spectrogram was
prepared using a model 661-B Sona-Graph manufactured by the Kay
Electric Company.
Results
Study area.— Field work was carried out seven kilometers southwest
of the town of Cuatro Cienegas de Carranza, Coahuila, Mexico. Cuatro
Cienegas lies in an intermontane basin on the east edge of the Mesa del
Norte. The basin measures about 30 by 40 km. An arid Chihuahuan
climate prevails, but large numbers of springs give rise to complex
systems of internal drainage within the basin. Minckley (1969) has
summarized biological research in the Cuatro Cienegas area.
The study area consisted of low dunes and flats. The dunes were
stabilized and of high gypsum content. The flats had powdery soil and
were only slightly elevated above the water table; it appeared that flood¬
ing was of seasonal occurrence. Zonation of the soil was lacking, and
vegetation was sparse. Flats were primarily vegetated by chenopods
(Allenrolfea occidentals (Wats.) Kuntze, Suaeda spp.) and unidentified
sedges. Dunes supported yucca ( Yucca sp.), mesquite ( Prosopis gland-
ulosa Torr.), ocotillo ( Fouquiera sp.), and two species of Opuntia.
Web and retreat structure.— Diguetia imperiosa constructed its
web in spaces between the branches of low plants, or suspended it be¬
tween adjacent plants. All webs observed were less than one meter above
the ground. Orientation of the web appeared to be non-random; 75 per
cent of the webs observed were suspended on the west facing side of
the supporting plants.
The web invariably was composed of four structural parts (Fig. 2).
Hereafter, the term “web” will refer to the entire web, and the term
“webbing” will refer to the various parts of the web. The parts of the
112
The Pan-Pacific Entomologist
Fig. 1. Adult female Diguetia imperiosa Gertsch and Mulailc from Cuatro
Cienegas de Carranza, Coahuila, Mexico. Photo by Carolyn Cavalier. Fig. 2. Web
of female Diguetia imperiosa near Cuatro Cienegas de Carranza, Coahuila, Mexico.
Guy lines (g), tangle webbing, and sheet webbing (s) are visible. Orb web of
Metepeira sp. is located in upper right portion of D. imperiosa web.
Vol. 49, No. 2, April 1973
113
Table 1 . Accessory materials used by Diguetia imperiosa in retreats
construction. Analysis based on dissection of 30 retreats collected 15
August 1968 near Cuatro Cienegas de Carranza, Coahuila, Mexico.
Material
No. of
retreats
% occurrence
Plant Material
Leguminosae: Prosopis glandulosa Torr.
24
80
Cactaceae: Opuntia spp.
3
10
Cyperaceae: unidentified sedge
4
13
Chenopodiaceae: Suaeda spp. and
Allenrolfea occidentalis (Wats.) Kuntze
7
23
Arthropod Material
Arachnida: Diguetia imperiosa exuvia
8
26
Salticidae
1
3
Araneidae
1
3
Insecta: Coleoptera: Oedemeridae: Oxacis sp.
14
47
Scarabeidae
9
30
Bostrichidae, Coccinellidae,
Curculionidae, Tenebrionidae
4
13
Hymenoptera: Myrmicidae: Myrmicinae
14
47
Orthoptera: Acrididae
4
13
Lepidoptera: unidentified moths
4
13
Diptera: Muscidae
1
3
Hemiptera: Coreidae, Corixidae,
Pentatomidae
3
10
Homoptera: Cicadellidae
4
13
Cercopidae, Psyllidae, Cicadidae
3
10
web were the guy lines, the tangle webbing, the sheet webbing, and the
retreat. The guy lines extended to the apices of the branches of the sup¬
porting plant and provided suspension for the web. They consisted of
multiple strands of heavy silk. The tangle webbing consisted of an ir¬
regular mesh of silk which filled the area above and below the sheet
webbing. The sheet webbing consisted of a fine silk mesh radiating
horizontally from the centrally positioned cocoon; it was oval to roughly
circular in shape and was no greater than 0.3 m 2 in area. Neither
tangle nor sheet webbing consisted of viscid silk. Each spider had one
retreat associated with its web. The retreat was of a characteristic con¬
ical shape, and was open at the lower end. It was suspended by strong
strands of silk which were connected to the guy lines and to plant
branches, and always hung vertically in the web. The internal diameter
of the retreat was sufficiently large to allow the spider to completely
114
The Pan-Pacific Entomologist
immature
i—i
5 mm
Fig. 3. Retreats of female, male, and immature Diguetia imperiosa. Fig. 4.
Retreat of female Diguetia imperiosa. Opened retreat shows egg sacs with silk
cover removed, a = accessory materials, s = silk mesh, e := egg case.
Vol. 49, No. 2, April 1973
115
Fig. 5. Sound spectrogram of stridulation of adult female Diguetia imperiosa,
recorded in laboratory 15 December 1968.
enter; access was from the underside of the sheet webbing. The re¬
treat was constructed of accessory materials, such as plant leaves, insect
bodies, and other objects, tightly bound together with a fine silk mesh
(Fig. 3). Mature females laid a series of egg sacs inside the retreat in
an overlapping, shingle-like arrangement (Fig. 4). The first egg sac
laid was placed near the upper, closed end of the retreat. The egg sacs
were disc-shaped and about 8 mm in diameter. They were formed by
two densely woven discs of silk weakly joined at their peripheries. Each
egg sac was covered with a coarse silk mesh with the strands spaced
widely enough to allow free passage of emerging spiderlings.
Retreats of mature females were larger and of more tightly woven
construction than the retreats of males. Retreats of immature spiders
were constructed of materials of smaller, more uniform size than the
retreats of mature spiders (Fig. 3).
To determine what accessory materials were used in retreat construc¬
tion, 30 retreats were dissected. Leaves and twigs of mesquite (Prosopis
glandulosa Torr.) and Chenopodiaceae were the commonest plant ma¬
terials, and oedemerid beetles (Oxacis sp.) and ants were the commonest
insects (Table 1).
Behavior of spiders in the web.— Diguetia imperiosa was primarily
nocturnal, remaining inside the retreat during the day. However, spiders
left the retreat at any hour, day or night, to capture prey. They also
left the retreat during the period of midday high temperature, which
116
The Pan-Pacific Entomologist
reached as much as 47° C during the field study. At this time the spiders
positioned themselves in the web so that they were in the shadow cast
by the retreat. Later in the afternoon the spiders always returned to
their retreat. It was not determined if spiders would leave the retreat
at midday on cool or overcast days. At dusk, spiders again left the
retreat and remained exposed on the underside of the sheet webbing
during the hours of darkness. To study web-building techniques of
D. imperiosa in the field, the webs of two adult females were removed
on 12 August. The spiders and their retreats were left in the supporting
plants. The spiders replaced the guy lines supporting the retreat first,
then erected guy lines to support the sheet and tangle webbing. Spinning
of sheet and tangle webbing followed; but three days later, when field
observations ceased, only sparse webs had been constructed. The in¬
complete nature of the sheet and tangle webbing suggested that prey
capture would have been severely reduced.
In the laboratory, spiders began to construct webs in the same man¬
ner as individuals in the field, but after a month they still had not
constructed complete webs.
Diguetia imperiosa reacted to web disturbance in three different
ways. Slight movements in any part of the web induced attack behavior.
Vol. 49, No. 2, April 1973
117
Disturbances of a greater magnitude, involving significant movement
of the entire web, caused spiders to withdraw into the retreat. This
response occurred when capture of spiders was attempted in the field.
Sudden severe disturbance to the whole web, or destruction of the web,
caused spiders to drop from the web and remain motionless on the
ground in a death feigning posture, in which the legs were drawn close
to the body, and no movement occurred for as long as 45 minutes.
Spiders later returned to the web area by climbing the nearest plant.
No drag lines were established in these abrupt falls from the web.
Two webs in the study area each contained a male-female pair of
spiders. In both of these webs the female had already deposited eggs,
and no mating activity was observed. When prey entered the webs of
these paired spiders, females attacked and handled the prey in the usual
manner, but males did not respond. When disturbed, in each case the
females withdrew into the retreat in the same way as single females,
but the males ran about the web. In one instance a male did attempt to
enter the female’s retreat after she had withdrawn, but he emerged almost
immediately.
Prey became trapped either in the tangle webbing or on the sheet
webbing. When prey entered the web, the spider ran out on the under¬
side of the sheet webbing toward the source of disturbance. In a few
instances, spiders shook the sheet webbing when approaching prey. If
prey remained trapped in the tangle webbing, spiders had to climb into
the tangle webbing to attack. In this situation, a spider would run to
the edge of the sheet webbing to ascend the tangle webbing; no use of
holes in the sheet webbing to pass from one side to the other was ob¬
served. In most of the prey captures observed in the field, the spiders
encountered and attacked prey on the sheet webbing. Spiders attacked
from the underside of the sheet webbing in these cases. The first pair
of legs were extended through the sheet webbing to hold the prey down
while it was bitten through the sheet webbing; the hold on the prey
was then released. When the prey ceased struggling, the spider cut a
hole in the sheet webbing with the chelicerae, and pulled the prey
through to the underside of the sheet webbing.
When prey capture occurred at night, feeding took place in an exposed
position out under the sheet webbing, but prey captured during the day
was carried to a location just beneath the retreat to be eaten.
When spiders captured large winged insects, a simple form of prey
wrapping occurred. The prey was suspended from a silk line and rotated
while a few turns of silk were used to bind the wings. Small prey was
118
The Pan-Pacific Entomologist
No. egg sacs
Fig. 7. Egg production of 30 female Diguetia imperiosa. Small dots represent
totals of individual retreats; large dots represent mean totals.
captured and eaten without the use of silk. Prey wrapping was also
observed when an insect became trapped in the web while the spider
was feeding on previously captured prey. The first prey was suspended
and wrapped; then the spider attacked the new prey.
When a nonprey item such as a piece of plant material fell onto the
sheet webbing, spiders discarded the item after a few seconds of ex¬
ploratory contact using the pedipalps and first pair of legs; a hole was
Vol. 49, No. 2, April 1973
119
cut in the sheet webbing with the chelicerae and the object was permitted
to drop to the ground.
Attack behavior of captive D. imperiosa did not differ from that ob¬
served in the field.
Stridulation. —Stridulation was observed in several adult female
D. imperiosa in the laboratory when the spiders were physically dis¬
turbed, as when grasped by forceps. The stridulatory mechanism in the
genus Diguetia consists of fine grooves on the outer face of the paturon
of the chelicerae, which serve as a file, and a row of small tubercles on
the prolateral side of the femur of the pedipalp, which serve as a scraper.
During stridulation, the chelicerae are spread and the pedipalps are
simultaneously rapidly raised and lowered. In D. imperiosa , the re¬
sultant sound occurred in bursts lasting for 75 milliseconds, separated
by 100 millisecond intervals. The frequency averaged about 10,000
Hertz (Fig. 5). Males had fewer and smaller femoral tubercles than
females, but no live males were available for recording stridulation.
Associates. —Two species of spiders were directly associated with
Diguetia imperiosa webs in the field. Immature spiders of an unde¬
termined species of the araneid genus Metepeira were found as a com¬
mensal in 25 per cent of the D. imperiosa webs examined in the field.
Metepeira constructed a small orb web in the upper tangle webbing
of the host spider. These orb webs were less than five centimeters in
diameter and were vertically oriented. No mature Metepeira were found
associated with D. imperiosa.
Two webs examined in the field each contained an individual of an
undetermined species of the genus Mimetus, spiders predaceous on web
building spiders. In both cases the D. imperiosa which had constructed
the web was missing.
No parasitization or predation of egg sacs was found.
Reproductive biology. —Descriptive terms of development are those
used by Hite et al. (1966). Eggs of D. imperiosa were white, spherical,
and varied from 0.45 to 0.55 mm in diameter. When eggs hatched a
first stage postembryo emerged. This first stage postembryo was covered
with a thin membrane through which the developing spider could be
seen. The first stage postembryo was yellowish white except for black
eyes, and ranged from 0.53 to 0.62 mm in length. The second stage
postembryo was free moving and possessed tarsal claws, but lacked
development of cheliceral fangs and body pigment. This stage ranged
from 0.80 to 0.90 mm in length. The next molt gave rise to spiderlings
with fully developed claws, chelicerae, and body pigmentation resem¬
bling that of the adult. The range in body size was the same as the
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The Pan-Pacific Entomologist
Table 2. Egg production by Diguetia imperiosa. Based on egg sacs
contained in seven retreats, each with five egg sacs, collected 14 August
1968. Number of eggs and young in each sac is represented.
Egg sac
(oldest to
newest)
Individual
#102
#103 #104
#130
#134
#135
#137
(first egg
sac laid)
70
236 181
262
321
415
190
200
217 261
206
269
367
148
178
162 253
208
222
268
110
167
152 200
125
244
241
151
(last egg
sac laid)
206
128 246
140
289
75
107
mean
164
179 228
188
269
273
141
F-Test Analysis
Source of
Sum
Mean
variation
DF
Squares
Square
F
P
total
34
197622
retreat
6
79848
13308
3.56
P < 0.01
position of
egg sacs
4
28040
7010
1.87
P < 0.05
error
24
89734
3738
second stage postembryo. Development to the spiderling stage occurred
within the egg sac. The first and second stage embryos were inactive,
developmental stages, while the spiderlings were capable of feeding and
silk production. Spiderlings escaped from the cocoon by splitting the
egg sac along the loosely joined peripheral seam. The emergence of
spiderlings from the retreats took place in the laboratory from 22
August to 23 September 1968. At this time emergence had occurred
in only 43 per cent of the 56 retreats containing eggs. No emergence
occurred after 23 September. Nonemerged spiderlings remained alive
inside the egg sacs until December 1968. After this time all had died.
Twenty-six per cent of the living females continued laying eggs until
23 September, 1968, whereas complete emergence from all egg sacs had
already occurred in four per cent of retreats containing eggs collected
on 14 and 15 August. Eggs laid in the laboratory failed to develop
beyond the first stage postembryo.
Vol. 49, No. 2, April 1973
121
Maturity of female spiders collected was determined either by the
presence of eggs in the accompanying retreat or by examining the ova¬
ries. In a sample of 30 mature females, the number of eggs laid per
female ranged from none to 1,675, and the mean was 734. The egg
sacs of 89 female retreats were counted. The number of egg sacs con¬
structed per female ranged from zero to ten, with a mean of 3.6. Only
20 per cent of the females laid more than five egg sacs (Fig. 6).
Considerable variability existed in the numbers of eggs laid in sub¬
sequent egg sacs by the same spider. Some females laid progressively
fewer eggs in later egg sacs, but others laid increasing numbers. Spiders
which laid the same number of egg sacs during the reproductive season
showed much variability in the egg number of corresponding egg sacs
in the sequence, as well as in total number of eggs laid. This was shown
by seven spiders which laid five egg sacs each (Table 2). Differences in
the total number of eggs laid was highly significant (F = 3.56; P <
0.01). Based on the relative position within retreats, number of eggs
laid in the corresponding egg sacs by the seven spiders, there was no
pattern of increase or decrease in egg laying by these spiders (F = 1.87;
P > 0.5). Similar results were found in a group of three spiders which
laid three egg sacs each; there was a significant difference in total egg
number laid by individual spiders (F = 10.7; 0.05 > P > 0.01), and
the group showed no pattern of differential egg laying in relation to
the sequence of egg sacs (F = 0.414; P > 0.05).
No correlation was found between size of female spiders (carapace
length) and total egg production, nor was there any indication of de¬
creasing fertility in the later egg sacs.
Egg counts carried out on a sample of 30 retreats containing eggs
showed that spiders laying over five egg sacs did not greatly increase
their egg production over those which had laid only five egg sacs (Fig.
7). But egg production decreased sharply in spiders laying fewer than
five egg sacs.
Of the 135 retreats collected, 89 had been built by females, 37 by
males, and five by immature spiders. Of the 82 Diguetia imperiosa
found with these retreats, 73 were mature females, four were mature
males, and five were immature.
Discussion
Since nearly all the Diguetia imperiosa collected were mature, the
spider probably has an annual life cycle, with one brood per year. A
few females captured at maturity lived up to two additional years in the
laboratory, but they did not molt, and became very inactive.
122
The Pan-Pacific Entomologist
Stridulatory mechanisms of several types are widespread in spiders.
In certain of the Theraphosamorphae stridulation occurs when the spider
assumes a defensive posture, and the sound is readily audible to humans
(Savory, 1928). Many araneomorph spiders have stridulatory organs,
but the nature of the sound produced is generally unknown. The strid¬
ulation of Diguetia imperiosa is faintly audible to some human observers.
As the stridulatory organ is found in both sexes, and since stridulation
can be elicited by disturbing the females, a defensive function is
indicated.
The lack of egg parasitization was surprising, since a fairly large
sample of retreats and egg cases was examined for parasites. Cazier
and Mortenson (1962) found both hymenopteran and coleopteran egg
parasites of Diguetia canities, and five out of 75 retreats of Diguetia
sp. from Yavapai County, Arizona sent to me by Dr. Cazier contained
hymenopteran parasites. Suitable parasites may be absent in the Cuatro
Cienegas Basin, or parasites may have been operating at a low level
when this study was done.
Acknowledgments
My thanks to Dr. Mont A. Cazier, who suggested this study and pro¬
vided help and advice in the field. Dr. Wendell L. Minckley furnished
helpful information concerning the Cuatro Cienegas Basin, and Dr.
Donald J. Pinkava made the plant determinations.
I am grateful to Dr. Stanley C. Williams, Dr. Robert I. Bowman, and
Dr. Bernard Goldstein for their suggestions and criticism regarding
the manuscript. My appreciation to Dr. Williams for the time he gen¬
erously spent discussing this study and related topics, and to Dr. Bowman
for providing facilities to make the sound spectrogram.
Thanks are also due Sandra Ortega for clerical help, and Carolyn
Cavalier for help with the photography.
Literature Cited
Cazier, M. A. and M. A. Mortenson. 1962. Analysis of the habitat, web design,
cocoon and egg sacs of the tube weaving spider Diguetia canities
(McCook). (Aranea, Diguetidae). Bull. S. Calif. Acad. Sci., 61: 65-88.
Eberhard, W. 1967. Attack behavior of diguetid spiders and the origin of prey
wrapping in spiders. Psyche, 74: 173-181.
Gerschman de Pikelin, B. S. and R. D. E. Schiapelli. 1962. La familia
Diguetidae en la Argentina. Physis, 23: 205-208.
Gertsch, W. J. 1949. American spiders. D. van Nostrand Co., N. Y., 285 p.
1958. The spider family Diguetidae. Amer. Mus. Novitates, 1904: 1-24.
Hite, J. M., W. J. Gladney, J. L. Lancaster, Jr., and W. H. Whitcomb. 1966.
Vol. 49, No. 2, April 1973
123
Biology of the brown recluse spider. Arkansas Agr. Exp. Sta. Bull.
711, 26 p.
Minckley, W. L. 1969. Environments of the Bolson of Cuatro Cienegas, Coahuila,
Mexico. Texas West. Press Sci. Ser. 2, 65 p.
Savory, T. H. 1928. The biology of spiders. MacMillan Co., N. Y., 376 p.
Williams, S. C. 1968. Scorpion preservation for taxonomic and morphological
studies. Wasmann J. Biol., 26: 133-136.
BOOK REVIEW
Ecological Studies 2. Integrated Experimental Ecology. Methods and Re¬
sults of Ecosystem Research in the German Solling Project. Edited by
Heinz Ellenberg. Springer-Verlag, New York, Inc. 214 p., 53 figs., 44 tables.
1971. $16.80.
One of the first International Biological Program (IBP) ecosystem studies was
initiated in 1967 by the West German IBP Working Group for the Study of Ter¬
restrial Biological Communities. A group of 50 scientists under the direction of
Professor Ellenberg undertook an interdisciplinary study in the High Solling
in the center of the Federal Republic of Germany. The area contains both forest
and grassland ecosystems. This volume presents preliminary reports from many
of the cooperative studies with emphasis on the methods used. A final report is
expected after the conclusion of the project in 1972.
This volume contains 28 reports produced by 33 scientists. The introductory
chapter by H. Ellenberg sets the background for the project by describing its
relationship to IBP, the structure and functioning of ecosystems, the organization
of the integrated research, and the research area and experimental plots. The
remaining reports are subdivided into four major parts: Primary Production;
Secondary Production; Environmental Conditions; and Range of Validity of the
Results.
Insects are dealt with in four reports in Part 2: Secondary Production. These
all involve food and energy turnover of various insects. Sampling techniques for
studies of five species of leaf feeding weevils are described by W. Funke, with
emphasis on the use of photo-eclectors (emergence traps). The report by K. Winter
presents the approaches used to study some of 40 species of Lepidoptera, most of
which are phytophagous on beech trees. Methods used to study energy flow in
phytophagous and predatory arthropods are the subject of the report by W. Funke
and G. Weidemann. The latter author also reports on the methods used to study
population dynamics, standing crop, and production in predatory arthropods of
the soil surface.
The basic thrust of the project is to develop a clear understanding of the bio¬
logical basis of productivity using an integrated approach involving scientists
from many disciplines. This volume presents only preliminary results and leaves
one anxiously awaiting the production, assimilation and integration of the final
reports. The main contribution of the present volume is the description of method¬
ology which provides a valuable background for all ecologists.— Robbin W. Thorp,
University of California, Davis, 95616.
124
The Pan-Pacific Entomologist
A New Species of Spinolochus from
North America
(Hymenoptera: Ichneumonidae)
Torolf R. Torgersen
U.S. Forest Service, Pacific Northwest Forest and Range Experiment Station,
Forestry Sciences Laboratory, Juneau, Alaska
Horstmann (1971) proposed the genus Spinolochus in his revision
of the European Tersilochinae and included in it a single species, S.
laevifrons (Holmgren) from northern Europe. A second species has
appeared in collections from Alaska, indicating that the genus is Holarc-
tic in distribution and represented by more than one species. The Alas¬
kan species of Spinolochus is described below.
Two females of the new species were collected by L. C. Beckwith near
Dry Gulch on the Kenai Peninsula southeast of Anchorage, Alaska. The
specimens were obtained from windowpane flight traps operated from
1 June to 25 September 1969.
Key to Females of Spinolochus
1. Ovipositor in dorsal view parallel-sided for most of length, then narrowing
abruptly to a subparallel-sided tip, in lateral view smoothly curved and taper¬
ing evenly to acute apex; median half of apical margin of clypeus weakly
convex; first abdominal tergite in lateral view with slight depression above
glymma . laevifrons (Holmgren)
Ovipositor in dorsal view parallel-sided to beyond midlength, gradually
flaring, then converging smoothly to rounded apex (Fig. la), in lateral view
with a raised preapical nodus (Fig. lb) ; median half of apical margin of
clypeus nearly truncate; first abdominal tergite without depression above
glymma, smoothly convex or nearly straight above_ distolatus Torgersen, n. sp.
Spinolochus distolatus Torgersen, new species $
(Fig. la, b)
Male. —Unknown.
Female.— Front wing 3.5-4.0 mm long, hyaline. Head broad, as wide across
eyes as thorax just below tegulae; width across eyes 1.12-1.14X width at temples;
profile of face from dorsal aspect elevated medially near antennal fossae. Antenna
with 26-29 segments. Clypeus sparsely punctate basally and apically, the median
half of its apical margin nearly truncate. Mesoscutum minutely corearious, sparsely
punctate; notaulus short and indistinct, anteriorly with rugulose area. Prepectal
carina extending dorsally, almost touching mesopleural margin before it curves
briefly to nearly parallel it; wrinkling posterior to prepectus near its dorsal ex¬
tension arranged in several clearly discernible ridges. Area below anterior end of
The Pan-Pacific Entomologist 49: 124-126. April 1973
Vol. 49, No. 2, April 1973
125
Fig. 1. Spinolochus distolatus n. sp.; a, dorsal aspect of ovipositor; b, lateral
aspect of ovipositor.
sternaulus rugulose, merging ventrad into coarse punctations then into smooth
but weakly mat sternal area. Anterodorsal corner of mesopleurum with rugae
clearly arranged in several rows; one to several rugae present in area of meso-
pleural fovea. Propodeum with areola impressed, not sharply defined; longitu¬
dinal carinae represented anterior to petiolar area by irregular rugae. Apical
transverse carina of propodeum well developed. First abdominal tergite in pro¬
file smoothly convex or nearly straight above glymma. Abdominal tergites 6 and 7
each with deep membraneous excision medially.
Black. Mandible yellow-brown, its teeth and base fuscous. Scape, pedicel,
palps, tegula, legs, hind coxa, abdominal segments 2-9, and ovipositor fusco-
testaceous. Basal 0.4-0.7 of hind coxa infuscate, and hind tarsus a little paler
than the rest of legs. Basal margins and/or median areas of abdominal segments
weakly infuscate or darker red.
In addition to the key characters given for separating S. laevifrons and
S. distolatus , the following may also be helpful in differentiating the
two. While the clypeus is nearly uniformly fuscous or black in S. disto¬
latus , 0.5 to 0.8 of the apical portion of the clypeus in S. laevifrons may
be yellow-brown to fusco-testaceous. The abdomen of S. laevifrons some¬
times is quite dark compared with S. distolatus. Also, setation on the
ovipositor sheath of S. laevifrons is rather sparse as compared with
the better developed setation on S. distolatus.
Holotype female, Dry Gulch [Kenai Penn.], Alaska, 25 August
1969, white spruce type, window flight trap, [L. C.] Beckwith, (U.S.
National Museum). Paratype female, Dry Gulch, Alaska, 28 August
1969, white spruce type, window flight trap, Beckwith (Townes Coll.,
Ann Arbor, Mich.).
Acknowledgments
The author expresses his thanks to Dr. Henry K. Townes of the Amer¬
ican Entomological Institute, Ann Arbor, Michigan, for his assistance
in preparing the above description.
126
The Pan-Pacific Entomologist
Literature Cited
Horstmann, K. 1971. Revision der europaischen Tersilochinen (Hymenoptera,
Ichneumonidae) Teil I. Veroff. Zool. Staatssaml. Miinchen, 15: 77-79.
SCIENTIFIC NOTE
Wing dimorphism in Carabus maeander (Coleoptera :Carabidae) .—The
hind wings of this species are usually strongly reduced but sometimes are large
and functional (Lapouge, 1929. Wytsman’s Genera Ins., 192: 1-747). However,
Lindroth (1961. Opusc. Entomol. Suppl., 20: 1-200) did not record long winged
forms for this species in Canada and Alaska.
I examined 49 specimens of this species taken in Quebec and I found four long
winged specimens from the following places: Argenteuil Co., lie Carillon, 16
August 1969 (1 male). Beauhamois Co., Baie-de-la-Faim, 27 August 1966 (1 male).
Vaudreuil Co., Rigaud, 15 May 1966 and 1 May 1972 (2 males). These long winged
forms are geographically restricted to the Montreal region.
Short winged specimens were taken from the following places: Abitibi Co.,
Amos, 10 June 1962, 13 May and 12 June 1968 (4) ; lie Nepawa, 18 June 1968 (1) ;
Lac D’Alembert, 27 May 1968 (4) ; Lac Figuery, 11 May 1968 (1) ; La Ferme,
19 May 1968 (1) ; Mancebourg, 13 June 1968 (4). Charlevoix-Est Co., Baie-Sainte-
Catherine, 27 June 1969 (1) ; Port-au-Persil, 22 June 1969 (1) ; Port-au-Saumon,
5 July 1964 and 18 July 1965 (2) ; Sainte-Mathilde, 13 June 1965 (1). Iles-de-
la-Madeleine Co., Grosse-Ile, 7 August 1970 (5) ; Ile-au-Loup, 5 August 1970 (1) ;
He du Cap-aux-Meules, 31 July 1970 (1) ; He du Havre-Aubert, 2 August 1970 (4).
Montmorency No. 1 Co., Saint-Joachim, 20 April 1968 and 5 June 1969 (2). Sa¬
guenay Co., Blanc-Sablon, 4 July 1971 (2) ; Brador, 7 June 1971 (1) ; Grandes-
Bergeronnes, 31 July 1964 (1) ; Harrington, 1 July 1971 (4). Temiscamingue Co.,
Notre-Dame-des Quinze, 21 May 1968 (1) ; Rollet, 12 and 24 May 1968 (3). Vaud¬
reuil Co., Rigaud, 15 May 1971 (1).
The geographical distribution of the two forms presents a distinct pattern where
concentration of the short winged form indicates an old centrum. Study of wing
dimorphism in Pterostichus anthracinus Illiger (Lindroth, 1945. Die fennoskan-
dischen Carabidae, 3: 1-911) has shown that the condition of the wings has a
hereditary base: short winged forms are dominant, behaving in the simple Men-
delian fashion. In consequence, long winged individuals are homozygotes. In a
mixed population of a dimorphic species, selection functions in accordance with
the completely different means of dispersal of the two forms. During stable periods,
when the habitat is exposed to small changes, the short winged forms are advan¬
taged and become more predominant. In unstable periods, the situation is reversed.
When a dimorphic species invade new areas, as climate grows better, for instance
in late-and post-glacial time, the long winged forms function as groups of pioneer¬
ing parachutists. Long wing forms of dimorphic species do fly sometimes. Flight
power has a positive selective importance in populations living in highly unstable
habitats. Stability, restriction and isolation of environments and stenotopy help
short winged forms. In fact, Carabus maeander lives in highly unstable habitats
such as borders of rivers and wet agricultural lands, in the Montreal region.-— Andre
Larochelle, College Bourget, C. P. 1000, Rigaud, Quebec.
Vol. 49, No. 2, April 1973
127
A New Species of Phobetus
(Coleoptera: Scarabaeidae)
Alan R. Hardy
Insect Taxonomy Laboratory, California Department of Agriculture,
Sacramento, 95814
Investigation of the sand dune habitat in the Southwest has resulted
in the discovery of several species of Scarabaeidae, some new (Hardy
1971, Howden and Hardy 1971). Recently a new species of Phobetus
has been taken in this habitat.
Phobetus was revised by Barrett (1935), and later reworked by Cazier
(1937b). With the exception of the description of Phobetus comatus
robinsoni by Saylor (1939), nothing has since been published on this
genus. The following observations concerning species belonging to
Cazier’s comatus group warrant note.
Phobetus saylori Cazier
This species was described from a unique specimen taken at Little-
rock, Los Angeles Co., California (Cazier 1937a). Cazier (1937b:84)
reported an additional specimen from the same locality.
This species has been found in large numbers at Pinyon Flats, River¬
side Co., California (also known as Seven Level Hills, Pines to Palms
Highway), about 15 miles southwest of Palm Desert. I have examined
additional specimens from Morongo Valley, San Bernardino Co., Cal¬
ifornia. Dates of collection are mostly in April, however the range
covers from February to May.
Cazier (1937b:77) has discussed the variability in the number of
antennal segments of Phobetus , but no mention is made of variability
in the number of segments of the club. Specimens from the type locality
of P. saylori have the antennal club composed of three segments, as
is typical of the other species of Phobetus. The examples from Morongo
Valley have the beginnings of an additional segment, as the last seg¬
ment of the flagellum before the club has become laterally elongated,
to about one-third the length of the other segments of the club. This
elongation is more pronounced in the Seven Level Hill population,
with the segment usually at least three-fourths the length of the club,
so that in fact the club could be considered to be four segmented.
Additional segments in the antennal clubs of the Pachydemini are
not unusual. The Palearctic genera Pachydema , Hemictenius , Tany-
The Pan-Pacific Entomologist 49: 127-131. April 1973
128
The Pan-Pacific Entomologist
proctus and Protoctenius have five segmented clubs, while Elaphocera
and Pachydemocera have seven segments (Medvedev 1952:17).
Following is a description of a new species of Phobetus with a five
segmented club.
Phobetus chearyi Hardy, new species
Holotype male. —Length 13 mm, width (elytra humeri) 5 mm. Head : Clypeus,
antennae and ventral surfaces flavotestaceous; vertex rufopiceous. Scape, pedicel,
mouthparts, and underside of reflexed anterior surface of clypeus with long white
hairs. Labrum bilobed, tumid, produced anteriorly. Clypeus with median longi¬
tudinal impression, flanked by round impression on either side. Clypeus with few
scattered fine punctures. Front and vertex laterally with scattered to close medium
punctures. Antennae with five segmented club. Prothorax : Pale flavotestaceous.
Disc glabrous, shining, with scattered fine punctures. Margins with numerous
long white hairs meeting over top of prothorax. Ventral surface densely clothed
with long white hairs. Scuteilum: With white hairs posteriorly; polished, flavo¬
testaceous; partially obscured by pro thoracic hairs. Elytra : Disc shining,
flavotestaceous, becoming brown at suture and along outer margins. Striae finely
punctate, intervals plane with scattered fine punctures, each elytron with few
scattered white hairs basally, glabrous apically. Outer margins with short, yel¬
lowish hairs, sutural margins with longer white hairs. Thorax : Underside densely
clothed with long white hairs, polished, shining, flavotestaceous. Abdomen:
Flavotestaceous, clothed with long white hairs, dense and scattered laterally,
medially glabrous except for transverse row of white hairs on midline of each
sternite. Pygidium with long white hairs scattered evenly over disc. Legs: Ante¬
rior femur ventrally with long white hairs, tibia bidentate, third tooth present
only as faint projection on outer margin of tibia; tibial teeth testaceous. Apical
spur shorter than tibial teeth. Tarsal claws deeply cleft, median tooth as long
as outer. Mesothoracic femur with scattered rows of long white setae. Tibia with
spiniform medial carina on outer surface, apically with two slender spurs. Meta-
thoracic femur with dense long white hairs on anterior margin, few scattered hairs
elsewhere. Tibia with broken spiniform carina on outer face. Corbel with tarsal
articulation extended, rising above plane of corbel. Corbel surrounded by fringe
of spinules; with two apical spurs, one on either side of channel extending from
posterior margin to tarsal articulatory socket. Male genitalia: Figure 4.
Allotype female. —Length 14 mm, width (elytral humeri) 5.5 mm. Differs from
holotype as follows: Front vertex less punctate, vertex with two well marked
depressions. Antennal club shorter, subequal to scape; basal segment only half
as long as the other four segments. Elytra proportionately broader (width/length
= 0.375, male; 0.421, female). Abdomen massive, much larger than male (see
Figs.).
Figs. 1-3. Phobetus chearyi. Fig. 1 . Holotype male. Fig. 2. Allotype female.
Fig. 3. Male antennal club showing five segments.
Vol. 49, No. 2, April 1973
129
130
The Pan-Pacific Entomologist
Variation in para types. —Length 11 mm-12.5 mm. Variation
appears to be slight, most notable differences are in the coloration and
width of the marginal bands of the elytra, the holotype falling at about
the midpoint of the range.
Diagnosis. —Best diagnosed on the basis of the five segmented an¬
tennal club, but the long pale pubescence and uniformly light coloration
make this species easily recognizable.
Holotype male (CAS #11,658), and seven male paratypes, Beach
Dunes, 32 mi. S. San Felipe, Baja California Norte, Mexico, 25
April, 1970, A. R. Hardy and B. S. Cheary. Allotype and 19 male para¬
types, same locality on 6-7 May. Known only from the type locality.
Paratypes are deposited in the following collections: California Acad¬
emy of Sciences, United States National Museum, Canadian National
Collection, Museum of Comparative Zoology, American Museum of
Natural History, Los Angeles County Museum of Natural History, and
the collections of the following individuals: Henry F. Howden, Robert
W. Woodruff, Robert W. L. Potts, Brian S. Cheary, and the author.
I take great pleasure in naming this species for Dr. Brian Cheary,
Vol. 49, No. 2, April 1973
131
in recognition of his help in collecting the type series, and his fine
work as a coleopterist.
The type series was collected on desert sand dunes that were within
a few hundred feet of the Gulf of California. In this area the desert
vegetation conies right to the beach at the edge of the gulf, where
through wind and wave action the sand as been piled into dunes ap¬
proximately 20 to 30 feet high, and for about one-quarter mile in width.
Typical desert vegetation characteristic of the Sonoran Desert, and
especially Creosote and Bunch Grass are spotted throughout the hollows
and washes of these dunes. The area is fairly rugged and at least five
miles of four-wheel drive terrain isolates it from the nearest road. The
beetles were collected as they flew into blacklights placed upon the
dunes. The individuals would alight upon the sheet and within a very
few seconds would rapidly take wing and disappear into the dark. It
was thus necessary to sit at each sheet for the flight period with a net,
in order to capture the specimens upon their brief visit. The period
of activity lasted no more than one-half hour any of the three evenings
spent on the dunes, and rapidly ceased twilight disappeared. The
lone female was found crawling on the sand at the base of a large bush,
and was not observed to fly, if indeed such a thing is possible for such
a large, apparently heavy individual. Extensive sifting of the dunes
failed to locate either adults or larvae, and no additional females were
collected, although special effort was made to find any present.
Literature Cited
Barrett, R. E. 1935. New species of North American Scarabaeidae II. Can.
Entomol., 67: 49-52.
Cazier, M. 1937a. Four new California Coleoptera. Pan-Pac. Entomol., 13:
115-118.
1937b. Revision of the Pachydemini of North America. Pomona J. Entomol.
Zool., 29: 73-87.
Hardy, A. R. 1971. The North American Areodina with a description of a
new genus from California. Pan-Pac. Entomol., 47: 235-242.
Howden, H. F. and A. R. Hardy. 1971. Generic placement and adult behavior
of the genus Leptohoplia Saylor (Coleoptera: Scarabaeidae). Proc.
Entomol. Soc. Wash., 73: 337-341.
Medvedev, S. I. 1952. Scarabaeidae, subfamily Melolonthinae, part 2 (cock¬
chafers). IN Fauna USSR. Zoological Institute of the Academy of
Sciences of USSR, New Series no. 52: 1-274. Moscow and Leningrad.
[In Russian]
Saylor, L. 1939. Seven new scarab beetles from California. Proc. Entomol.
Soc. Wash., 41: 54-58.
132
The Pan-Pacific Entomologist
New Species of Triclioptera
D. G. Denning
2016 Donald Drive, Moraga, California 94556
During the past several years new species of caddis flies have been
collected in western United States. Six of these have been selected for
description here. Distributional records of several inadequately known
species are given. The hitherto unknown immature stages of two species
of Farula are included. Unless indicated otherwise, types of the new
species are in the author’s collection.
Glossosomatidae
Glossosoma sequoia Denning, new species
This species belongs to the alascense-oregonense section of the Ripae-
glossa line and is closely related to C. merica Denn. Distinguishing
characters are the eighth sternite tubercle, the cerci, claspers and aede-
agus shield. The coiled thoracic process, which occurs in the males of
some species of Glossosoma, also contains characters of diagnostic
value.
Male. —Length 8.5 mm. Wings fuscus, except for pterostigma, quite glabrous;
head, thorax light brown. The spiral bulbous portion of the prealar coiled thoracic
process dark brown. Genitalia as in Fig. 1. Fifth sternite (Fig. 1A) with narrow
auriculate-like lobes. Seventh sternum with prominent spatulate dark brown process.
Eighth sternum tubercle short, subacute, blackish (Fig. IB). Tenth tergite com¬
pletely covered by typical hood-like ninth segment. Cercus and tenth tergite consist
of pair of lobes separated on meson by a membraneous sheath; cercus divided into
ventrad digitate lobe and dorsad wide, truncate lobe; mesal lobe (tenth tergite)
darkened distally, irregularly dentated (Fig. 1C). Claspers short, subtriangular,
ventral margin straight, apex acute, dorsal margin dark bearing minute setae.
Aedeagus shield arcuate, ventral process massive and joined to base of clasper,
about midway to apex it becomes bifid and tapers to an acute ventrad curved apex;
from ventral aspect (Fig. ID) apices acute, convergent. Membraneous aedeagus
bearing internal pair of minute sclerotized processes, best seen from ventral aspect
(Fig. ID).
Holotype male , Lodgepole, Sequoia National Park, California,
6,800 feet, 27 August 1969, black light, R. G. and M. C. Malin. Deposited
in the Entomology Collection, University of California, Riverside,
California.
The Pan-Pacific Entomologist 49: 132-143. April 1973
Vol. 49, No. 2, April 1973
133
PSYCHOMYIIDAE
Tinodes gabriella Denning, new species
Members of the genus are cosmopolitan, widespread, and occasionally
abundant. This new species represents the tenth North American species.
All known species are found only in the western states and adjoining
Mexico. This species may be distinguished from other Tinodes by the
lateral or the ventral aspect of the claspers, by the mesal prong of the
clasper, and by the aedeagal sheath. Tinodes gabriella is related to T.
provo Ross.
Male. —Length 8.5 mm. Wings fuscus, head and thorax darker, appendages
similar. Spurs 2-4-4, same color as legs. Genitalia as in Fig. 2. Ninth sternum
trianguloid, largely covered by eighth sternum, dorsal corner connected to aedeagal
sheath. Apex of ninth tergum terminated in membraneous sheath. Cerci fusiform,
extending caudad beyond remainder of genitalia. Clasper basal segment somewhat
rectangular, ventral margin slightly shorter than dorsal; from ventral aspect (Fig.
2A) apices blunt, short acute spine present along mesal margin near apex; apical
segment with dorsal and ventral margins almost parallel, apex obtuse; from ventral
aspect, apices convergent. Mesal blade of clasper curved ventrocaudad, apex
truncate, pair of prominent acute spines, subequal in size at base (Fig. 2). Aede-
agus encased in a long, slender tubular sheath, curved ventrad and armed with
prominent spines (Fig. 2B) ; aedeagus elongate, apex acuminate (Fig. 2C).
Holotype male , Graveyard Canyon, East Fork, San Gabriel River,
Los Angeles County, California, 9 May 1970, J. A. Honey. Type
will be deposited in the Entomology Collection, Los Angeles County
Museum of Natural History, Los Angeles, California.
Because the distribution of Tinodes species is poorly known, the
following distributional records are given.
Tinodes consueta McLachlan
This ubiquitous species in California is often present in abundance.
It is here recorded from Gold Beach, Oregon, 30 June 1967, Kenneth
Goeden, and Medford, Oregon, 1 July 1966, L. G. Gentner.
Tinodes provo Ross and Merkley
The species occurs in Utah, Arizona and California. It is here re¬
corded from Gooding County, Idaho, 10 April, R. L. Newell. In the
Aqua Caliente Indian Reservation, Riverside County, California, P. H.
Arnaud collected several males and females on 24 and 25 February
1970. At 5000 feet elevation, T. W. Fisher collected a male in Inyo
County, California on 9 March 1966; it probably also occurs in Nevada.
134
The Pan-Pacific Entomologist
FIG3 HYDROPSYCHE CORA
Fig. 1 . Glossosoma sequoia Denning, male genitalia, lateral view. 1A, fifth
sternite lobe, lateral view. IB, eighth sternite tubercle. 1C, cercus and tenth tergite,
lateral view. ID, aedeagus apex, ventral view. Fig. 2. Tinodes gabriella Denning,
male genitalia, lateral view. 2A, apices of basal segment of clasper, ventral view.
2B, aedeagus sheath, ventral view. 2C, aedeagus apex. Fig. 3. Hydropsyche cora
Denning, male genitalia. 3A, lateral view of male genitalia. 3B, tenth tergite apex,
dorsal view. 3C, aedeagus lateral view. 3D, apex of aedeagus, dorsal view. 3E,
dorsal ovoid plate, lateral view. Fig. 4. Farula honeyi Denning, male genitalia,
lateral view. 4A, male genitalia, dorsal view. 4B, clasp ers, ventral view. Fig. 5.
Farula honeyi Denning, female genitalia, 5A, apex eleventh tergite, dorsal view.
5B, spermatheca ventral view, 5C, fifth sternite, lateral aspect.
Vol. 49, No. 2, April 1973
135
Tinodes SIGODANA Ross and Merkley
To date collections are available only from Los Angeles County,
California. It has recently been collected from June to July, 1970 and
1971 by J. A. Honey in the San Gabriel Mountains, Los Angeles County,
California.
Tinodes powelli Denning
New records from southern California are available. Riverside
County: near Palm Desert, 26 June 1970, 23 July 1969, Saul Frommer;
Aqua Caliente Indian Reservation, 24 February 1970, P. H. Arnaud;
Los Ninas Ranch, Murrieta Road, 18 July 1971, J. A. Honey; San Diego
County: Scissors Crossing, 7 October 1971, J. A. Honey; Orange
County: Holy Jim Creek, 25 April 1970, J. A. Honey. Utah: Wash¬
ington County, Leeds Canyon, July 1967, W. J. Hanson. The species
is now known to occur from Baja California to Utah.
Tinodes siskiyou Denning
Described from Jackson County, Oregon, it has now been collected in
Wallowa County, Oregon, 1 July 1967, Kenneth Goeden. California
records are now available: Mendocino County, 9 March 1968, J. R.
Heifer; Sonoma County, 17 June 1972, D. G. Denning, along a heavily
shaded shallow cool water creek; Lake County, Big Canyon Creek, June
17 and October 21, 1972, Paul Peterson.
Tinodes cascadia Denning
Known only from Oregon. A new Oregon locality is: Douglas County,
Oregon, 13 miles East Reedsport, collected resting on a wet seepage
cliff, 1 June 1966, J. D. Vertrees and Joe Schuh.
Tinodes belisa Denning
Known from Oregon south to Butte, Lake and Sonoma County,
California.
Hydropsychidae
Hydropsyche cora Denning, new species
This new species is related to H. venada Ross. It is easily discernible
from other Hydropsyche in the configuration of the tenth tergum, the
position of the spur associated with the membraneous dorsolateral pro¬
cess of the aedeagus, the dorsal plates and the short, thick peculiar profile
of the aedeagus.
136
The Pan-Pacific Entomologist
of male genitalia. 6B, dorsal view of male genitalia. 6C, ventral view of claspers.
6D, female genitalia, lateral view. 6E, spermatheca, ventral view. Fig. 7. Farula
malkini Ross. 7A, female genitalia, lateral view. 7B, spermatheca, ventral view.
7C, head of larva, dorsal view. 7D, larval labrum. 7E, larval prothorax, meso-
thorax and metathorax, dorsal view. 7F, foreleg of larva. 7G, pupa, apex of ab¬
domen. 7H, pupal mandible. 71, sixth, seventh, eighth tergite of pupa, showing
mesal plates. Fig. 8. Farula honeyi Denning, pupa, 8A, apex of abdomen, dorsal
view. 8B, abdominal segments, six, seven and eight, dorsal view.
Vol. 49, No. 2, April 1973
137
Male.— Length 7.5 mm. Wings dark gray, head, thorax and antennae dark
brown, legs yellowish. Genitalia as in Fig. 3. Ninth segment annular, lateral
projection ovate, tergum directed caudad. Tenth tergum with distinct dorsal
angulation; from dorsal aspect, Fig. 3B, apex divided into pair of short subacute
processes, slightly convergent. Clasper apical segment short, digitate, slightly
narrowed from basal segment. Aedeagus, Fig. 3C, with base at right angle to
ninth segment, basal half large and irregular, distal portion considerably narrowed;
apex divided into dorsal and ventral branches with small membraneous area in
between bearing minute extrusible spines; dorsomesal membraneous area bearing
pair of short black acute spines near base (Fig. 3D) and ovoid dorsal plates (Fig.
3E) appear dorsad to apex.
Holotype male , Valley Forge Canyon, San Gabriel Mountains, Los
Angeles County, California, 26 May 1971, J. A. Honey. Type to be
deposited in the Entomology collection, Los Angeles County Museum
of Natural History, Exposition Park, Los Angeles, California.
Limnephilidae
Farula Milne
The genus Farula is known only from Washington, Oregon and Cal¬
ifornia. There are seven species ascribed to the genus, including the
two new species described here. Members of the genus are seldom col¬
lected and are rare in collections. Previous to the collection of the new
species F. honeyi , and F. petersoni , all members of the genus were known
only from Washington and Oregon. Farula species are generally col¬
lected in cool high humidity areas from relatively low elevations to 5,000
feet. One new species described here is from southern California and
represents a great extension southward of this interesting genus. To be
consistent with described species the same lettering system is used as
that employed by Ross (1950) in describing the complicated male
genitalia.
Farula lioneyi Denning, new species
Male.— Length 6 mm. Wings dark gray, densely setose; head, thorax, femora
blackish, antennae and legs brownish, covered with dense dark setae. Maxillary
palpi porrect, slightly shorter than labial palpi, third segment slightly longer than
second, light yellowish. Head between antennae and palpi with thick yellowish
setae. Genitalia as in Fig. 4. Dorsal lobe of ninth tergite, process “a”, slender
and undulating in lateral aspect, distal half bifurcated, slender, subacute (Fig. 4A)
entire structure slightly sclerotized, difficult to discern. Process “b”, (probable
ventral lobe of ninth tergite), long, slender, arcuate from lateral aspect, apex
flattened, acute; from dorsal view (Fig. 4A) distal portion widened and convergent.
Cercus “c”, slender, digitate, directed dorsocaudad. Process “d”, (probable tenth
tergite), directed ventrocaudad, lateral branch barely discernible from lateral
view; from dorsal view (Fig. 4A) bifid distally, apices divergent, mesolateral
arm projected laterad, short and stocky; transparent membraneous sheath extends
138
The Pan-Pacific Entomologist
FIG 10 PARTHINA VIERRA
FI G 1 2 LEPIDOSTOMA LflCINATUM
Fig. 9. Parthina vierra Denning, male genitalia. 9A, male genitalia, lateral
view. 9B, tenth tergite, dorsal view. 9C, clasper, ventral view. 9D, aedeagus,
right lateral view showing internal dorsal rod, and the slender ventral rod. 9E,
aedeagus, left lateral view, showing internal bifid sclerotized structure. Fig. 10,
female genitalia, lateral view. 10A, eleventh tergum, dorsal view. 10B, spermatheca,
ventral view. Fig. 11. Parthina linea Denning. 11A, aedeagus, right lateral view,
showing the dorsal and ventral internal rods. 11B, left lateral view, bifid internal
structure. Fig. 12. Lepidostoma lacinatum Flint. 12A, male genitalia, lateral
aspect. 12B, dorsal view, tenth tergite. 12C, clasper, caudoventral view. 12D,
clasper apex, mesodorsal view.
from apex of lateral arm to cercus and margin of ninth segment. Claspers, “cl”,
heavily sclerotized, tapered to dorsad curved acute apex; from ventral aspect (Fig.
4B) claspers widely separated, acute, directed caudad; process “e” originates near
base, best discernible from ventral aspect, digitate, lightly sclerotized, setose, homol-
Vol. 49, No. 2, April 1973
139
ogy unknown. Aedeagus “ae”, semimembraneous, difficult to discern except from
lateral aspect (Fig. 4) slender throughout, apex abruptly curved ventrad.
Female.— Length 6 mm. Similar to male in general structure and color. Tenth
tergite narrow and elongate, division between tenth and eleventh tergite obscure
(Fig. 5) eleventh tergite declivent, apex nearly truncate, from dorsal aspect (Fig.
5A) bifid. Spermatheca (Fig. 5B) occupying most of ninth and tenth sternites,
mesal aperture long, narrow. Fifth sternite cephalid corner developed into obtuse
process and invaginated small clear, elongate area (Fig. 5C).
Pupa.— Length 5 mm. General structure similar to that of F. malkini (Fig. 8A).
Apex of mandibles acute, slender, no serrations. As in F. malkini labrum bears six
long black setae. Apex of abdomen and mesal plates of tergites six, seven and
eight, almost identical to F. malkini (Fig. 8B).
Holotype male, San Gabriel Mountains, Valley Forge Canyon,
Los Angeles County, California, 26 May 1971, J. A. Honey. Allotype,
same data as for holotype. Paratypes, 13 males, 8 females, same data as
for holotype; 1 male, Angeles Crest Highway, San Gabriel Mountains,
Los Angeles County, California, 2 June 1970, J. A. Honey. Holotype,
allotype and 2 male and 2 female paratypes deposited in Entomology
Collection, Los Angeles County Museum of Natural History, Los Angeles,
California.
I take pleasure in naming this Farida in honor of the collector, J. A.
Honey, who has collected many very interesting Trichoptera. The col¬
lection site consisted of “a very small trickle of water running down
almost verticle rocks” at an altitude of 4,800 feet. Valley Forge Canyon
is “a small, very steep canyon,” the stream forms “many small falls and
pools,” in the summer “it is reduced to a trickle.”
Farula petersoni Denning, new species
This species is related to F. malkini , but in many details of the male
genitalia it differs greatly from that species.
Male.— Length 6 mm. Wings gray, pubescence sparse. Basal segment of antenna
whitish except for mesal brown stripe entire length. Labium also whitish. Maxil¬
lary palpi porrect, whitish, pubescence dark and quite heavy. Pronotum with two
pairs of semicircular white areas; mesoscutellum whitish, remainder black. Gen¬
italia as in Fig. 6. Dorsal lobe of ninth tergum, process “a” (Fig. 6A) lightly
sclerotized, digitate, apex oblique and bearing dense fringe of long brownish setae
in lateral aspect; from dorsal aspect (Fig. 6B) bifurcated from base, each lateral
lobe slender, directed caudad. Process “b”, sinuate in lateral aspect (Fig. 6A)
acuminate, apex acute and curved ventrad; viewed dorsally rod-like structures acute,
curved laterad then mesad, gradually convergent (Fig. 6B). Cercus “c”, long,
slender, about same width throughout, setae sparse. Process “d”, stout, heavily
sclerotized, directed caudad beyond cerci, dark subacute apex curved laterad;
basolateral branch barely discernible; from dorsal aspect (Fig. 6B) basal branch
directed caudolaterad, apex attached to lightly sclerotized and pigmented sheath
140
The Pan-Pacific Entomologist
dorsad to cercus and ninth tergum, apices of mesal branches acute and divergent.
Claspers “cl”, considerably narrowed distally, apex acute, curved sharply ventrad;
from ventral aspect (Fig. 6C) directed caudad, apices acute, curved strongly
mesad; process “e” originates near base, digitate, slender, apex subacute, directed
caudoventrad; filamentous branch “f” (Fig. 6A) arise from base and extend caudad
slightly beyond process “d”. Aedeagal shield lightly sclerotized, trough-like, apical
emargination rounded from ventral aspect (Fig. 6C).
Female. —Practically identical to male in size, wing pubescence and general
coloration. Genitalia as in Fig. 6D. Tenth segment annular, division between
it and eleventh tergum obscure. Eleventh tergum declivant, ventrocaudal apex
rounded, apex bifid from dorsal aspect. Spermatheca (Fig. 6E) occupy portion
of ninth and most of tenth sternites; mesal aperture wide, elongate.
Holotype male, unnamed creek one mile NW Trinidad, Humboldt
County, California, 1 May 1972, elevation 50 feet, Paul A. Peterson.
Allotype female, same data as for holotype, except 3 June 1971. Para-
types 3 males, 2 females, same data as for allotype.
This species is named in honor of the collector, Paul A. Peterson, who
collected the species from a small creek “about 3 feet wide and 1 foot
deep, with a rocky, moss-covered bottom and scattered gravel-filled
pools.” “The creek ends abruptly in a 30 foot waterfall emptying into
the Pacific Ocean.”
Farula rainieri Milne
Until recently known from 1 male collected at Mt. Rainier, Washing¬
ton. Schmid, 1968 recorded the species from Chinook Pass, Washington,
Skagit County, Washington and Marion County, Oregon. On 15 June
1969, R. W. Baumann collected 1 male at a creek at Christmas Falls,
Mt. Rainier, Washington.
Farula malkini Ross
Previously known from 2 males collected in Benton County, Oregon,
it was recently recorded in large numbers from Oak Creek in Benton
County by Anderson and Wold. The species was collected in emergence
traps in April and May. Here recorded from Douglas County, Oregon,
18 miles east of Reedsport, Loon Creek Road, 27 May 1964, 3 males,
2 females, and on 1 June 1966, 1 male, 2 females, J. D. Vertrees and
Joe Schuh. Adults, larval and pupal cases collected from a wet seepage
cliff. Cases collected when few mature larvae or pupae were present,
the majority of cases contained dead and disintegrating larvae and
pupae.
Female. —-Similar to male in color and size. Genitalia as in Fig. 7A. From
lateral aspect tenth segment annular; eleventh tergum obtuse, declivent; from
Vol. 49, No. 2, April 1973
141
dorsal aspect, lateral apices widely cleft, subacute; bursa copulatrix triangular,
mesal aperture narrow, long (Fig. 7B).
Larva.— Head uniformly dark brown except for irregular cleared eye area, near
apical corner inconspicuous slender antenna (Fig. 7C) epicranial suture distinct,
from dorsal aspect (Fig. 7D) ; labrum with row of several short setae. Prothorax
with no trace of a transverse furrow; dark brown, completely and heavily scle-
rotized, distinct mesal fracture; mesothorax completely sclerotized, light brown,
distinct mesal fracture, apical margin with mesal emargination; metathorax with
pair of brown sclerotized irregular-shaped lateral plates and pair of oval semi-
membraneous grayish areas hearing three minute setae (Fig. 7E). Forelegs (Fig.
7F) with femora widened, apical margin short, only slightly longer than tibia.
Based on one partially intact larva, remainder too decomposed for description.
Pupa.— Apex of abdomen (Fig. 7G) with long, finger-like lobes, narrowed
distally, largely devoid of setae. Mandibles acute (Fig. 7H) no serrations; labrum
with group of six long dark setae. Sixth, seventh and eighth tergites with mesal
plates (Fig. 71) ; sixth tergite with pair of irregular oval plates near apical margin
and pair of narrow elongate plates at posterior corner; seventh and eighth tergites
with pair of irregular oval plates near anterior margin, each bearing several minute
hooks; abdomen without lateral fringe of hair.
Case.-— Case of mature larva and pupa 5-6 mm long, cylindrical and slightly
curved, tapering gradually from 0.5 mm to almost 1 mm in diameter. Constructed
of minute sand grains, all similar in size and cemented together to form smooth
exterior.
Odontoceridae
Partliina vierra Denning, new species
The genus has been represented, by a single species, P. linae. This
second species may be recognized by the configuration of the tenth
tergite, the short apical segment of the clasper and the internal pro¬
cesses of the aedeagus.
Male.— Length 7.5 mm. General color dark brown, wings densely setose, re¬
flexed fold of the subcostal cell of forewing filled with intermixture of dark and
light colored scales; head and thorax dark brown, mesoscutellum whitish, setose
sparse; maxillary and labial palpi light brown, setose; tibial spurs densely covered
with short brown setae. Genitalia as in Fig. 9. Ninth segment widest laterally
with obtuse cephalad and short truncate distal projections, dark line extends through
segment, tergum considerably narrowed, distinct semicircular clear area near
sternum (Fig. 9A). Tenth tergite composed of pair of lateral convex plates, distal
corner deeply emarginate, resultant acute prongs directed ventrad; from dorsal
aspect (Fig. 9B) lateral plates widely separated, apex obliquely truncate, slightly
divergent. Cerci digitate, bearing several very long setae. Clasper basal segment
long, cylindrical, gradually tapered distally, basal portion characteristically dark¬
ened; apical segment short, truncate (Fig. 9A) ; from ventral aspect (Fig. 9C)
apical segment acute, curved mesad. Basal portion aedeagus sclerotized, distal
portion semimembraneous; from left lateral aspect (Fig. 9E) internal bifid, light
brown sclerotized structure discernible; from right lateral aspect (Fig. 9D) internal
dorsal rod present, distally acuminate and sharply curved ventrad, near base acute
142
The Pan-Pacific Entomologist
ventral angulation present, located ventrad long, slender, acuminate rod reaches
almost to apex; aedeagus slightly longer than combined ninth segment and claspers.
Female. —Genitalia as in Fig. 10. General appearance similar to male except
absence of scales on forewings. Tenth segment large, oval, light brown sclerotized;
eleventh tergum lightly sclerotized, from dorsal aspect (Fig. 10A) narrowly sep¬
arated on meson, apex rounded; bursa copulatrix (Fig. 10B) long and narrow.
Holotype male , Uvas Creek, Uvas Canyon, Santa Clara County,
California, 3 July 1971, D. G. Denning; allotype, same data as for
holotype; paratypes, 4 males, 3 females, same data as for holotype.
Parthina linea Denning
In the original description the internal structures of the aedeagus
were not figured. When viewed from the left lateral aspect, Fig. 11B,
the internal structure divided into a dorsal process widened distally, and
an acute short ventral process, from right lateral aspect, Fig. 11 A,
dorsal rod acuminate and curved slightly ventrad, ventral rod long,
slender, acuminate.
Parthina linea was described from specimens collected in Tuolumne
County, California, and from near McMinnville, Oregon. Additional
records are: 27 males, 6 females west of Selma, Josephine County,
Oregon, 6 May 1964, Joe Schuh; 10 males, 1 female Macalester Wash
near Martinez, Yuma County, Arizona, 30 March 1961, C. A. Toschi
Tauber; this water is backed up from the Colorado River and contains
numerous cattails and algae; 1 male Santa Clara County, California,
1 June 1960, S. D. Smith.
Lepidostomatidae
Lepidostoma lacinatum Flint
This species has not been recorded since it was described from Du¬
rango and Sinaloa, Mexico by Flint, 1967. The collection in Arizona
represents an extension northward of about 700 miles. One male col¬
lected at Miller Canyon, Huachuca Mts., Santa Cruz County, Arizona,
17 August 1971, 7,000 feet, J. A. Honey.
The Arizona specimen suggests the species may be quite variable
(Fig. 12). Major differences are: the short, robust tenth tergite, apices
truncate, not distally narrowed (Fig. 12A) from dorsal aspect (Fig.
12B) apices widely separated; aedeagal prongs only slightly conver¬
gent; apex of apicodorsal lobe of clasper curved ventrad, lateral surface
with a dense mat of long yellowish setae which obscures details of the
structure, from caudoventral aspect (Fig. 12C) apex acute; from meso-
Vol. 49, No. 2, April 1973
143
dorsal aspect (Fig. 12D) the acute hooked ventral arm of the mesodorsal
process reaches almost to margin. ,
Literature Cited
Anderson, N. H. and J. L. Wold. 1972. Emergence Trap Collections of Tri-
choptera from an Oregon Stream. Can. Entomol., 104: 189-201.
Flint, 0. S. 1967. Studies of Neotropical Caddis Flies, VI: on a collection from
northwestern Mexico. Proc. Entomol. Soc. Wash., 69: 162-176.
Ross, N. H. 1950. Synoptic notes on Some Nearctic Limnephilid Caddis-flies
(Trichoptera, Limnephilidae). Amer. Midland Natur., 43: 410-429.
Schmid, F. 1968. Quelques Trichopteres Neartiques nouveaux ou peau connus.
Natur. Can., 95: 673-698.
BOOK REVIEW
The Moths of America North of Mexico. Fascicle 21, Sphingoidea. Ronald
W. Hodges. E. W. Classey Ltd. and R. B. D. Publications, Inc. 158 p., 14 color
plates, 19 figs., xii. 1971. $24 (subscription price $19.60). North American
Distributor: Entomological Reprint Specialists.
This is the first to be published in a projected series of 41 Fascicles treating
“The Moths of America North of Mexico.” The concept of covering the more than
10,000 species with color illustrations of all species and their major color forms
is truly a monumental one. The present fascicle on the Sphingoidea provides an
excellent advertisement for the high quality of the series. The heavy paper, large
clear print, beautifully reproduced life size color illustrations, and excellent line
drawings are a credit to all involved with the production of this fascicle. A brief
history of the project with photos of those involved appears in an introductory
section.
Keys to 40 genera and 115 species of adults and partial keys to genera based
on pupae and mature larvae are presented. Species treatments vary in complete¬
ness. They include synonymies, adult and larval diagnoses, geographic and sea¬
sonal distributions, food plants and habits. Infraspecific variation is discussed and
often illustrated, but subspecies are not treated as such. Text figures of genitalia
are provided for representatives of most genera. A few name changes are apparent
even to the non-lepidopterist. Celerio and Protoparce are treated as synonyms of
Hyles and Manduca respectively.
The 14 plates containing 199 color photos are followed by two plates of ex¬
cellent, clearly labelled structural drawings by the author’s wife. The fascicle
concludes with indices to animal and to plant names.
The cost of the series seems a bit high for the average entomologist, but the
quality of the publication and the impressive color plates make it a worthwhile
investment for dedicated lepidopterists and a manditory reference for all insti¬
tutions.— Robbin W. Thorp, University of California, Davis, 95616.
144
The Pan-Pacific Entomologist
The Behavior of Microhemhex nigrifrons
(Hymenoptera: Sphecidae)
John Alcocic and Allen F. Ryan
Department of Zoology, Arizona State University, Tempe, 85281, and
Departments of Psychology, Physiology, & Biophysics, University of Washington,
Seattle, 98195
The behavior of only a few species of Microbembex has been reported
in any detail (Evans, 1966, for M. monodonta (Say) ; Goodman, 1970,
for M. californica Bohart). This paper describes the behavior of a pre¬
viously unstudied member of the genus, M. nigrifrons (Provancher).
Over two summers (1971, 1972) we made a series of observations on
this species in central Washington. The study site was located approx¬
imately five miles south of Interstate 10 on Dodson Road within ten
miles of Royal City. A series of low dunes scattered through this area
are inhabited by M. nigrifrons and other bembicine wasps. The dunes rise
out of a flat plain covered with sage and other desert scrub brush with
buckwheat ( Eriogonum sp.) common on the fringes and crests of some
dunes.
Nesting behavior.- —-This species of Microbembex is extremely similar
behaviorally to M. monodonta , M. californica, and all the South Amer¬
ican species studied by Evans (personal communication). It nests
primarily on the lower fringes of sand dunes and sand banks often
burrowing into a slope. Upon completion of a burrow, a task that may
require two to three hours early in the season (June), the female per¬
forms an initial closure similar to that described for M. monodonta by
Evans (1966: 370). The wasp walks quickly away from the entrance
kicking sand back toward the burrow. After going out 15-25 cm, it
then returns and repeats the process in another direction over and over
again until the nest is surrounded by a series of radiating lines. The
burrow often, but not always, descends at a shallow angle for about
one-half its length and then drops much more steeply before levelling
off just before the cell (Fig. 1). All nests excavated were single celled.
The egg is laid upright in a vertical position in the empty cell. (One
nest held an egg and one small dead beetle.) Provisioning females
apparently select any dead arthropod available: Arachnida: Scor-
pionida 1 (the tail), Araneida 5; Insecta: Ephemeroptera 3, Orthop-
tera 15, Hemiptera 8, Neuroptera 1, Trichoptera 1, Lepidoptera 12
(adult and larval forms), Diptera 24, Hymenoptera 15, Coleoptera 31
The Pan-Pacific Entomologist 49: 144-148. April 1973
Vol. 49, No. 2, April 1973
145
Fig. 1. A diagram of two nests of M. nigrifrons illustrating the variation in
length and depth as well as nest design that existed at the Dodson Rd. site.
(adult and larval forms). In addition a few females were seen hovering
over and then touching small brown seeds and other bits of dried veg¬
etable material perhaps inspecting them as possible prey items.
Three females were observed shortly after they had taken a food item.
Two clung upside down to the underside of a branch in the dunes re¬
arranging the prey prior to flying back to the nest. The other was
oriented vertically head down on a stick. Females returning to the nest
were harassed by other members of their species and one successful prey
stealing was observed. The average interval (N = 8) between provision¬
ing trips to the nest was 21 minutes (range = 7-44). In every respect
the behavior of M. nigrifrons is highly similar, if not identical, to that
of M. monodonta.
Nest DIMENSIONS and sand moisture.— Evans (1966) noted con¬
siderable variation in cell depth and burrow length both at the same site
and between nesting locations. He demonstrated that part of the dif¬
ferences between burrows was related to the size differences between
females. Moreover, Evans speculated that variation between locations
could be due to differences in the moisture content of the sand noting
that the deepest nests were found in very dry Kansas dunes.
Because preliminary excavations of some nests in mid 1971 revealed
exceptionally long and deep burrows this aspect of nesting behavior
146
The Pan-Pacific Entomologist
Table 1. The relation between the nest dimensions of M. nigrifrons
and the depth of moist sand in the dunes.
10 June
12 July
12 August 1972
14 Sept. 1971
Average depth of moist
sand (cm below surface) 5
11
17.5
5
Nests excavated
Average burrow length
12
12
10
10
(cm) 1
28.3
31.4
48.5
18.7
Range
Average cell depth
(15-63)
(26-40)
(42-57)
(14-35)
(cm) 2
13.4
18.9
31.8
14,0
Range
(11-23)
(14-23)
(25-38)
(9-20)
Average distance between moisture
line and cell (cm) 3
10.3
8.1
14.3
9.0
1 Correlation of burrow length with average depth of moist sand, r — .76, P <[ -01.
2 Correlation of cell depth with average depth of moist sand, r — .54, P <C .01.
3 F test of mean depths of cells below moisture line, F^ 3 40 ^ = 16.8, P <[ .01.
was chosen for more study. On four occasions (14 September 1971,
10 June, 12 July and 12 August 1972) a sample of 10-12 nests was
excavated. In the Dodson Road dune area the moisture level in the
dunes was very clear cut; extremely dry loose sand gave way to moist
compact sand anywhere between 3.5-20 cm beneath the surface de¬
pending on the time of year. Table 1 presents data on the correlation
between average depth of moist sand and the dimensions of the nests.
It is abundantly clear that the lower the moisture level, the deeper and
longer the burrows.
The adaptive significance of this behavior is not obscure. It is to
the wasp’s advantage to locate its cell in an area where the egg and larva
will not desiccate prior to formation of a cocoon. This response does
indicate a degree of behavioral flexibility with wasps in this population
capable of constructing a burrow anywhere from 14-63 cm long depend¬
ing on environmental conditions. But even this variation can be readily
achieved if the wasp is simply programmed to dig a nest with the cell
about 10 cm beneath the moisture line (approximately 80% of the sam¬
ple had cells 7—15 cm below the line). However, the matter is somewhat
more complex than this. Wasps building nests when the moisture level
was very low placed their cells significantly deeper below the line than
those nesting when moist sand was closer to the surface of the dunes
(Table 1).
Orientation to the nest. —Evans (1966) remarks that M. mono-
Vol. 49, No. 2, April 1973
147
donta exhibits an astonishing ability to locate its nest entrance despite
great disturbance to the surrounding terrain. He describes experiments
by himself and others in which landmarks close to and far from the
nest were moved without affecting the orientation of the wasp in the
slightest.
We report here a very simple experiment on the homing ability of
M. nigrifrons. It was apparent from casual observation that females had
no difficulty finding their nest although the sand over and around the
entrance might be very much disturbed by our activities. To test land¬
mark learning by this species a ring of whitish stones 2-3 cm in diameter
were placed in a circle about 15 cm from a nest entrance on 9 August
1972. In addition, a twig 8 cm long was placed inside the ring. The
female was digging the nest when this was done.
On 12 August an observer returned to the site to find that there were
now two females nesting inside the ring. The stones and twig were
moved 25 cm to one side. A small grey pebble was placed where the
twig had been to mark the location of the burrows for the benefit of
the observer. The depressions left by the small stones were covered
with fine sand.
One female returned with prey from her first provisioning trip
after displacement of the landmarks and flew directly to the displaced
ring. There she began to dig at a spot within the ring that corresponded
to the site where the entrance would have been had the rocks not been
moved. After 10 minutes of alternately digging and flying up, the female
left the ring and flew to the true nest entrance. After digging there only
briefly, she returned to the displaced ring, dug for a short time, flew
back to the true nest site, opened the burrow and entered. Upon leaving
the nest the wasp performed an elaborate and prolonged closure similar
to an “initial closure.” Five minutes later, she returned with another
food item. The wasp flew directly to the displaced ring and dug there
briefly before flying to the actual nest which she opened and entered.
Nest closure was performed normally. On her third provisioning trip,
the female went directly to the nest entrance and entered the burrow.
Nest closure was normal.
The second female, returning from her first provisioning trip follow¬
ing displacement of the landmarks, went straight to the spot in the
displaced ring where her entrance would have been had the stones not
been moved. After about 5 minutes of digging, she flew to her true
burrow and began work there. Like the other wasp, she returned to
the ring and made several trips back and forth before finally opening
the burrow and entering. Nest closure was performed normally.
148
The Pan-Pacific Entomologist
It seems clear that M. nigrifrons will use landmarks close to the nest
to locate the entrance. When these cues are very conspicuous it is pos¬
sible to disorient the wasp by moving them. However, the effect is only
temporary, unlike the permanent inability of Philanthus triangulum L.
to find its nest when a ring of pine cones was displaced some distance
from the entrance (Tinbergen, 1951). It may be that M. nigrifrons
learns a variety of cues and can rely on alternate landmarks if major
ones are removed or displaced.
The fact that the first female appeared to be more disoriented than
the second one (digging for a longer period of time at the false nest
site and performing an elaborate closure once the real nest had been
found) is interesting. It seems probable that the second wasp had built
her nest after the first one and therefore may have been less accustomed
to and dependent upon the ring of stones as orientation guides. Much
more work is necessary on this aspect of Microbembex behavior.
Acknowledgments
This study was partly supported by National Science Foundation
Grant GB-28714X. We thank Dr. H. E. Evans for his advice and Dr.
R. M. Bohart for his kindness in identifying the wasp for us.
Literature Cited
Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps.
Harvard Univ. Press, Cambridge, 526 pp.
Goodman, W. G. 1970. Observations on the behavior and biology of Microbembex
californica Bohart (Hymenoptera: Sphecidae). Pan-Pac. Entomol.,
46: 207-209.
Tinbergen, N. 1951. The study of instinct. Oxford Univ. Press, Oxford, 228 pp.
RECENT LITERATURE
Cerambycidae of North America. Part VI, No. 1. Taxonomy and Classifi¬
cation of the Subfamily Lepturinae. E. Gorton Linsley and John A. Chemsak.
University of California Publications in Entomology. Volume 69. University
of California Press, Berkeley. 188 p., 41 figs., 2 plates. 1972. $5.50.
This volume is part of a continuing series toward a monograph of the long-horned
beetles of America north of Mexico. It treats the tribes Desmocerini, Necydalini,
and the 22 genera of Lepturini with lateral spines or tubercles on the pronotum
and/or with entire eyes. Those Lepturini without pronotal tubercles and emarginate
eyes will be treated subsequently in a second number of Part VI. Exquisite half¬
tone illustrations by Celeste Greene and distribution maps are provided for many
representative species.— Robbin W. Thorp, University of California, Davis, 95616
Vol. 49, No. 2, April 1973
149
A New Subspecies of Plebejus acmon
(Lepidoptera: Lycaenidae)
Carll Goodpasture
Department of Entomology, University of California, Davis 95616
Plebejus acmon (Westwood & Hewitson) and Plebejus lupini (Bois-
duval) are closely related, variable species characterized by conspicuous
sexual dimorphism, a well marked orange or pink hind wing band, and
use of Eriogonum as foodplant. A recent biosystematic study of this
complex and of an additional related species, P. neurona (Skinner),
suggests that biological relationships are unusually complex. Morpho¬
logical intergradation, apparent character displacement, and wing pig¬
mentation polymorphism complicate analysis of variation. Discussions
of regional morphological and biological differentiation within and
intergradation between P. acmon group members is presented elsewhere
(Goodpasture, 1973a).
The purpose of the present paper is to describe a new entity of the
P. acmon-P. lupini species complex from the southern Rocky Mountain
region. The new entity is considered conspecific with P. acmon because
of morphological intergradation with this species. Recognition of this
entity as a subspecies clarifies patterns of variation in this complex.
Plebejus acmon texanus Goodpasture, new subspecies
Male. —Forewing length (straight-line distance from base of costa to apex of
Mi) of type series x = 11.2 mm db .45 mm 1 2 (N — 13) ; Type 12.3 mm. Dorsal
wing surface (Fig. 12).—General wing color created by more or less continuous
sheet of iridescent cyanic scales overlying dark scale subsurface. Overlay color
purple of low saturation and brilliance. Approximate hue 3 of iridesence of cyanic
overlay Purple-Blue Purple. Black band at apical margin of primary moderately
broad with few black scales extending basally along veins. Transition of black
marginal band on primaries with cyanic overlay scales gradual. Numerous dark
scales scattered over wing surface. Indistinct row of dark scales present at apex
of discal cell. Distal insulae of secondaries well separated from terminal line.
Submarginal band on secondaries orange, appearing rose when viewed from certain
angles, bordered basally by few dark scales. Ventral wing surface. —Dark gray
ground color; black spots encircled by distinct ring of white scales; submarginal
orange band broad. Genitalia. —Uncus lobes as seen in dorsal view blunt, not
strongly spatulate; in lateral view, small and quadrate as in Figs. 2 and 5.
Female. —Forewing length of type series x — 11.0 mm ± .61 mm (N = 14);
1 Standard deviation.
2 Color terminology follows the Munsell book of color (Anonymous, 1942. Munsell Color Co., Inc.,
Baltimore, Md., 145 pp.).
The Pan-Pacific Entomologist 49: 149-159. April 1973
150
The Pan-Pacific Entomologist
Figs. 1-9. Genitalia of species of the Plebejus acmon complex: Uncus of male,
lateral view (1-3) dorsal view (4-6). Sterigma of female (7-9), dorsal (upper),
Vol. 49, No. 2, April 1973
151
Durango,
Guanajuato
I
112
•N ,-N
/ •'/
./ I
/ V •
> v
104
Map 1. Distribution of Plebejus acmon texanus.
Type 11.7 mm. Dorsal wing surface. —Basal one-fifth covered by cyanic overlay.
Color and iridescence of overlay as in male. Ventral wing surface. —As in male
except ground color gray with brownish cast. Genitalia. —Sterigma heavily scle-
rotized, swollen and pointed apically; terminal plate extending basally onto dorsal
surface of sterigma as in Fig. 8.
Holotype male and allotype, 1 mi. S. Hillside, Yavapai Co., Arizona,
30 September 1969, J. A. Scott. Deposited in the Los Angeles County Mu¬
seum of Natural History, Los Angeles, California. Topoparatypes: 12 $,
13 2, 30 September 1969; 2 2, 21 September 1971, C. E. Goodpasture.
Geographical distribution (Map 1).—Arizona, New Mexico, western
Texas, westcentral Mexico, parts of Colorado, Utah, Nevada, and extreme
<e
ventral (lower), and lateral (center) views. Figs. 1, 4, 7. Plebejus acmon acmon.
Figs. 2, 5, 8. P. acmon texanus. Figs. 3, 6, 9. P. lupini. ul = uncus lobe, tp =
terminal plate.
152
The Pan-Pacific Entomologist
Figs. 10-14. Wing upper surface patterns of males of the Plebejus acmon com¬
plex. Fig. 10. P. acmon acmon Briceberg, Mariposa Co., California, 5 May 1969,
K. C. Hughes. Fig. 11. P. acmon lutzi Pole Mtn., Albany Co., Wyoming, 4 July
1970, C. D. Ferris. Fig. 12. P. acmon texanus Holotype. Fig. 13. P. lupini monticola
Sierra Pelona Road, Mint Canyon, Los Angeles Co., California, emgd. 12 May 1970, C.
Vol. 49, No. 2, April 1973
153
Map 2. Distribution of Plebejus acmon texanus (open arrows), P. acmon lutzi
(filled arrows), and P. acmon texanus-P. acmon lutzi intermediates (divided
arrows) in Colorado. Where arrows point to one locality, more than one phenotype
occurs. Shading roughly denotes areas above 7,000 feet.
southeastern California. Occurring most commonly in mountains below
about 9,000 feet. In southeastern and eastcentral Arizona, Brown (1965)
notes Upper Sonoran through Transition zones; elevations 4,000-7,500
feet. In Colorado, P. acmon texanus occurs at elevations between about
6,000 and 7,000 feet. At higher elevations, phenotypic blending with P.
acmon lutzi occurs (Map 2).
E. Goodpasture. Fig. 14. P. lupini lupini Echo Lake, El Dorado Co., California, 8,000
ft., 10 July 1970, C. E. Goodpasture. Note that darkness of photograph represents
darkness of wing color: lightest blue (Fig. 13) = lightest gray; darkest purple (Fig.
14) = darkest gray. Figs. 15-18. Wing patterns of females: uppersurface Figs. 15
and 18, undersurface Figs. 16 and 17. Fig. 15. P. acmon texanus National forest
boundary, road to Mt. Lemmon, Pima Co., Arizona, 8 April 1966, J. A. Scott.
Fig. 16. P. acmon texanus 21 mi. S Alpine, Brewster Co., Texas, 20 September
1970, J. A. Scott. Fig. 17. P. acmon acmon Monticello Dam, Yolo Co., California,
emgd. 1 May 1970, C. E. Goodpasture. Fig. 18. P. lupini lupini 5 mi. N Sun Pass,
Klamath Co., Oregon, 18 July 1971, C. E. Goodpasture. All figures X 2.2.
154
The Pan-Pacific Entomologist
Fig. 19. Variation in sterigma of females of Plebjus acmon from Schillings
Spring, Conejos Co., Colorado. Morphology as in P. acmon (a) ; P. a. texanus-
P. a. lutzi intermediate (b) ; P. a. texanus (c) ; similar to P. lupini (d).
Material examined (600 specimens).—Locality data is given for
representative material from throughout the range of P. acmon texanus.
Genitalia slide mounts were made of all specimens listed. Specimens
intermediate in genital morphology between P. acmon texanus and P.
acmon lutzi are indicated.
Arizona: Apache Co.: Alpine, 7 June 1937, 1$ ; Turkey Creek Rd., White Mts.,
20-21 June 1935, 1$. Cococino Co.: 7 mi. SW Williams, 26 July 1970, 1$ ; Pine
Flat Campground, 11 April 1963, 1$. Greenlee Co.: Hannagan Meadow, White
Mts., 22-27 June 1937, 4$, 3$. Mojave Co.: Hualapai Mts., 6 September 1964,
3$. Pima Co.: Mt Lemmon, 8 April 1966, 1 2 ; 20 mi. NE Tucson, 4,200 ft., 23
October 1960, 1$. Yavapai Co.: 1 mi. S Hillside, 30 September 1969, 2$. Cal¬
ifornia: San Bernardino Co.: New York Mts.: 1 June 1968, 4$, 1$ ; Keystone
Canyon, 7 mi. SE Ivanpah, 5,000 ft. and 5,500 ft., 23 September-8 October, 1 $,
2 2;% mi. SE Horse Thief Springs, Kingston Range, 4,400-4,800 ft., 29 June 1969,
2 $ ; Gilroy Canyon, Providence Mts., 21 April-30 May 1968, 2 $. Colorado :
Alamosa Co.: Great Sand Dunes Nat’l. Monument, 12 June 1945, 2$ (interme¬
diate). Conejos Co.: % mi. SE Schilbngs Spring, 9 July 1967, 5 2 (2 interme¬
diate) ; W end of Alamosa Reservoir, 9 July 1967, 2 2- Custer Co.: Baldy Peak,
12,000 ft., 29 July 1970, 2 $ ; Hermit Pass Rd., 22 July 1968, 1 $, 12; 1.5 mi. NW
Westcliffe, 25 July 1965, 3$, 12- Fremont Co.: Bear Creek, 22 June 1970, 12 ;
Hillside School, 16 August 1968, 2 2 ■ Gunnison Co.: 9 mi. N Crested Butte, 26-29
July 1966, 1$ (intermediate) ; Gothic, 31 August 1961, 1$ (intermediate) ; Taylor
River, 10 July 1961, 1$ (intermediate). Huerfano Co.: 7 mi. W Malachite, 4 July
1970, 1 $. Jefferson Co.: 1 mi. W Golden, 10 September 1965, 2$ ; Phillipsberg,
4 September 1961, 1 $ (intermediate). Larimer Co.: Round Butte, 29 June 1964,
1 $, 12- Montezuma Co.: Mesa Verde Nat’l. Park, 22 July-1 September, 1 $ , 12-
Vol. 49, No. 2, April 1973
155
Fig. 20. Seasonal distribution of Plebejus acmon texanus.
Pueblo Co.: SW of Pueblo, 31 July 1962, 1 $. Saguache Co.: Mosca Pass Trail,
8,500 ft., 10 July 1970, 3$ ; 1 mi. SW Villa Grove, 17 June 1966, 1 $. New Mexico:
Mora Co.: 8.5 mi. E Wagon Mound on Hwy 120, 16 August 1970, 1$. McKinley
Co.: 4 mi. S Fort Wingate, 10 July 1966, 1$. Sandoval Co.: Sandia Mts. 8,500 ft.,
3 June 1945, 1$. Sante Fe Co.: Cerrillos, 17 August 1970, 12. Torrance Co.:
Manzana Mts., 2 June 1945, 12- Nebraska: Phelps Co.: Atlantic, 23 July 1964,
1$. Nevada: Douglas Co.: 1.5 mi. E Glenbroolc, 4 June 1970, 1$. Clark Co.:
Sheep Range, 3-4 mi. NW Mormon Well, 1-2 July 1969, 1# 12. Lincoln Co.:
10 mi. W Caliente, 6,200 ft., 3 July 1969, 1 $ . White Pine Co.: 1 mi. S Blind Spring,
W end Snake Range, 5,800 ft., 29 July 1969, 1$. Texas: Brewster Co.: 21 mi.
S Alpine, 8 August-20 September, 3$, 42. Jeff Davis Co.: Fort Davis, 13 May
1962, 12. Utah: Washington Co.: Zion Nat’l. Park, 15 June 1938, 1$, 12-
MEXICO: Chihuahua: 17 mi. W Chihuahua and 7 mi. E Pedernales, 7,900 ft.,
11 July 1964, 2 2. Durango: 22 mi. E El Salto, 8,000 ft., 22 July 1964, 1$.
Guanajuato: 10 mi. N Guanajuato, 4 July 1954, 1 $.
Seasonal distribution (Fig. 20).—Multivoltine; collection records
indicate that flight season extends from early spring to late summer.
Data from Fig. 20 suggest multimodality of seasonal abundance with one
peak in spring and another in late summer. Separate spring and summer
flight periods ( March-April and mid-August-September) are reported
for southeastern Arizona; elevation 4,000-6,000 feet (Brown, 1965).
FOODPLANTS.— Certain perennial, bush-like Eriogonum species. Ovi-
position has been observed on Eriogonum wrightii var. wrightii Torr.
ex Benth. at Hillside, Yavapai Co., Arizona and on E. racemosum Nutt.
Table I. Major differentiating characters for the Plebejus acmon-P. lupini complex. Color terminology follows the
Munsell book of color (Anonymous, 1942).
Character
a. acmon and a. lutzi
a. texanus
l. lupini and l. monticola
Male genitalia:
Shape of lobe of uncus in
lateral view.
Uncus in dorsal view
Female genitalia:
Shape of sterigma.
Sclerotization.
Wing pigmentation (dorsal) :
1. Marginal band on FW of
males.
2. Basad extension of marginal
band along FW veins of
males.
3. Marginal band-overlay tran¬
sition of males.
4. Approximate hue of irides¬
cence of cyanic overlay.
Small and angulate (Fig. 1).
Sharply pointed (Fig. 4).
Tubular (Fig. 7).
Entirely membranous or partly
sclerotized. Terminal plate in¬
complete, confined to ventral
apex of sterigma.
Reduced to a thin terminal line
(Fig. 10) in acmon. Broad in
lutzi, very broad in melanic lutzi. 1
Absent or faint. Prominent in
melanic lutzi.
Abrupt (Figs. 10, 13). Gradual
in melanic lutzi.
Purplish Purple-Blue to Purple-
Blue Purple. Purple-Blue in some
Nevada, Utah and Idaho lutzi.
Small and quadrate (Fig. 2).
Intermediate (Fig. 5).
Apically bulbous, sharply pointed
(Fig. 8).
Terminal plate complete, extend¬
ing basad along dorsal surface of
sterigma.
Broad (Fig. 11) or broader than
as in Fig. 12.
Prominent (Fig. 12).
Gradual (Fig. 12).
Purplish Purple-Blue to Purple-
Blue Purple.
Large and strongly quadrate
(Fig. 3).
Blunt (Fig. 6).
Apically heart to spade shaped
(Fig. 9).
As in texanus except basad exten¬
sion sometimes lacking.
Broad. Very broad in melanic
lupini.
Prominent in lupini. Faint in
monticola.
Gradual in lupini. Abrupt in
monticola.
lupini as in texanus: tending
toward Blue Purple-Blue E. of
the Sierra Nevada, monticola:
Blue Purple-Blue to Purple-Blue
in S. Calif. (=type monticola),
Greenish Blue-Green to Blue-
Green in some Kern & L. A. Co.
specimens (=type chlorina) ;
usually as in texanus and lupini
in San Diego Co. south.
on
Os
The Pan-Pacific Entomologist
Table I. (Cont.)
Character
a. acmon and a. lutzi
a. texanus
l. lupini and l. monticola
5. Distal insulae of HW of
males.
6. Color of submarginal band
on HW of males.
7. Border proximal to submar¬
ginal band.
8. Dark scales at apex of FW
discal cell in males.
9. Dark scales scattered over
wing surface.
10. Color of forewing in males.
Separate from terminal line.
Fused with terminal line in
melanic lutzi.
Pink in acmon. Orange in lutzi.
Absent in acmon. Usually present
in lutzi.
Absent in acmon. A few scales
in lutzi.
Absent or few. Many in melanic
in lutzi.
Purple of high saturation and
brilliance in acmon. Purple to
Blue, usually of high saturation
and brilliance in lutzi.
Separate from terminal line.
Orange.
Present
Distinct scale patch.
Many.
Purple of low saturation and
brilliance.
Usually fused with terminal line
in lupini. Usually separate in
monticola.
Orange.
Present in lupini except at low
elevations E. of the Sierra
Nevada. Usually present in
monticola.
Absent in monticola. lupini as
in lutzi.
Absent or few. Many in melanic
lupini.
lupini as in texanus. monticola:
as in lupini to Blue (type monti¬
cola) or Blue Green (type chlor-
ina) of high saturation and bril¬
liance, sometimes largely brown
in Tehachapi Mts., S. Calif.
25 to 0 in lupini. 90 to 0 in
monticola, highest in L. A. Co.,
Calif. (=type monticola).
11. Range in percent of wing 90 to 0 in acmon, highest in over- As in lutzi.
surface covered by overlay in wintering and spring generations
females. (=type cottlei) 25 (Fig. 15) to
0 (Fig. 18) in lutzi.
<
O
tr*
1 Melanic specimens of P. acmon lutzi and P. lupini lupini are extensively dark scaled and are similar or identical in wing characters. Known from higher elevations
and more northern latitudes.
.. 49, No. 2, April 1973 157
158
The Pan-Pacific Entomologist
at Mesa Verde National Park, Montezuma Co., Colorado. Associations
of adults with E. effusum Nutt, in Colorado and E. corymbosum var.
velutinum Reveal & Brotherson in New Mexico suggests that these plants
are also utilized. More complete data on foodplant use are given else¬
where (Goodpasture, 1973b). The type series was collected in asso¬
ciation with E. wrightii var. wrightii.
Discussion
Morphological character combinations that differentiate P. acmon
texanus from other members of the P. acmon-P. lupini complex are
summarized in Table I. Variability of the diagnostic characters listed
is least in morphology of female genitalia and greatest in uppersurface
wing patterns of males. It should be noted that females lack diagnostic
external characters. Species designations of P. acmon group females can
not be made without examination of the genitalia. Subspecies assign¬
ments within P. acmon and P. lupini usually require series of both sexes.
In external phenotype P. acmon texanus closely resembles P. lupini
lupini in uppersurface wing color (characters 6 and 10 of Table I),
forewing marginal band (3, 4 and 5), color of hindwing submarginal
band (8) and presence of a dark border proximal to hindwing sub¬
marginal band (9). These two subspecies differ primarily in intensity
of expression of characters. For example, on the uppersurface of the
primaries, discal scale spot and basad extension of marginal band scales
along veins are strongly expressed in P. acmon texanus and weak or
absent in all but strongly melanic P. lupini lupini. It is of interest that
at least one high elevation population (Mono Pass, California) of P.
lupini lupini is composed largely of individuals pigmentationally in¬
distinguishable from P. acmon texanus. Of ten male wing pigmentation
characters listed in Table I, at least nine are shared by some specimens
of both P. acmon texanus and P. lupini lupini, whereas approximately
six characters are shared with P. lupini monticola, and P. acmon lutzi
and only one or two with P. acmon acmon.
Genital morphology in P. acmon texanus appears intermediate between
that of P. acmon and P. lupini. Genital characters most consistently
differentiating these two species are in males, the uncus, and in females,
the extensible ostium bursa (sterigma). The sharply pointed terminal
plate of P. acmon texanus is unique among P. acmon group members.
Plebejus acmon texanus is treated as a subspecies of P. acmon rather
than of P. lupini which it most closely resembles because of the occur¬
rence of intergradation with P. acmon in at least some areas between
Vol. 49, No. 2, April 1973
159
the ranges of the two entities. In Colorado, several populations include
individuals that are morphologically intermediate between P. acmon
texanus and P. acmon lutzi. At these localities variation in genital mor¬
phology is unusually great. For example, at Schillings Spring, scle-
rotization of the extensible organ in females ranges from complete as
in P. acmon texanus to incomplete as in P. acmon acmon and P. acmon
lutzi (Fig. 13). Localities in Colorado where pigmentational and/or
genitalic intermediates have been collected are shown in Map 2. A
“blend zone” following an altitudinal as well as a geographic gradient
is indicated by the data plotted. Whether or not this apparent zone of
intergradation and marked variation occurs in other areas between the
ranges of P. acmon texanus and P. acmon lutzi is unknown.
At the extreme western edge of the distribution of P. acmon texanus ,
a different situation may exist. Small series of P. acmon examined from
desert mountains of southeastern California include both P. acmon
texanus and P. acmon acmon but not intermediate phenotypes. Further
collecting at these and other localities at the margins of the distribution
of P. acmon texanus is needed to help clarify these situations.
Acknowledgments
Appreciation is expressed to the following persons and institutions
for the loan of specimens without which this study could not have been
undertaken: J. P. Donahue (Los Angeles County Museum of Natural
History), C. D. Ferris (Laramie, Wyoming), G. A. Gorelick (Azusa,
California), Chris Henne (Pearblossom, California), K. C. Hughes and
R. E. Stanford (Los Angeles, California), J. A. Powell (University of
California at Berkeley), C. A. Selcerman (Los Angeles, California), J. A.
Scott (formerly University of California at Berkeley), Oakley Shields
(University of California at Davis).
Literature Cited
Brown, K. S. 1965. Some comments on Arizona butterflies. J. Lepidopt. Soc.,
19: 107-115.
Goodpasture, C. E. 1973a. Biology and systematics of the Plebejus ( Icaricia )
acmon group. J. Kansas Entomol. Soc. (In press).
1973b. Foodplant specificity in the Plebejus ( Icaricia ) acmon group. J. Lepi¬
dopt. Soc. (In press).
160
The Pan-Pacific Entomologist
New Ephemerellidae from Madagascar and Afghanistan 1
(Ephemero p tera)
Richard K. Allen
California State University, Los Angeles, 90032
A collection of Ephemerellidae from W. Wittmer of the Naturhis-
torisches Museum, Basal, Switzerland, included an undescribed genus
and species represented by two female imagoes. A third female imago
was examined at the University of Utah, Salt Lake City, with the per¬
mission of G. F. Edmunds, Jr. The forewing characters of Manohyphella
n. gen. are similar to Teloganella Ulmer, Teloganodes Ulmer, and Ephem-
erellina Lestage, and it is herein included, with the above genera, in
the subfamily Teloganodinae. Two undescribed species of Ephemerella
Walsh were also included in a collection of nymphs from Afghanistan
which were loaned to the author by G. F. Edmunds, Jr. One belongs
in Drunella Needham and is the eighteenth species in the subgenus
known to occur in Asia. The other is a representative of Serratella
Edmunds and constitutes a new Asian record for the subgenus as the
taxon was previously reported from only North America.
Manohyphella Allen, new genus
Small mayflies with robust body and proportionately narrow wings. Forewings
with long connected or unconnected intercalaries. Intercalaries of forewing re¬
duced in number with only one between IMP and MP 2 , only one between MP 2
and CuA, and only three between CuA and CuP (Fig. 2). Hindwings propor¬
tionately small and costal projection sharp and at apex of wing (Fig. 3).
Type species. —Manohyphella keiseri Allen, new species
This is the third genus of Ephemerellidae to be reported from the
Ethiopean Region. Ephemerellina was described from South Africa,
and is now known from Australia and eastern China. Allen and Ed¬
munds (1963) reported a record of Ephemerella (Eurylophella ) Tiensuu
from Madagascar, but they considered it questionable and possibly
due to a labeling error. Manohyphella is distinguished from the other
described Teloganodinae by the character of the hindwing. The costal
projection of Ephemerellina is in the anterior half of the wing, the
projection is blunt and symmetrical, and there are usually between
fifteen and twenty crossveins in the wing (Fig. 5). In Teloganella and
1 The research on which this paper is based was supported by National Science Foundation Grant
GB-35591, and the University of Utah collections on which parts of this paper are based were obtained
with National Science Foundation grant support to G. F. Edmunds, Jr.
The Pan-Pacific Entomologist 49: 160-164. April 1973
Vol. 49, No. 2, April 1973
161
Fig. 1. Ephemerella ( Serratella ) subsolana Allen, n. sp., mature nymph, dorsal
view.
Teloganodes, the costal projection is blunt and asymmetrical, and there
are usually less than five crossveins in the wing (Fig. 4). The above
two genera are further characterized by well-developed prominent cross¬
veins arising from the posterior margin of the costal projection and
162
The Pan-Pacific Entomologist
Figs. 2-3. Manohyphella keiseri, wings. Fig. 2, forewing. Fig. 3, hindwing.
Fig. 4. Teloganella, hindwing. Fig. 5. Ephemerellina, hindwing. Figs. 6-8.
Ephemerella ( Drunella ) kabulensis, nymphal structures. Fig. 6, thorax and ab¬
domen, dorsal view. Fig. 7, head, front view. Fig. 8, head, side view. Figs. 9-10.
Ephemerella ( Serratella ) subsolana, nymphal structures. Fig. 9, maxilla. Fig.
10a, right fore leg. Fig. 10b, tarsal claw.
Vol. 49, No. 2, April 1973
163
extending to the subcosta and the radius (Fig. 4). The costal projection
of the hindwing of Manohyphella is in the posterior half of the wing,
the projection is sharp and symmetrical, and there are only two cross¬
veins in the wing. The crossveins originate near the posterior margin
of the costal projection, as in Teloganella and Teloganodes , but they are
poorly developed and barely discernible (Fig. 3).
Etymology.— Manohyphella is from the Greek words manos mean¬
ing rare and hyphe meaning web, and from the Latin diminutive ella.
Manohyphella keiseri Allen, new species
Female imago. —Length: body 5.0-6.0 mm; forewing 10.0 mm. General color red¬
dish brown to black. Head dark brown. Pronotum reddish brown; pronotum with
median elevated longitudinal ridge; mesonotum reddish brown with dark mark¬
ings along sutures; metonotum with prominent posterior submedian flap-like
structures; forewings hyaline, stigmatic area and base of wings brown (Fig. 2) ;
longitudinal veins black, hindwings hyaline, base of wings marked with brown;
longitudinal vein brown (Fig. 3) ; legs light brown. Abdomen reddish brown
without distinctive markings. Caudal filaments yellowish brown.
Holotype female imago , Ranomafana, Madagascar, 12 September
1958, F. Keiser, in collection Naturhistorisches Museum, Basal, Switz¬
erland. Paratopotype, female imago, same data and deposition as
holotype. Paratype, female imago, Farimbony Riv., at R. N. 2, Prov.
Tamatave, Madagascar, 15 October 1971, G. F. and C. A. Edmunds,
F. Emmanuel, in collection University of Utah, Salt Lake City.
Etymology. —Named in honor of F. Keiser, the collector of the
holotype.
Ephemerella (Drunella) kabulensis Allen, new species
Mature nymph.— Length: body 9.5-10.5 mm; caudal filaments 6.0-7.0 mm.
General color light brown. Head brown with dark transverse band across ante¬
rior margin of compound eyes; head with moderately developed paired occipital
tubercles (Fig. 7), with moderately developed median tubercle on frons between
compound eyes (Fig. 8), and with short lateral genal projections. Thorax light
brown; thoracic nota without tubercles, but often with scattered long setae; legs
brown; ventral (leading) edge of fore femora with tubercles; anterior margin
of fore femora with longitudinal ridge near middle of segment and median trans¬
verse ridge extending from longitudinal ridge to anterior margin of segment; apical
tibial projection straight and moderately developed; tarsal claws with 1 or 2 sub-
basal denticles. Abdominal trega brown; terga 2-10 with paired submedian tuber¬
cles; tubercles moderately developed on terga 2-9, small on tergum 10; terga 2-9
often with scattered long setae; tergum 1 with row of setae along posterior margin
and tergum 10 with median tuft of setae (Fig. 6) ; abdominal sterna light brown.
Caudal filaments light brown.
164
The Pan-Pacific Entomologist
Holotype mature nymph , Kabul River near Pagham, Kabul Prov¬
ince, Afghanistan, 26 August 1967, M. Nazim, in collection University
of Utah, Salt Lake City. Paratopotypes, 68 nymphs, same data as holo¬
type, 10 nymphs in collection California State University, Los Angeles,
and 5 each in collections of California Academy of Sciences, San Fran¬
cisco, and the Canadian National Collection, Ottawa; 13 nymphs, 25
May 1967, M. Nazim; 5 nymphs, 9 August 1967, G. Sharafi et al. ; 2
nymphs, 8 April 1967, M. Nazim; 7 nymphs, 29 June 1967, M. Nazim.
Paratypes, 14 nymphs, Kabul River nr. Dara Pagnan, Kabul Prov.,
Afghanistan, 8 April to 26 July 1967, M. Nazim; 28 nymphs, Kabul
River nr. Dara Pagnan, Kabul Prov., 7 July to 1 August 1967, G. Sharifi
et al.; 7 nymphs, Pang Shir River, 5 km. N. Charikar, Parwan Prov.,
Afghanistan, 9 July 1967; 5 nymphs, Ciakhak Maidan, Wardek Prov.,
Afghanistan, 15 July 1967, G. Sharifi et al. ; 5 nymphs, Ghalkaii, Gardez,
Paktia Prov., Afghanistan, 28 July 1967, G. Sharifi et al., all paratypes
except as noted in collection University of Utah.
Etymology. —The name is based upon the type locality, the Kabul
River.
Ephemerella (Serratella) subsolana Allen, new species
Mature nymph. —Length: body 6.0-7.0 mm; caudal filaments 4.5-5.S mm.
General color light brown. Head light brown with Y-shaped brown marking (Fig.
1) ; head without tubercles or projections; maxillary palpi moderately well-devel¬
oped and three-segmented (Fig. 9). Thoracic nota light brown with pale median
longitudinal stripe (Fig. 1) ; femora light brown, pale at apices; anterior surface
of fore legs with spines (Fig 10a) ; tibiae and tarsi pale; tarsal claws with 8-10
denticles (Fig. 10b). Abdomen brown; abdominal terga without tubercles, but
with paired submedian elevated ridges on terga 2-7; ridges covered with heavy
short spicules (Fig. 1) ; tergum 8 with scattered spicules, and terga 8-9 with
marginal spicules; abdominal sterna brown, darker posteriorly. Caudal filaments
brown basally, light apically.
Holotype mature nymph, Kabul River, Kotasungi, Kabul Prov¬
ince, Afghanistan, 13 May 1967, M. Nazim, in collection University
of Utah, Salt Lake City.
Etymology. —The name is from the Latin word subsolanus mean¬
ing eastern.
Literature Cited
Allen, R. K. and G. F. Edmunds, Jr. 1963. New and little known Ephemerel-
lidae from southern Asia, Africa, and Madagascar (Ephemeroptera).
Pac. Insects, 5: 11-22.
Vol. 49, No. 2, April 1973
165
A Reclescription of the Scorpion Vejovis flavus
(Scorpionida: Vejovidae)
Michael E. Soleglad
3927 Polack Street, San Diego, California 92110
In 1900, Nathan Banks listed the new scorpion Vejovis flavus Banks.
Unfortunately, Banks’ description was comprised of only a few state¬
ments embedded in a key. To the best of my knowledge these statements
are the only descriptive information on V. flavus. Evidently, Banks’
knowledge of V. flavus was based on a single female specimen from
Albuquerque, New Mexico.
In recent years Vejovis confusus Stahnke of Arizona and southern
Nevada has been confused with V. flavus. Due to this association with
V. confusus, V. flavus has been considered a member of the eusthenura
group of Vejovis. The findings of this study verify that V. flavus is a
member of the eusthenura group although displaying some interesting
deviations from the group.
I am grateful to Dr. Herbert W. Levi of the Museum of Comparative
Zoology, Harvard University, Cambridge, Massachusetts, for the loan
of the type specimen.
Vejovis flavus Banks
(Figs. 1-6; Table 1)
Diagnosis. —A unique member of the eusthenura group, somewhat
related to Vejovis confusus of Arizona and southern Nevada from which
it can be separated by the following characters; Inferior median keels
delicately crenulate on all caudal segments instead of smooth on segments
I-II; chelal carinae distinct, superior, inner accessory, and inner carinae
mildly crenulate as opposed to smooth; pectinal tooth count greater,
21-22 for female, in contrast to 13-17.
Holotype female. —Entire specimen dark orange, undoubtedly caused by pro¬
longed preservation. Eyes and aculeus dark brown to black. No other detectable
markings. Structure: Measurements as given in table 1. Carapace: Posterior
surface covered with coarse granules, ocular triangle smooth; longer than wide,
anterior region noticeably narrowed (Fig. 2) ; shorter than fifth caudal segment.
Anterior edge essentially straight; median tubercle situated somewhat anterior,
in ratio 3.1/8.6; median eyes and tubercle small, one-fifth width of carapace at
that point. Posterior lateral eye reduced considerably, anterior two approximately
same size. Preabdomen: All tergites granulated on extreme posterior edges;
seventh tergite with two pairs of serrate keels. Sternites smooth with slit-like
stigmata; sternite V lacking keels. Cauda: Basal segment wider than long. Seg-
The Pan-Pacific Entomologist 49: 165-170. April 1973
166 The Pan-Pacific Entomologist
Fig. 1. Vejovis jlavus Banks, holotype female, dorsal view.
ments I-IV: Dorsal and dorsal lateral keels serrate, ending in elongate posterior
spine. Lateral keels crenulate on segment I, crenulate on posterior third of II,
posterior quarter of III, and absent on IV. Inferior lateral keels crenulate; in¬
ferior median keels delicately crenulate on posterior two-thirds of segment I and
on entirety of II-IV. Segment V: Dorsal keels granulate; lateral keels granulate
on anterior half; inferior lateral keels serrate; inferior median keel crenulate to
serrate. Intercarinal spaces essentially agranular. Telson : Vesicle not as wide
or deep as fifth caudal segment width; ventral surface with mild granulation.
Very weak subaculear nodule at base of aculeus; aculeus short with average curve
(Fig. 3). Pectines : Well developed (Fig. 4) ; three times as long as widest point.
Pectinal tooth count 22/21 for female; middle lamellae 13. Basal piece separated
on anterior half, length/width ratio 1.8/2.6. Genital operculum completely fused.
Chelicerae : Lower edge of movable finger smooth with weak serrulae on distal
portion; lower distal tooth of movable finger noticeably longer than upper counter¬
part. Other dentition standard for Vejovinae. Pedipalps: Femoral carinae serrate;
inner face with 7-9 serrate granules. Dorsal carinae of tibia crenulate; inner
ventral carina crenulate to serrate, exterior ventral carina smooth; inner proximal
projection reduced, armed with 7-9 crenulate granules. All eight carinae of chela
developed; inner, inner accessory, and superior carinae subtly crenulate, other
carinae smooth. Movable finger shorter than either carapace or fifth caudal segment.
Teeth in single -straight row, little to no scalloping (Fig. 5). Supernumerary teeth
number six on fixed finger and seven on movable finger. Trichobothrial pattern
(Fig. 6) with counts standard for genus: 26 chela, 19 tibia, and 3 femur. Walking
legs : Single file of delicate spines flanked by longer setae on tarsus venter. Carinae
of patellae delicately crenulate.
Vol. 49, No. 2, April 1973
167
Fig. 2. Vejovis flavus Banks, holotype female, carapace.
Male. —U nknown.
Type data.— Holotype female from Albuquerque, New Mexico. Date
and. collector information are missing from the original specimen label.
Specimen is in remarkably good condition. Base color is dark, un-
Fig. 3. Vejovis flavus Banks, holotype female, telson, lateral view.
168
The Pan-Pacific Entomologist
Fig. 4. Vejovis flavus Banks, holotype female, pectines.
doubtedly caused by prolonged preservation. Subtle setation has been
essentially obliterated throughout the years.
Distribution.— Known from type locality only.
Comments. —Brief as Banks’ description was, I found a couple of
discrepancies in comparing it with the holotype female. In the key,
Banks states that V. flavus has smooth chelal palms. However, the holo¬
type is equipped with carinae on its chelae, some even crenulate. At
this point in the key Banks is contrasting V. flavus with Vejovis punc-
tipalpi (Wood) a species whose chelae are heavily crested, much more
so than V. flavus. Also, Banks’ states that the inferior median keels of
the first caudal segment are plain whereas on the holotype they are
weakly crenulate on the posterior two-thirds.
Although a member of the eusthenura group, V. flavus presents two
Fig. 5. Vejovis flavus Banks, holotype female, chela, lateral view.
Vol. 49, No. 2, April 1973
169
Fig. 6. Vejovis flavus Banks, holotype female, trichobothrial pattern of chela
(top row), tibia (middle row), and femur (bottom row). Exterior (left column),
dorsal (left middle column), ventral (right middle column), and interior (right
column) views. Letter abbreviations: E and e, exterior; D and d, dorsal; I and i,
interior; V, ventral; M, median; B and b, basal; t, terminal; st, subterminal;
sb, subbasal.
170
The Pan-Pacific Entomologist
Table 1. Measurements (in millimeters) of Vejovis flavus Banks.
Holotype
(female)
Total length
33.9
Carapace, length
5.0
Width at lateral eyes
2.7
Width at caudal edge
4.4
Preabdomen, length
12.1
Postabdomen, length
16.8
Caudal segment I (length/width/depth)
2.3/2.6/2.0
Caudal segment II (length/width/depth)
2.7/2.5/2.0
Caudal segment III (length/width/depth)
3.0/2.5/2.0
Caudal segment IV (length/width/depth)
3.5/2.4/2.0
Caudal segment V (length/width/depth)
5.3/2.2/1.9
Telson, length
4.7
Vesicle (length/width/depth)
2.7/1.6/1.3
Aculeus, length
2.0
Pedipalp, length
15.2
Femur (length/depth)
4.0/1.2
Tibia (length/depth)
4.1/1.6
Chela, length
7.1
Palm (length/width/depth)
2.9/1.9/1.6
Fixed finger, length
3.4
Movable finger, length
4.4
Pec tines
Teeth
22/21
Middle lamellae
13
interesting deviations from the definition given by Williams (1970).
The carinal structure of the chelae is not characteristic of the eusthenura
group which normally has reduced chelal palms, and the carinae, if
present, are usually underdeveloped and smooth. The pectinal tooth
count of the female is somewhat large for this group, exceeding all the
other known species, male as well as female. By projecting from a max¬
imum total length/pectinal tooth count ratio for the eusthenura group,
one may suspect that the female holotype is not fully matured.
Literature Cited
Banks, N. 1900. Synopses of North-American Invertebrates. Amer. Natur.,
34(401) : 421-427.
Williams, S. C. 1970. New Scorpions Belonging to the Eusthenura Group of
Vejovis from Baja California, Mexico (Scorpionida: Vejovidae). Proc.
Calif. Acad. Sci., 4th ser., 37(12) : 395-417.
Vol. 49, No. 2, April 1973
171
Unisexual Generation of Anclricus atrimentus
(Hymenoptera: Cynipidae)
D. Charles Dailey and Christine M. Sprenger
Sierra College, Rocklin, California 95677
The previously undescribed unisexual generation of Andricus at¬
rimentus Kinsey (1922), a cynipid gallmaker occurring on Quercus
douglasii Hooker & Arnott has formerly been considered identical to
Andricus kingi Bassett from Quercus lobata Nee, both Lepidobalanus
or white oaks. This was probably due to the similarity of their uni¬
sexual generation galls (Weld, 1957). Both are red, conical, detachable
leaf galls. Weld reared insects and supplied unisexual generation galls
of Andricus atrimentus to Houard (1946). Due to the coriaceous meso-
scutum of the unisexual generation, the species is hereby removed from
its present assignment in Dros Kinsey and returned to its original place¬
ment in Andricus Hartig. Kinsey’s description of the tarsal claws being
simple in the bisexual generation insects is erroneous. Toothed con¬
dition of the claws is consistent with the unisexual generation, its occur¬
rence on white oaks, and its placement in Andricus.
Female wasp. — Head: coriaceous; cheeks not widened behind eyes, slightly
narrower than thorax, occiput not concave; malar space 0.4 eye height, not grooved;
antennae filiform, segment one longer than two, segment three longer than four, 14
segmented with 13 = 14 or 13 with 13 twice as long as 12. Thorax: mesoscutum
polished coriaceous, smoother posteriorly, longer than broad, sparsely pubescent; an¬
terior parallel lines indistinct, less than one-half length of mesoscutum; notaulices
complete, polished, wider posteriorly; median groove distinct, only slightly longer
than width of notaulix at the scutellar suture; parapsidal lines indistinct. Scutellum
reticulate posteriorly, sparsely pubescent with smooth area behind foveae; foveae
smooth, polished, bare. Propodeum rugose. Wing surface and hind margin pubes¬
cent, aerolet Ho t° Vl2 length of first cubital cell, occasionally obsolete in plesio-
types; cubital and radial cells open, radial cell 3.5 times as long as high; Ri vein
arcuate. Tarsal claws toothed. Abdomen: slightly longer than head plus thorax,
slightly longer than high. Tergite II pubescent at base, II-VII micropunctate, more
than two tergites visible in dorsal view. Ventral spine nine times as long as wide,
six times as long as high, sparsely pubescent. Color; body uniform dark amber to
brown; wing veins yellow to yellow-brown; legs and antennae yellow-brown, latter
darker distally. Size: 2.0 mm. Range of 10 specimens 1.7 to 2.2 mm body length
(average of 10 specimens 2.0 mm). Total number of specimens 40.
Gall.— Monothalamous, conical, flat-based, detachable, on lower leaf margin.
Sides nearly straight, flaring slightly at base. Basal larval cell oval in side view,
1.6 mm in diameter, 0.8 mm high, pubescent. Upper surface of larval cell coriaceous
with sparse pubescence; brown, darker towards center. Red and yellow striped
The Pan-Pacific Entomologist 49: 171-173. April 1973
172
The Pan-Pacific Entomologist
Figs. 1, 2. Andricus atrimentus. 15 X Life Size. Fig. 1. Unisexual generation
gall. Fig. 2. Bisexual generation gall.
when fresh, turning to brown and yellow with age (Fig. 1). Size: Height 3.6 to
4.6 mm (average of 10 galls 3.9 mm) ; width 3.0 to 4.6 mm (average of 10 galls
3.3 mm). Total number of specimens 20.
Systematics. —The unisexual generation of Andricus atrimentus
can be distinguished from Andricus kingi by the following character¬
istics. Andricus atrimentus has the head not broadened behind the
eyes and slightly narrower than thorax, mesoscutum and scutellum
sparsely pubescent, galls red and yellow striped with lateral edges flat
or down curved. Andricus kingi has the head distinctly narrower than
thorax and head broadened behind eyes, mesoscutum and scutellum
pubescent, galls red with lateral edges recurved. Rearings of A. kingi
by Rosenthal and Koehler (1971) and the senior author (unpublished)
also confirm separate identity of the bisexual generations of these two
species.
Life history. —Five years of field observations by the senior author
on an isolated Quercus douglasii at Davis, Yolo County, California
have shown that the A. atrimentus bisexual generation females emerge
between 1 and 30 April over a two week period and usually oviposit
in the lower leaf surface near the margin. Within 60 days small tan
to pink blisters appear at oviposition sites. During the next 30 days
galls develop the final conical shape (Fig. 1). Pupation occurs during
late October to November and emergence of the unisexual generation
females occurs on warm days in late winter. They then oviposit in leaf
Vol. 49, No. 2, April 1973
173
buds and when the buds open the leaves already contain small blisters.
By mid March the integral leal galls of the bisexual generation usually
reach full size and the outer white layer develops its pink tinted tissue
paper texture. The blue-black larval cell is connected to the outer layer
by numerous blue-black filaments which become white distally (Fig. 2).
Host.— Quercus douglasii ; similar galls noted on Quercus dumosa
Nuttall and Quercus turbinella californica Tucker but not confirmed by
reared insects. Numerous attempts to rear the unisexual generation of
A. atrimentus on Quercus lobata have invariably been unsuccessful.
Distribution.— Apparently throughout the range of the host plants
from foothills of Coast Range and west slope of Sierras from Mendocino
County to Mohave Desert border.
Plesiotypes.- —American Museum of Natural History, New York;
United States National Museum, Washington, D. C.; California Acad¬
emy of Sciences, San Francisco; University of California at Davis;
Weld collection, in possession of Robert J. Lyon, Los Angeles City
College, Los Angeles, California.
Literature Cited
Houard, C. 1946. Les collections cecidologiques du laboratoire d’entomologie
du Museum d’Histoire Naturelle de Paris: Galls des Etats-Unis et du
Mexique. Ann. Entomol. Soc. France, 112: 25-26.
Kinsey, A. C. 1922. New Pacific Coast Cynipidae. Amer. Mus. Natur. Hist. Bull.,
46: 279-280.
Rosenthal, S. S. and C. S. Koehler. 1971. Heterogony in some gall-forming
Cynipidae with notes on the biology of Neuroterus saltatorius. Ann.
Entomol. Soc. Amer., 64(3) : 565-570.
Weld, L. H. 1957. Cynipid Galls of the Pacific Slope. Privately Printed, Ann
Arbor, Mich. 64 p.
RECENT LITERATURE
Animals of the Arctic. The Ecology of the Far North. Bernard Stonehouse.
Holt, Rinehart and Winston, Inc., New York. 172 p., over 200 color photos. 1971.
$10.95.
This book is a non technical treatment of animal ecology in the Arctic with
beautiful color illustrations, but with only brief references and illustrations of
insects.— Robbin W. Thorp, University of California, Davis, 95616.
The Caddis Flies, or Trichoptera, of Illinois. Herbert H. Ross. Entomological
Reprint Specialists, Los Angeles. 326 p., 961 figs., and preface to reprint edition
by author. Reprinted 1972. $9.95.
A reprint of Dr. Ross’ classic faunal study of Trichoptera. —Robbin W. Thorp,
University of California, Davis, 95616.
174
The Pan-Pacific Entomologist
A New Species of Diplolepis from California
(Hymenoptera: Cynipidae)
D. Charles Dailey and Linda Campbell
Sierra College, Rocklin, California 95677
A previously undescribed cynipid wasp from California, Diplolepis
inconspicuis n. sp., produces an inconspicuous fusiform twig or bud
gall on Rosa calif ornica Chambers and Schlecht.
Biology. —The original gall is an inconspicuous fusiform enlarge¬
ment of an apical or lateral stem or bud which yields a single adult
D. inconspicuis in February or March (Figs. 1 & 2). After emerging,
the female oviposits in a young stem or bud. If this gall is oviposited
in by a female inquiline Periclistus arefactus McCracken and Egbert
which emerge from old galls in the succeeding months, the new gall and
gallmaker fail to develop normally (Fig. 3). The resulting enlarged,
tan, woody to pithy, inquiline-modified gall bears lines which are aborted
leaf margins and/or abscission scars of aborted leaflets (Fig. 4). It
yields only inquilines and parasites. Because the inquiline-modified
galls of D. inconspicuis closely resemble the normal galls of D. arefacta
(Gillette) (1894), occurring from Colorado to Nevada, McCracken
and Egbert (1922) mistakenly assumed this California gall was pro¬
duced by D. arefacta and named the inquiline Periclistus arefactus.
Weld’s (1957) note on the inquiline-modified galls suggests the original
gall was a single celled stem swelling subsequently modified by the
inquiline. Our rearings have confirmed this.
Diplolepis inconspicuis Dailey and Campbell, new species
Holotype female. — Head : Antennae black, 14 segmented; malar space 0.5
times eye height. Thorax : Mesoscutum coriaceous, pubescent; median groove
smooth, one-third length of mesoscutum at scutellum as distinct and wide as
notaulicies; notaulicies complete, pitted, slightly incurved at scutellum; scutellum
entirely rugose, humped above extended curvature of mesoscutum, foveae ridged;
mesopleuron with dorsal and ventral polished areas occupying about one-half and
one-fourth of mesopleuron respectively, separated by mesolateral rugose area;
wings with clouded radial area, radial cell closed, areolet about one-sixth length
of cubital cell; cubital cell open proximally. Legs amber, tarsi brown, hind femur
tapering suddenly near distal end. Abdomen: Propodeum rugose, first abdominal
tergite polished coriaceous, sparsely pubescent medially; tergites II-VI progres¬
sively more coriaceous, pubescent posteriorly, abdomen equal to combined head
and thorax length; hypopygium extending beyond abdomen. Color: Head and
thorax black, abdomen red. Length: 3.9 mm.
The Pan-Pacific Entomologist 49: 174-176. April 1973
Vol. 49, No. 2, April 1973
175
Figs. 1-4. Galls of Diplolepis inconspicuis n. sp. Fig. 1. Unaborted dissected
stem gall. Fig. 2. Normal but broken stem gall and bud gall. Fig. 3. Immature
inquiline aborted bud gall. Fig. 4. Mature inquiline aborted gall which yields
Periclistus arefactus. All galls are shown twice life size.
Holotype female , and gall, Mt. Diablo, Contra Costa County, Cal¬
ifornia C. Dailey #1204, emerged 12 February 1972. Deposited in
U. S. National Museum.
Paratype females and galls, Folsom Lake, Placer County (22,6 galls),
Mt. Diablo, Contra Costa County, (10 2, 60 galls), Fairfield, Solano
County, (4 2,20 galls), Pleasant Valley, Solano County, (42,29 galls),
Marysville, Yuba County, (12, 115 galls), Sacramento, Sacramento
County, (72, 12 galls). All specimens from California. Variation:
Abdomen from 1.0 to 1.1 times combined head and thorax length.
Length: 3.3 to 4.6 mm. Average of 10 specimens 3.9 mm.
Sex ratio. —No males have been reared.
Paratype insects and galls are deposited in U. S. National Museum,
Washington, D. C., California Academy of Science, San Francisco,
California; Weld collection in possession of R. J. Lyon, Los Angeles,
California; University of California, Davis; University of Saskatchewan,
Saskatoon, Alberta, Canada; and the senior author’s collection.
The galls of D. inconspicuis and D. nodulosa (Beutenmuller) (1909)
are often indistinguishable though the latter are occasionally more
spiny. The polished dorsal mesopleural area and longer hypopygium
of D. inconspicuis females readily separate them from females of D.
nodulosa, which have a small smooth dorsal mesopleural area and
hypopygium not extending beyond the last abdominal segment.
176
The Pan-Pacific Entomologist
Acknowledgments
The authors are grateful to Mr. J. D. Shorthouse of the University
of Saskatchewan for the loan of D. nodulosa insects and galls and
Barbara Young for the illustrations.
Literature Cited
Beutenmuller, W. 1909. Descriptions of the new Cynipidae. Entomol. News,
20: 247.
Gillette, C. P. 1894. New Cynipidae. Can. Entomol., 26: 157.
McCracken, I. and D. Egbert. 1922. California gallmaking Cynipidae with
descriptions of new species. Stanford Univ. Biol. Sci., 3: 62.
Weld, L. H. 1947. Cynipid Galls of the Pacific Slope. Printed privately, Ann
Arbor, Mich., 64 p.
BOOK REVIEW
Biological Control. Edited by C. B. Huffalcer. Plenum Press, New York. 511 p.,
97 figs., 51 tables. 1971. $19.50.
This hook is a collection of papers presented by 33 contributors to a symposium
of the Ecological Society of America and the Association for the Advancement of
Science in December 1969. Each of the 20 papers is treated as a chapter with its
own bibliography. These chapters are arranged in four sections. The book termi¬
nates with a complete index to all authors cited in the text of each paper including
secondary authors, and a complete subject index.
The first five chapters (Section 1) deal with the theory and evaluation of bio¬
logical control including: the “Pesticide Syndrome” and its remedy through in¬
tegrated control; the role of natural enemies in natural control; the adaptability
of introduced parasitoids; models, life tables and experimental techniques for
evaluating natural enemies.
Chapters 6-10 (Section 2) discuss recent successful examples of biological
control of weeds, coccids, greenhouse pests, and the winter moth in Canada. The
emphasis is on recent examples so some of the better known examples of weed
and pest insect control (e.g. Opuntia in Australia, Klamath weed and cottony
cushion scale in California) are omitted.
Chapters 11-13 (Section 3) treat examples of naturally-occurring biological
control in America north of Mexico, in natural and agricultural ecosystems. The
recent appreciation of naturally-occurring biological control is an important de¬
velopment in pest management thinking.
Section 4 (Chapters 14-20) considers the role of biological control in a systems
approach to pest control. The majority of these chapters discuss the development
of various integrated control programs.
This book is a well balanced treatment of documented examples and the theory
of biological control with an emphasis on the role of biological control in inte¬
grated control programs. Since integrated control had its inception in California,
it is appropriate that about half of the authors are associated with campuses of
the University of California. The book is an outstanding reference on the theory
and practice of applied ecology in pest management.— Robbin W. Thorp, University
of California, Davis, 95616.
Vol. 49, No. 2, April 1973
177
Thomas Wrentmore Cook (1884—1962)
Paul H. Arnaud, Jr. and Mary Manning Wale
California Academy of Sciences, San Francisco, 94118, and 5933 Camden Street,
Oakland, California 94605
Thomas Wrentmore Cook was born on 6 December 1884 in Sydney,
Australia, and died in Oakland, California on 17 February 1962. He
was the eldest of the four children of George Wrentmore Cook and
Sarah Helsby Cook. His paternal grandfather, Edward Wrentmore Cook,
born in Somersetshire, emigrated to Australia in the 1850’s and even¬
tually built up a large bakery business in Sydney. Dr. Cook’s maternal
grandmother, nee Jane Hutton, came from Cambridge, England. She
was the daughter of a cabinet maker who responded to the British
government’s efforts to bring skilled craftsmen to Australia. The
family then made the six months’ voyage around the Cape of Good
Hope. In Sydney Jane met Thomas Helsby, a native of Lancashire
and they were married. The young bridegroom followed the trade of
stone mason; ultimately he became a building contractor.
Thus, at the time of his birth Dr. Cook was a second generation
Australian, something relatively rare in the year 1884. He attended
Sydney Technical High School where he studied drafting, but at the
insistence of his Grandfather Helsby (who had little use for hook learn¬
ing) began and completed a five-year apprenticeship as a machinist and
instrument maker. This was a bitter blow to the sixteen year old hoy
who already had intellectual aspirations. His interest in natural history
was also very strong and had been so since earliest childhood. A period
spent in Brisbane when he was between twelve and fourteen years of
age was especially influential in this respect. The sub-tropical environ¬
ment, the proximity to the Great Barrier Reef, the beautiful and exotic
birds—sulphur-crested cockatoos, budgerigars, and such—left an im¬
print vividly felt throughout his life.
After completing the above-mentioned apprenticeship Dr. Cook
worked for six months in the Fiji Islands, helping to construct a sugar
mill. Again, his surroundings were of intense interest. The East Indians
who made up the work force, the native Fijians, the tropical environ¬
ment all stimulated his curiosity. He returned briefly to Sydney, then
once more set out to see the world after the fashion of so many adven¬
turous young men. Hearing that there was a great demand for machinists
in San Francisco, then rebuilding after the earthquake of 1906, he took
The Pan-Pacific Entomologist 49: 177-181. April 1973
178
The Pan-Pacific Entomologist
Vol. 49, No. 2, April 1973
179
passage for that city, arriving 18 April 1907. Twenty dollars (hidden
in a money belt), the clothes he was wearing, and a book on shells were
his sole possessions. Everything else had been stolen aboard ship.
As things turned out Dr. Cook ended up working in Salinas at the
Spreckels Sugar Mill instead of in San Francisco, simply because he
heard of immediate job openings there. It was in Salinas that he met
and married Mary Jensen. This marriage ended in divorce in 1922.
He acquired U. S. citizenship in 1912 and during the First World
War was at the Naval Ship Yard, Vallejo, California. Following this
Dr. Cook was employed as an instrument maker by the University of
California, Berkeley, working for various scientists, but particularly
for the Engineering Department. He had unusual manual dexterity and
from rough two-dimensional sketches could visualize objects in three
dimensions.
At this point a member of his family offered to finance his way
through dental school. Dr. Cook was not especially keen about dentistry,
but saw this as a means of accomplishing his long held ambition of
attending college. For some time he had been taking night classes in
order to prepare himself for the entrance examinations to the University.
He passed the matriculation examination and in 1922 received his
D. D. S. degree from the University of California, winning membership
in the dental honor society. He was given a post as dental surgeon at
the University’s Cowell Hospital in Berkeley; at the same time he opened
a private practice in Berkeley. He also attended classes in anthropology
at the University. This was a subject which had long interested him,
probably having its inception in contacts as a youth with Australian
aborigines. The unusual energy and drive possessed by Dr. Cook are
shown in his ability to carry on all of these activities simultaneously.
He received a B. A. in 1925 and an M. S. in 1926, degrees in physical
anthropology. His Master’s thesis, based upon observation of Univer¬
sity students, was entitled “Dental Arches; a Study in Dento-Physical
Anthropology.”
Eater, Dr. Cook also began taking courses in entomology with Pro¬
fessor E. 0. Essig at the University of California, Berkeley. In the
early 1930’s he moved to Davis where he continued to follow the prac¬
tice of dentistry. He enrolled at this branch of the University of Cal¬
ifornia, completing his B. S. degree in entomology in 1932 under the
direction of Professor W. B. Herms. He then went to Harvard Univer¬
sity, commencing graduate studies involving the ants of California,
under Dr. William Morton Wheeler. This period at Harvard was dis-
180
The Pan-Pacific Entomologist
appointing because he did not receive the kind of guidance he had
expected.
In the spring of 1933 he moved to Washington, D. C. and began
working on a problem in physical anthropology at the Smithsonian
Institution under the general guidance of Gerrit S. Miller. The fruit
of this labor appeared in an article, “The Human Chin and Human
Tooth Change” (1933, Int. J. Orthodont. Dent. Child., 19: 730-734).
In Washington Dr. Cook met Mary Barbara Manning, then on the
staff of the Library of Congress. They were married on 19 April 1933
and began a journey to Australia, visiting England—-the land of his
ancestors, and France. In Marseilles they boarded a ship for the six
weeks’ voyage to Sydney. There were brief stops at Port Said, Colombo,
and Melbourne, where they visited the Museum and viewed the natural
history collection.
Plans for settling in Sydney where Dr. Cook’s mother and brother
lived were changed because of the depressed economic conditions in
Australia and the length of time it would have taken to obtain a dental
license. In the fall of 1933 he returned to Berkeley and resumed his
dental practice. At this time, following his insatiable desire for knowl¬
edge he began taking courses at the Pacific School of Religion. He
had long been interested in William Frederick Bade, the renowned
authority on the Old Testament, and was eager to attend his lectures.
Dr. Cook also took courses with Theodore McCown and James Muilen-
berg and finally completed the requirements for the degree of Bachelor
of Theology in 1939. The title of his thesis was “A Study of Some of
the Universal Religious Tendencies in Pre-Literate Man; a Problem
in Anthropology.” The thesis included the case history of Crashing
Thunder, a young Indian boy, and his initiation into the Peyote cult,
antedating by several years the current interest in psychedelic drugs.
Because of the Depression dentistry was bringing in just a bare living
and America’s defense efforts seemed to hold more possibility for finan¬
cial security. Accordingly Dr. Cook took a civil service position as
instrument maker with the U. S. Government. In August 1941 he sailed
for Hawaii, with the idea of having his family follow at a later date,
and was assigned duty at Pearl Harbor Navy Yard. He was working
there on the fateful day of December 7th but escaped injury except
for some fragments of metal striking his eye. Civilian workers were
frozen to their jobs and it was two years before be could obtain a re¬
lease to return to the mainland and rejoin his family. Meanwhile he
was transferred from the Navy to the Corps of Army Engineers which
had its headquarters at Punahou School. He greatly enjoyed the chal-
Vol. 49, No. 2, April 1973
181
lenge to his ingenuity posed by the task of improvising all sorts of
equipment which could not be obtained from the mainland.
Late in 1943 after Dr. Cook’s return to the Bay Area he recommenced
the practice of dentistry with great success. By the time he had reached
the age of seventy he was semi-retired but carried on a limited practice
at his residence. In March 1961 he suffered a mild stroke, the harbinger
of the massive strokes causing his death on 17 February 1962. Until
three weeks before his decease he had a clear, active mind and was still
interested in his many fields of knowledge.
It was about 1948 that Dr. Cook decided to put into book form the
data he had gathered on California ants. The better part of five years
was spent completing his volume entitled The Ants of California. Pri¬
vately published in 1953, it consists of 462 pages and 94 illustrations.
The drawings for the illustrations were made by Marian A. Kendall.
During the last fourteen years of his life Dr. Cook concentrated on
assembling a highly specialized library on the ants of the world. He
had bound in 51 volumes a virtually complete set of the publications
of William Morton Wheeler. His collection of books on entomology
as indicated in a note by Arnaud (1972, Pan-Pac. Entomol., 48: 63)
was donated to the California Academy of Sciences by his widow.
Dr. Cook’s collection of 4,397 pinned specimens of ants, 213 vials of
ants (uncounted as to number) and 27 Schmitt boxes of miscellaneous
insects was donated to The Oakland Museum in 1964. The ants were
transferred to the Los Angeles County Museum of Natural History, Los
Angeles in 1971.
Dr. Cook is survived by his widow Mary Manning Cook (now Mrs.
George Wale), and two children from this marriage, Thomas Manning
Cook of Berkeley and Barbara Wrentmore Cook (Mrs. Gilbert Barnes)
of Indianapolis. From his first marriage he is survived by two sons,
Donald Wrentmore Cook of San Marino, California, and Ralph A.
Cook of Salinas, California. He is also survived by five grandchildren
and two great grandchildren.
Thus ended the career of a versatile, many-sided man who was above
all a naturalist.
182
The Pan-Pacific Entomologist
SCIENTIFIC NOTES
A Biological Note on Two Species of Ageniella from California
(Hymenoptera:Pompilidae).- —Very little information is available on any
aspect of behavior for members of the nominate subgenus of Ageniella. The prey
records and fragmentary nesting information given here for two western species
are thus of considerable interest. On 24 October 1966, Wasbauer found a female
Ageniella blaisdelli (Fox) on a small, isolated bank of bare, consolidated sand on
the Sacramento River levee near Sacramento, California. The bank had a nearly
vertical face toward the river, about fifteen feet from the water. It was surrounded
by cottonwood trees ( Populus fremontii) , short grass and a stand of California
mugwort ( Artemisia vulgaris var. heterophylla) . When first seen at about 1:00
p.m., Pacific Daylight Time, the wasp was on the top of the bank, moving rapidly
with characteristic wing twitching and investigating burrow entrances presumably
of other Hymenoptera. It ran under a fallen cottonwood leaf at the base of the
bank and proceeded immediately to a burrow entrance near the top of the vertical
face. It entered tire burrow, remained ten minutes and then repeated the pro¬
cedure, remaining in the burrow twenty minutes. As it left, a female Astata
occidentalis Cresson (Det. R. M. Bohart) entered the burrow, without prey, re¬
mained only a matter of seconds and flew off. In the meantime, the Ageniella
female had returned to the cottonwood leaf and pulled a paralyzed spider from
under it. It alternately ran and flew in short hops directly to the burrow entrance,
four feet away. Its progress was so rapid that the manner of prey transport could
not be ascertained. It deposited the spider just outside the entrance, entered, re¬
appeared head first and pulled the spider in by the end of the abdomen. It had
been inside the burrow about five minutes when the Astata returned, again without
prey. Both wasps were collected shortly thereafter, as they left the burrow. On
excavation, the burrow which was 1 cm in diameter, was found to enter the soil
at a 45° angle. It was straight for 15 cm and was not traced further. At 1.3 cm
in from the entrance was a horizontal lateral burrow which extended 2.5 cm and
terminated in a slightly enlarged cell, 1.2 cm long. The cell had not been closed.
The spider found in the cell was an immature Pardosa sp. (Lycosidae) (Det. R.
Leech) with all of its legs amputated at the coxae, except the right anterior which
was severed near the apex of the tibia. A wasp egg was not found.
Another female of Ageniella blasdelli was found by Leech at Stove Pipe Camp¬
ground, 6100 feet, in the Chowchilla Mountains, Mariposa County, California, on
6 August 1971. It was dragging a spider across a dusty Forest Service road in
mid-afternoon in full sunlight. The spider, in this case, was an immature female
lycosid, Tarentula sp. prob. kochi (Keyserling) (Det. R. Leech). All eight legs
had been amputated at the coxae.
There is a female of Ageniella (A.) coronata Banks in the collection of the
California Academy of Sciences, pinned with a female clubionid spider, probably
Liocranoides sp. (Det. R. Leech). All the legs of this spider had been amputated.
The label reads only “Mendocino Co. Cal. VII-20-23. E. R. Leach.” Almost all
insects similarly labeled by the late Edwin R. Leach were actually taken at the
junction of Yale Creek with Rancheria Creek, about two miles south of the old
site of Yorkville Post Office on Route 28 (i.e., some 14 airlines miles northwest
of Cloverdale).
Evans [1969, Entomol. Soc. Amer. Misc. Publ., 3: 107-108] has summarized
Vol. 49, No. 2, April 1973
183
the published biological information on Nearctic species of Ageniella. He states
that two members of the subgenus Ageniella , A. accepta (Cresson) and A. conflicta
Banks, both closely related to A. blaisdelli, seek out natural crevices in the soil
and that A. accepta is reported to dig a short burrow from a crevice. Thus it is
likely that all three species in this group, and possibly all Nearctic species in the
subgenus Ageniella , normally nest in such situations.— Marius S. Wasbauer,
California Department of Agriculture , Sacramento, 95814; and Hugh B. Leech,
California Academy of Sciences, San Francisco, 94118.
Recent Annoyance to Man in Utah by Triatoma protracta (Hemiptera:
Reduviidae)In the southwestern United States conenose bugs, Triatoma and
Paratriatoma, feed predominantly upon the blood of wild mammals, but will also
feed upon human blood. Allergic reactions to their feeding vary from barely
perceptible local disturbances at the site to severe systemic reactions (James and
Harwood, 1969, Herm’s Medical Entomology, 6th Ed. 131-132). Their role as
vectors of sylvatic Trypanosoma cruzi Chagas in mammals is well established but
their ability to transfer it to< man is questionable. Naturally infected Triatoma p.
protracta (Uhler) have been reported in Utah from Kane County (Wood, 1956,
Bull. S. Calif. Acad. Sci., 55: 180) and Wayne County (Ryckman, 1962, Univ. Calif.
Publ. Entomol., 27: 115).
In 1965 a 60 year old man living near Springdale, Washington County, con¬
tacted me for information concerning treatment for his severe hypersensitivity to
bites of Triatoma. Local and systemic reactions to the salivary injections of the
triatomes necessitated repeated treatments by resident physicians. Since the orig¬
inal inquiry, he has reported at least 20 bite reactions of varying severity 7, 3, 1,
1, 5, and 3 times per year successively from 1966 through 1971. These occurred
during the summer periods of active dispersal of the insects. His tabulation of
bite sites indicates this subspecies feeds on any exposed area of the body. The wife
either was not fed upon by Triatoma or did not react to the bites.
The home, situated on a hill, at an elevation of 1,219 m, produced 19 males, 29
females and 2 fifth instar nymphs of T. p. protracta during the six year period.
The nymphs indicate a nearby colonizing site. This was probably a rock wall in
front of the house since the home was on an intact concrete slab and there was no
opportunity for triatomes to come up through the floor as at the San Joaquin Ex¬
perimental Range in California (Wood, 1951, Bull. S. Calif. Acad. Sci., 50: 106).
The yearly totals for 1966 through 1971 were 15, 7, 12, 1, 9, and 6 with monthly
totals for May through October of 3, 8, 15, 14, 9, and 1. The months of greatest
annoyance were July and August. For 1966 through 1968, two cats may have been
attractants since the larger number of bugs (34) was collected inside the home
during this time. From 1969 through 1971 without cats, the same person collected
16 triatomes inside the home. From 1966 through 1968, the number of bite re¬
actions was 11 and from 1969 through 1971, 9. Most bugs fed at night. One fed by
day on 2 August 1966 resulting in “3 bites on the back” while resting on the
sofa in the living room.
Locations of capture for 43 of 50 triatomes received revealed a 30:8:5 ratio for
bedroom (10 d, 18$, 2 fifth instar nymphs), living room (4$, 4$), kitchen
(2$, 3$ ) complex as compared with a 14:2:3 ratio for 40 T. p. protracta at the
SJER. Time of capture included 3 at dawn, 17 in daylight, 2 at dusk and 20 at
night.
184
The Pan-Pacific Entomologist
The recta of 48 triatomes (21 $, 25 2, 2 fifth instar nymphs) were removed by
me and crushed in sodium citrate solution for examination for trypanosomes. Forty-
five samples were negative and three were positive. Chagas’ trypanosome was
found in the feces of 2 $ collected 8 and 23 July 1970 and 1 $ 30 September 1971
when examined 4 to 6 days after capture. Since all trypanoform (trypomastigote)
and crithidiform (epimastigote) parasites appeared dead, no attempt was made
to infect laboratory mice. Springdale, therefore, constitutes a new locality for
occurrence of Chagas’ zoonosis in Utah. Of a total of 110 Triatoma p. protracta
examined by me from various localities in Utah to 1972, 5 or 4.5% were infected.—
Sherwin F. Wood, Los Angeles City College, Los Angeles, California 90029.
BOOK REVIEW
Insect Vision. G. A. Mazolchin-Porshnyakov. Translated from the Russian by
R. and L. Masironi. Translation edited by T. H. Goldsmith. Plenum Press, New
York. 306 p., 125 figs., 29 tables. 1969. $22.50.
This translation is an important contribution to our knowledge of insect vision,
especially in its treatment of the author’s own works and other Russian literature
not readily available to English speaking scientists. Originally published in 1958,
the author revised and updated the book for the English translation.
The book starts with considerations of the structure and neurology of compound
eyes, then discusses the theories of mosaic versus diffraction image formation. Other
chapters treat the photomechanical phenomena, electrical responses, resolving
power, and perception of polarized light and color by compound eyes, the struc¬
ture and function of ocelli, light orientation, and ultraviolet perception.
This monograph presents a fascinating and thought provoking coverage of visual
physiology in insects and will remain a standard reference in this field.— Robbin
W. Thorp, University of California, Davis, 95616.
EDITORIAL NOTICE
The editorship of the Pan-Pacific Entomologist will change hands beginning
with the July 1973 issue (Vol. 49, No. 3). New manuscripts should be submitted
to Dr. John Doyen, Division of Entomology, University of California, Berkeley,
California 94720.
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ENTOMOLOGICAL EXCHANGES AND SALES
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for $1.50 per line. Ads will be arranged in order of their receipt. Contact the
Advertising Manager, W. H. Lange, Department of Entomology, University of Cali¬
fornia, Davis, California 95616.
FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca,
Department of Biology, University of Arizona, Tucson, 85721.
FOR SALE. Over 900 species of butterflies from the Malaysian and Indonesian
Jungles, very rare moth (14" wing span), Scorpion, giant sized Indonesian flying
lizard, centipedes, stick insects, all types of rhinoceros beetles and rare beetles
in the families Scarabaeidae, Lucanidae, Mordellidae, Cerambycidae, etc. Write
Sim Yam Seng, 21, Joon Hiang Road, Singapore, 19, Singapore.
THE PAN-PACIFIC ENTOMOLOGIST
Information for Contributors
Papers on the systematic and biological phases of entomology are favored, including
articles up to ten printed pages on insect taxonomy, morphology, behavior, life history,
and distribution. Excess pagination must be approved and may be charged to the author.
Papers are published in approximately the order that they are received. Immediate publi¬
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a printed page may be published as space is available, in Scientific Notes.
Manuscripts for publication, proof, and all editorial matters should be addressed to the
Editor, Robbin W. Thorp, Department of Entomology, University of California, Davis,
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Essig, E. 0. 1926. A butterfly migration. Pan-Pac. Entomol.,
2 : 211 - 212 .
1958. Insects and mites of western North America. Rev.
ed. The Macmillan Co., New York. 1050 pp.
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Vol. 49
JULY 1973
No. 3
THE
Pan-Pacific Entomologist
DOUTT—The genus Paranaphoidea Girault (Hymenoptera: Mymaridae) .... 185
DAILEY AND SPRENGER—Synonymy of Andricus gigas and the bisexual
generation of Andricus crenatus (Hymenoptera: Cynipidae) _ 188
DOUTT—The genus Paratrichogramma Girault (Hymenoptera: Trichogram-
matidae) _ 192
SLOBODCHIKOFF—Behavioral studies of three morphotypes of Therion
circumflexum (Hymenoptera: Ichneumonidae) _ 197
WHEELER AND WHEELER—The ant larvae of the tribes Basicerotini and
Dacetini: second supplement (Hymenoptera: Formicidae: Myrmi-
cinae) _ 207
DOUTT—The genus Polynemoidea Girault (Hymenoptera: Mymaridae) .... 215
DOUTT—The fossil Mymaridae (Hymenoptera: Chalcidoidea) _ 2211
GREGARICK, SCHUSTER AND TOFTNER —Macrobiotus montanus from
California (Tardigrada: Macrobiotidae) _ 229
GRISSELL—New species of North American Torymidae (Hymenoptera) _ 232
BROWN AND ALLEN—A new Comadia from the San Joaquin Valley of
California (Lepidoptera: Cossidae) _ 240
BENEDETTI—Notes on the biology of Neomachilis halophila on a Cal¬
ifornia sandy beach (Thysanura: Machilidae) _ 246
ADAMS—Coniopterygidae from Chile, with three new species (Neuroptera) 250
CHANDLER AND NUTTING—A new species of Hamotus from galleries
of Microcerotermes septentrionalis in western Mexico (Coleoptera:
Pselaphidae) (Isoptera: Termitidae) _ 255
GREGARICK, SCHUSTER AND TOFTNER—Descriptive morphology of
eggs of some species in the Macrobiotus hufelandii group (Tardigrada:
Macrobiotidae) ...._ 258
MILLER—Mealybugs of San Miguel Island, California (Homoptera: Coc-
coidea: Pseudococcidae) _ 264'
GARDNER—A new milliped of the genus Tynomma from California (Lysi-
opetalidea: Lysiopetalidae) _ 270
AYALA—Two new subspecies of the Drosophila willistoni group (Diptera:
Drosophilidae) _ 273
SCIENTIFIC NOTES _ 196, 257, 279
BOOK REVIEWS ___ 228, 239, 280-283
BOOK NOTICE ____ 245
RECENT LITERATURE _ 191, 206, 220, 249
SAN FRANCISCO, CALIFORNIA • 1973
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
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Title of Publication: The Pan-Pacific Entomologist.
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This issue mailed 19 October 1973
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ALLEN PRESS, INC. »*7n T *° LAWRENCE, KANSAS
The Pan-Pacific Entomologist
Vol. 49
July 1973
No. 3
The Genus Paranaphoidea Girault
(Hymenoptera: Mymaridae)
Richard L. Doutt 1
University of California, Berkeley
Girault twice described Paranaphoidea as a new genus; first in 1913,
and then curiously without explanation again in 1914. Students of the
Mymaridae have been misled by his descriptions, for this resulted in
the wrong placement of the genus in the subfamily Mymarinae by An-
necke and Doutt (1961). The two subfamilies of Mymaridae are dif¬
ferentiated fundamentally by the manner in which the abdomen and
thorax are joined. In the Alaptinae the mesophragma projects strongly
into the abdomen which is sessile and broadly joined to the thorax.
It is otherwise in the Mymarinae to which Paranaphoidea presumably
belonged on the basis of Girault’s statements such as “phragma appar¬
ently absent,” “like Anaphoidea ,” and “This genus, for the present, I
consider allied with Anaphoidea ” (Girault, 1914). His choice of the
name Paranaphoidea, of course, further suggested that the two genera
were closely related.
For years no one saw Girault’s specimens, he published no illustra¬
tions, and his word descriptions were obviously inadequate. Apparently
this caused Ogloblin (1935) also to misinterpret the genus when he
described Paranaphoidea silvana and P. clavata. These two species seem
better placed in Patasson, which is now the proper name for Anaphoidea.
Through the courtesy of E. C. Dahms, Curator of Entomology at
the Queensland Museum, I have had the opportunity to study there
the Girault types. Paranaphoidea clearly belongs in the subfamily
Alaptinae. The mesophragma is well developed and strongly projects
into the abdomen which is sessile and broadly joined to the thorax.
The tarsi are 4-segmented so the genus is in the tribe Anagrini accord¬
ing to the classification proposed by Annecke and Doutt (1961). The
venation is much like that of Stethynium which is a member of the
same tribe. The posterior wings are very broad for the family, nearly
1 Present address: 9240 S. Riverbend Avenue, Parlier, Ca. 93648.
The Pan-Pacific Entomologist 49: 185-187. July 1973
186
The Pan-Pacific Entomologist
Fig. 1. Wings of Paranaphoidea densiciliata Girault.
like those in the Eulophidae yet distinctly pedicellate at base (Fig. 1).
Contrary to Girault’s description the strigil is present. The ovipositor is
strongly developed, and in some species the valvifers extend far forward
beneath the thorax in a membranous pouch. The antennae are inserted
close to the inner margins of the compound eyes about the middle of
the face. The scape and pedicel of the female antennae are compar¬
atively short; in some species the scape is serrated but smooth in others;
the funicle is 6-segmented; the club 2-segmented. The forewings are
somewhat truncate apically, the marginal fringes short, the discal cilia
entirely lacking beneath the sinuate venation.
The most extraordinary species is P. elongata which Girault (1923)
in a very curious, privately published paper described as “the largest
known Mymarid.” It does possess an exceptionally elongate abdomen
and a tremendously long ovipositor for a Mymarid, and all of its char¬
acters such as tarsomeres, flagellomeres and even its wing venation seem
abnormally elongated.
The type species is P. egregia Girault from which Girault seems to
have separated the species P. caudata and P. intermedia on little more
than color differences. In an unpublished manuscript Girault indicated
that Anaphoidea harveyi Girault belongs to this genus.
The genus is presently constituted as follows, although the proper
Vol. 49, No. 3, July 1973
187
status of P. caudata and P. intermedia deserves further study and the
correct inclusion of P. harveyi and P. nigriclava needs confirmation.
Paranaphoidea Girault
Paranaphoidea Girault, 1913. Mem. Queensland Mus., 2: 115.
Paranaphoidea Girault, 1914. Proc. Entomol. Soc. Wash., 16(2) : 112.
Type. — Paranaphoidea egregia Girault. Orig. desig.
auripes Girault, 1925. Some gem-like or marvellous inhabitants of the woodlands
heretofore unknown and by most never seen nor dreamt of. Privately publ.,
3 pp., p. 2. (Type: Mulgowie, Queensland).
caudata Girault, 1915. Mem. Queensland Mus., 3: 161. (Type: Cooktown,
Queensland).
densiciliata Girault, 1938. Rev. Entomol., 9: 391. (Type: Canterbury, Victoria).
densiciliata nostra Girault, 1938. Rev. Entomol., 9: 391. (Type: Chinchilla,
Australia).
egregia Girault, 1913. Mem. Queensland Mus., 2: 115. (Type: Nelson, Queensland).
elongata Girault, 1923. Loves Wooed and Won in Australia. Privately publ. 3 pp.,
p. 2. (Type: Wynnum, Australia).
eucalypti Girault, 1925. Insec. Insc. Menstr. 13: 94 (Type: Banyo, Queensland).
harveyi (Girault).
Anaphoidea harveyi Girault, 1912. Mem. Queensland Mus., 1: 151-152. (Type:
Nelson, Queensland).
intermedia Girault, 1915. Mem. Queensland Mus., 3: 161. (Type: Gordonvale,
Queensland).
nigriclava Girault, 1915. Mem. Queensland Mus., 3: 179. (Type: Burnett Heads,
Queensland).
ponderosa Girault, 1915. Mem. Queensland Mus., 2: 115-116. (Type: Nelson,
Queensland).
tennysoni Girault, 1920. Insec. Inscit. Menstr. 8: 98. (Type: Cairns, Queensland).
Literature Cited
Annecke, D. P. and R. L. Doutt. 1961. The Genera of the Mymaridae. Hy-
menoptera: Chalcidoidea. S. Afr. Dep. Agr. Tech. Serv., Entomol.
Mem., 5: 1-71.
Girault, A. A. 1913. Australian Hymenoptera Chalcidoidea—II. Supplement.
Mem. Queensland Mus., 2: 107-129.
1914. Descriptions of new chalcid-flies. Proc. Entomol. Soc. Wash., 16(2):
109-119.
1923. Loves Wooed and Won in Australia. Privately publ., Brisbane. 3 pp.
Ogloblin, A. A. 1935. Dos especies nuevas del genero Paranaphoidea Gir. (Hym.
Mymaridae). Rev. Entomol. (Rio de Janeiro) 5(2): 149-153.
188
The Pan-Pacific Entomologist
Synonymy of Andricus gigas and the Bisexual Generation of
Andricus crenatus
(Hymenoptera: Cynipidae)
D. Charles Dailey and Christine M. Sprenger
Sierra College, Rocklin, California 95677
Field observations and rearings have shown the bisexual generation
of Andricus crenatus Weld (1952) is Andricus gigas Kinsey (1922).
Because Kinsey erroneously described a toothed tarsal condition, Weld
(1951) transferred A. gigas to Dryocosmus Giraud. Due to the actual
toothed tarsal claws of both generations, the species is now returned to
the original placement in Andricus Hartig. The valid name again
becomes Andricus gigas.
Biology.- —The senior author made collections (November 1969)
of A. crenatus from a Quercus douglasii Hooker & Arnott in Rumsey
Canyon, Colusa County, California. The galls numbered 10 to 20 per
leaf and comprised about 90% of the galls on the tree (Fig. 2). The
unisexual generation females emerged in February and were bagged
on a Q. douglasii in Davis, California for controlled rearing.
The tan, conical, bisexual generation galls produced occurred on the
staminate flowers and occasionally in leaves (Fig. 1). Those at the
base of staminate flowers were often hidden by bud scales. They were
collected 14 April 1969 and yielded insects through 17 April 1969 while
being maintained indoors. Galls were also collected from the original
tree by the senior author on 16 April 1969. The ecology and description
of the galls suggested that the bisexual generation of A. crenatus was
A. gigas.
Systematics. —The insects from the April 1969 rearings matched
the original description of A. gigas with the exception of having a
toothed rather than simple tarsal claw. Two circumstances suggested
A. gigas Kinsey might in reality have a toothed tarsal claw and thus
not belong in Dryocosmus.
First, the occurrence of a bisexual Dryocosmus on a California white
oak is unusual for the following reasons. Nearly all American Dryocos¬
mus occur on oaks of the subgenus Erythrobalanus (red oaks), but
A. gigas was recorded from a Lepidobalanus (white) oak. No other
described species of Dryocosmus from California occur on the white
oaks. Since the name A. gigas represents a bisexual generation, which
are generally smaller, less conspicuous, and more poorly known than
the unisexual generation, the unknown alternate unisexual generation
The Pan-Pacific Entomologist 49: 188-191. July 1973
Vol. 49, No. 3, July 1973
189
Figs. 1, 2. Andricus gigas. Fig. 1. Bisexual generation gall, 6X- Fig. 2. Uni¬
sexual generation gall, 40 X.
of the species has either not been found or probably belongs to another
genus.
Second, all of Kinsey’s adults were imperfect, lacking full antennae.
As of 1971 none of the type specimens in the American Museum of
Natural History had a tarsal claw (J. G. Rozen, personal communi¬
cation) . Five other species described by Kinsey in the same 1922 paper,
which were erroneously described as having simple tarsal claws, were
correctly transferred by Weld (1951) to genera having toothed tarsal
claws. The species transferred were Andricus atrimentus Kinsey, Andri¬
cus pedicellatus Kinsey, Bassettia ligni Kinsey, Neuroterus varians
Kinsey, and Neuroterus decipiens Kinsey.
Though Weld’s manuscript Dryocosmus key lists Quercus lobata
Nee as an additional host and gives a larger maximum adult A. gigas
size of 2.8 mm, his field notes (# 1157a) and book (1957) give no
definite indication that a species matching the original A. gigas de¬
scription, including simple tarsal claw, has been collected subsequently.
This suggested the need to confirm the tarsal condition of A. gigas.
Paratype (labeled Cotype) galls without emergence holes were dis¬
sected and one teneral adult female was extracted. The fore and mid
right legs were removed and mounted on a slide. The tarsal claws are
toothed. Thus A. gigas does not belong in Dryocosmus.
The assumption, from the previous paragraph, that the cotype gall,
which matches the original description, is an A. gigas gall needs re¬
viewing. The dissected female is yellow-brown on the thorax, probably
due to teneral coloration, and has distinct, complete notaulices while
the original description indicates that the notaulices should be “fine
but evident only posteriorly, less evident, discontinous or absent an¬
teriorly where the area is finely coriaceous.” The paratype insects
examined have discontinous notaulices which do reach the pronotum,
190
The Pan-Pacific Entomologist
but are weak. Tulloch (1929) stated, “An examination of the immature
pupal stages of Polistes reveals the presence of these prescutal sutures
which gradually disappear as the chitin hardens before attaining the
adult condition.” This may explain the more marked notaulices on
the teneral adult.
These conditions all indicate the alternate generation of A. gigas
is a senior synonym of the bisexual generation of A. crenatus.
There has been a great deal of confusion about the correct terminology
to designate the thoracic furrows and lines of Hymenoptera. Tulloch
(1929) reviewed the confusion in terminology used for thoracic lines
in Hymenoptera and indicates the lateral-most lines on the Hymenoptera
mesonotum medial to the suture are correctly referred to as parapsidal
lines or furrows. Some Hymenoptera, including cynipids, also have
longitudinal furrows correctly called notaulices, medially of the parap¬
sidal furrows. The latter usually have been referred to incorrectly as
parapsidal furrows by cynipid taxonomists. Eady and Quinlan (1963)
use both terms correctly.
The wing venation terminology employed by Kinsey (1929) and Weld
(1957) varies markedly from that used by Snodgrass (1935). Snod¬
grass, using the Comstock-Needham system, places the cubitus vein
posterior to the median. Kinsey and Weld, however, refer to the median
vein as being posterior to the cubitus. This terminology was possibly a
carry-over from some older system. It is important to standardize the
terminology for wing venation of cynipids with the Comstock-Needham
system to avoid further confusion. Eady and Quinlan (1963) have
corrected cynipid terminology to conform with the Comstock-Needham
system and others working with cynipids should follow this corrected
system.
Type deposition. —One plesiotype female, dissected from paratype
gall, in the American Museum of Natural History, New York, New York.
Acknowledgments. —The authors are grateful to Mr. Robert Lyon
of Los Angeles City College for access to the Weld collection and field
notes and for reviewing this paper, to Dr. Paul H. Arnaud of the Cal¬
ifornia Academy of Sciences for the loan of paratype galls and insects,
and to Dr. Jerome G. Rozen, Jr. of the American Museum of Natural
History for the loan of paratype galls which yielded a teneral adult,
making this paper possible.
Literature Cited
Eady, R. D. and J. Quinlan. 1963. Handbooks for the Identification of British
Insects, Hymenoptera: Cynipidae, Key to Families and Subfamilies
Vol. 49, No. 3, July 1973
191
and Cynipinae (Including galls). Royal Ent. Soc. London, 7 pt. 1A:
81 p.
Kinsey, A. C. 1922. New Pacific Coast Cynipidae. Amer. Mus. Nat. Hist.
Bull., 46: 279-295.
1929. The Gall Wasp Genus Cynips. Indiana Univ. Stud., 16(2, 3, 4) : 577 p.
Snodgrass, R. E. 1935. Principles of Insect Morphology. McGraw-Hill, New
York, 667 p.
Tulloch, G. S. 1929. Proper Use of Terms “Parapsides” and “Parapsidal
furrows” Psyche, 36: 376-382.
Weld, L. H. 1951. p. 600-654 in Muesebeck, C. F. W., K. V. Krombein, and
H. K. Townes. Hymenoptera of America North of Mexico. U. S. Dep.
Agr., Monogr. 2, 1240 p.
1952. New American Cynipid Wasps from Galls. Proc. U. S. Nat. Mus.,
102: 315-342.
1957. Cynipid Galls of the Pacific Slope. Privately Printed; Ann Arbor,
Mich., 64 p.
RECENT LITERATURE
Behavior of Marine Animals. Current Perspectives in Research, vol. i: In-
verterrates. vol. ii: Verterrates. Edited by Howard E. Winn and Bori L.
Olla. Plenum Press, New York. 503 p. (1-244 -f- xxix; 245-503 -(- xviii). 1972.
$32.00 ($16.00 each).
The articles on behavior of several groups of arthropods in Vol. I will be of
special interest to insect behaviorists.— Robbin W. Thorp, University of California ,
Davis, 95616.
Insects. Ross E. Hutchins. Prentice-Hall, Inc., Englewood Cliffs, N. J. 324 p.
illus. 1966. Prisim Paperback edition, 1972. $3.50.
Pagination and text in the paperback edition are the same as in the original
edition, but the smaller format reduces the size of type and photos. Many illus¬
trations appear different, but are merely reduced, inverted or put on an angle to
better fit the smaller page. A few illustrations (e.g. on pages 82, 114, etc.) are
different, but without loss of quality.— Robbin W. Thorp, University of California,
Davis, 95616.
Love and Hate. The Natural History of Behavior Patterns. Irenaus Eibl-
Eibesfeldt. Translated by Geoffrey Strachan. Holt, Rinehart and Winston, Inc.,
New York. 276 p., 63 figs. 1972. $7.95.
An ethologist examines the roots of some behavior patterns in man in light of
some social interactions among higher vertebrates and a few social insects.— Robbin
W. Thorp, University of California, Davis, 95616.
192
The Pan-Pacific Entomologist
The Genus Paratrichogramma Girault
(Hymenoptera: Trichogrammatidae)
Richard L. Doutt 1
University of California, Berkeley
The genus Paratricho gramma is easily separated from the 71 other
valid genera in the family Trichogrammatidae as recognized by Doutt
and Viggiani (1968). Its female antennal formula of a single annellus,
a single flagellomere, and a solid club is unusual and when this char¬
acter is combined with the distinctive wing venation the genus is readily
set apart. Similarly the males are almost unmistakable, and would not
be easily confused with other members of the family because of the
peculiar bottle-shaped second funicle segment.
There are two species described from Australia, namely P. Cinderella
Girault, which was illustrated by Doutt and Viggiani (1968), and P.
fusca Girault. The type of P. fusca is apparently lost, for Girault de¬
posited it in the Queensland Museum with the number “Hy/803” and
indicated (Girault, 1912) that it was mounted “in xylol-balsam with
the type female of Polynema spenceri Girault and two trichogram-
matids.” I have examined a slide in the Queensland Museum which
does have the type of P. spenceri, a male of Trichogramma australicum ,
and a chalcid body with several fragmented legs but no tarsi, no head,
no antennae, and no wings. The slide is labelled “Paratrichogramma
fusca Girault 2 Hy/803”, and the locality data match the original
description. In red ink the label also has the number 3415, which cor¬
responds to the number given P. fusca by Girault in an unpublished
revision of his collection. However, in red ink on the slide label under
the designation “Paratrichogramma fusca ” Mr. Girault has also writ¬
ten “destroyed.” All of this evidence indicates that the type specimen
of P. fusca was originally on this particular slide, but now it must be
considered as missing or destroyed. This is especially unfortunate
because Girault’s original description is too brief and inadequate to
form a satisfactory concept of the missing specimen. Perhaps the most
useful and reliable distinction is that Girault considered P. fusca to
have forewings more densely ciliated discally than P. Cinderella with
about 18 long lines of cilia instead of 5 short lines from the apex as
in P. Cinderella.
1 Present address: 9240 S. Riverbend Avenue, Parlier, Ca. 93648.
The Pan-Pacific Entomologist 49: 192-196. July 1973
Vol. 49, No. 3, July 1973
193
Figs. 1-4. Paratrichogramma pretoriensis. Fig. 1. Male antenna. Fig. 2. Male
wings. Fig. 3. Female antenna. Fig. 4. Female forewing.
There are 4 specimens of P. Cinderella at the Queensland Museum
and one at the U.S.N.M. The distribution from these individuals is
indicated as Queensland and Thursday Island (Torres Strait). This
constituted the extent of our information on the genus until a few
Paratricho gramma were recognized in collections from California and
South Africa. These represent new species and are described as follows:
Paratrichogramma pretoriensis Doutt, new species
Female. —Length 0.45 mm; body color golden brown, head yellow gold, eyes
and ocelli bright red, antennae light brown, coxae dark, trochanters white, femora
194
The Pan-Pacific Entomologist
Figs. 5-8. Paratrichogramma californica. Fig. 5. Female antenna. Fig. 6.
Female wings. Fig. 7. Male antenna. Fig. 8. Male forewing.
and tibiae white with median area dark, banded, apex of tibiae and third tarsomere
dark, wings hyaline, venation brown. Head, frontal view slightly wider than long,
face deeply impressed, roughly sculptured; eyes large; toruli low on face, ocelli
large, prominent; mandibles with two pointed teeth, third blunt; eyes without
spicules; antennae as Fig. 3 with long scape, large pedicel, single annellus, single
flagellomere with stalk-like peduncle attachment to single club. Scutum and
scutellum with light, reticulated sculpture; thorax broad as abdomen, slightly
over half length of abdomen, phragma large, wide, parallel sided, broadly truncate
at apex, extending to middle of abdomen. Ovipositor small, not exserted. Fore
femora longer than tibiae; foretarsi with short basitarsus; middle and hind tibiae
Vol. 49, No. 3, July 1973
195
longer than femora, basal tarsomeres as long or longer than succeeding segments.
Wings as Fig. 4 with broad marginal vein and sessile stigma; center of wing disc
with rough leathery texture.
Male.— Body color similar to female, slightly darker. Wings dusky at base;
head, legs, antennae colored as female. Face impressed, eyes smaller than female;
antennae as in Fig. 1 with peculiar bottle-shaped second funicle. Thorax as wide
as abdomen, half length. Scutum with reticulate sculpture only on anterior portion,
scutellum smooth. External genitalia large. Wings as in Fig. 2.
Paratrichogramma pretoriensis is readily differentiated from P. Cin¬
derella by the broader marginal vein and more abundant discal cilia
of the forewings; and from P. californica by the reticulated sculpturing
of the scutum.
Holotype female and allotype from suction trap 1 (1029), Pretoria,
South Africa, February 1958. D. P. Annecke. Paratype male from
suction trap, Pretoria, South Africa, March 1958. D. P. Annecke.
Holotype and allotype to be deposited at Plant Protection Research
Institute, Pretoria. Paratype at Division of Biological Control, Uni¬
versity of California.
Paratrichogramma californica Doutt, new species
Female. —Length 0.45 mm, body dark brown, except propodeum golden, head
bright golden yellow, eyes and ocelli crimson, antennae pale gold, legs light brown
to smoky becoming pallid at tips of tibiae and basal tarsomeres, but hind coxae
dark brown, wings hyaline, smoky near base, veins brown. Eyes with inner margins
straight, eyes extend three-fifths length of face; toruli low on face; face deeply
impressed, sculptured by transverse ridges; mandibles with two distinct teeth
and truncate shoulder; antennae with long scape, pedicel slightly shorter than
single funicle segment, club obliquely truncate (Fig. 5). Body compact, stout;
scutum without distinct reticulated sculpture as occurs in P. pretoriensis and P.
Cinderella. Thorax slightly shorter than abdomen, phragma large, extending nearly
to middle of abdomen. Ovipositor about half length of abdomen, small hypogynium
at base, not exserted. Legs somewhat slender, rather long for compact body, hind
coxae especially elongate, fore femora and fore tibiae about equal length. Wings
as in Fig. 6.
Male. —Body color pattern basically as female but darker, with very dark band
across abdomen. Head with transverse sculpturing similar to female, eyes smaller,
extending just half length of face. Antennae (Fig. 7) with peculiar bottle-shaped
second funicle, club has suggestion of fused basal segment. Scutum with reticu¬
lations on anterior-lateral areas. Phragma extending posteriorly three-fifths length
of abdomen. Wings as in Fig. 8.
Holotype female, allotype, one paratype female Hub, California,
27 August 1956, from Atriplex bracteosa. C. E. Kennett. One paratype
female, from Atriplex bracteosa , 4 mi. W Kearney Park, Fresno Co.,
Calif. 13 September 1951. C. E. Kennett and C. B. Huffaker. Type series
at Division of Biological Control, University of California.
196
The Pan-Pacific Entomologist
Literature Cited
Doutt, R. L. and G. Viggiani. 1968. The classification of the Trichogramma-
tidae (Hymenoptera: Chalcidoidea). Proc. Calif. Acad. Sci., 35(20):
477-586.
Girault, A. A. 1912. Australian Hymenoptera Chalcidoidea—I. Mem. Queens¬
land Mus., I: 66-189.
SCIENTIFIC NOTE
Leptocoris rubrolineatus, an Occasional Predator of the California
Oakworm, Phryganidia californica (Hemiptera: Rhopalidae; Lepidoptera:
Dioptidae).—In coastal California, the western boxelder bug, Leptocoris rubro¬
lineatus Barber, occurs in large numbers on bigleaf maple, Acer macrophyllum
Pursh. The adult bugs are often common on nearby vegetation, including coast live
oak, Quercus agrifolia Nee, on which feed larvae of the California oakworm,
Phryganidia calif ornica Packard.
Boxelder bugs have heretofore been recorded feeding only on plant hosts (Met¬
calf, Flint, and Metcalf, 1962, Destructive and Useful Insects, 4th ed. McGraw-
Hill, N. Y.). While investigating mortality of the California oakworm, I noticed
Leptocoris adults feeding on oakworm pupae. The oakworm forms a hanging
chrysalid with no surrounding cocoon. A careful search of a coast live oak trunk
near the campus of California State University, Hayward, on 2 July 1970 revealed
109 oakworm pupae, of which 8 were being fed upon by adult Leptocoris. The
tree was 12 m from the trunk of a bigleaf maple swarming with boxelder bugs.
A search of oaks 45 m and 51 nr from this maple revealed few Leptocoris, none of
which were feeding on oakworm pupae.
I collected 10 adult Leptocoris and placed them in a plastic petri dish with 50
oakworm pupae. They fed readily on the pupae though no alternate food was
available. Pupae upon which the bugs fed did not survive. In the field, for up
to 45 minutes each, I followed individual boxelder bugs that had completed feed¬
ing on oakworm pupae, and in no case did they feed on additional pupae. To
estimate the extent of predation by Leptocoris, I examined oakworm pupae from
several oaks growing within 20 m of maples, but I was unable to accurately dis¬
tinguish feeding sites of Leptocoris from oviposition punctures of parasitic Hy¬
menoptera, which were also common in the area.
Harville (1955, Microent. 20: 83-166) makes no reference to Leptocoris pre¬
dation in his discussion of natural mortality of Phryganidia. Feeding on oakworm
pupae by Leptocoris is probably of little significance in oakworm population
dynamics because of its localized nature though it is interesting to note a normally
phytophagous insect feeding on animal matter.— David J. Horn, Department of
Entomology, The Ohio State University, Columbus, 43210.
Vol. 49, No. 3, July 1973
197
Behavioral Studies of Three Morphotypes of
Therion circumflexum
(Hymenoptera: Ichneumonidae)
C. N. Slobodchikoff
Department of Biological Sciences, Northern Arizona University, Flagstaff, 86001
Therion is a genus of wasps that are parasitoids of Lepidoptera
caterpillars. The female lays an egg into the body cavity of a host
caterpillar; the egg develops into a larva that feeds on the host’s non-
vital tissues. Eventually the host pupates, at which time the parasitoid
larva kills the host, pupates inside the host’s pupal shell, and emerges
from the pupal shell as an adult wasp. It has recently been shown that
all North American Therion can be considered as members of a single
species, T. circumflexum (L.), with distinct groups of populations that
can be segregated into ten separate morphological types or morphotypes
(Slobodchikoff, 1971). The purpose of the present paper is to report
on the behavior of three of the ten morphotypes.
The behavior of the three morphotypes, CIRCUMFLEXUM, CAL-
IFORNICUM, and MORIO was studied both in the field and in the
laboratory. When this study was begun, it was expected that minor
differences would be observed between morphotypes in most of the
behavioral categories described in the present paper. However, obser¬
vation and experimentation demonstrated that all three morphotypes
have the same behavioral response patterns in every category except
host selection.
Materials and Methods
The materials and methods used in the behavioral studies were the
following. CALIFORNICUM adults were observed while contained
in a sleeve cage, 0.5 m on each side with a fine wire mesh. CIRCUM¬
FLEXUM and MORIO adults were observed while contained in trans¬
parent plastic shoeboxes and in half-gallon ice cream cartons covered
with a piece of 2 mm-thick glass. The shoeboxes had their ends cut
off and substituted by a fine wire mesh. The top of each shoebox had
a 5 X 6 cm mesh to insure proper ventilation and had three circular
holes, each measuring 2.5 cm in diameter, for introducing food, water,
and experimental items. These circular holes were normally plugged
with cotton. Wasp parts were weighed on a Mettler balance to the
nearest 0.05 mg. Potter’s clay was used for constructing the model of
The Pan-Pacific Entomologist 49: 197-206. July 1973
193
The Pan-Pacific Entomologist
the insect in flight. The model was baked in a Braun oven at 40° C for
24 hours, then allowed to cool for 24 hours. The response to light was
tested with an American Optical Company microscope illuminator and
a gooseneck desk lamp with a 100 watt bulb. Temperature was mea¬
sured by a Celsius thermometer and by a hygrothermograph. Tem¬
perature in the laboratory remained at 20 ± 2° C, relative humidity
averaged 40 per cent. Mating was observed in part while the wasps
were caged in a 90 X 30 mm glass vial with a cotton plug at the top.
Results and Discussion
Habitats studied. —Extensive field observations of CIRCUMFLEX-
UM’S behavior were made at Lily Lake, Marin County, California.
Lily Lake is situated in the California Coast Ranges, and is surrounded
by hills and ridges with narrow canyons containing seasonal streams.
The habitat around Lily Lake may be characterized as grassland along
the tops of ridges, oak-chaparral woodland among the slopes of the
hills, and mixed evergreen-redwood forest along the canyon floors.
CIRCUMFLEXUM was found only in the oak-chaparral areas. In¬
dividuals of this morphotype were typically seen flying around oaks
and in the grass surrounding oak trees. Specimens of CALIFORNICUM
were collected from similar habitats in areas near Lily Lake. Obser¬
vations of the behavior of CALIFORNICUM were made at Hallelujah
Junction, Lassen County, California, and at a cotton field near Kerman,
Kern County, California. Hallelujah Junction has a Great Basin flora
and fauna. Therion individuals were found flying around willows
along a stream. The cotton field near Kerman was, at the time the
observations were made, an experimental plot for control of bollworm
(Heliothis zea (Boddie)) by Bacillus thuringiensis Berliner, and had not
been sprayed with insecticides for the previous two years.
The behavior of MORIO was observed at Verdi, Washoe County,
Nevada, in a patch of willows along the Truckee River. The habitat
around Verdi may also, like Hallelujah Junction, be characterized as
Great Basin. Only three females and four males were found at this
locality during two days of observation. No other Therion were found
in surrounding willow patches for half a mile in either direction along
the Truckee River. The Verdi colony is probably an isolated one with
a low population density.
In California, the three morphotypes are often found in the vicinity
of willows. CALIFORNICUM is also found in cotton and alfalfa
fields. In addition to willows, CIRCUMFLEXUM is found near oaks
Vol. 49, No. 3, July 1973
199
in oak-woodland habitats. MORIO has only been recorded from hab¬
itats containing willows. In all observed cases, the Therion lived in
isolated colonies. At a given locality, all the Therion individuals were
found within a small area, often less than one square mile in size. Out¬
side of this area, no Therion were found, although the habitat was quite
similar in surrounding areas.
Flight.— Therion males and females characteristically fly with the
abdomen elevated and the hind legs outstretched (Fig. 1). Sometimes
the middle and front pairs of legs are extended to the sides, with the
femora pointed laterally away from the thorax and the tibiae and tarsi
pointed downward. As the wasp flies, the abdomen may be seen to
occasionally move up or down. This movement follows a change in the
position of the thorax relative to a horizontal plane. If the thorax dips
forward, so that the head is lowered and the propodeum is raised, the
abdomen moves down. If the thorax dips backwards, so that the head
is raised and the propodeum is lowered, the abdomen moves up. If an
imaginary plane is extended through the thorax while the wasp is in
flight, so that half of the thorax is above this plane and half is below,
the abdomen is held at a 20 degree angle above the plane and the legs are
held at a 20 degree angle below the plane.
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Table 1. Measurements of body parts of a male circumflexum.
Part
W eight
mg
Dimensions
mm
(1 X h X w)
Volume
mm 3
Head
2.75
1.3 X 1.7 X 2.0
4.42
Thorax and wings
12.40
6.0 X 3.0 X 3.0
54.00
Abdomen
9.00
16.0 X 2.2 X 0.8
26.40
Hind legs
2.55
17.5 X 0.5 X 0.5
4.38
The wasp in flight may be considered as three separate arms balanced
around a center of gravity. The thorax, head, wings, and first two
pairs of legs comprise one arm. The hind pair of legs comprise the
second arm, and the abdomen comprises the third arm. On the basis
of observations of the movement of these arms, the center of gravity
of the wasp in flight may be predicted to be at the point where the arms
join, near the hind end of the propodeum. If the head pitches forward
or backward, the center of gravity shifts. To compensate for the shift,
the abdomen moves either up or down, bringing the center of gravity
back to the end of the propodeum.
To see whether the center of gravity of a wasp is actually at the end
of the propodeum, an experiment using a clay model was performed.
A male CIRCUMFLEXUM, freshly killed, was cut into the following
parts: (1) the head; (2) the thorax, wings, and first two pairs of legs;
(3) the abdomen, including the petiole; and (4) the hind legs, includ¬
ing the coxae. These four parts were weighed and were measured. The
volume of each part was calculated. The data obtained from these
measurements are summarized in Table 1. A scale model was made,
attempting to reproduce the actual contours of the wasp as closely as
possible. The scaling factor for the weight was 1 X 10 3 , and the factor
for the dimensions was 0.5 X 10. The abdomen and legs were fixed at
20 degree angles with respect to the horizontal plane. When the model
was suspended by a string tied around the juncture of the abdomen,
legs, and thorax, the model balanced in the position that the wasp uses
for level flight.
Resting and grooming. —In both the laboratory and in the field,
Therion adults may frequently be observed at rest clinging to an object
in a head-upright position perpendicular to the ground, and either re¬
maining motionless or cleaning portions of the body. Cleaning consists
of a series of actions. First one antennal flagellum, then the other, is
drawn through the space between the strongly curved front tibial spur
Vol. 49, No. 3, July 1973
201
Fig. 2. Spongy empodium between tarsal claws of T. circumflexum. Scanning
electron micrograph, 420X- Fig. 3. Tarsal pad and spongy empodium, tarsal
segments 4-5 of T. circumflexum. Scanning electron micrograph, 170X-
and the first tarsal segment. The right front leg cleans the right fla¬
gellum, while the left front leg cleans the left one. The tibial spur and
the front tarsal segments are then brushed against the mouthparts, each
leg brushing the mouthparts on its own side. Following this, the tarsal
segments and tibial spur of the middle leg are brushed against the end
of the tibia and the tarsal segments of the front leg. The same action
is repeated by the hind legs, brushing against the tibia and tarsal seg¬
ments of the middle leg. Then the wings are held out flat on either side
of the abdomen, with their upper surfaces parallel to the dorsum of the
wasp. The tibia of one hind leg brushes against the surface of the other
two wings. After cleaning the wings, the hind legs are used to brush
the posterior half of the abdomen. This is accomplished by simulta¬
neously moving the tarsal segments and tibial spurs of each hind leg
along the sides of the abdomen toward the posterior end. The abdomen-
brushing action is usually repeated 2-3 times. Cleaning behavior con¬
cludes with the brushing of the tibial spur and lower surfaces of the
tarsal segments on one hind leg against the upper surface of the other
leg’s tarsal segments, then a corresponding action for brushing the
segments of the other leg.
Therion adults often cling to blades of grass or small twigs during
resting in the field. In the laboratory they are capable of clinging to
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The Pan-Pacific Entomologist
glass and smooth plastic as well as to materials such as twigs, paper
towels, and wire meshes. Clinging to smooth surfaces is facilitated by
a large, spongy, empodium (Fig. 2) on the pretarsus and by four tar¬
sal pads per leg. Each tarsal pad lies in the intersegmental space be¬
tween tarsal segments, and appears to overlap the segment distal to it
(Fig. 3). The pad consists of spongy ridged tissue that, when pressed
against a smooth surface, has a suction effect. The empodium also
functions as a suction pad.
Response to light. —Both males and females of the three morpho-
types exhibit positive phototaxis. If given an intensity gradient of light,
the wasps will move to the area of greatest intensity. A simple exper¬
iment was performed to establish this. Eight males and ten females
of CIRCUMFLEXUM were confined in a plastic cage. The cage was
placed in a darkened room at 9 p.m., PDT, for one hour. A micro¬
scope illuminator, set at low intensity (300 ft. candles at the end of
the cage nearest light, 100 ft. candles at the middle of the cage), was
placed one foot from one end of the cage. The illuminator was posi¬
tioned so that the beam of light would pass along the length of the cage.
At the end of one hour, the illuminator was switched off. Initially,
the Therion were distributed fairly uniformly in the cage, all in rest¬
ing positions. However, within five seconds of application of the light
stimulus, movement was elicited. Within 30 seconds, seven males and
all ten females had crawled or flown to the side of the cage closest to
the light. Within one minute, all the individuals were along that side.
This experiment was repeated three times on separate days. In each
case, all individuals were along the side closest to the light within one
minute. The wasps showed no habituation to the light, and remained
at the side of the cage nearest to the light until the light was switched
off.
The same experiment was performed with four males and three fe¬
males of MORIO, under similar experimental conditions. In three
replicates, on separate days, the same results as with CIRCUMFLEXUM
were obtained. CALIFORNICUM males and females were found to
respond in a similar fashion to a desk lamp after a period of darkness.
If a diffuse light, such as that produced by an overhead white fluores¬
cent lamp, was switched on after a period of darkness, and the intensity
gradient was comparatively low, the wasps of all three morphotypes
became active within one minute. However, they did not congregate
at any given side of the cage.
Response to temperature. —Both CALIFORNICUM and CIRCUM¬
FLEXUM become increasingly active with temperatures higher than
Vol. 49, No. 3, July 1973
203
ca. 15 degrees C. CALIFORNICUM adults at Hallelujah Junction and
near Kerman begin to appear about 7:30 or 8:00 a.m., PDT, when the
temperature begins to exceed 17 degrees C. CALIFORNICUM adults
near Lily Lake, where temperatures are generally cooler, become active
later in the day, around 9:30 or 10:00 a.m., PDT, when the temperature
begins to exceed 15 degrees C. On sunny warmer mornings, CIRCUM-
FLEXUM adults at Lily Lake become active earlier than on cooler sunny
mornings. The same response to temperature has been reported for
eastern MORIO by Tothill (1922).
Mating.— The mating behavior of CALIFORNICUM and CIRCUM-
FLEXUM was observed in the laboratory, but not in the field. The
mating behavior of MORIO was not observed. CALIFORNICUM was
observed mating on two occasions, one while the individuals were con¬
fined in the plastic cage described above, and one while the individuals
mating were confined in a glass vial.
Both morphotypes go through the same set of actions for mating.
Tothill (1922) reports that MORIO mates by pairing in flight and
falling to the ground. No pairing in flight was observed for the two
morphotypes studied, but that may be a consequence of laboratory
confinement. In both morphotypes, the male approaches the female
from the rear and moves his antennae along the posterior half of her
abdomen 2-5 times. He then climbs onto her dorsum, curls his abdomen
around so that the tip of the genital capsule is pointing anteriorly, and
couples with her.
The male and female remain coupled for 1-2 minutes (55 and 104
seconds for CALIFORNICUM, 70 and 78 seconds for CIRCUM-
FLEXUM), following which the female immediately flies off. The
male, however, remains motionless for about another minute (45 and
62 for CALIFORNICUM, 48 and 75 seconds for CIRCUMFLEXUM).
In all four observed matings, the male assumed the characteristic position
taken by wasps at death. The wings were partially spread to the sides
and depressed, the abdomen was partially curled underneath itself,
and the legs were spread to the sides. The CIRCUMFLEXUM indi¬
viduals were field collected and it is not known if any sperm was trans¬
ferred during the observed matings, but the CALIFORNICUM indi¬
viduals were laboratory reared, and gave rise to female progeny. Tothill
(1922) reports that in two observed matings of MORIO, the individuals
remained coupled for five and fifteen minutes respectively.
Feeding and drinking.— In the laboratory males and females readily
accept either a diet of clover honey mixed with tap water in a 1:1 pro¬
portion or a diet of pure clover honey. These diets provide sufficient
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The Pan-Pacific Entomologist
nutrients for the wasps to produce viable progeny. With either diet,
water is the only other requirement. Wetting a piece of cotton in the
cage is a convenient way of providing the water, since the wasps can
easily extract the absorbed liquid. Water has to be provided daily.
When three laboratory-reared, newly emerged individuals of CALI¬
FORNICUM were fed diluted honey but were deprived of pure water,
all three wasps were found dead at the end of 36 hours. The general
characteristics of drinking behavior in ichneumonids have been dis¬
cussed by Townes (1958). Tothill (1922) found that adults of MORIO
in the eastern United States eat pollen grains of goldenrod in the field,
and will eat sugar mixed with water, the fresh inside of a banana skin,
or pollen grains of goldenrod in the laboratory. The adults of CAL-
IFORNICUM and CIRCUMFLEXUM probably also normally feed
on pollen grains.
Host searching and selection. —When searching for hosts, females
of all three morphotypes characteristically fly in slow zigzags not more
than 1.5 meters above the ground. Periodically, a female will alight
on a twig, leaf, or grass stem and explore the surrounding area within
a 15 cm radius. During such explorations, the antennae are moved up,
down, and from side to side, frequently touching the substrate on which
the female is walking. Following a local exploration, the female resumes
her zigzag pattern of flight.
The factors involved in host selection have been described elsewhere
(Slobodchikoff, 1971). Odor provides an important initial cue that
allows the wasps to find a potential host. Once the host is located, a
female uses tactile cues provided by the surface of the caterpillar. The
hosts of CIRCUMFLEXUM and CALIFORNICUM have no secondary
setae on the integument, while the hosts of MORIO have dense patches
of secondary setae. A female of all three morphotypes will approach
a potential host and extend her antennae to the caterpillar’s surface,
briefly touching the caterpillar. If a MORIO female encounters an
integument bare of patches of secondary setae, she loses all further
interest in the caterpillar as a potential host. Similarly, females of
CIRCUMFLEXUM and CALIFORNICUM lose all interest in a po¬
tential host, even in the presence of the proper odor, if an antennal
scan of the surface encounters anything other than a wrinkled integu¬
ment free of secondary setae. Even when a female wasp is deprived
of suitable hosts for prolonged periods of time, caterpillars with im¬
proper cues are rejected as hosts.
Oviposition. —Oviposition behavior begins when the wasp encounters
a suitable host. In all three morphotypes studied, oviposition follows
Vol. 49, No. 3, July 1973
205
Figs. 4^7. Oviposition behavior sequence of female T. circumflexum, morpho-
type MORIO, in response to a woolly bear caterpillar, Estigmene acrea Drury.
In this particular sequence, the female wasp misjudged the position of the cater¬
pillar. See text for further explanation.
the same pattern. With her antennae held above her head, the female
approaches to within 1.0-1.5 cm of the larva. She taps either the larval
integument (if the larva is relatively hairless) or several larval hairs
(if the larva is hairy) a few times with her antennae (Fig. 4). Then
the antennae are raised above the head again. The abdomen is curled
under the thorax so that the tip of the ovipositor is below and slightly
in front of the head (Fig. 5). The abdomen fits between the hind coxae,
which are pressed against the sides of the abdomen. The wasp typically
cants her body to either her right or her left side, so that the dorsum
of the wasp is held at approximately 45 degrees relative to the substrate
on which the wasp is standing (Fig. 6). This allows her to rapidly
extend the abdomen without interference from the substrate. If she
decides to sting the host, the entire abdomen is rapidly thrown forward
and the ovipositor pierces the larval integument. With the extension
of the abdomen, the thorax is pulled up and back, and the first two pairs
of legs are lifted from the surface of the substrate (Fig. 7). Actual egg-
laying, from the time the abdomen is thrust forward, takes less than
one second. The ovipositor penetrates to the body wall opposite the
point of entry. This was first recorded by Tothill (1922) as character¬
istic of egg-laying by MORIO. After ovipositing, the female usually
cleans herself.
206
The Pan-Pacific Entomologist
Conclusions
The similarities and differences of behavioral patterns can be inter¬
preted in terms of selection pressure and partitioning of the environ¬
ment. All three morphotypes are sympatric, in the broad sense, through
at least part of their range (Slobodchikoff, 1971). However, populations
of the three morphotypes are usually small, with each population dis¬
tributed over less than two square miles. Populations of the different
morphotypes generally do not overlap, so that functionally the morpho¬
types are microallopatric. At the same time, populations of different
morphotypes may be separated by as little as two miles, and may occur
in the same type of habitat. Fluctuations in population size may bring
two different morphotypes into contact, in which case it would be ad¬
vantageous to insure maximum survival of progeny. Selection would
be expected to have a disruptive effect on host preferences, and a neutral
effect on all behavioral patterns not directly connected with the survival
of offspring.
Literature Cited
Slobodchikoff, C. N. 1971. Patterns of differentiation and evolution in the
genus Therion (Hymenoptera: Ichneumonidae). Univ. Calif., Ph.D.
dissertation.
Tothill, J. D. 1922. The natural control of the fall webworm ( Hyphantria
cunea Drury), in Canada together with an account of its several
parasites. Can. Dep. Agr. Tech. Bull., No. 3, 107 p.
Townes, H. K. 1958. Some biological characteristics of the Ichneumonidae
(Hymenoptera) in relation to biological control. J. Econ. Entomol.,
51: 650-652.
RECENT LITERATURE
Hewiston on Butterflies 1867-1877. With a preface by Dr. L. G. Higgins. E. W.
Classey, Ltd., Middlesex, England. 200 p. 1972. $12.50. North American Dis¬
tributor: Entomological Reprint Specialists.
A reprinted collection of four articles containing descriptions of exotic Lepi-
doptera, mostly Neotropical.— Robbin W. Thorp, University of California, Davis ,
95616.
The Environment, The Establishment and The Law. Harmon Henkin, Martin
Merta, and James Staples. 223 p., illus., 1971.
A review of the DDT controversy centering on the 1968-1969 hearings conducted
in Madison, Wisconsin with many direct quotes from the testimony given. Appen¬
dices include the ruling, a model pesticide law and a glossary defining many words
unfamiliar to the layman including “entomologist.”- —Robbin W. Thorp, University
of California, Davis, 95616.
Vol. 49, No. 3, July 1973
207
The Ant Larvae of the Tribes Basicerotini and Dacetini:
Second Supplement
(Hymenoptera: Formicidae: Myrmicinae)
George C. Wheeler and Jeanette Wheeler
Laboratory of Desert Biology, Desert Research Institute,
University of Nevada System, Reno, 89507
Subsequent to the publication of our first supplement on the larvae
of the subfamily Myrmicinae (Wheeler and Wheeler 1960a) 1 we have
received from other myrmecologists so much additional material that
it has become necessary to publish another supplement.
Tribe Basicerotini
Brown and Kempf (1960: 165).—Differences between larvae of
Basicerotini and Dacetini are based on our 1954 paper.
Genus Basiceros Schulz
Brown and Kempf (1960: 171).—Larval characters after our 1954
description.
Genus Rhopalothrix Mayr
We have changed our minds about Rh. gravis Mann (1954: 117) : we
now regard our material as mature and classify its profile as aspididri-
form (Wheeler and Wheeler 1960b: 103, 106).
Genus Eurhopalothrix Brown and Kempf
Profile myrmiciform. Head large. Body hairs numerous, short to
long; of two types: (1) flexuous and denticulate and (2) flexuous,
denticulate, ending in a sharp-pointed bulb, on the dorsum of AI to AVI
(two per somite). Cranium subcordate. Antennae minute. Head hairs
short to long, denticulate and moderately numerous. Mandibles lepto-
thoraciform. Maxillae with a constriction between palp and galea.
In our 1960b key (p. 109) Eurhopalothrix would run to Orectogna-
thus , from which it may be distinguished by its lack of bifid hairs.
Eurhopalothrix australis Brown and Kempf
(Fig. 1)
Length (through spiracles) about 2 mm. Profile myrmiciform (i.e., stout and
rather elongate; diameter greatest at AIV and AV; slightly attenuated anteriorly;
1 We shall cite each of our papers the first time it is referred to with our names as authors; in sub
sequent citations we shall use date.
The Pan-Pacific Entomologist 49: 207-214. July 1973
208
The Pan-Pacific Entomologist
Fig. la-e. — Eurhopalothrix australis, a, Head in anterior view, Xlllj b, left
mandible in anterior view, X267; c, larva in side view, X35; d, e, two types of
body hairs, X263. Fig. 2a-e. Colobostruma sp. a, Head in anterior view, XlH;
b, left mandible in anterior view, X267; c, d, two types of body hairs, X267; e,
larva in side view, X42. Fig. 3a-e .—Mesostruma browni. a, Larva in side view,
Vol. 49, No. 3, July 1973
209
thorax stout and arched or bent ventrally, but not differentiated into neck; pos¬
terior end broadly rounded). Anus ventral. Head on anterior end and of about
same diameter as Tl. Spiracles small. Integument of ventral surface of thorax
and AI and All with few short transverse rows of minute spinules. Body hairs
numerous and uniformly distributed, denticulate. Of two types: (1) 0.04-0.15 mm
long, on all somites, shorter without and longer hairs with alveolus and articular
membrane; (2) about 0.15 mm long, with stout shaft and slightly swollen bulb
just below sharp-pointed tip, two on dorsal surface of each AI-AVI. Cranium
somewhat broader than long and feebly cordate. Antennae minute, each with
three sensilla, each bearing a minute spinule. Head hairs moderately numerous,
short to long (0.038-0.075 mm), denticulate and slightly curved. Labrum bilobed,
short and broad; each lobe with three or four minute hairs and/or sensilla on
anterior surface, with minute spinules in short rows and with one isolated and two
contiguous sensilla on ventral border, with coarse isolated spinules on the lateral
border and with cluster of three sensilla on posterior surface; entire posterior
surface spinulose, spinules minute and in numerous rows, rows transverse in middle
half. Mandibles leptothoraciform (i.e., moderately narrow, tapering gradually
and curving gradually to apical tooth; anterior surface produced medially into
blade with two subapical teeth), with three or four denticles on posterior surface.
Maxillae large, each divided by constriction between palp and galea, distal portion
spinulose and with conoidal apex; palp digitiform, with five sensilla (two apical
and encapsulated and three subapical and bearing one spinule each) ; galea digiti¬
form and bearing two apical sensilla. Labrum very thick, short, broad and feebly
bilobed; anterior surface densely spinulose, spinules in transverse rows; each palp
with five sensilla (two apical and encapsulated, three lateral and with one spinule
each) ; isolated sensillum between each palp and opening of sericteries, latter a
transverse slit. Hypopharynx densely spinulose, spinules long and in numerous
transverse rows. (Material studied: six larvae from New South Wales, courtesy
of Rev. B. B. Lowery.) In alcohol mature larvae cling to each other by means of
the interlocked hairs.
Tribe Dacetini
Our latter-day taxonomists say that we should use all possible kinds
of characters in classification. Here is a possible chemical character
of the tribe Dacetini: when larvae are cleaned in 10% KOH, the in¬
soluble meconium turns red. We have found this to be true of the
following species: Colobostruma sp., Epopostruma alata, E. quad-
rispinosa, Mesostruma browni, Orectognathus antennatus, O. mjobergi ,
O. nigriventris , Strumigenys perplexa.
<-
Xl8; b, head in anterior view, X88; c, left mandible in anterior view, Xl77; d,
e, two types of body hairs, X267. Fig. 4 .—Strumigenys perplexa. Left mandible
in anterior view, X290.
210
The Pan-Pacific Entomologist
Genus Acanthognathus Mayr
G. C. and J. Wheeler (1969: 110).—Generic characterization.
Wilson (1962: 414).—The danger of piercing the skin of the larvae
is avoided thus: “a pair of small teeth near the base of the mandibles
are used to grip and carry brood, thus by-passing the apical teeth (Mann,
1922: later confirmed by W. L. Brown, in litt., 1961).” But we can
find no such statement in Mann (1922).
Acanthognathus rudis Brown and Kempf
G. C. and J. Wheeler 1969: Description and figures.
Genus Colobostruma Wheeler
Profile orectognathiform. Anus with a small posterior lip. Head
large. Body hairs sparse and short; of two types: (1) smooth, deeply
bifid with the tips curling away from each other and (2) unbranched
and denticulate. Frons and clypeus bulging. Antennae small. Head
hairs few and short to long; unbranched and either smooth or denticu¬
late. Mandibles leptothoraciform.
In our 1960b key the genus Colobostruma runs to 0rectognathus,
from which it can be distinguished by the lack of denticles on its bifid
hairs.
Colobostruma sp.
(Fig. 2)
Length (through spiracles) about 2.1 mm. Profile orectognathiform (i.e.,
abdomen rather stout, diameter greatest at AIV and AV, attenuated anteriorly;
thorax rather slender and curved ventrally, forming indistinct neck; posterior
end broadly rounded). Head large. Anus ventral, with posterior lip. Leg, wing
and gonopod vestiges present. About ten differentiated somites. T2 spiracle
about twice diameter of remainder. Integument of entire venter and of dorsal
surfaces of posterior somites with minute spinules in short transverse rows, else¬
where spinules minute and isolated. Body hairs sparse and short. Of two types:
(1) 0.044-0.125 mm long, curved and denticulate, on ventral and lateral surfaces
of thorax and AI-AVIII and on all surfaces of AIX and AX: (2) about 0.14 mm
long, smooth, deeply bifid, branches strongly divergent and tips curled, on dorsal
and lateral surfaces of all somites except AIX and AX. Cranium subhexagonal:
frons and clypeus bulging. Antennae small, each with three sensilla, each bearing
one rather long spinule. Head hairs few, 0.03-0.15 mm long, smooth or with minute
denticles. Labrum feebly bilobed; narrowed dorsally; each lobe with three minute
hairs and—medially—minute spinules in short transverse rows on anterior sur¬
face, with one isolated and two contiguous sensilla and numerous spinules on
ventral border, with four isolated and three contiguous sensilla on posterior sur¬
face; entire posterior surface densely spinulose, spinules minute and in numerous
Vol. 49, No. 3, July 1973
211
short transverse rows. Mandibles leptothoraciform (i.e., moderately narrow; taper¬
ing gradually and curving gradually to apical tooth; anterior surface produced
medially into blade which bears two subapical teeth), the teeth large and directed
medially. Maxillae large, lobose and adnate; each palp a skewed peg with four
apical and one lateral sensilla; each galea digitiform, with two apical sensilla.
Labium thick; anterior surface spinulose, spinules minute and in transverse rows;
each palp a short frustum, with four apical and one lateral sensilla; isolated sen-
sillum between each palp and opening of sericteries, latter a transverse slit. Hypo-
pharynx densely spinulose, spinules rather long and in numerous subtransverse
rows, rows so close together that spinules overlap. (Material studied: eight larvae
from New South Wales, courtesy of Rev. B. B. Lowery.)
Genus Daceton Petty
Brown and Wilson (1959: 290 ).—“Daceton possesses, in addition
to truly primitive features, characters that appear to represent signif¬
icant specializations away from the main line of dacetine evolution, viz.,
in sculpturing, worker polymorphism, cephalic articulation, and larval
morphology (see Brown 1953a; and Wheeler and Wheeler, 1954).”
Daceton armigerum (Latreille)
The mature workers of this species are highly polymorphic; the
mature worker larvae range from 7 mm to 12 mm in length (through
spiracles). Sexual larvae are 12.3—12.9 mm (through spiracles); the
head hairs about twice as numerous, otherwise very similar to the
worker larva. (Material studied: numerous larvae from Buenos Aires—
25 km S. of Pucallpa, Peru.) Wilson (1962: 413-414) discussed brood
care, feeding, and transport in this species.
Genus Epopostruma Forel
Epopostruma alata Forel
Length (through spiracles) about 3.3 mm. Very similar to E. sp. (1954: 128),
except in following details. Integument of venter and of dorsal surface of posterior
somites with minute spinules in short transverse rows, elsewhere spinules shorter
and less numerous. Body hairs: (1) 0.038-0.125 mm long; (2) 0.075-0.163 mm
long. Head hairs longer (0.075-0.175 mm long). Mandibles stouter and with
medial teeth larger and directed more nearly medially. (Material studied: eight
larvae from New South Wales, courtesy of Rev. B. B. Lowery.)
Epopostruma frosti (Brown)
Length (through spiracles) about 5.3 mm. Very similar to E. sp. (1954: 128)
except as follows. Thorax and AI more slender. Anus with posterior lip. Integu¬
ment of venter of neck and of dorsal surface of posterior somites with minute
spinules in short transverse rows. Body hairs more numerous. Of three types:
212
The Pan-Pacific Entomologist
(1) 0.05-0.125 mm long, few, with single shaft and few denticles, on venter of
anterior somites; (2) 0.075-0.15 mm long, with denticulate hooked tip, on re¬
mainder of venter of abdomen; (3) 0.05-0.4 mm long, with short-bifid tip, branches
sparsely denticulate, on dorsal and lateral surfaces of all somites. Cranium with
straight sides. Head hairs 0.03-0.125 mm long, usually with few denticles. Man¬
dibles with apex more curved, teeth larger and more widely separated. (Material
studied: 14 larvae from South Australia, courtesy of Rev. B. B. Lowery.)
Epopostruma quadrispinosa For el
Length (through spiracles) about 4.0 mm . Very similar to E. sp. (1954: 128)
except as follows. Body hairs (1) 0.05-0.125 mm long, unbranched, with hooked
tip, on venter of each somite: (2) 0.05-0.125 mm long. Head hairs 0.025-0.125
mm long. Mandibles with apex more curved, teeth larger and more widely sep¬
arated. (Material studied: 15 larvae from New South Wales, courtesy of Rev.
B. B. Lowery.)
Genus Mesostruma Brown
Profile aspididriform. Body hairs sparse. Of two types: (1) with single den¬
ticulate shaft; (2) deeply bifid, with branches denticulate. Head hairs sparse
and smooth or with a few denticles. Mandibles leptothoraciform, with two medial
teeth.
In our 1960b key Mesostruma would run to Alistruma from which
it can not be separated at present. We are giving below a complete
description of M. browni , because we regard M. laevigatus, which we
described (1954: 130), as a sexual larva.
Mesostruma browni Taylor
(Fig. 3)
Length (through spiracles) about 4.0 mm. Profile aspididriform (i.e., moderately
stout; no neck, but thorax and first two abdominal somites strongly curved ven-
trally; diameter greatest at AV, decreasing to AI, then increasing slightly to T2,
decreasing rapidly to diameter of head; dorsal profile C-shaped, ventral J-shaped;
anus ventral). Anus without lip. Spiracles small, T2 largest. Integument of venter
of anterior somites and dorsa of posterior somites with minute spinules in short
transverse rows. Body hairs sparse. Of two types: (1) 0.038-0.125 mm long, on
all surfaces of AX and on venter of remaining somites, single shaft with minute
denticles; (2) 0.063-0.15 mm long, deeply bifid, with branches denticulate, on
dorsal and lateral surfaces of all somites except AX. Cranium subhexagonal in
anterior view, as broad as long; occiput feebly concave. Antennae very small,
each with three sensilla, each bearing a rather long spinule. Head hairs few,
0.025-0.125 mm long, smooth or with few denticles. Labrum small; breadth twice
length; subrectangular, with ventral comers rounded, medial border feebly con¬
cave; each lobe with three or four sensilla on anterior surface near ventral border,
with cluster of three sensilla on ventral border and with three contiguous sensilla
on posterior surface; entire posterior surface spinulose, spinules minute, dorsal
stouter and ventral finer. Mandibles leptothoraciform (i.e., moderately narrow;
Vol. 49, No. 3, July 1973
213
tapering gradually and curving gradually to apical tooth; anterior surface pro¬
duced medially into blade with two subapical teeth). Maxillae short and lobose;
palp chair-shaped, with one lateral (bearing spinule) and four apical (two en¬
capsulated and two bearing one spinule each) sensilla; galea a frustum with two
apical sensilla. Labium with numerous short rows of minute spinules on anterior
surface; palps similar to maxillary palps, but shorter; isolated sensillum between
each palp and opening of sericteries; the latter a transverse slit. Hypopharynx
with few short arcuate rows of spinules. (Material studied: six larvae from Can¬
berra, ACT, courtesy of Rev. B. B. Lowery.)
Genus Neostruma Brown
Neostruma mustelina (Weber)
Brown 1959; 9—“In one natural nest, an entomobryid was found
with larvae feeding on it.”
Genus Orectognathus Mayr
Orectognathus antennatus F. Smith
Length (through spiracles) about 4.2 mm. Similar to 0. clarki (1954: 126)
except as follows. Neck more slender, remainder of abdomen more swollen. Body
hairs (1) 0.05-0.2 mm long, longer with fewer denticles and tending toward
single hook dorsally, ventrally shorter and more denticulate, flexuous; (2) 0.06-
0.15 mm long, few, deeply bifid, with few denticles on each branch. Cranium sub-
rectangular, with dorsal border feebly concave. Head hairs shorter (0.025-0.125
mm long). Labrum broader ventrally; ventral border of each lobe with two iso¬
lated and two contiguous sensilla. Each mandible with apical tooth sharper and
medial teeth larger. (Material studied: seven larvae from New South Wales,
courtesy of Rev. B. B. Lowery.)
Orectognathus mjobergi Forel
Length (through spiracles) about 4.2 mm. Similar to O. clarki (1954: 126)
except as follows. Body hairs of two types: (1) 0.025-0.075 mm long, bifid with
branches denticulate, on dorsal and lateral surfaces of all somites; (2) 0.06-0.24
mm long, with single denticulate shaft (sometimes with denticulate side branch),
largely confined to the venter but with transverse band around middle of each
somite. Cranium subrectangular, with occiput feebly concave. Entire posterior
surface of labrum spinulose, spinules minute and in short transverse rows. (Ma¬
terial studied: numerous larvae from New South Wales, courtesy of Rev. B. B.
Lowery.)
Orectognathus nigriventris Mercovich
Immature larva. —Length (through spiracles) about 2.5 mm. Similar to O.
clarki (1954: 126) except as follows. Body hairs of three types: (1) 0.05-0.25 mm
long, shortest ventrally, slightly curved to flexuous, with minute denticles; (2)
0.1-0.3 mm long, deeply bifid, few on dorsal and lateral surfaces of each somite;
214
The Pan-Pacific Entomologist
(3) about 0.3 mm long, uncinate, with flexuous shaft, two to six on T2-AX. Head
very large. Head hairs 0.025-0.275 mm long, slightly curved and smooth or with
few denticles. (Material studied: numerous immature larvae from New South
Wales, courtesy of Rev. B. B. Lowery.)
Orectognathus rostratus Lowery
Length (through spiracles) about 3.5 mm. Similar to 0. clarki (1954: 126)
except as follows. Integument of venter of anterior somites and of dorsum of
AX with minute spinules in short transverse rows. Body hairs (1) 0.06-0.18 mm
long, bifid, with denticles; (2) 0.025-0.23 mm long, on ventral surface; (3) about
0.3 mm long, with stout shaft and small single hook, two on dorsum of each AI-AV.
Cranium subrectangular hut with occipital border feebly concave. Head hairs
0.025-0.175 mm long, slightly curved and usually with few denticles. (Material
studied: six larvae from New South Wales, courtesy of Rev. B. B. Lowery.)
Genus Strumigenys F. Smith
Strumigenys perplexa F. Smith
(Fig. 4)
Length (through spiracles) about 1.9 mm. Similar to S. louisianae (1954: 136)
except as follows. Body hairs shorter: (1) 0.025-0.06 mm long, on venter of each
somite; (2) 0.038-0.1 mm long; (3) about 0.15 mm long. Head hairs shorter
(0.013-0.075 mm long). Labrum with breadth only l 1 /-* times length. Mandibles
with all teeth blunter, medial teeth small, close together and divergent. Maxillary
palp represented by cluster of three sensilla. Each labial palp represented by cluster
of four sensilla. (Material studied: numerous larvae from New South Wales,
courtesy of Rev. B. B. Lowery.)
Literature Cited
Brown, W. L. 1959. A revision of the dacetine ant genus Neostruma. Breviora,
No. 107, 13 p.
Brown, W. L., and W. W. Kempf. 1960. A world revision of the ant tribe
Basicerotini. Stud. Entomol., 3: 161-250.
Brown, W. L., and E. 0. Wilson. 1959. The evolution of the dacetine ants.
Quart. Rev. Biol., 34: 278-294.
Mann, W. M. 1922. Ants from Honduras and Guatemala. Proc. U. S. Nat. Mus.,
61: 1-54.
Wheeler, G. C. and J. Wheeler. 1954. The ant larvae of the myrmicine tribes
Basicerotini and Dacetini. Psyche, 61: 111-145.
1960a. Supplementary studies on the larvae of the Myrmicinae. Proc. Entomol.
Soc. Wash., 62: 1-32.
1960b. The ant larva of the subfamily Myrmicinae. Ann. Entomol. Soc. Amer.,
53: 98-110.
1969. The larvae of Acanthognathus. Psyche, 76: 110-113.
Wilson, E. O. 1962. Behavior of Daceton armigerum (Latreille), with a clas¬
sification of self-grooming movements in ants. Bull. Mus. Comp. Zool.
Harvard Coll., 127 : 404^421.
Vol. 49, No. 3, July 1973
215
The Genus Polynemoidea Girault
(Hymenoptera: Mymaridae)
Richard L. Doutt 1
University of California, Berkeley
The finding of a fossil species of Polynemoidea in Oligocene amber
from Chiapas, Mexico, and the recognition of a synonymous genus
from Tasmania has motivated this synopsis.
Polynemoidea Girault
Polynemoidea Girault, A. A. 1913. Mem. Queensland Mus., 2: 116.
Selenaeus Waterhouse, C. 0. 1914. Trans. Entomol. Soc. London, 1914 pp. 536-
537. (New synonymy ).
Type of genus: Polynemoidea varicornis Girault
Polynemoidea varicornis Girault, A. A. 1913. Mem. Queensland Mus., 2: 116.
Selenaeus turneri Waterhouse, C. 0. 1914. Trans. Entomol. Soc. London, 1914,
p. 537. (New synonymy ).
The type species, P. varicornis Girault, was described from a female
collected at Hobart, Tasmania. I have not seen this specimen which is
reportedly in the South Australian Museum at Adelaide under type
number I. 1228. However, I have examined two female specimens in
the Girault collection at the Queensland Museum, Brisbane. These are
together on one slide. The labels are in Girault’s handwriting and one
reads U Polynemoidea varicornis Girault 2 Cotype.” The word “Type”
is above the word “Cotype” but is crossed out. The other label reads
“Bred from wood. Hobart, Tas. H. W. Lea.” Since a type was designated
it is highly probable that these two female specimens are paratypes.
They are from the same locality (Tasmania) as the type and certainly
fit the original description.
The species P. varicornis is very distinct from the other members of
the genus in several respects. The scape is serrated; funicle segments
2, 3 and 4 are remarkably elongate and much lighter in color than
other antennal segments (Fig. 1). The ovipositor is greatly exserted
(Fig. 2). The first tarsal segment is especially long (Fig. 3), and the
forewings tend to be pointed (Fig. 4). In body size it is the largest
species in the genus.
It is interesting that a single mymarid female also collected in Tas¬
mania was described as Selenaeus turneri Waterhouse. This specimen
1 Present address : 9240 S. Riverbend Avenue, Parlier, California 93648.
The Pan-Pacific Entomologist 49: 215-220. July 1973
216
The Pan-Pacific Entomologist
Figs. 1-4. Polynemoidea varicornis. Fig. 1. Antenna. Fig. 2. Body pro¬
portions. Fig. 3. Fore tibia and tarsi. Fig. 4. Forewing.
is in the British Museum (Natural History), London, and is identical
with P. varicornis Girault. This synonymous species was thought by
the collector to be associated with some homopterous galls, but there
is so far no definite host relationship established for any species in
the genus Polynemoidea.
Polynemoidea lincolni Girault
Polynemoidea lincolni Girault, A. A. 1913. Mem. Queensland Mus. 2: 116-117.
The type specimen is a single female collected by sweeping jungle
vegetation, Nelson (Cairns) Queensland, July 10, 1913 (A. P. Dodd).
The original description is accurate except that all the funicle segments
are dark and segments 1 and 2 are subequal (Fig. 5). The ovipositor
originates at the base of the abdomen and is slightly exserted. The
wings are broad (Fig. 9). The thorax is without sculpturing and the
dark brown or black mesoscutum strikingly contrasts with the yellow
scutellum and propodeum. The eyes and head are black. The mandibles
have two teeth and a blunt truncation.
Vol. 49, No. 3, July 1973
217
Figs. 5-8. Antennae. Fig. 5. Polynemoidea lincolni. Fig. 6. P. particoxae.
Fig. 7. P. domestica. Fig. 8. P. mexicana.
Polynemoidea particoxae Girault
Polynemoidea particoxae Girault, A. A. 1938. Rev. Entomol. (Rio de Janeiro),
9: 390-391.
This species is related to P. lincolni but funicle segments 1 and 2 are
somewhat more slender (Fig. 6) and the forewing is narrower (Fig. 10).
The head is black, eyes purplish black. The mandibles have 3 distinct
teeth. The scutum and scutellum are black, the propodeum is yellow.
The forecoxae are dark but the middle and hind pairs are light, tro¬
chanters pale, femora dark, tibia and tarsi light. The abdomen is short,
dark; the ovipositor originates at base of abdomen and is only slightly
exserted. Both P. lincolni (Fig. 9) and P. particoxae (Fig. 10) have
forewings with long, slender marginal and stigmal veins.
218
The Pan-Pacific Entomologist
Figs. 9-12. Forewings. Fig. 9. Polynemoidea lincolni. Fig. 10. P. particoxae.
Fig. 11. P. incerta. Fig. 12. P. domestica.
Vol. 49, No. 3, July 1973
219
POLYNEMOIDEA INCERTA Girault
Polynemoidea incerta Girault, A. A. 1938. Rev. Entomol., Rio de Janeiro, 9: 391.
The type specimen is a fragmented male. The antennal clubs are
missing. There are eleven flagellar segments present on one antenna.
The marginal vein is extraordinarily long for this genus (Fig. 11).
This single male is doubtfully retained in this genus although it does
belong in the Tribe Anaphini.
Polynemoidea domestica Girault
Polynemoidea domestica Girault, A. A. 1931. A new habit in an old insect, Homo
pudicus and new Eurytomidae. Privately published, Brisbane, Australia, 4 pp.
(p. 3).
The type specimen is a single fragmented female collected on a win¬
dow of Girault’s home, Indooroopilly, Queensland, 1929. The body
is dark brown, legs lighter brown. Eyes crimson. Antenna as in Fig. 7,
scape slender. Scutum with distinct reticulate sculpturing. Wings with
short marginal and stigmal veins (Fig. 12).
Polynemoidea mexicana Doutt, new species
Female. —Body, viewed laterally, compressed, hypogynium absent. Thorax
elongate, about equal to abdomen. Color golden brown. Head with very short
dimension on anterior-posterior axis, but very long dimension on dorsal-ventral
axis. Antennae as in Fig. 8. Scape moderate length, normal; pedicel somewhat
enlarged, elongate. Six funicle segments, first is smallest, increasing in width
distally. Club distinctly 3-segmented, long. One antenna with segments somewhat
compressed (possibly artifact of fossilization process). Legs normal, wing char¬
acteristics mostly hidden by bubbles and fractures in amber block, but discal
ciliation similar to P. domestica. Ovipositor exserted but not remarkably so, thus
similar to P. domestica, but with different antennal characters.
Holotype female , Las Cruces landslide, Chiapas, Mexico.
Described from single female embedded in amber. This specimen
is at the University of California Museum of Paleontology and has the
type number 12865, loc. B-5104 (1957). The B-5104 amber is from
the Las Cruces landslide, Chiapas, Mexico. This locality is about 23 km
(airline) east-southeast of the town of Simojovel, about 6.8 km south¬
east of the headquarters of the Rancho Santo Domingo, about 1.4 km
northwest of Rancho San Jose de Buenavista. The landslide is on the
south slope, near the southeast end of the major ridge known locally
as Nichcalan (also as Cerro Balumtun). The amber was found in a
dense blue limestone near the base of the slide. The marine invertebrate
fauna from adjacent beds indicates an age in the range from late Oli-
220
The Pan-Pacific Entomologist
gocene to early Miocene. (This locality description from Fennah, R. G.
1963. U. C. Publ. Entomol. 31(1) : 45.)
Key to Females of Australian Species of Polynemoidea
1. Ovipositor very long, scape asperate, basal tarsal segments much longer than
any succeeding segments _ varicornis Girault
Ovipositor not long, only slightly exserted, scape not asperate_ 2
2. Thorax with coarse, reticulate sculpturing, thorax entirely dark brown,
ovipositor originating in middle of abdomen _ domestica Girault
Thorax without coarse, reticulate sculpturing, thorax black and yellow,
ovipositor originates at base of abdomen_ 3
3. Mandibles with three distinct teeth, mesoscutum and scutellum black, meta¬
thorax yellow, forecoxae dark, middle and hind coxae pale _
_ particoxae Girault
Mandibles with two teeth and blunt truncation, mesoscutum black, scutellum
and propodeum yellow, all coxae pale _ lincolni Girault
RECENT LITERATURE
A series of paperbacks dealing with environmental problems have been received
from the publisher, Houghton Mifflin Company, Boston (no price information
was included). The first three are collections of reprinted articles by many en¬
vironmentalists including such well known ecologists and entomologists as: J.
Ralph Audy, Barry Commoner, Paul R. Ehrlich, Frank E. Engler, and Eugene P.
Odum. The fourth book deals with the DDT controversy.
Environ/mental. Essays on the Planet as a Home. Edited by Paul Shepard
and Daniel McKinley. 308 p., illus., 1971.
The emphasis of this book is on the human overpopulation problem and its
social and psychological effects on man. Each of the 20 reprinted articles is
preceded by a brief introduction by the editors and concludes with a set of ref¬
erences. The book concludes with biographical notes on the authors and an ex¬
tensive list of additional readings.
The Politics of Neglect: The Environmental Crisis. Edited by Roy L. Meek
and John A. Straayer. 256 p., 1971.
The 28 reprinted articles are arranged into six chapters considering problems
from population to pollution. Each chapter is preceded by an introductory state¬
ment by the editors. Tables, footnotes, figures and references are omitted from
the articles, but a terminal bibliography of general references is provided.
Vol. 49, No. 3, July 1973
221
The Fossil Mymariclae
(Hymenoptera: Chalcidoidea)
Richard L. Doutt 1
University of California, Berkeley
The Mymaridae are very minute wasps which develop solely as pri¬
mary parasitoids within the eggs of other insects. The fossilized speci¬
mens found in amber are very similar morphologically to species living
today, and it is probable that the fundamental habits of the mymarids
as parasitoids have remained unchanged for at least 30,000,000 years.
These small insects are frequently found entangled in sticky material
placed on boards suspended in trees, so it is understandable how their
general search for hosts in arboreal habitats could easily have led to
occasional entrapment in gums or resins, and thus eventually to fossil-
ization in amber.
The Mymaridae were first recognized as fossils in amber by Loew
in 1847. They have been found in Canadian (Cretaceous), Baltic (early
Oligocene), Mexican (late Oligocene and early Miocene), and Sicilian
(Miocene) ambers. Species have also been described from the recent
copal of Madagascar and Zanzibar. This paper includes comments
on all of the known fossil members of this interesting family of chalcids,
but it is primarily concerned with a series of Mymaridae found in late
Oligocene or early Miocene amber from the Simojovel area, Chiapas,
Mexico. According to Langenheim (1969) the leguminous genus Hy-
menaea is the probable source of the Chiapas amber. The modern
species, H. courbaril , is known for its abundant secretion of resin that
accumulates in the soil around the tree. Langenheim states that in
Mexico today, //. courbaril grows commonly along rivers that enter
the ocean in mangrove-fringed estuaries. It is believed that the Chiapas
amber was deposited in such an estuarine environment.
According to the classification proposed by Annecke and Doutt
(1961), the family Mymaridae consists of two subfamilies and five
tribes. Both subfamilies are well represented in the fossil record (Ap¬
pendix). However, most of the fossil species in each subfamily are in
a single tribe. In each case, Alaptini and Ooctonini, all members of
the tribe are characterized by possessing five tarsomeres. This is a
more primitive condition than the four tarsomere formula exhibited
by members of the other tribes in each subfamily. Of the 16 species
■’■Present address: 9240 S. Riverbend Avenue, Parlier, Ca. 93648.
The Pan-Pacific Entomologist 49: 221-228. July 1973
222
The Pan-Pacific Entomologist
recognized in the fossil record only Alaptus globosicornis Girault,
Alaptus psocidivorus Gahan, and Palaeomymar duisburgi (Stein)
are known to be extant.
Subfamily Alaptinae
Genus Alaptus Westwood
The species of Alaptus are noteworthy because of their minuteness.
Adult specimens of these wasps usually do not exceed 0.3 mm in total
length, and are among the smallest of all known insects. The few species
of Alaptus which have been studied biologically develop within the
eggs of Psocoptera.
In 1909 Meunier found and described an Alaptus in recent copal
from Zanzibar. This first account of a fossilized Alaptus is based on
a single male specimen for which Meunier proposed the name Alaptus
fructuosus. It is difficult to compare this species with existing forms
because Meunier’s description is inadequate in certain critical details,
and our present system of classification is based primarily on female
rather than on male characters. Therefore the correct placement of
any Alaptus male at the species level is largely speculative.
The series of Mymaridae in the Simojovel amber contains at least
two species of Alaptus. There are four female specimens (University
of California Museum of Paleontology numbers 12773, B-1402; 12778,
B-1402; UCMP 12631, B-4112; and UCMP 12632, B-4117) which are
indistinguishable from Alaptus psocidivorus Gahan, a species which
exists today in California as a parasite of the psocid, Peripsocus cal-
ifornicus Banks, on Toyon, Heteromeles arbutifolia M. Roem.
One female Alaptus (UCMP 12897, B-5106) is morphologically very
close to the species A. globosicornis Girault. Although the first funicle
segment is longer than typical A. globosicornis , and the general habitus
is less compact, these differences are insufficient to justify naming this
single specimen as a representative of a new species. This conservative
view is justified because A. globosicornis appears to be a highly vari¬
able species, and Girault (1912) not only indicated that modifications
in the original description of the species were necessary but that at
least two “varieties 55 existed, which he termed hawaiiensis and austra-
liensis. One specimen in the present series of fossil Alaptus (UCMP
12633, B-7043) is difficult to place, but it is probably also A. globosi¬
cornis or closely related.
There are five male Alaptus in the Simojovel amber that seem to
represent two species. Two specimens are very dark colored and have
Vol. 49, No. 3, July 1973
223
comparatively short antennal segments (UCMP 12898, B-5106 and
UCMP 12634, B-7041). The other specimens (UCMP 12625; UCMP
12636, B-4112; and UCMP 12638, B-4112) are light brown in color with
more elongate legs and antennae. It is possible that these males relate re¬
spectively to A. globosicornis and A. psocidovorus, but there is no
way at present to associate them with certainty.
Genus Litus Haliday
Meunier (1901) described Litus elegans from a female found in
Baltic (early Oligocene) amber, and later (1909) he described Litus
bene ficus from the recent copal of Madagascar. The Simojovel amber
contains one female Litus which is distinctly different from Meunier’s
species, and is also unlike any of the species which are known to exist
today. It is described as follows:
Litus mexicanus Doutt, new species
Female. —Length 0.54 mm. Antenna as in Fig. 1. Scape long, moderately ex¬
panded, funicle segments heavily setaceous, segments 2 to 6 each with single
short paddle-shaped structure located dorsally and apically, club large, elongate,
with many short setae. Head distorted in this specimen but ommatidia large. Con¬
dition of thorax precludes analysis of surface sculpturing. Basitarsal segment
of foreleg long, strigil prominent, calcar large, stout, curved and bifid. Middle
legs long and slender. All tibiae with strong setae. Forewings broken but general
shape and venation normal for genus. Hindwing maculate, with single line of
small discal setae. Ovipositor originates at base of abdomen, large, prominent,
markedly exserted.
The expanded scape, paddle-shaped sensoria, and the strongly de¬
veloped and exserted ovipositor distinguish L. mexicanus from other
species of Litus.
The type specimen is in the University of California Museum of
Paleontology, Berkeley, with the UCMP number 12812, B.-7041. From
Oligocene amber, Chiapas, Mexico.
Species of Litus occur today in South America and in the Caribbean
area, but this fossil is the first record from Mexico. The true host
relationships of Litus are not known, although L. cynipseus Haliday
was found in a colony of the ant, Lasius niger , where it may have been
parasitizing eggs of coleopterous myrmecophiles.
Genus Malfattia Meunier
Meunier (1901) described this genus from a single specimen with
four segmented tarsi and nine segmented antennae. The genus has
224
The Pan-Pacific Entomologist
been synonymized with Litus by Ashmead (1904) although he doubted
whether this was correct. The four segmented tarsi would make such
placement erroneous. Meunier believed that his specimen was related
to one from Sicilian amber discussed by Malfatti, and accordingly
named it in his honor. Malfattia remains as a very poorly known and
doubtfully placed genus until Meunier’s specimen can be re-examined.
Subfamily Mymarinae
Genus Lymaenon Walker
The species Lymaenon henneberti (Meunier) described from Baltic
amber in 1905 is the only known fossil in this genus. There are many
modern species in Lymaenon (formerly Gonatocerus) , and they attack
a wide variety of hosts in many different habitats.
Genus Arescon Walker
Meunier (1901, 1905) described two species of fossil Mymaridae in
the genus Leimacis, but properly they should be placed in the genus
Arescon. Both species, Arescon armata (Meunier) and A. baltica
(Meunier) are from Baltic amber. Nothing is known of the host re¬
lationships of our modern species.
Genus Ooctonus Haliday
Brues (1937) found a Mymarid in Canadian amber which he de¬
scribed as Ooctonus (?) minutissimus but gave no indication why he
believed the generic placement to be doubtful. This specimen is Cre-
Vol. 49, No. 3, July 1973
225
taceous, and is therefore the oldest known fossil member of the family.
Ooctonus is reared from eggs of leafhoppers and is not considered to
be a primitive form.
Genus Palaeomymar Meunier
Palaeomymar Meunier, 1901. Ann. Soc. Sci. Bruxelles. 25(2) : 288-289.
Type: Palaeomymar duisburgi (Stein) 1877, Mitt. Miinchen entomol. Ver.
1: 30. New Combination.
Palaeomymar succini Meunier, 1901, Ann. Soc. Sci. Bruxelles. 25(2) :
289.
Petiolaria anomala Blood and Kryger, 1922. Entomol. Mon. Mag. 58:
229.
Mymaromma Girault, 1920. Insec. Inscit. Menstr. 8: 38. New synonym.
Type: Mymaromma goethei Girault.
Petiolaria Blood and Kryger, 1922. Entomol. Mon. Mag. 58: 229. New synonym.
Type: Petiolaria anomala Blood and Kryger.
Mymaromella Girault, 1931. A new habit in an old insect, Homo pudicus and new
Eurytomidae. Privately published, Brisbane, Sept. 1, 1931. p. 4 New
synonym.
Type. — Mymaromella mira Girault.
The above synonymy is solely the result of this comparative study
of fossil Mymaridae, and it has the virtue of solving the problem of the
correct placement of Palaeomymar which has been troubling students
of Mymaridae for many years. The riddle began in 1868 when Duisburg
described and illustrated, but did not name, a female Mymarid in amber
which he believed belonged to the genus Mymar. Later Stein (1877)
in studying another specimen in amber believed that it agreed in all
the essentials with Duisburg’s insect and accordingly named it Mymar
Duisburgi.
In 1901 Meunier placed M. Duisburgi Stein into the genus Eustochus ,
created the new genus Palaeomymar for five specimens of the series
discussed by Duisburg in 1868, and stated that representatives of the
genus Mymar were not known in the fossilized state.
In 1922 Blood and Kryger discovered a modern mymarid of a very
unusual type in England and Denmark. They designated it as a new
genus and species, namely Petiolaria anomala Blood and Kryger. They
were not aware that the same genus occurred in Australia and had been
described there two years earlier as Mymaromma by Girault.
In 1948 Bakkendorf independently found the 1868 paper by Duisburg
and correctly concluded that the amber insects seen and described
therein were conspecific with Petiolaria anomala Blood and Kryger.
Bakkendorf did not mention the works of Stein (1877) nor of Meunier
(1901), but his recognition of Duisburg’s specimen as P. anomala
226
The Pan-Pacific Entomologist
was correct and has furnished the clue to the present placement of
Palaeomymar. There is now ample evidence that the species pictured
by Duisburg (1868), described and named by Stein (1877), discussed
by Meunier (1901) under Eustochus, and identified by Bakkendorf
(1948) as Petiolaria anomala Blood and Kryger is actually a single
species, based on characters of the female sex. The males of this same
species were not recognized as such by Meunier who instead proposed
the genus Palaeomymar for them.
It is very probable that Meunier’s specimens of Palaeomymar are
identical with the males of Mymaromma ( Petiolaria ) anomala. One
is especially convinced of this when Debauche’s (1948) illustrations
of the structural characters of Mymaromma are compared with the
figures of the same structures published by Meunier (1901). The male
antennae of Mymaromma are distinctive and these same unique fea¬
tures are seen in the antennae figured by Meunier for Palaeomymar.
Similarly, the female antennae of Mymaromma anomala with its char¬
acteristic seven segmented funicle is mirrored in the figure given by
Meunier for what he termed Eustochus duisburgi. Although Meunier’s
drawing of the forewing of Palaeomymar is not the best example of
entomological draftsmanship it is nevertheless unmistakably the wing
of the insect which has been known as Mymaromma.
The foregoing evidence seems sufficient to justify the proposed
synonymy. It is of interest that Palaeomymar is well represented by a
large series of specimens in the Chiapas amber at the University of
California Museum of Paleontology. These specimens deserve very
careful study.
Genus Anaphes Haliday
This genus contains two fossil representatives which were described
by Meunier (1901) from Baltic amber. These are Anaphes schell -
wieniens and A. splendens.
The Mexican amber contains a male which is probably an Anaphes.
This lone male in the Simojovel material can be placed only tentatively
because the classification of the group is based upon female characters.
The specimen with the UCMP number 12640, B-7046 is very dark,
which is characteristic of many Anaphes , and it has the general habitus
and wings of that genus.
Genus P olynemoidea Girault
Until its discovery in the Chiapas amber, no member of this genus
had ever been found outside the southern hemisphere, and it is a genus
Vol. 49, No. 3, July 1973
227
that is seldom seen by mymarid specialists today. The five extant spe¬
cies are all Australian, and the closely allied genera such as Idiocentrus
and Notomymar are also antipodal. Thus the specimen in the Simojovel
series is of considerable interest, and was described as Polynemoidea
mexicana Doutt (1973). It bears the UCMP number 12865, B-5104.
Acknowledgment
I am grateful to Joseph H. Peck, Jr., Senior Museum Scientist, Uni¬
versity of California Museum of Paleontology, for his considerable
assistance with specimen data. This material was collected and prepared
under NSF Grants NSF-G10705; NSF-G 20016.
Appendix
List of fossil Mymaridae with amber sources.
Alaptinae:
Alaptini
Alaptas fructuosus Meunier, 1909. Zanzibar copal, recent.
A. globosicornis Girault, 1912. Mexican, late Oligocene, early Miocene.
A. psocidivorus Gahan, 1927. Mexican, late Oligocene, early Miocene.
Litus beneficus Meunier. 1909. Madagascar copal, recent.
L. elegans Meunier. 1901. Baltic, early Oligocene.
L. mexicanus Doutt. 1973. Mexican, late Oligocene, early Miocene.
Anagrini
Malfattia Molitorae Meunier. 1904. Baltic, early Oligocene; Sicilian, Miocene.
(This species is doubtfully placed in Anagrini)
Mymarinae:
Ooctonini
Arescon baltica (Meunier). 1901. Baltic, early Oligocene.
A. armata (Meunier). 1905. Baltic, early Oligocene.
Lymaenon henneberti (Meunier). 1905. Baltic, early Oligocene.
Ooctonus minutissimus Brues. 1937. Canadian, Cretaceous.
Palaeomymar duisburgi (Stein). 1877. Baltic, early Oligocene.
Palaeomymar sp. (undetermined). Mexican, late Oligocene, early Miocene.
Anaphini
Anaphes schellwieniens Meunier. 1901. Baltic, early Oligocene.
A. splendens Meunier. 1901. Baltic, early Oligocene.
Polynemoidea mexicana Doutt. 1973. Mexican, late Oligocene, early Miocene.
Anaphes sp. (undetermined). Mexican, late Oligocene, early Miocene.
Literature Cited
Annecke, D. P. and R. L. Doutt. 1961. The genera of the Mymaridae. Hy-
menoptera: Chalcidoidea. S. Afr. Dep. Agr. Tech. Serv. Entomol. Mem.,
5: 1-71.
Ashmead, W. H. 1904. Classification of the chalcid flies. Mem. Carnegie Mus.,
1(4) : 225-555.
228
The Pan-Pacific Entomologist
Bakkendorf, 0. 1948. A comparison of a mymarid from Baltic amber with a
recent species, Petiolaria anomala (Micro-Hym.). Entomol. Medd.,
25: 213-218.
Blood, B. N. and J. P. Kryger. 1922. A new mymarid from Brockenhurst.
Entomol. Mon. Mag., 8: 229-230.
Brues, C. T. 1937. Insects and arachnids from Canadian amber. Univ. Toronto,
Geol. Ser., 40: 44.
Debauche, H. R. 1948. Etude sur les Mymarommidae et les Mymaridae de la
Belgique (Hymenoptera Chalcidoidea). Mem. Mus. Roy. Hist. Natur.
Bruxelles 108: 1-248.
Doutt, R. L. 1973. The genus Polynemoidea (Hymenoptera: Mymaridae).
Pan-Pac. Entomol., 49(3) : In press.
Duisburg, V. 1868. Zur Bernstein-Fauna. Schrift. Konigl. Phys.-Okon. Gesellsch.
Konigsberg., 9: 23-28.
Girault, A. A. 1912. Australian Hymenoptera Chalcidoidea—I. Mem. Queens¬
land Mus., 1: 123-124.
Langeniieim, J. H. 1969. Amber: a botanical inquiry. Science, 163(3872) :
1157-1169.
Meunier, F. 1901. Contribution a la Faune des Mymaridae “ou Atomes Ailes”
de l’Ambre. Ann. Soc. Sci. Bruxelles, 25: 282-292.
1905. Sur deux Mymaridae de l’Ambre de la Baltique. Misc. Entomol., Rev.
Entomol. Int., 13(1) : 1-4.
1909. Sur deux Mymarinae du copal recent de Madagascar et de Zanzibar.
(Hym.). Bull. Soc. Entomol. France, 1909, pp. 145-149.
Stein, J. P. E. F. 1877. Drei merkwiirdige Bemstein-Insekten. Mitt. Munch.
Entomol. Ver., 1: 28-30.
BOOK REVIEW
Australian Butterflies. Charles McCubbin. Thomas Nelson (Australia) Ltd.,
Melbourne. 206 p., fully illustrated in color, xxxi. 1971. $30.00. North Amer¬
ican Distributor: Entomological Reprint Specialists.
This quarto size book is fully and beautifully illustrated with water color paint¬
ings by the author of the butterflies (Papillionoidea) and skippers (Hesperioidea)
of Australia. Among the most valuable and distinctive aspects are the many color
illustrations of immatures, food plants and habitats as well as descriptions of life
history and behavior of these Australian Lepidoptera.
The introductory chapter includes valuable information for the amateur on
classification, the position of Lepidoptera among insects, the general life history
of butterflies and skippers, collecting and curating methods and materials.
This is an extremely important butterfly book with special appeal to all inter¬
ested in exotic Lepidoptera and their natural history. —Robbin W. Thorp, University
of California, Davis, 95616.
Vol. 49, No. 3, July 1973
229
Macrobiotus montanus from California
(Tardigrada: Macrobiotidae)
Albert A. Grigarick, Robert 0. Schuster and Elizabeth C. Toftner
University of California, Davis 95616
Macrobiotus montanus Murray, 1910, previously unknown from
California, was collected at Auburn, Placer County, California. It
was the most common species of Macrobiotus occurring in the moss
Pseudobraunia californica (Lesg.) Broth, and the lichen Parmelia
flaventior Stirton, both found growing on rocks and small trees. 1
The collections from which specimens of M. montanus were pre¬
pared for study were made in October and November of 1972. The
tardigrades were washed from samples submerged in water for a few
hours and recovered from a 325 mesh screen in a small amount of water.
They were fixed in boiling 95% ethanol and either slide-mounted in
Hoyer’s or prepared for study by scanning electron microscopy. The
latter specimens were dehydrated in absolute isopropanol, transferred
to amyl acetate and critical-point dried using C0 2 at 980 psi. Specimens
affixed to stubs with silver print were coated with 100A each of silver
followed by gold and were examined using the Cambridge Stereoscan
electron microscope at the Davis Campus Facility for Advanced
Instrumentation. 2
Macrobiotus montanus was described originally from specimens
collected at 6,000 ft., Nun’s Veil Mtn., New Zealand. Ramazzotti (1972)
records the species from Africa, Europe, the Galapagos Islands and
North America. Individuals of this California population are rede¬
scribed and illustrated in detail because the information obtained
by electron microscopy is new and will allow more critical comparisons
to be made with other populations.
The eggs of this species (Figs. 1A; 2C, D) are 77/x to 87/x in diameter
and the hemispherical processes are about 6/x high, 9/x across and sep¬
arated from each other by 0.8 /jl to 3.0^,. At the apices of the processes,
and beneath their surfaces some regular structural features exist that
may be mistaken for pores if observed with the phase microscope (Fig.
2D-p). The inter-process area of the shell is wrinkled and may have
six to eight minute pores (obvious at 2,000 to 5,000 diameters).
Mature embryos and juveniles can be recognized using the key to
1 Identifications provided by William A. Weber, University of Colorado Musuem, Boulder.
2 Support for use of the Stereoscan was provided by Faculty Research Grant D141.
The Pan-Pacific Entomologist 49: 229-231. July 1973
230
The Pan-Pacific Entomologist
Bill
&
Fig. 1. A. Mouthparts within embryonated egg. B. Mouthparts of adult (speci¬
men 50(V in length).
Macrobiotus in Ramazzotti (1972). However, in larger individuals
macroplacoid II is visibly shorter than I or III (Fig. IB) and these
specimens will lead to M. carsicus in this key. To arrive at M. montanus
in this key, the macroplacoids must be considered as being subequal
in length and of oval shape, or observation must be limited to the
smaller specimens. The cuticles of individuals ranging in size from the
smallest (290 /a) to the largest (550/x) were examined using the Stereo¬
scan and the specimens were later slide-mounted to confirm their iden¬
tity from conventional characters. All of the specimens possessed 10
buccal lamellae (Fig. 2E), the cuticle is somewhat wrinkled and while
occasional small pores were found, they were not a conspicuous or
regular feature. Distinct cribrate structures are present in the inter-
segmental creases (Fig. 2F—i). In an area dorsolateral to legs III the
cribrate structures were observed frequently to be greatly enlarged.
For all but the smallest individuals, a distinct granular area is present
dorsally at the bases of legs IV (Fig. 2F-e).
Macrobiotus montanus is assigned to the “intermedius group” by
Ramazzotti (1972). Present groupings may not reflect phylogenetic
relationship because within this group M. hufelandi has 10 buccal la¬
mellae as does M. montanus but M. areolatus has 12. Cuticular pores
are characteristic of M. hufelandi but are inconspicuous or absent from
M. areolatus and M. montanus . The accumulation of additional infor-
Vol. 49, No. 3, July 1973
231
Fig. 2. C. Scanning electron photograph of egg, 600X- D. Structure of rup¬
tured egg recorded by phase microscopy, 375X. E. Scanning electron photograph
of buccal opening, 1,585 X- F. Scanning electron photograph of cuticular mor¬
phology of dorsum, 1,975 X-
mation resulting from electron microscopy will better define the species
of Macrobiotus and will hopefully provide information by which their
relationships can be re-evaluated.
Literature Cited
Murray, J. 1910. Tardigrada. British Antarctic Exped. 1907-9 under command
of Sir E. H. Shackelton, C.V.O. Reports on the Scientific Investigations
1 to 5 (83-185 pis. XIV-XXI).
Ramazzotti, G. 1972. II Phylum Tardigrada. Mem. 1st. Ital. Idrobiol. Dott
Marco de Marchi, Pallanza Italy, 28: 1-732.
232
The Pan-Pacific Entomologist
New Species of North American Torymidae
(Hymenoptera)
E. E. Grissell
University of California, Davis 95616
Material representing four undescribed species of Torymidae has
recently been made available through the investigations of several
entomologists. I am describing them at present to make the names
available for biological studies currently underway.
Two of the undescribed species are in the genus Monodontomerus
Westwood. No species have been described from North America since
Gahan (1941) last characterized and revised the genus. Dr. B. D. Burks
(United States National Museum, Washington, D. C.) first sent material
and suggested I describe a species of Monodontomerus reared from the
douglas-fir tussock moth, Hemerocampa pseudotsugata McDunnough.
Another Monodontomerus , reared from an anthidiine bee, Dianthidium
heterulkei Schwarz, was compared for me with U. S. National Museum
material by Dr. Burks and determined as new. Two species of Torymus
Dalman are also described. One has been reared from a cecidomyiid
gall, Walshomyia cupressi Gagne, on Cupressus and the other from a
cynipid gall, Disholcaspis cinerosa (Bassett), on Quercus.
The holotype, allotype, and some paratypes of each new species will
be deposited in the United States National Museum. Paratypes will
also be placed in the Canadian National Collection, Ottawa, and the
Entomology Museum, University of California, Davis. Additional
paratype deposition is indicated where needed.
Monodontomerus saltuosus Grissell, new species
Female holotype. — (Chromosome number, 2n rz 10). Body length 4.0 mm,
ovipositor 1.4 mm. Head, scape, thorax, coxae, and femora dull metallic blue-
green to black (when viewed at some angles), abdomen dark blue-green to almost
black, flagellum black, tibiae and tarsi burnt orange, mandibles and wing veins
dark brown, wings hyaline except brown stain surrounding stigmal vein and ex¬
tending from postmarginal vein about one-fourth distance to posterior wing
margin. Head, thorax, coxae, femora, and gastral terga IV-VI with fine, even
reticulations, mesepimeron, median frenal area, and tergum I polished. Areas
with thick, appressed pubescence include: head, dorsum of thorax except frenal
area, coxae except median areas, femora, and gastral terga III-V laterally; long
erect pubescence present at lateral propodeal margins; eyes with erect, conspic¬
uous pubescence. Face as in Fig. 3; ocellocular distance slightly less than lateral
ocellus diameter (5:6) ; antennal funicle cylindrical, segments subquadrate;
The Pan-Pacific Entomologist 49: 232-239. July 1973
Vol. 49, No. 3, July 1973
233
complete malar sulcus present; mandibles tridentate; clypeus slightly protuberant,
but not extending to imaginary line connecting lateral corners of oral fossa.
Dorsellum of metanotum (Fig. 14) convex with fine, median carina; propodeum
with triangular median depression somewhat truncate at posterior fourth, median
carina becoming crooked and obscure posteriorly, anteriorly the median carina
with obscure fork; forewing with postmarginal vein about one-half as long as
marginal; hindfemur with denticle arising abruptly (Fig. 8). Abdomen about
as long as head and thorax combined, terga without median posterior emargin-
ations; ovipositor 0.72 times length of abdomen.
Male allotype. — (n = 5). Length 3.2 mm. Similar to female except an¬
tennal toruli slightly lower on face.
Holotype female, Iron Mountain, 24 miles east of Placerville,
El Dorado County, California, August 1970, ex Hemerocampa pseu-
dotsugata, C. G. Thompson collector; allotype, 3 2,3$ paratypes same
data; 42, 4$ same data except coll. 3 November 1971 em. January
1972, D. L. Dahlsten collector. Additional paratypes all same host data
and county: 43 2, 9 $ Plummer Ridge, 22 mi. SE Placerville, coll. 2
November 1971, em. 10 November to 23 January 1972, D. L. Dahlsten
collector; 32, 1$ same data except coll. August 1970. Additional
specimens not paratypes: 5 2, 1$ Sandia Mtns., near Albuquerque,
New Mexico, coll. August and November 1959, A. Rivas, Yasinski
collectors, same host data as holotype. Paratypes will be placed in the
collection of the Division of Biological Control, University of California,
Albany as well as those institutions listed in the introduction.
Variation.— Females vary in length from 2.5 mm (ovipositor 0.69
times as long as abdomen) to 4.2 mm (ovipositor 0.62 times abdomen).
Males range in length from 2.1 to 3.2 mm. In several specimens the
scape, fore and midfemora are infused with yellow rather than the nor¬
mal metallic bluish green color. This may have been the result of
killing individuals in a teneral condition. In other respects this species
appears to be quite constant.
Systematics. —Monodontomerus saltuosus will most readily run to
M. subobsoletus Gahan (= minor Ratzeburg) in Gahan’s 1941 key
to Monodontomerus. It agrees with M. minor in several respects in¬
cluding coloration, sculpturing, facial proportions (except position of
antennal toruli), proportion of wing veins, slender hindfemoral tooth,
and relative length of ovipositor. Monodontomerus saltuosus differs
from M. minor in both sexes particularly by the convex dorsellum with
a weak median carina in M. saltuosus (Fig. 14) as opposed to the
deeply concave dorsellum with pronounced median carina of M. minor
(Fig. 13) ; M. minor also has the propodeal carina markedly forked
anteriorly and M. saltuosus is not noticeably forked. Monodontomerus
234
The Pan-Pacific Entomologist
11 M. dementi
12 M. montivagus
Figs. 1-6. Heads of Monodontomerus and Torymus, facial view. Figs. 7-10.
Hindfemora, lateral view. Figs. 11-12. Male scape; A. lateral view; B. ventral
view. Figs. 13-14. Propodea, posterior view.
saltuosus may be separated in addition by the posterior apex of the
scutellum which is pitted; the apex is smooth in M. minor. A character
which works well for females but not males, is the relative position of
the antennal toruli. In M. saltuosus they are somewhat higher than in
M. minor , causing the lower face to be proportionately longer in M.
saltuosus (cf. Figs. 3 and 4).
Vol. 49, No. 3, July 1973
235
Monodontomerus dementi Grissell, new species
Female holotype.- —(2n=12). Body length 3.2 mm, ovipositor 2.6 mm. Head,
scape, thorax, coxae, abdomen, and femora except apices, bright metallic blue,
flagellum black, tibiae, tarsi, and apices of femora mahogany to burnt orange,
mandibles and wing veins dark brown, wings hyaline except slight suggestion of
brown stain surrounding stigmal area. Head, dorsum of thorax except frenum,
and metapleuron shagreened, mesepisternum and hindcoxa with distinct punctation,
femora and propodeum laterally distinctly reticulate, mesepimeron dorsally, frenum,
dorsellum, and median propodeum polished, terga I-V polished dorsally but with
extremely fine reticulations. Recumbent pubescence on head, dorsum of thorax
except frenum, metapleuron, coxae except median areas, femora, terga I and III—
VI laterally, longer erect pubescence at lateral margins of propodeum, eyes with
inconspicuous erect pubescence. Face as in Fig. 1; ocellocular distance greater
than lateral ocellus diameter (12:8) ; antennal funicle cylindrical, segments sub¬
quadrate; complete malar sulcus present; mandibles tridentate; clypeus projecting
slightly, extending to imaginary line connecting lateral corners of oral fossa.
Dorsellum convex with fine median carina which projects slightly posteriorly;
propodeum with triangular median depression ending at posterior %, median
carina distinct entire length, anteriorly with an almost imperceptible fork; fore¬
wing with postmarginal vein about one-half as long as marginal; hindfemur with
tooth arising gradually (Fig. 7). Abdomen slightly longer than head and thorax
combined, terga without median posterior emarginations; ovipositor about 1.6
times length of abdomen.
Male allotype. — (n = 6). Length 2.7 mm. Similar to female except scape
markedly transversely flattened, distally expanded, and bowed slightly lengthwise
(Fig. 11), undersurface appearing polished but with fine reticulations, upper
surface strongly shagreened.
Holotype female , 1 mile west West Thumb, Yellowstone National
Park, Wyoming, emerged 30-31 September 1971 ex cell collected 5
September 1971 of Dianthidium heterulkei heterulkei, S. L. Clement
collector. Allotype male and 6 female paratypes same data as holotype;
29, 2$ paratypes same data but coll. 16-24 July 1971, em. 31 July
1971; 16 2,7 $ paratypes laboratory reared on Megachile pacifica Pan¬
zer from original field collected material.
Variation.— The primary variation of this species lies in the size of
laboratory reared individuals. A size factor of greater than 2 to 1
between largest and smallest specimen resulted from over or under¬
crowding of the gregarious parasitic larvae on host larvae in gelatin
capsules. Field collected individuals (9 females) range in length from
2.4 to 4.5 mm, while their laboratory reared progeny (17 females) range
from 2.2 to 5.5 mm. One dwarf specimen has the ovipositor 1.1 times
the length of the abdomen. The range for ovipositor lengths of other
specimens is 1.5 to 1.8 times the abdominal length. Males range from
236
The Pan-Pacific Entomologist
2.3 to 3.9 mm. Small specimens tend to have a greater ocellocular:
lateral ocellus diameter ratio (15:9) than do larger specimens (13:11).
Systematics. —Monodontomerus dementi will key to M. montivagus
Ashmead in Gahan’s key (1941) and resembles that species in most
respects except for the following. In M. dementi the body is blue
whereas in M. montivagus it is olive green, M. dementi (Fig. 1) has
the malar space equal to one-half the eye height (x = 0.50, range 0.44-
0.58, n = 20) as compared to one-third in M. montivagus (Fig. 2) (x
= 0.32, range 0.29-0.35, n = 20), M. dementi has the scape consis¬
tently metallic blue whereas M. montivagus has the scape yellow, and
M. dementi has the ovipositor about equal to the body length as com¬
pared to M. montivagus which has the ovipositor barely longer than
the abdomen. An additional character in the males is the scape of M.
montivagus which is usually more arcuate in lateral view (Fig. 12A)
and less expanded in ventral view (Fig. 12B) as compared with M.
dementi which is more evenly bowed in lateral view (Fig. 11 A) and
noticeably expanded in ventral view (Fig. 11B).
I take pleasure in naming this species after Stephen L. Clement, a
fellow student of the Hymenoptera.
Torymus frankiei Grissell, new species
Female holotype.— Length 3.5 mm, ovipositor 3.8 mm. Head, thorax, coxae,
femora, and abdomen metallic green, scape and tibiae yellow-orange infused with
varying amounts of metallic green, flagellum black, tarsi pale straw-yellow, wing
veins brown. Face and frons completely shagreened, thoracic dorsum except frenum
transversely rugose with effaced setigerous punctures, frenum medially trans¬
versely rugose, laterally longitudinally rugose, propodeum, terga I—II dorsally and
inner face of hindcoxa appearing polished to lightly reticulate, outer face of hind-
coxa strongly reticulate. Face as in Fig. 5; ocellocular distance less than lateral
ocellus diameter (8:10) ; antennal funicle cylindrical, pedicel:Fl:F2 ratio as
6.5:7:6.5, Fl-5 longer than wide, F6-7 subquadrate; frenal groove distinct, frenum
one-third length of scutellum; mesepimeron twice as high as wide; propodeum
with two submedian foveae subtended mesally by carinae which connect to pos¬
terior of propodeum; hindcoxa dorsally smooth, without setae, with dorsal carina
extending onto outer coxal face, hindfemur with angulate denticle subapically
beneath (Fig. 10) ; costal cell anteriorly with almost complete setal row beneath,
basal vein with setae, cubital vein bare basally, with several setae at intersection
of basal vein, area beneath marginal vein distinctly setose, no obvious setal rows
radiating from stigma, stigmal vein subsessile. Terga I-IV obviously incised
apically; ovipositor 2.1 times length of abdomen.
Male allotype. —Length 2.3 mm. About as in female except as follows: body
coppery green, scape dark, pedicel:Fl:F2 ratio as 10:12:10.5, Fl-3 slightly longer
than wide, F4-7 subquadrate. Punctures on thoracic dorsum obscure, indicated
by setae; hindfemoral denticle obscure, appearing almost absent.
Vol. 49, No. 3, July 1973
237
Holotype female, 2 miles east Ft. Bragg, Mendocino County,
California, emerged 16 April 1968 ex galls collected 1 March 1968
of Walshomyia cupressi on Cupressus pygmaea (Lemmon) Sargent,
G. W. Frankie collector; allotype, 72, and 10$ paratypes with same
data except em. 12 March to 18 April 1968. Additional paratypes:
13 2, 19 $ Ft. Bragg, same data as holotype except coll. 6 March to
25 May 1967, em. 8 April to 14 June 1967. Other specimens not para¬
types: 9 2, 5$ Carson Ridge, Marin Co., Calif., coll. 2-10 April em.
22 April to 7 May, ex Walshomyia cupressi on Cupressus sargenti
Jepson. Paratypes of this species will be placed in the Entomology
Museum, Texas A & M University, College Station, as well as those
institutions mentioned in the introduction.
Variation.— Females of T. frankiei range in size from 2.5 mm (ovi¬
positor 2.2 times length of abdomen) to 3.5 mm (ovipositor 2.1 times
abdomen). Males vary in length from 1.9 to 2.7 mm. Color ranges
from entirely coppery green (some males) to partially green and copper
to all green (most specimens). In other respects this species appears
quite stable in characters which typically tend to vary in Torymus such
as relative funicular lengths and ocelli ratios.
Systematics.— Torymus frankiei clearly belongs to a group of species
characterized by T. fullawayi (Huber) by reason of its rugose thorax,
the frenum differently sculptured than the rest of the scutellum, the
large propodeal foveae, and the denticulate hindfemur. It is nearest
to T. denticulatus (Breland) in lacking the median propodeal carina
but differs from that species in having a setose costal cell and the hind-
coxa dorsally smooth with the carina extending onto the outer coxal
face. In T. denticulatus the costal cell is almost bare and the coxa has
a completely dorsal carina.
This species is named for Dr. Gordon W. Frankie in recognition of
his work with Torymus.
Torymus memnonius Grissell, new species
Female holotype.— Length 4.4 mm, ovipositor 4.1 mm. Head, coxae, femora,
and abdomen black, scape orange, fore and midfemora smoky, hindfemur orange
infused with black, tarsi white tinged orange, wing veins brownish white, stigma
and parastigma dark brown. Face and frons shagreened, thoracic dorsum, dorsellum,
propodeum, hindfemur, and hindcoxa evenly reticulate (size of reticulation vary¬
ing somewhat with location), abdomen evenly punctate but appearing reticulate
except tergum I polished dorsally. Face as in Fig. 6; ocellocular distance equal
to lateral ocellus (6:6) ; antennal funicle cylindrical, pedicel:Fl:F2 ratio as 7:10:10,
FI—5 longer than broad, F6-7 subquadrate. Frenal groove absent (or seen at
lateral margin only), frenum indicated by slightly smaller reticulation than an-
238
The Pan-Pacific Entomologist
terior of scutellum and by absence of setae; mesepimeron 2.3 times as high as
wide, slightly narrowed dorsally; propodeum with large pits one-fifth length of
propodeum along anterior margin, two weak submedian carinae one-half length
of propodeum on posterior margin, spiracle 1.5 times own long diameter from
posterior margin; hindcoxa dorsally weakly carinate and without setae, hind-
femur greatly expanded distally and with obvious denticulate angle beneath (Fig.
9) ; costal cell bare except anteriorly beneath with proximal row one-fifth length
of cell and distal row both above and beneath one-third cell length, cubital vein
without setae basally, basal vein setose, area beneath marginal vein setose for
distal two-thirds, one obscure setal row radiating from stigma distally; stigmal
vein sessile. Terga I-IV obviously incised apically, laterally with numerous long
recumbent setae; ovipositor 2.0 times length of abdomen.
Male allotype. —Length 3.1 mm. About as in female except body black with
bluish tinge, scape and tibiae bluish black. Pedicel:Fl:F2 ratio 6:8:8, all seg¬
ments at least slightly longer than wide. Propodeum with anterior median carina
one-half length of propodeum, posterior submedian carina one-third length of
propodeum.
Holotype female, H. F. Heep Farm, 11 miles south Austin, Travis
County, Texas, emerged 1 July 1971 ex galls collected 21 May 1971 of
Disholcaspis cinerosa on Quercus virginiana Miller, G. W. Frankie
collector; allotype, 23 2, 20 $ same data as holotype except numerous
collection and emergence dates. Additional paratypes: 79,6$ Texas
A & M University, College Station, Brazos Co., Texas, same host data
as holotype, various collection and emergence dates. Paratypes of
this species will be placed in the Entomology Museum, Texas A & M
University as well as those institutions listed in the introduction.
Variation. —Size range in females is 3.6 mm (ovipositor 3.0 times
length of abdomen) to 4.6 mm (ovipositor 2.7 times abdomen). The
ovipositor ranges from 2.0 to 3.0 times as long as the abdomen. Males
range from 2.1 to 3.1 mm. In some specimens of both sexes the black
body color may be tinged with blue or green. The propodeal carinae
of both sexes vary from absent to weakly developed. No specimens
have complete carinae.
Systematics. — Torymus memnonius is very similar to T. melanocerae
(Ashmead) but differs in having the hindfemur expanded and angulate
beneath (slightly bulged in T. melanocerae but not angulate), in lack¬
ing the frenal groove (distinct in T. melanocerae), and in having the
body and femora black ( T . melanocerae is blue to blue-green).
Acknowledgments
I wish to express my appreciation to Dr. B. D. Burks for the loan of
specimens and for suggestions concerning the systematic placement of
the 2 new Monodontomerus. In addition Dr. D. L. Dahlsten (University
Vol. 49, No. 3, July 1973
239
of California, Gill Tract, Albany, California), Dr. G. W. Frankie (Texas
A & M University, College Station, Texas), and Mr. S. L. Clement
(University of California, Davis, California) have contributed reared
specimens for this study. Mr. C. Goodpasture (University of California,
Davis) has determined the karotype number of the two Monodontomerus
in connection with his research. In addition he has offered constructive
criticism of this manuscript. I am indebted to these gentlemen.
Literature Cited
Gaiian, A. B. 1941. A revision of the chalcid-flies of the genus Monodontomerus
in the United States National Museum. Proc. U. S. Nat. Mus., 90:
461-482.
BOOK REVIEW
Annual Review of Ecology and Systematics. Volume 2. Edited by R. F.
Johnston. Annual Reviews Inc., Palo Alto, California. 510 p., 60 figs., 11 tables.
1971. $10.00.
This volume contains 19 articles by 21 contributors with the majority of articles
devoted to aspects of ecology and only three relating to systematics.
For the entomologist with varied interests many articles will have general appeal.
Only the article by R. van den Bosch on biological control is devoted exclusively
to insects. The article by D. H. Janzen considers the ecological processes of seed
predation by insects and other animals with special emphasis on the chemical
aspects of host specificity. The colonization of small bodies of water held by
leaves, flowers, or tree holes (= phytolemata) by organisms (ranging from bacteria
to insects and even a few frog tadpoles), and community structure in this micro¬
habitat are discussed by B. Maguire, Jr. The role of animals including insects in
the breakdown of litter receives some attention from M. Witkamp in Iris article
on soils as ecosystem components. The article on laboratory stream research by
C. E. Warren and G. E. Davis, although not dealing specifically with insects,
presents a valuable discussion of the application of studies in these closed or partly
closed systems in relation to natural ecosystems. Feeding strategies are discussed
in two articles: the theoretical aspects and models are treated by T. W. Schoener;
while D. H. Morse contrasts the adaptive strategies among insectivorous birds.
Two of the three systematic articles deal with phenetics while the third discusses
karyotypes. The relationship between character sets in the subject of the article
by J. S. Farris. Classificatory strategies and group measures in the cluster concept
are discussed by W. T. Williams. The value of the karyotype in systematics is
increasing as techniques are developed to detect new ways in which karyotypes
differ and this topic is reviewed by R. C. Jackson.
This collection of review articles contains something of value for a wide audience
interested in ecology and systematics.— Robbin W. Thorp, University of California ,
Davis , 95616.
240
The Pan-Pacific Entomologist
A New Comadia from the San Joanquin Valley of California
(Lepidoptera: Cossidae)
R. M. Brown and R. P. Allen
Martinez, California 94553 and California Department of Agriculture
Sacramento, 95814
Barnes and McDunnough (1911) proposed Comadia to include four
species taken from Hypopta (Hubner) and designated H. bertholdi
(Grote) the type species. Barnes and Benjamin (1923) described six
additional species and subsequently described C. polingi , the last species
to be named (Barnes and Benjamin 1927). Comadia has thus stood
until the present. Interest in the genus was revived with the discovery
of lepidopterous larvae in the roots of Alkali Blite, Suaeda fruticosa
(L.) Forsk. by the junior author about 1950.
Preliminary study by Harry K. Clench of Carnegie Museum, Pitts¬
burgh, Pennsylvania, indicated this to be a new species, described
herein. We thank Mr. Clench for suggesting the name used and releas¬
ing his notes, without which this paper would have been more difficult.
Comadia suaeclivora Brown and Allen, new species
Male. — Head: Antennae bipectinate to end; longest pectinations three times
shaft diameter; shaft white dorsally to end. Palpi upturned close to front, not
reaching middle of eye; dark laterally. Front and vertex creamy-white with scat¬
tered brown scales. Tongue absent. Thorax: Vestiture long, loose, creamy-white,
lighter ventrally; collar and posterior edge of thorax dark brown; partagia darker
and edged in darker brown. Legs with long hair-like scales; prothoracic coxae
and femora with considerable dark brown scaling; mesothoracic and metathoracic
legs lighter; metathoracic tibiae and first two segments of tarsi swollen; two
pairs of spurs present; tarsi ventrally spined; tarsal segments with brown rings
in middle. Forewing dorsally white along costa to subterminal line, white in¬
vading upper and distal part of cell; eleven dark brown costal spots; base of cell
Ma-Cui white; caudal edge of median vein to Cm white; both sides of A 2 white;
end of discal cell dark brown; basal area of cells Ma-Ma and Cm-Cm dark brown;
distal of subterminal line fuscous; finge white with brown checks at end of veins,
ochreous patches distad of cell, below origin of Cu 2 and central portion of cell
Ai-Az. Forewing ventral, costa with basal three-fifths dark brown, remainder
white with costal spots as above; dark subapical patch; subterminal line present,
distally dark between veins; remainder of wing light gray, fringed as above. Length
of wing, 15 mm. Hindwing dorsally white basad, distad light gray; veins darkened;
fringe white. Hindwing ventrally creamy-white with dark brown scales; fringe white.
Abdomen: Dorsally creamy-white cephalad, caudad darker; extending well beyond
hind wings; ventrally creamy-white with evenly scattered brown scales. Length
of body 16 mm. Male genitalia with harpe upturned, square at posterior edge,
The Pan-Pacific Entomologist 49: 240-245. July 1973
Vol. 49, No. 3, July 1973
241
Fig. 1. Male genitalia of C. suaedivora, a) left harpe, b) tegumen and uncus
(ventral), c) tegumen and uncus (lateral), d) aedeagus, (all 22.5X)- Fig. 2.
Last two larval segments of C. suaedivora. Fig. 3. Pupa of C. suaedivora , a)
ventral, b) lateral.
242
The Pan-Pacific Entomologist
inner surface lightly excavated; harpe with large hook shaped process at base
just below costa (Fig. la) ; transtilla reduced; gnathos spoon shaped just above
transtilla; teguraen, broad, hood-shaped, four to six strong setae at inner anterior
corners of base; uncus small, turned down in short strong hook (Fig. Ib-c).
Female. —Antennae ventrally serrate; serrations shallowly bifid, each with one
seta about as long as free part of projection. Wing pattern similar to that of male,
somewhat darker; subterminal line reduced. Abdomen heavier and considerably
longer than male. Length of forewing 17 mm.; of body 22 mm .
Holotype male , (California Academy of Sciences, type #11608),
Tipton, Tulare County, California, 17 May 1958, R. P. Allen. Allo¬
type, 3 mi. N. Corcoran, Kings County, California, 20 May 1972, R. M.
Brown. Paratypes, 80 males, 12 females; California, Fresno Co.: Ker¬
man 15 14 May 1962, R. P. Allen; 1$ 9 May 1962, H. E. Gleason;
10 mi. S. Kerman 25 3 June 1960, R. P. Allen; Tranquility 1 S 14 May
1962, H. E. Gleason. Kings Co. : 3 mi. N. Corcoran, 3 S 20 May 1972,
R. M. Brown. Tulare Co.: Earlimart, 3c? 14 May 1962, 1# 21 May
1962, 1 9 31 May 1962, D. Rush; Tipton, 1 S 15 May 1953, 15 10
May 1956, 15 5 15 May 1956, 33 5 17 May 1958, 49 7 May 1956
(reared), 1 9 27 May 1959 (reared), 15, 19 30 May 1959 (reared),
19 5 June 1959 (reared), 2 5,19 1 May 1960 (reared), 15 10 June
1960, 115 18 May 1961, 3 5,29 21 March 1969 (reared), R. P. Allen.
Type disposition. —The holotype and allotype will be permanently
deposited in the collection of the California Academy of Sciences,
San Francisco. One pair of paratypes will be deposited in each of the
following collections; Carnegie Museum, Pittsburgh, Pennsylvania;
Los Angeles County Museum of Natural History, Los Angeles, Cal¬
ifornia; Canada Department of Agriculture, Ottawa, Ontario; Amer¬
ican Museum of Natural History, New York, N. Y.; and the collec¬
tion of the senior author. The remaining paratypes will be placed in
the collection of California Department of Agriculture, Sacramento,
California.
Remarks. —Barnes and Benjamin (1923) divided Comadia into
three groups based on thoracic vestiture as follows: (1) Thoracic
scales broad, ( engelhardti B&B) ; (2) Thoracic scales hair-like, spat-
ulate at end (see below) ; (3) Thoracic scales long, narrow to end
(henrici Grt. & menfredi Neum.).
The second group at present is the largest and the most difficult.
This group now contains six species and three subspecies: C. bertholdi
bertholdi Grt., Colorado; C. bertholdi fusca B&B, Durango, Colorado;
C. bertholdi edwardi Neum. & Dyar, Colorado; C. intrusa B&B South¬
ern California; C. dolli B&B Clark Co. Nevada; C. stabilis B&B Trout
Vol. 49, No. 3, July 1973
243
Fig. 4*. A cluster of larvae in the hollowed crown of the host, Suaeda fruticosa .
Fig. 5. Mature larvae showing lack of setae and the prominent anal horn.
Creek, Juab Co. Utah; C. polingi B&B Bert, Otero Co. New Mexico;
C. suaedivora Brown & Allen Tipton, California; C. subterminata B&B
Redington, Arizona.
Comadia suaedivora like the other species, shows considerable vari¬
ation. The amount of dark shading is the most variable character;
only the few white areas described seem to be stable. The majority
of females in the type series are smaller than the males. These prob¬
ably were dwarfed while being reared under artificial conditions. The
two females taken at light are the same size or larger than the males.
244
The Pan-Pacific Entomologist
Comadia suaedivora can be told from C. dolli, C. stabilis, C. intrusa
and C. polingi by the greater amount of dark suffusion and the whiter
costa. The C. bertholdi group begins to approach it with C. b. fusca
being the closest. From this subspecies it can be told by the lack of a
prominent dark spot on the forewing below Cu 2 - From C. subterminata
it can be told by its smaller size and narrower, less rounded forewings,
plus the lack of reticulations. This is an arbitrary placement until
further study is made.
Biology
Larvae of C. suaedivora bore in the crown and roots of Suaeda fru-
ticosa a low, woody halophytic shrub common to the semialkaline soil
found in the San Joaquin Valley. Larvae are gregarious and completely
hollow the woody area in which they feed (Fig. 5), usually killing the
host. They are easily found by digging out and breaking open the
crown and roots of the host. Various stages in development of larvae
found together in the host indicates a possible two year larval cycle.
The known range of the moth extends from southern Madera County
to northern Kern County, including portions of Fresno, Kings, and
Tulare counties. A large population at Tipton, Tulare Co. has declined
over the years due to host removal by development of cotton storage
yards, and perhaps in part by the junior author’s intermittent digging
of plants in search of larvae. The host does not readily extend itself.
Adults fly from early May to mid June. The males are more attracted
to light than females. Rearing by the junior author lends support to
this: 15 adults (6 males and 9 females) were reared with a ratio of
40% males to 60% females; in the field 77 specimens were taken (75
males and 2 females) with a ratio of 97.4% males to 2 . 6 % females. If
there was an equal attraction for light between the sexes, a more equat-
able ratio should be found. The most extensive flights have been ob¬
served on warm, overcast nights during the latter part of May.
Eggs are deposited in a tight cluster glued to the host, probably near
the crown. They have raised surface reticulations forming square and
rectangular depressed cells. Eggs are 2.0 mm in height and 1.0 mm
in diameter.
Larvae attain a length of 30 mm. When alive they have rose-lavender
highlights which fade rapidly when specimens are placed in alcohol.
The head is partly covered by the first thoracic segment. Thoracic
legs are short, with a strong claw; abdominal legs are represented by
crotchets arranged in a uniordinal series on segments A 3 through Ag.
Vol. 49, No. 3, July 1973
245
Setae are sparse and short. The most striking feature of the larvae is the
heavily sclerotized horn on the dorsal surface of the anal flap (Fig.
2 & 4). This horn is dark brown and does not extend above the level
of the back. The purpose of the horn is open to speculation; it may be
pushed against the wood to help the larvae travel inside the host.
Pupae (Fig. 3) are about 15 mm long, dark brown and heavily
spined. The spines suggest that the pupa is active and able to travel.
Considerable digging in the vicinity of infested plants has produced
no pupa in the field, but they have been obtained by rearing. Reared
adults leave their pupal cases protruding conspicuously above ground
after emergence. This indicates the larvae, upon leaving the host, prob¬
ably constructs a subterranean cell prior to pupation. Prior to emer¬
gence of the imago, the pupa makes its way to the ground surface.
Rivers (1897) in a description of the life history of H. bertholdi Grt.,
points out that the pupa also extends above ground prior to adult
emergence.
Literature Cited
Barnes, W. and J. H. McDunnough. 1911. Revision of the Cossidae of North
America. Contrib. Natnr. Hist. Lepidoptera, 1: 26-29.
Barnes, W. and F. H. Benjamin. 1923. Cossidae, Cossinae. Contrib. Natur.
Hist. Lepidoptera, 5: 88-96.
1927. A new Cossid from New Mexico (Lepid., Cossidae). Pan-Pac. Entomol.,
4: 67.
Rivers, J. J. 1897. Some Facts In The Life History of Hypopta Bertholdi Grote.
Psyche, 8(249) : 10.
BOOK NOTICE
Environment and Man. Richard H. Wagner. W. W. Norton and Co., Inc. 491 p.,
196 figs., 9 tables. 1971. $7.50.
This book, produced on recycled paper, is intended as a general introduction
to man-environment problems requiring little background in the sciences. The
stress is on understanding basic problems. Additional readings are listed at the
end of each of the 23 chapters and an Appendix to professional and lay journals
containing articles of interest is provided for those who wish to keep up with this
rapidly expanding area of information.
The book is divided into phases of reciprocal influence between man and en¬
vironment: the environment in shaping man; man shaping the environment; the
man-made city environment shaping man; and finally the ultimate problem of
balancing resources and population to allow coexistence of all organisms. It
presents an easy to read, well balanced overview of man-environment problems.—
Robbin W. Thorp, University of California, Davis, 95616.
246
The Pan-Pacific Entomologist
Notes on the Biology of ISeomachilis halophila
on a California Sandy Beach
(Thysanura: Machilidae)
Robert Benedetti 1
Hopkins Marine Station of Stanford University, Pacific Grove, California
Neotnachilis halophila Silvestri, originally described from beaches
near San Francisco, is common under rocks and in crevices in the
high intertidal zone on California beaches (W. Evans, pers. comm.).
I have found it most abundant just on the seaward side of the last ter¬
restrial vegetation in areas covered with rocks too high on the beach
to be disturbed by most high tides, and especially among rocks piled
in such a manner that a small space occurs between rock and sand.
Little is known of the natural history of this species. The present study
deals with population structure, activity rhythm, and food. All obser¬
vations were carried out near Pacific Grove, California, at sites on Pt.
Pinos and on Mussel Point near the Hopkins Marine Station in April
and May 1972.
Population structure. —Specimens (135) were collected at Pt.
Pinos 19 and 20 May. In the laboratory they were anesthetized with
ether, sexed, and measured under a dissecting microscope. Females were
determined by the presence of the elongate ovipositor extending from the
eighth and ninth segments (Lubbock, 1873). Length was measured
from the tip of the head to the posterior end of the abdomen, excluding
caudal cerci and filament (Fig. 1). Males and females occur in approxi¬
mately equal numbers. Some females are larger than the largest males.
A marked discontinuity in size is seen in the population. Many juve¬
niles, no longer than 3.5 mm, are present, but the next shortest size
class begins at 8.1 mm. In a Californian lepismid studied by Lasker
(1956), seasonal breeding was reported, eggs being found only in
April and May. The discontinuity observed in the present study sug¬
gests a similarly limited breeding period in late winter or early spring.
The numerous juveniles were found along with the much larger adults.
Activity pattern. —Preliminary observations indicated that Neo-
machilis remained at rest during the day, mainly on undersides of rocks.
At night they were more active, and were found crawling on sand be¬
neath rocks and on upper surfaces of rocks. Activity patterns of the
1 Mailing address: 3502 N. 19th, Tacoma, Washington 98406.
The Pan-Pacific Entomologist 49: 246-249. July 1973
Vol. 49, No. 3, July 1973
247
13.9-14.5
13.3-13.8
12.7-132
12 . 1 - 12.6
N=|I5
ZII. 4-120
2
^10.8-11.3
£10.0-10.7'
3 9.4-10.0'
o
uj 8.8-9.3
N
w 8.1-8.7
3.6-8.C
2.5-3.5
T ~ I ' I "H
'i
0 10 30 50
NO. OF FEMALES
NO. OF MALES WHOLE POPULATION
Fig. 1. Population structure of Neomachilis at Pacific Grove, California. 19-
20 May.
population were determined in two 24-hour periods of regularly re¬
peated field observation, one carried out at Pt. Pinos and the other at
Mussel Point. Observations in darkness were made with a red-filtered
flashlight. At Pt. Pinos, two areas of about thirty square feet each
were marked off and examined every two or three hours for location
and number of Neomachilis. At Mussel Point one site of approximately
thirty square feet was similarly observed. At each observation time
animals were categorized as either 1) stationary on undersurfaces of
rocks, 2) crawling on sand beneath and around rocks, or 3) crawling
on the tops of rocks (Fig. 2). The increase in total number, decrease
in number on undersurfaces of rocks, and increase in number on sand
and on tops of rocks, all indicate that the activity of Neomachilis in¬
creases with the coming to darkness, reaches a peak shortly before
sunrise, and declines rapidly at dawn (Fig. 2). The early morning
activity peak coincides with the period of heaviest dew fall (S. Johnson,
pers. comm.).
Food and feeding. —Food studies were carried out on animals col¬
lected at Pt. Pinos on 25 and 26 May and immediately preserved in
95% alcohol. In the laboratory animals were decapitated and the whole
alimentary canal removed as described by Lasker (1956). Intestinal
contents were removed, mounted in glycerine, and examined under a
243
The Pan-Pacific Entomologist
3 ^
DAY
T
a*
100
MAY 15-16^1972
POINT PINOS 80
Q 60
z
O 40
u_
o 20
z
. Q
8:00
z MAY 26-27
O 1972
MUSSEL
POINT
KEY
TOTAL NO. SEEN
UNDER ROCK
TOP OF
ROCK
12:00 16:00 20:00 24:00 4:00 8:00 12:00 16:00
TIME
Fig. 2. Activity pattern of Neomachilis at Pacific Grove, California.
compound microscope. Major categories of food were determined
qualitatively. Six samples of ten animals each were taken over a 16.5
hour period to observe any changes in quantity and nature of the gut
contents.
A fair variety of material occurs in the gut of Neomachilis with uni¬
cellular green algae, yeast, and pine pollen being the materials most
often found. The unicellular algae are derived from and encrusting
lichen growing on the upper surfaces of the rocks under which Neo¬
machilis is found. The probable source of the pollen is the Monterey
Pine (Pinus radiata ), the most abundant tree near Pacific Grove shores,
and a species which produces large quantities of wind-borne pollen
in late winter and early spring. The source of the yeast is unknown.
The other materials found represent common high beach detritus. In
22 % of the animals, sporozoans (Gregarinidae) were found in the mid
gut, in numbers from two to thirty-six. The largest numbers of animals
with guts partly or wholly filled were found in early morning, indicating
that feeding tends to occur during periods of greatest activity.
Vol. 49, No. 3, July 1973
249
Summary
Neomachilis inhabits the spaces beneath rocks of the upper beach
and shore areas just below the land vegetation. Males and females occur
in equal numbers, but large females are larger than large males. Ac¬
tivity is largely nocturnal, and greatest activity occurs just before dawn.
Common foods include unicellular green algae (from lichens), yeast,
pine pollen, and vegetable detritus. Most feeding occurs during the pre¬
dawn period of greatest activity.
Acknowledgments
Appreciation is expressed to Dr. Pedro Wygodzinsky for identification,
Dr. W. Evans for aid in the preparation of the manuscript, and Drs.
Isabella and Donald Abbott for technical guidance during the study.
Literature Cited
Lasker, R. 1956. The nutrition of the silverfish Ctenolepsima lineata with
special reference to cellulose digestion. Unpublished Ph.D. thesis,
Stanford University, Stanford, California.
Lubbock, J. 1873. Monograph of Collembola and Thysanura. pp. 201-250, The
Ray Society, London, pp. 278.
RECENT LITERATURE
The Survival Equation: Man, Resources, and His Environment. Edited by
Roger Revelle, Ashok Khasla and Maris Vinovskis. 511 p., illus. 1971.
Each of the 38 reprinted articles starts with a brief introduction by the editors.
Some of the articles are edited and abridged by the editors, but most retain their
tables and references. The volume is copiously illustrated with good quality photos.
These are without identifying legends, but a terminal list of credits is provided.
250
The Pan-Pacific Entomologist
Coniopterygidae from Chile, with Three New Species
(Neuroptera)
Phillip A. Adams
California State University, Fullerton, 92634
In his excellent revision of the Coniopterygidae, Meinander (1972)
lists only one species from Chile. Analysis of material in my collection
brings the number of Chilean species to five, three of which are new.
The specimens were collected by Luis E. Pena, and remain in my col¬
lection unless otherwise noted. P. Wygodzinsky (American Museum of
Natural History), has graciously allowed me to borrow the type of
Pampoconis latipennis.
The genitalic terminology of Meinander (1972) is followed, using
these abbreviations: ect—ectoproct, gs—gonarcus, p—penis, pm—
paramere, st—stylus.
Pampoconis latipennis Meinander
(Fig. 2f)
Aisen, Puerto Ibanez, 12-17 January 1961. Genitalia of this speci¬
men closely resemble those of the type. The ectoproct margins are
smoothly curved in both specimens, not angulate as in Meinander’s
Figure 90A; his outline apparently follows the protruding setose sub-
anal plate and the medial surface of the stylus, rather than the incon¬
spicuous ectoproct margin. The posterior penis plates are fused ven-
trally almost to the apex; on the Aisen specimen, a ligulate membranous
lobe protrudes posteriorly to the ninth sternite and a dark oval sclerite
lies between the antennal bases. Previously recorded from El Bolson,
Rio Negro [Neuquen], Argentina, approximately 400 miles north (holo-
type $, examined).
Pampoconis punctipennis Adams, new species
(Fig. 1)
Male.— Head with unsclerotized area including antennal sockets, extending in
long tongue to clypeus; narrow median dark y-shaped band from vertex supports
antennal articulations. Eye small. Antennae 3.8 mm long, 36 segments about 1.5
times longer than wide, densely pale pilose. Apical labial palpomere spatulate.
Body sclerites dark fuscous, legs paler. Wings (Fig. le). Membrane pale; in
forewing, distal veins and wing margin dark-spotted. Forewing length 5.6 mm.
Male genitalia with ectoprocts (Fig. la-b) bearing 5-6 stout marginal spines
(modified setae). Ninth sternite with lateral tufts of long setae, and posterior
ligulate sclerite. Penis a pair of sclerites, rodlike anteriorly, vertically flattened
The Pan-Pacific Entomologist 49: 250-254. July 1973
Vol. 49, No. 3, July 1973
251
Fig. 1. Pampoconis punctipennis, male: a, b, terminalia, lateral and ventral;
c, d, penis and styli, ventral and lateral; e, wings. Scale for a-d.
and spinose posteriorly. Styli articulate on ectoproct, bearing dorsal knob with
anteriorly directed spine, parameres long and rodlike.
Holotype male , Santiago, Rio Colorado, Chile, October 1958 (Cal¬
ifornia Academy of Sciences, San Francisco, in alcohol).
This species differs from the other two Pampoconis species in wing
maculation, in presence of sclerite posteriorly to the ninth sternite, and
in shape of the styli.
Pampoconis uncinatus Adams, new species
(Fig. 2a-e)
Male. —Head with large unsclerotized area, lacking median sclerotized band
of P. punctipennis. Eye height 0.33 mm. Antennae 2.5 mm, 26 segmented. Seg-
252
The Pan-Pacific Entomologist
Fig. 2a-e, Pampoconis uncinatus, male: a, b, terminalia, lateral and ventral,
c, d, penis, lateral and ventral; e, wings. Fig. 2f, Pampoconis latipennis, stylus,
lateral. Scale for a-d, f.
ments about 1.5 times as long as wide, sparsely pale pilose. Body sclerites brown,
scuta with darker spot each side, first and second axillary sclerites black. Abdomen
weakly sclerotized except for terminalia; underlying tissue pink. Wing veins
uniformly brown, membrane paler. Forewing length, 4.65 mm. In forewing, Rsi
curves sharply anteriad, CuP not sinuous. Genitalia with ectoproct (Fig. 2a)
bearing single posterior marginal spinose seta; a tuft of setae on lateral apices
of ninth stemite. Styli (Fig. 2b) articulating ventrally on ectoproct, with apical
hook extending about one third of stylus length; parameres rodlike. Penis (Fig.
2c, d) 0.364 mm long, with anterior rods; a pair of posterior vertical spiny plates
bear serrate-margined lateral ridges and broad dorsolateral expansions.
Female. —As for male except: Antennae 1.85 mm, 27 segmented; eye height
0.22 mm; terminal maxillary palpomere slightly swollen. More robust, with
shorter legs. Forewing length 4.34 mm. Genitalia resemble those of P. latipennis.
Holotype male, and allotype, Talca, El Radal, Chile, 900 m, 28-
30 November 1957 (California Academy of Sciences, in alcohol).
Vol. 49, No. 3, July 1973
253
b
f-
0.25mm
9 -
Fig. 3. Incasemidalis meinanderi, male: a, b, c, terminalia, lateral, ventral,
and apical; d, e, internal armature, lateral and dorsal, f, scale for a-c; g, scale
for d, e.
This species is similar to P. latipennis Meinander, but differs in
having a pronounced forward curve in end of Rsi, non-sinuous CuP,
in location of the ectoproct spine, in shape of stylus (which is a larger,
sickle-shaped simple hook in P. latipennis Fig. 2f) and in structure of
penial apex. The genitalia are less massive than in a male P. latipennis ,
which had a 0.465 mm penis and a 4.08 mm forewing.
Incasemidalis meinanderi Adams, new species
(Fig. 3)
Male. —Head gray-brown, similar in shape to that of 7. peruviensis Meinander
but eyes are relatively larger (height of eye, 0.28 mm; bottom of eye to top of
postocular lobe, 0.43 mm). Antennae 39-40 segmented, 2.8 mm long, resembling
those of I. peruviensis. Body sclerites dark brown; abdomen pale except for
terminalia. Wings with membrane light brown, paler posteriorly. Venation much
as in 7. peruviensis, but in both wings radial crossvein intersects Rs-j-MA at its
fork. Forewing length 4.02 mm, width 1.57 mm; hindwing length 3.29 mm, width
1.50 mm. Male genitalia with ectoprocts ventrally rounded and bent inward;
ninth sternite strongly sclerotized and bifid apically. Gonarcus bears arched
nearly rectangular median projection. Parameres thin downcurved hooks attached
to anterior rods. Penis dorsally forms thin vertical blade attached to ventral side
of gonarcus projection; ventrally widening into curved band attached to apex
of ninth sternite. Tenth sternite not apparent.
254
The Pan-Pacific Entomologist
Holotype male, Coquimbo, El Calabozo, Illapel, Chile, 1700 m,
21-23 November 1959 (California Academy of Sciences).
This specimen conforms to the generic description by Meinander in
all respects except that no tenth sternite is present. It differs from both
I. colombiensis and I. peruviensis in the shape of the gonarcus lobe,
and in the simple bandlike penis. I take pleasure in naming the species
after Martin Meinander, the eminent specialist in Coniopterygidae.
Semidalis kolbei Enderlein
This species is abundantly represented from 23 localities extending
from Coquimbo (Nague, Los Vilos) in the north to Magellanes (Natales)
in the south, and from the coast to the Andes. Dates range from Sep¬
tember through March. Two female specimens represent the first record
from Argentina (Neuquen, San Martin de los Andes, 11 March 1955,
F. W. Walz).
Literature Cited
Enderlein, G. 1906. Monographie der Coniopterygiden. Zool. Jahrb. Abt. Syst.
Oekol. Geogr. Tiere, 23: 173-242.
Meinander, M. 1972. A revision of the family Coniopterygidae. Acta Zool. Fenn.
136: 1-357.
Vol. 49, No. 3, July 1973
255
A New Species of Hamotus from Galleries of
Microcerotermes septentrionalis in Western Mexico 1
(Coleoptera: Pselaphidae)
(Isoptera: Termitidae)
Donald S. Chandler and William L. Nutting
University of Arizona Tucson, 85721
The proposed species can be placed by Park’s (1942) monograph
in the subgenus Hamotoides, Group XIV, where it keys out to Hamotus
( Hamotoides ) veracruzensis Park. This new species is easily separated
from H. veracruzensis and all others in the group by the concave tergite
V of both sexes.
Hamotus (Hamotoides) cavus Chandler and Nutting New Species
Stramineous to castaneous, stramineous pubescence composed of a moderate
number of long setae. Head subtruncate at base; eyes prominent, containing ap¬
proximately 37 facets; posterior emargination of eye densely pubescent, projecting
across one-third eye width; two small vertexal foveae between eyes, large, oval me¬
dian fovea between antennal tubercles; single median gular fovea; mandibles four¬
toothed; third segment of maxillary palpi with moderate groove through entire
length; antennae with distinct, three segmented club.
Pronotum only slightly inflated anterior to foveae, its anterior edge truncate;
median fovea not much wider than transverse sulcus connecting median and lateral
foveae, transverse sulcus deep with distinct edges, lateral foveae large.
Elytron widened apically, rounded and notched at lateral apical edge and with
two basal foveae; sternites and tergites gradually narrowing apically, sternites
I-IV with lateral foveae, sternite V without, tergites I-IV margined laterally, I-III
with two close lateral foveae on each side, IV-V with one lateral fovea on each
side, tergite V concave.
Males with small spine on inner side of protibiae (fig. 1), spine difficult to see
through dense pubescence.
Holotype male. —length 2.40 mm; head length 0.53 mm, width across eyes
0.51 mm, width across antennal insertions 0.25 mm, 0.29 mm from head base to
median antennal fovea, vertexal foveae 0.08 mm from eyes, median antennal fovea
0.12 mm from vertexal foveae; antennal segment lengths and widths: I 0.17 X
0.09 mm; II 0.08 X 0.08; III 0.07 X 0.07; IV 0.07 X0.07; V 0.06 X 0.08; VI 0.07
X 0.08; VII 0.05 X 0.08; VIII 0.05 X 0.08; IX 0.10 X 0.12; X 0.17 X 0.16; XI
0.24 X 0.19; third segment of maxillary palpi 0.31 mm long, 0.12 wide.
Pronotum length 0.48 mm, 0.24 mm wide at base, 0.52 mm wide across lateral
foveae; median and lateral foveae 0.11 mm from base, lateral foveae 0.21 mm
from median fovea.
Elytron 0.77 mm long, 0.56 mm wide near apex; fovea 0.06 mm apart on elytron;
widths between lateral foveae of tergites: I 0.53 mm; II 0.65; III 0.59; IV 0.53;
1 University of Arizona Agricultural Experiment Station Journal Series #2061.
The Pan-Pacific Entomologist 49: 255-257. July 1973
256
The Pan-Pacific Entomologist
Fig. 1. Anterior view male protibia.
Fig. 2. Left lateral view male genitalia.
V 0.34; tergite V in form of rounded, thick rhomboid, 0.24 mm long; sternite
widths between lateral foveae: I 0.73 mm; II 0.89; III 0.76; IV 0.57; sternite
V 0.39 mm wide.
Protibia 0.51 mm long, spine 0.37 mm from base. Male genitalia asymmetrical
(fig. 2).
Female.— Similar to male but lacking spine on protibiae.
Holotype male, 3 mi N Barra de Navidad, Bahia de Coastecomate,
Jalisco, Mexico, 9 August 1964, in galleries of Microcerotermes sep-
tentrionalis, W. L. Light and G. C. Nutting.
Paratypes. —4 males, 15 females, same data as holotype. Slide
mounted holotype male to be deposited at the Field Museum of Natural
History, Chicago.
At least 40 of these beetles were observed when the extensive super¬
ficial galleries of Microcerotermes septentrionalis Light were broken
into. The colony, probably numbering in the tens of thousands, oc¬
cupied several feet of a sound log, about 1 foot in diameter and 12 feet
long, in a dense thorn forest area. About two dozen pairs of Nasuti-
termes nigriceps (Haldeman) ( Termitidae ) were also established in
cells within abandoned peripheral workings of the Microcerotermes
colony. These incipient colonies consisted of pairs with no brood and
some with a small number of eggs, nymphs and soldiers. Further, one
incipient colony of Incisitermes platycephalus (Light) (Kalotermitidae)
was found in the same situation.
Park (p. 310) mentions that other Hamotus species have been col¬
lected with termites. Little can be said about any kind of relationship
between the new species and any of the termites beyond the fact that
Vol. 49, No. 3, July 1973
257
they were in the galleries with Microcerotermes. In such a rich asso¬
ciation the possibility of their acting as predators seems especially good.
Literature Cited
Park, 0. 1942. A study in neotropical Pselaphidae. Northwestern Univ. Studies
Bio. Sci. Medicine, No. 1, 403 p.
SCIENTIFIC NOTE
Lectotype designation for Gerris buenoi Kirkaldy (Heteroptera: Ger-
ridae).—Recently we wanted to examine the type material of Gerris buenoi
Kirkaldy (1911, Entomol. News, 22: 246) in connection with our work on the
aquatic Hemiptera of California. Kirkaldy’s description was published posthu¬
mously through the efforts of J. de la Torre-Bueno. The description is incomplete
to the extent that the number of specimens examined, their locality data, and de¬
position are not given. Kirkaldy apparently had both macropterous and apterous
males and females. Torre-Bueno, in his note appended to Kirkaldy’s description,
states that E. P. Van Duzee supplied the material on which the description was
based, and that the bugs bore a manuscript label “ Gerris sulcatus Uhler.” Further¬
more, Torre-Bueno said that the “species is widely distributed and ranges from
British Columbia to the Atlantic states.”
Drake and Harris (1934, Ann. Carnegie Mus., 23: 179-240) apparently did not
see type material of buenoi when they published their revision of the North Amer¬
ican species, although they stated that the “type is in the Kirkaldy collection, U. S.
National Museum.” A search of this collection did not reveal its presence, and
there is no record of the “type” ever having been deposited there.
Paul Arnaud, of the California Academy of Sciences, San Francisco, has located
in the Academy collection at least a portion of the material that was before Kirk¬
aldy. It consists of three macropterous females all bearing E. P. Van Duzee col¬
lection labels. One specimen has a handwritten label il Limnotrechus sulcatus
Uhl.” plus the following locality data: Ft. Collins, Colorado, July 12, 1898, E. P.
Van Duzee collector. We are selecting this specimen as lectotype of Gerris buenoi
Kirkaldy. The data on the other two females are “N. Colo., May 10, 1898” and
“Montreal West, May 4, 1903.” The lectotype agrees with the usual interpretation
of buenoi. — A. S. Menice, Systematic Entomology Laboratory, U.S.D.A., Washing¬
ton, D. C. 20560 and J. T. Polhemus, University of Colorado Museum, Boulder.
258
The Pan-Pacific Entomologist
Descriptive Morphology of Eggs of Some Species
in the Macrobiotus hufelandii Group
(Tardigrada: Macrobiotidae)
Albert A. Grigarick, Robert 0. Schuster
and Elizabeth C. Toftner
Department of Entomology, University of California, Davis, 95616
The identification of species of Macrobiotus is frequently dependent
more on characteristics of the egg than on features of the adults. The
existing descriptive morphology of these eggs has resulted mainly from
observations by light microscopy before the development of phase con¬
trast and with scant attention being paid to details now recognized as
important for species discrimination. Light microscopes can provide
usable images only to 1,000 X and can’t resolve small but useful details.
Phase systems provide additional information but still can’t detect mi¬
nute detail, distinguish between surface and subsurface features, or pro¬
vide much information concerning surface topography. Vague and in¬
consistent terminology used to communicate information has lessened
the value of light microscope observations. Illustrations are often of
poor quality and written descriptions are usually incomplete and are
not always compatible with the drawings.
The scanning electron microscope (SEM) provides capabilities for
observing new information about these eggs and for photographically
recording this information in a manner that can be communicated pre¬
cisely. In addition, information so recorded can be easily quantified
and analyzed by existing systems. We have developed a simple method
for recording and analyzing information in a reproducible and com¬
municable form and have attempted to identify this information with
species of Macrobiotus in the hufelandii group.
PROCEDURE
To be able to study eggs of reasonably consistent quality, a stan¬
dardized procedure was used for the preparation of all samples. The
eggs, in small quantities of water, were fixed with boiling 95% ethanol
and later dehydrated in absolute isopropanol. The isopropanol was
replaced by amyl acetate and the specimens critical-point dried (An¬
derson, 1951) using CO 2 at 69 K cm 2 . Silver conductive paint was used
to affix the eggs to SEM stubs and the specimens were vacuum coated
with 100 A of silver followed by 100 A of gold.
The Pan-Pacific Entomologist 49: 258-263. July 1973
Vol. 49, No. 3, July 1973
259
Each egg was photographed using a Cambridge Stereoscan Mark II
microscope. The photographs were taken at approximately 1,000 diam¬
eters and these images were used to determine egg diameter, the number
of processes per hemisphere, the lengths and widths of processes and the
amount of serration of the apical discs.
A Metals Research Instrument Corporation “Quantimet 720” Image
Analyzing Computer (IAC) was used to detect and record features of
the photographic enlargements of the egg shells. High contrast 3,000
diameter enlargements of each egg were examined using a 25 mm lens
on the video scanner and the information was displayed as a format of
300 X 300 picture points (PP), with each PP representing a length
of 0.13 mm of the 3,000X photograph. Expressed as micra, an area of
13/a X 13/a (169 square micra) of egg surface was analyzed, and each
PP represented a length of 0.043 /a or an area of 0.0018 sq. micra.
The following characteristics of each egg were recorded with the IAC:
1) total pore area; 2) total number of pores; 3) pore size distribution
expressed as the number of pores with diameters greater than 0.258/a,
0.344/a, 0.430/a, 0.516/a, 0.602/a, 0.688/a, and 0.774/a; 4) total perim¬
eter of pores.
The hufelandii group of Macrobiotus, further limited to those for
which the pharynx contains two macroplacoids but no microplacoid,
contains the following species: hufelandii Schultze, hufelandii recens
Cuenot, pseudohufelandii Iharos, hibiscus Barros, anderssoni Richters,
and inermis Binda and Pilato. Fifty eggs of this group from world¬
wide localities were examined. The literature rvas surveyed for descrip¬
tive information and drawings of the eggs of these species. Our photo¬
graphs of eggs best agreeing with published information were assumed
to represent those species. New information provided by this study
is presented 1) as photographs, and 2) as detailed description provided
by SEM and IAC study.
Many of the eggs could not be assigned to described species because
they differed from presently accepted parameters of size, number of
processes, or sculpture of the shell. The application of numerical tax¬
onomy programs to accurate measurements of multiple characters wil]
eventually be used to delimit species and populations. The potential
of a combined SEM-IAC approach to secure information suitable for
statistical studies has been demonstrated by applying 2-way analysis
of variance and a mean separation by Duncan’s multiple range test for
the small samples now available. For three species with multiple spec¬
imens the analysis of variance used only four replications of selected
characters for the entire egg and three replications for characters of
260
The Pan-Pacific Entomologist
Table 1. Mean values for measurements of eggs of Macrobiotus based
on SEM and IAC data. Mean values of characters of different species
associated with the same letter are not significantly different from each
other at the ** 1%, or * 5% levels according to Duncan’s multiple range
test.
nr. hibiscus
n = 2
hibiscus
n 6
• rH
a
o
c n
03
■3 11
ctf C
• pH
• pH
T3
C
CO
D , |
*3 II
? hufelandii
n = l
hufelandii recens
n = 1
ENTIRE EGG
Egg diameter in /***
70
58b
81a
73ab
82
53
Number of processes/
hemisphere**
143
117c
34a
81b
62
89
Process diameter in /***
5
4b
10a
6b
8
5
Process length in /** *
6
5b
16a
7b
10
7
Discal serrations
9
9
10
16
4
10
Disc open or closed apically
closed
closed
closed
closed
open
closed
169 SQ. MICRA OF EGG
SURFACE
Number of pores**
41
142b
76ab
26a
0
168
Pore area in sq. /**
19
33b
36b
13a
0
25
Pore perimeter in /***
59
148b
109b
35a
0
121
Pore diameter distribution
>0.258/*
40
146
71
24
0
137
>0.344/*
40
103
66
22
0
107
>0.430/*
38
81
60
22
0
62
>0.516/*
32
66
55
19
0
36
>0.602/*
25
52
46
15
0
16
>0.688/*
18
40
34
12
0
11
>0.774/*
14
28
30
9
0
7
the egg surface although the numbers of individuals indicated in table
1 are greater.
DISCUSSION
Macrobiotus hufelandii Schultze, 1833 (table 1, fig. 3)—the
original description of hufelandii didn’t include the egg stage. Doyere
(1840) first described it as spherical and studded with processes swol¬
len at the top but didn’t mention whether the shell had pores. Most
Vol. 49, No. 3, July 1973
261
authors presently agree that the egg of hufelandii has a smooth shell
with a circle of pores around the base of each process.
Eight eggs agreeing with this description were available from col¬
lections made in Sweden. These eggs are most distinguished by the
reduced number of pores. The smallest pore area was 2 sq. fi per 13
fi sq. with pore diameters in the range of 0.43 [jl or less.
Macrobiotus sp. ? hufelandii (table 1, fig. 5)—a single example
of an egg of hufelaiidii from California differs from the above examples
in having a wrinkled shell and a total absence of pores. The processes
are hollow with internal openings basally to each of the ridges radiat¬
ing outward from the processes. The space between the inner and outer
shell walls thus contains an appreciable amount of air space although
the shell is not porous. If further study doesn’t provide intergrades be¬
tween the forms, the Californian population will need to be recognized
as a distinct entity.
Macrobiotus hufelandii recens Cuenot, 1932 (table 1, fig. 1) —
the egg of this species in no way resembles the illustration by Cuenot
but agrees exactly with that illustrated by Ramazzotti (1972) as an
egg probably representing recens. The former recens egg has about
twice as many processes of lesser diameter and processes without distal
discs. The variation is inordinate and the species represented in fig¬
ure 1 may eventually be recognized as distinct. Of the kinds of eggs
considered here, this one has the greatest total number of pores and
largest pore perimeter.
Macrobiotus anderssoni Richters, 1907 (table 1, fig. 2)—examples
of this species agree with the figure accompanying the original de¬
scription. The small number of long processes with microdentate rings
and the extremely porous shell surface is distinctive.
Macrobiotus hibiscus Barros, 1942 (table 1, fig. 4)—six eggs of
hibiscus were from four widely separated localities. . . Tanzania, Italy,
Finland and California. The eggs are small with many widely spaced
small processes. The interprocess area is comprised of pores differing
greatly in size.
Macrobiotus sp. nr. hibiscus (table 1, fig. 6)—a few eggs of a
general hibiscus configuration appear to represent undescribed species.
One example of these is a large egg with many closely spaced processes
and few pores in the reduced shell area. Any understanding of the sig¬
nificance of these various eggs is dependent on the accumulation of suf¬
ficiently large numbers to allow a statistical analysis of their variation.
262
The Pan-Pacific Entomologist
Vol. 49, No. 3, July 1973
263
Macrobiotus pseudoliufelandii Iharos, 1966 —we haven’t seen an
example of this egg which is described as having squat cone-shaped
processes about 5 fi high and 5/x wide at the base. The description in¬
dicates some 35—40 processes per hemisphere but the illustration has
over 80. The shell is supposedly smooth.
Macrobiotus inermis Binda and Pilato, 1971 —One extremely
flattened egg was examined following removal from a slide preparation.
The original description of this egg was essentially correct except the
shell surface has about 45 minute pores in each area bordered by the
bases of any four processes.
Closely spaced, short cone-shaped processes with small terminal discs
have walls that are apparently smooth but a granular texture is evident
at magnifications above 5,000X.
The eggs in inermis and pseudoliufelandii appear to be extremely
similar. Examination of eggs of the latter species by SEM will be re¬
quired to resolve the question of possible synonymy.
Literature Cited
Anderson, T. F. 1951. Techniques for the Preservation of Three-dimensional
Structure in Preparing Specimens for the Electron Microscope, Trans.
N. Y. Acad. Sci., Series II, 13: 130-134.
Barros, R. de. 1942. Tardigrados do Estado de Sao Paulo, Brazil II, Genero
“Macrobiotus”, Rev. Brasil. Biol., 2: 257-269.
Binda, M. G. and G. Pilato. 1971. Nuovo contributo all conoscenza dei Tardi-
gradi di Sicilia. Boll. Sedute Accad. Gioenia Sc. Nat., Catania IX:
623-633.
Doyere, M. P. L. N. 1840. Mem. Sur Les Tardigrades, Ann. Soc. Nat. 14 p. 269.
Iharos, G. 1966. Beitrage zur kenntnis der Tardigraden-Fauna Osterreichs.
Acta Zool. Academiae Scinetiarum Hungaricae XII: 123-127.
Ramazzotti, G. 1972. IL. Phylum Tardigrada Mem. 1st. Ital. di Idrobiol. 28,
732 p.
Richters, F. 1907. Antarktische Tardigraden. Zool. Anz. Leipzig 31: 915-916.
Schultz, C. A. S. 1833. Beschriebung des Macrobiotus hufelandii. Isis von
Oken pp. 708-713.
4 *
Figs. 1-6. Scanning electron microscope photographs of eggs of Macrobiotus
1. M. hufelandii recens Cuenot, 700'X (Mt. Aspiring National Park, South Island,
New Zealand). 2. M. anderssoni Richters, 470X (Abel Tasman National Park,
South Island, New Zealand). 3. M. hufelandii Schultze, 500X (Oland, Sweden).
4. M. hibiscus Barros, 675X (Muonio, Finland). 5. M. ? hufelandii, 455 (Auburn,
California). 6. Undescribed sp. nr. hibiscus 500X (Triangulation Station,
Chatham Islands, Rangatira Island, New Zealand).
264
The Pan-Pacific Entomologist
Mealybugs of San Miguel Island, California
(Homoptera: Coccoidea: Pseudococcidae)
Douglass R. Miller
Systematic Entomology Laboratory, USD A, ARS, Beltsville, Maryland 20705
In an earlier paper (Miller, 1971) I described the mealybug fauna
of Santa Cruz Island, and discussed the California Island biota, in¬
cluding brief statements on island name terminology, endemism, and
disjunctions.
In July 1970 I collected on San Miguel Island for nine days. Al¬
though this was far from an adequate amount of time to collect speci¬
mens of each mealybug species on the island, I think it was sufficient
to permit me to adequately compare the origins and relationship of the
pseudococcid faunas of San Miguel and Santa Cruz Islands.
San Miguel is approximately 14 square miles in land area, is 26 miles
from the nearest point on the mainland, is three miles from the nearest
island (Santa Rosa), and is 21 miles from Santa Cruz Island (Philbrick,
1967). The highest elevation, 831 feet, is on San Miguel Mountain.
The general topography of the island has been described by Bremner
(1933) as follows: “Most of the surface of the island is a wave-cut ter¬
race from 400 to 500 feet in elevation above which rise two hills. . . .
(San Miguel and Green Mountains).A lower terrace from 20
to 100 feet in elevation borders the south coast.The island has been
eroded but little since the terraces were cut. Two streams heading on
the flanks of Green Mountain and flowing northward and southward
respectively, have cut deep narrow canyons.Another steep narrow
canyon heads on the terrace near the ranch house and drains northward
to Cuyler Harbor. The island when first visited by Europeans, was cov¬
ered with a dense growth of brush.Cattle and sheep were placed on
the island.and allowed to multiply without restriction, with the re¬
sult that much of the vegetation was destroyed. The strong prevailing
winds from the northwest then formed dunes of drifting sand that now
cover most of the surface.” Sand beaches are common and surround
about % of the island. The only trees are several recently introduced
palms. The flora contains no taxa which occur exclusively on San Mi¬
guel, although there are nine plant species, subspecies, or varieties
which have been recorded on San Miguel and at least one other Cali¬
fornia Island but are not known on the mainland (Raven, 1967). The
herpetofauna includes one salamander and two lizards, none of which
The Pan-Pacific Entomologist 49: 264-269. July 1973
Vol. 49, No. 3, July 1973
265
are insular endemics (Savage, 1967). The terrestrial mammal fauna
consists of a bat, an endemic subspecies of deer mouse, and an en¬
demic race of gray fox. The fox species is an insular endemic (von
Bloeker, 1967). Marine mammals are abundant on the beaches and
off-shore waters of San Miguel. In recent years four or five species of
seals and sea lions have been reported (Bartholomew, 1967) in ad¬
dition to the sea otter (Allanson, 1955). A general study of the insect
and bird faunas of San Miguel has not been published to my knowledge.
The recent geologic history of the California Islands is not well under¬
stood. San Miguel apparently Avas last submerged during the First In¬
terglacial period of the Pleistocene (Orr, 1967). The time of the last
land connection to the mainland is not well established—late Pliocene
to mid-Pleistocene (Orr, 1967), mid-Pleistocene (Valentine and Lipps,
1967), or late Pleistocene and sub-Recent (Weaver and Doerner, 1967).
All seem to agree that if a connection actually occurred, it was as a
peninsular extension of the Santa Monica Mountains. After the last
continental land connection, there is evidence that San Miguel was
joined to the other Northern Channel Islands at least during the late
Pleistocene (Valentine and Lipps, 1967).
San Miguel Island Mealybugs
The following list includes 12 genera and 13 species. The localities
may be found on the accompanying map (Fig. 1). Plant names are
as listed by Munz and Keck (1965). The only previous mention of
a mealybug on San Miguel was by Cockerell (1938) who stated, “A
kind of mealy-bug, perhaps new, was found on Astragalus miguelensis .”
The unnamed mealybug was most likely Pseudococcus obscurus.
A monostherium
li.chtensioid.es (Cockerell)
1. Green Mountain, 11 July 1970, Artemisia calijornica (foliage)
2. Harris Point, 10 July 1970, A. calijornica (foliage)
Chorizococcus
abroniae McKenzie
1. Cuyler Harbor, 12 July 1970, Franseria sp. (decumbent branches)
Distichlicoccus
salinus (Cockerell)
1. Bay Point, 12 July 1970, Distichlis spicata (on leaf blade)
2. Cuyler Harbor, 9 July 1970, Distichlis spicata (on leaf blade)
Heterococcus
arenae Ferris
1. San Miguel Mountain, 11 July 1970, grass (in leaf sheath)
266
The Pan-Pacific Entomologist
n
0 | 1/2
milei
I
S
Fig. 1. San Miguel Island mealybug collection sites 1. Bay Point 2. Cuyler
Harbor 3. Green Mountain 4. Harris Point 5. San Miguel Mountain 6. Simonton
Cove 7. Ranch south of Cuyler Harbor.
Paludicoccus
distichlium (Kuwana)
1. Cuyler Harbor, 9 July 1970, Distichlis spicata (in leaf sheath)
Phenacoccus
colemani Ehrhorn
1. Harris Point, 10 July 1970, Dudleya sp. (on roots)
gossypii Townsend and Cockerell
1. Simonton Cove, 10 July 1970, Haplopappus sp. (on foliage)
Pseudococcus
obscurus Essig
1. Bay Point, 12 July 1970, Coreopsis gigantea (on roots)
2. 1 mi. S. Bay Point, 12 July 1970, Astragalus sp. (on roots)
3. Cuyler Harbor, 12 July 1970, Coreopsis gigantea, Eriogonum sp., Franseria
chamissonis (on roots)
4. Green Mountain, 11 July 1970, Lotus sp., Mesembryanthemum sp. (on
roots)
5. Ranch S. of Cuyler Harbor, 9 July 1970, Astragalus sp., Baccharis sp.,
Lupinus sp. (on roots)
6. San Miguel Mountain, 11 July 1970, Mesembryanthemum sp. (on roots)
7. Simonton Cove, 10 July 1970, Abronia sp., Astragalus sp., Eriogonum sp.
(on roots)
8. 0.5 mi. S. E. Simonton Cove, 10 July 1970, composite (on roots)
Puto
yuccae (Coquillett)
1. Bay Point, 12 July 1970, Atriplex semibaccata (on roots)
Vol. 49, No. 3, July 1973
267
2. Cuyler Harbor, 9 and 10 July 1970, Abronia sp., Atriplex semibaccata,
Eriogonum sp., Mesembryanthemum sp. (on foliage)
Rhizoecus
gracilis McKenzie
1. Bay Point, 12 July 1970, composite (on roots)
Spilococcus
keiferi McKenzie
1. Cuyler Harbor, 9 July 1970, Eriogonum sp. (on foliage)
T ridiscus
distichlii Ferris
1. San Miguel Mountain, 11 July 1970, Distichlis spicata (in leaf sheath)
T rionymus
smithii (Essig)
1. Green Mountain, 11 July 1970, grass (in leaf sheath)
2. 0.5 mi. S. E. Simonton Cove, 11 July 1970, grass (in leaf sheath).
Discussion
Based on the work of McKenzie (1967) there are approximately 40
species of mealybugs which have been collected on hosts that occur on
San Miguel Island and are found in the coastal mainland areas adja¬
cent to the Northern Channel Islands. Of the 40 species, only 13 have
been collected on San Miguel. Three of the 13 are distributed through¬
out most of California, three occur in saline regions along the coast, two
occur in the coastal mountain ranges and beaches, two in the coastal
ranges, southern deserts, and Sierra-Nevada Mountains, two in the
coastal ranges and the Sierra Nevada, and one exclusively on beaches.
All mealybug species that occur on San Miguel very probably are pres¬
ent in the coastal mountains or beaches of the immediate California
mainland. It is therefore likely that the San Miguel pseudoccocid fauna
is a depauperate aggregation of mealybug species which are closely al¬
lied to the adjacent mainland fauna. Based on these assumptions, it
is logical to conclude that San Miguel has been colonized from the ad¬
jacent California mainland via the other Northern Channel Islands.
Perhaps the most interesting aspect of the San Miguel mealybug fauna
is displayed by Pseudococcus obscurus. This species was encountered
in great abundance at nearly every collecting site. Although it is a
common California mealybug, I have never before seen it in such num¬
bers or in so many different kinds of habitats as on San Miguel. Al¬
though it is tempting to suggest that this expanded insular diversity is
caused directly by the availability of vacant feeding sites, alternative
explanations such as lack of natural enemies are possible.
268
The Pan-Pacific Entomologist
Comparison of the Mealybug Faunas of San Miguel
and Santa Cruz Islands
There are several noticeable differences in the Santa Cruz and San
Miguel mealybug faunas. (I) The Santa Cruz fauna contains nearly
twice as many mealybug species as that of San Miguel. Of the 13 spe¬
cies present on San Miguel, only two are unrecorded on Santa Cruz,
whereas of the 23 species on Santa Cruz, 12 are unknown on San Miguel.
The reason for these differences seems evident since Santa Cruz is larger
in size, is closer to the mainland, has a greater habitat diversity, and
apparently has been continuously emergent for a longer period of time.
(2) Although Santa Cruz supports a much larger number of pseudo-
coccid species, individuals are relatively rare. On San Miguel however,
the species number is small, but individuals, particularly of Pseudo¬
coccus obscurus, are abundant. Furthermore, on Santa Cruz there is
no evidence of unusual ecological diversity, but on San Miguel P. ob¬
scurus is not only exceptionally abundant, but also occurs on many
previously unrecorded hosts. These differences very possibly are un¬
related to insular phenomena. Many pseudococcids prefer sandy, well
drained soils, particularly those species that occur in the moist coastal
regions. On Santa Cruz this type of habitat is present but uncommon,
whereas on San Miguel nearly the entire island is covered by sand. Pre¬
sumably, once an oligophagous species such as P. obscurus has suc¬
cessfully colonized San Miguel and has adapted to a sandy habitat, it
can easily spread and become abundant. On Santa Cruz once an oli¬
gophagous species has become established, in order to successfully in¬
habit large geographic areas, it must adapt not only to many hosts, but
also to varied types of habitats, many of which are not very successfully
colonized even on the mainland. (3) On Santa Cruz, there are five
pseudococcid species which do not occur on the closest mainland but
do occur farther north in California. On San Miguel no disjunctions
were found; all species apparently occur on the adjacent mainland. (4)
On Santa Cruz the genustspecies ratio was about 1:2; on San Miguel
the ratio was approximately 1:1.
There are two similarities in these insular faunas. (1) Both lack the
endemic taxa encountered in other parts of their respective biotas. (2)
Both are depauperate aggregations of species similar to the pseudo¬
coccid fauna on the adj acent mainland.
Summary
Twelve genera and 13 species of mealybugs are known on San Miguel
Island, none of which are endemic. The pseudococcids comprise a de-
Vol. 49, No. 3, July 1973
269
pauperate aggregation of species similar to the mealybug fauna of the
adjacent mainland. One species, Pseudococcus obscurus, appears to be
much more abundant and to occupy more diverse habitats on San Mi¬
guel than on the mainland or on Santa Cruz Island.
Acknowledgments
I thank Mr. Robert 0. Schuster and Dr. Albert A. Grigarick, Uni¬
versity of California, Davis, for making arrangements for this trip to
San Miguel, for assisting in the collection of scale insects while on the
island, and for criticizing the manuscript. I also thank Louise M. Rus¬
sell, Dr. Arnold S. Menke and Dr. Willis W. Wirth, Systematic Ento¬
mology Laboratory, ARS, USDA, for reviewing and criticizing this
manuscript.
Literature Cited
Allanson, A. 1955. Sea otters on San Miguel. Pac. Discovery, 8: 24—25.
Bartholomew, G. A. 1967. Seal and sea lion populations of the California
Islands. Pp. 229-244 in R. N. Philbrick (ed.), (For complete citation
see Philbrick).
Bremner, C. St. J. 1933. Geology of San Miguel, Santa Barbara County, Cali¬
fornia. Santa Barbara Mus. Nat. Hist. Occas. Papers, (2) : 1-23.
Cockerell, T. D. A. 1938. San Miguel Island, California. Sci. Monthly, 46(2) :
180-187.
McKenzie, H. L. 1967. Mealybugs of California with taxonomy, biology and
control of North American species. Univ. Calif. Press, Berkeley, 525 pp.
Miller, D. R. 1971. Mealybugs of Santa Cruz Island, California. Pan-Pac.
Entomol., 47(4) : 293-303.
Munz, P. A. and D. D. Keck. 1965. A. California Flora. Univ. Calif. Press,
Berkeley, 1681 pp.
Orr, P. C. 1967. Geochronology of Santa Rosa Island, California. Pp. 317-325
in R. N. Philbrick (ed.), (For complete citation see Philbrick).
Philbrick, R. N. 1967. Introduction. Pp. 3-8 in R. N. Philbrick (ed.), Pro¬
ceedings of the Symposium on the Biology of the California Islands.
Santa Barbara Botanic Garden, Calif., 341 pp.
Raven, P. H. 1967. The floristics of the California Islands. Pp. 57-67 in R. N.
Philbrick (ed.), (For complete citation see Philbrick).
Savage, J. M. 1967. Evolution of the insular herpetofaunas. Pp. 219-227 in
R. N. Philbrick (ed.), (For complete citation see Philbrick).
Valentine, J. W. and J. H. Lipps. 1967. Late Cenozoic history of the Southern
California Islands. Pp. 21-35 in R. N. Philbrick (ed.), (For complete
citation see Philbrick).
von Bloeker, J. C., Jr. 1967. Land mammals of the Southern California Is¬
lands. Pp. 245-263 in R. N. Philbrick (ed.), (For complete citation
see Philbrick).
Weaver, D. W. and D. P. Doerner. 1967. Western Anacapia—A summary of
the Cenozoic history of the Northern Channel Islands. Pp. 13- 20 in
R. N. Philbrick (ed.), (For complete citation see Philbrick).
270
The Pan-Pacific Entomologist
A New Milliped of the Genus Tynomma from California
(Lysiopetalidea: Lysiopetalidae)
Michael R. Gardner
Pacheco, California
The genus Tynomma contains the only representatives of the crested
millipeds in Northern California. The two species recognized by Buckett
and Gardner (1969) are found abundantly in a variety of habitats.
Tynomma mutans (Chamberlin) occurs around San Francisco Bay, and
the larger Tynomma magnum Buckett and Gardner occurs further south
along the Monterey Coast. A third species has now been distinguished
in a collection of millipeds sent to me for determination by Dr. Evert
I. Schlinger for the California Insect Survey, University of California,
Berkeley. A key is presented below to separate these species.
Key to the Species of Tynomma Loomis
1. Transition to full number of crests occurring on segments 11 or 12; body
small (14-22 mm) ; lacking coxal lamina on mid-body legs of male-
_ mutans (Chamberlin)
Transition to full number of crests occurring on segments 14 or 15; body
large (30-40 mm) ; with distinct flattened process on alternate coxae of
mid-body legs in male___ 2
2. Transition to full crest number occurring on segment 14; gonopod with sol-
enomerite curving cephalad, prongs equal _ magnum Buckett and Gardner
Transition to full-crest number occurring on segment 15; gonopod with sol-
enomerite produced laterad, proximal prong much reduced _
- schlingeri Gardner, new species
Tynomma schlingeri Gardner, new species
Diagnosis. —Differing from magnum and mutans in changing to full crest num¬
ber on segment 15 rather than 14 or 12, and in the details of the gonopods (figs.
1 and 2).
Holotype male.— Head dark brown, mottled dorsally, light brown frontally,
gnathochilarium depigmented; antennae brown; tergites medium brown, with whit¬
ish middorsal stripe, and pale brown below third crest on each side; legs pale
brown. Head with surface lightly rugose; coronal suture short, not extending be¬
low vertex; frontal area wide, sunken mesally; clypeus smooth, with 12 stout setae
ventrally; antennae long, with article three longest, followed in length by articles
two, five, four, six, one and seven; all antennal articles clothed in short setae;
four terminal sense cones in apex of seventh antennal article; eye with 37 ocelli
in rows of 9, 8, 7, 6, 4, 3 and 1 ocelli, respectively. Body segments 61; collum nar¬
rower than head, with 20 crests; segments 2 through 13 with five primary crests,
including two suprarepugnatorial crests; segments 15 and succeeding with six pri-
Ti-ie Pan-Pacific Entomologist 49: 270-272. July 1973
Vol. 49, No. 3, July 1973
271
Figs. 1-2. Tynomma schlingeri, holotype male. Fig. 1. Gonopods, posterior
aspect. Fig. 2. Left gonopod, lateral aspect.
mary crests, including three suprarepugnatorial crests. Setae of primary dorsal crests
equalling one-half length of crest on anterior segments and one-third crest length
on middle body segments; setae moving to caudal end of crest on sixth segment,
projecting mesocaudad on that and succeeding segments, not enlarged caudally;
secondary crests large, subequal to primary crests on anterior segments, becoming
gradually reduced caudad, nearly obsolete on posterior third of body, two second¬
ary crests between two dorsal primary crests, one between others. Body tapering
gradually caudad; epiproct with lateral margins straight, exceeding anal valves,
with one pair of paramedial dorsal setae and two pairs of lateral marginal setae;
spinnerets prominent, produced caudad just below caudal margin or epiproct; a
pair of setae subtending spinnerets; anal valves smooth, with one pair of setae
removed from anal lips; anal lips narrow, slightly protruding; hypoproct lentic¬
ular, with two pairs of setae. Legs seven segmented, with transverse sutures on
tarsi of mid-body legs; podites from longest to shortest as follows: 7, 3, 4, 6, 5,
1, 2; legs one and two six-segmented, reduced; about half length of normal walk¬
ing legs, with large, erect setae on mesal surface of podites; tarsi with dense comb
mesally; second coxa with small, acute anterior process and large cylindrical pos¬
terior gland; third and succeeding legs with velutinous pad on mesal face of tar¬
sus; legs 3 through 24 with eversible coxal glands distally; beginning with leg 16,
alternate legs with coxa produced in subconical caudal process; beginning with
272
The Pan-Pacific Entomologist
leg 48, and continuing to leg 80, those legs with caudal process also with rounded,
sclerotized anterior lamina; succeeding legs normal. Gonopods anteriorly fused,
forming a single subtriangular ventral lamina with sinuous lateral margins, acute
apex, and median vertical carina; posteriorly, produced separately in long, suh-
cylindrical, narrow pieces greatly exceeding anterior lamina; distally expanded,
with rounded subapical posterior lamina, wide mesal lamina with small, crescent¬
shaped solenomerite, and rounded anterolateral lamina. Length: 30 mm.
Paratype female. —As in male except for sexual characters.
Specimens examined. Holotype male.— California; Stanislaus
County: Del Puerto Canyon, el. 1200 ft., Frank Raines Park, 13
April 1973 (E. I. Schlinger). Paratypes. 4 females, same data as holo-
type. All types will be placed in the type collection of the California
Academy of Sciences, San Francisco.
Discussion. —This distinctive species is closely related to magnum as
indicated by several features. Both are large, have a short anterior
gonopod and long posterior gonopod, rounded laminae on alternate
coxae, sutures on tarsi, and transition to full crest number occurring on
adjacent segments.
The type locality for schlingeri is an isolated canyon in the inner coast
ranges. In addition to the present species, an undescribed species of
Casey a (fam. Caseyidae) occurs there. Neither species has been col¬
lected elsewhere.
Literature Cited
Buckett, J. S., and M. R. Gardner. 1969. Revision of the chordeumid milliped
genus Tynomma Loomis from California. Pan-Pac. Entomol., 45: 204-
216.
Vol. 49, No. 3, July 1973
273
Two New Subspecies of the Drosophila willistoni Group
(Diptera: Drosophilidae ) 1
Francisco J. Ayala
Department of Genetics, University of California, Davis, California 95616
The genus Drosophila (Drosophilidae, Diptera) is characterized by
the existence of groups of two or more sibling species, i.e., species
morphologically so similar as to be practically indistinguishable.
Dobzhansky (1956) has speculated that the evolution of external
morphology has reached in the genus Drosophila a high degree of
perfection and that the adaptive evolution proceeds largely through
physiological channels. This hypothesis has recently been supported by
evidence showing that sibling species of Drosophila often differ in
their allozymes, that is, in enzymes coded for by different alleles at the
same genetic locus. Ayala and Powell (1972) have shown that allozymes
can be used as species diagnostic characters.
Since little morphological differentiation often exists between closely
related species, it is not surprising that the taxonomic category of sub¬
species has rarely been used in Drosophila , in spite of extensive study
of the genus. Whenever races or subspecies have been distinguished in
Drosophila , the distinction has, in most cases, been based on the oc¬
currence of incipient reproductive isolation and/or differences in the
frequencies of chromosomal inversions. In this paper I present evidence
showing that one pair of subspecies can be distinguished in each of the
species, Drosophila willistoni and D. equinoxialis. The subspecies can
be identified by their allozyme patterns, and these are used in their
formal description. I believe this is the first time that allozymes have
been used in the formal description of a taxon.
Reproductive Isolation
The Drosophila willistoni group contains at least six sibling species
endemic in the New World tropics (Spassky et al ., 1971). Two of
the siblings, D. insularis Dobzhansky and D. pavlovskiana Dobzhansky
and Kastritsis, are narrow endemics; the former on some islands of the
Lesser Antilles and the latter in Guyana. The other four species, D.
willistoni Sturtevant, D. equinoxialis Dobzhansky, D. paulistorum
Dobzhansky and Pavan, and D. tropicalis Dobzhansky and Pavan have
1 Work supported by NSF grant GB 30895.
274
The Pan-Pacific Entomologist
wide geographic distributions, which overlap through Central America
and the northern half of continental South America.
The distribution of D. willistoni extends from northern Mexico and
south Florida, through Central America, the Caribbean islands, and
continental South America down to La Plata in Argentina. D. willistoni
adults were collected in 1954 by Prof. C. Pavan and again in 1972 by
Prof. D. Brncic near Lima, Peru. The Lima flies from either sample
exhibit incipient reproductive isolation from flies collected in Colombia,
Venezuela, Trinidad and Brazil. Crosses having Lima flies as the male
parent produce fertile hybrids of both sexes. Females from Lima
crossed to males from the other localities yield fertile females but sterile
male hybrids. The populations near Lima are therefore assigned to a
new subspecies, D. willistoni quechua Ayala.
The known distribution of D. equinoxialis extends from central
Mexico, through Central America, the greater Antilles, and the northern
half of continental South America, including the Amazon basin. Flies
from Hispaniola, Puerto Rico and Costa Rica, exhibit incipient repro¬
ductive isolation from flies collected in eastern Panama or anywhere
in continental South America. The populations from Hispaniola, Puerto
Rico, and Costa Rica are therefore included in a second new subspecies,
D. equinoxialis caribbensis Ayala. Crosses between D. e. equinoxialis
and D. e. caribbensis produce fertile females but sterile males.
Enzyme Differentiation
Using techniques for starch gel electrophoresis and enzyme assay
described elsewhere (Ayala et al., 1972), we have studied genetic varia¬
tion at 23 to 30 gene loci coding for enzymes. The relevant results are
summarized in Table 1. At each of 5 out of 25 loci studied in both
subspecies there are substantial differences in the enzyme patterns, and
therefore in the genotypic constitution of D. w. willistoni and D. w.
quechua. Also at each of 5 out of 25 loci studied in both subspecies
substantial differences exist between D. e. equinoxialis and D. e. carib¬
bensis. Using the method of Ayala and Powell (1972) it is possible to
calculate the probability of correct diagnosis of the subspecies of a
single individual of known genotype. This probability ranges from
0.99998 to 0.974 for each of the five diagnostic loci in D. willistoni ;
using jointly the five loci, the probability of incorrect diagnosis of the
subspecies of a single individual is 3.4 X 10~ 14 . The probability of cor¬
rect attribution for each of the five diagnostic loci ranges in D .
equinoxialis from 0.9997 to 0.967; using jointly the five loci the prob-
Vol. 49, No. 3, July 1973
275
Table 1. Allelic frequencies at several diagnostic loci in two pairs
of subspecies of Drosophila. Alleles designed by their relative electro¬
phoretic mobility; some rare alleles have been omitted from the Table.
Sample size is the number of wild genes sampled; i.e., twice the num¬
ber of wild-collected individuals studied, except for sex-linked loci
which exist in males in single dose. The symbols for the loci refer to
the enzymes coded for by them, as follows: Est = esterase; Odh =
octanol dehydrogenase; Xdh — xanthine dehydrogenase; Acph = acid
phosphatase; Mdh = malate dehydrogenase; G3pdh = glyceraldehyde-3-
phosphate dehydrogenase; Hk = hexokinase.
Subspecies
Sample
size
Locus and alleles
Probability
of correct
diagnosis
of the
subspecies
A.
Est-2
98
100
102
104
D. w. willistoni
7012
.003
.041
.941
.006
.999
D. w. quechua
108
.231
.769
.000
.000
Est-4
98
100
102
104
D. w. willistoni
9692
.004
.146
.838
.011
.989
D. w. quechua
114
.000
1.000
.000
.000
Est-7
96
98
100
102
105
D. w. willistoni
6819
.025
.147
.563
.211
.049 .974
D. iv. quechua
59
.847
.000
.153
.000
.000
Odh-1
96
100
104
D. iv. willistoni
1088
.039
.882
.071
.994
D. w. quechua
64
.016
.000
.984
Xdh
95
97
98
100
101
D. tv. willistoni
2320
.000
.007
.114
.468
.322 .99998
D. w. quechua
64
.547
.422
.000
.000
.000
B.
Est-4
98
100
102
D. e. equinoxialis
2682
.150
.769
.081
.967
D. e. caribbensis
2166
.954
.036
.001
The Pan-Pacific Entomologist
276
Table 1. ( Continued)
Subspecies
Sample
size
Locus and alleles
Probability
of correct
diagnosis
of the
subspecies
Acph-1
88
94 100
102
104
106
108
D.
e.
equinoxialis
702
.000
.013 .172
.000
.811
.000
.014 .986
D.
e.
caribbensis
2062
.635
.027 .113
.005
.019
.176
.010
Mdh-2
94
106
D.
e.
equinoxialis
1900
.994
.000
.9997
D.
e.
caribbensis
1572
.0006
i .989
G3pdh
92
96
100
D.
e.
equinoxialis
32
.000
.063
.906
.977
D.
e.
caribbensis
754
.019
.939
.029
Hk-1
96
100
D.
e.
equinoxialis
428
.082
.914
.975
D.
e.
caribbensis
988
.982
.009
ability of misclassification of a single individual of known genotype is
8.0 X 10- 11 .
It should be noted that in the formal descriptions which follow, the
frequency with which individuals exhibit a given band is calculated on
the basis of genotypic frequencies. These frequencies can be calculated
from the data given in Table 1 by assuming Hardy-Weinberg
equilibrium.
Drosophila willistoni Sturtevant
This species becomes the nominate subspecies, Drosophila willistoni
willistoni Sturtevant.
Drosophila willistoni quechua Ayala, new subspecies
Morphologically indistinguishable from Drosophila willistoni willistoni as de¬
scribed by Sturtevant (1921), but differing from it by allozyme patterns in
electrophoretic assays for following enzymes. (1) Xanthine dehydrogenase: in
buffer system of pH 9.0, bands migrating anodally less than bands exhibited by
D. w. willistoni. (2) Esterase-2-, in buffer system of pH 8.65, bands migrating
anodally less than bands exhibited by D. w. willistoni. (3) Octanol dehydro-
Vol. 49, No. 3, July 1973
277
genase-l : in buffer system of pH 9.0, nearly always (more than 99.9 percent of
individuals) exhibits band with further anodal migration than common bands of
D. w. ivillistoni. (4) Esterase-4 : in buffer system of pH 8.65, band migrating
anodally less than most common band exhibited by D. w. willistoni. (5)
Esterase-7 : in buffer system of pH 8.65 nearly always (98 percent of the in¬
dividuals) with a band migrating anodally less than common bands of D. w.
willistoni.
Holotype male, laboratory reared, original stock from PERU: De¬
partment of Lima, near Lima, collected by net sweeping over fruit
baits in June 1972 by Prof. Danko Brncic, reared at Department of
Genetics, University of California, Davis by F. J. Ayala, killed 6-IV-
1973. Allotype and 497 paratypes (207 males, 290 females), same data
as holotype, with paratypes also dated 10-IV-1973, either point mounted
or preserved in alcohol. Holotype and allotype, point mounted, de¬
posited in the California Academy of Sciences, Department of Entomol¬
ogy, Type Number 11786. Paratypes deposited in the collections of the
California Academy of Sciences, National Museum of Natural History,
Washington, D.C., and University of California collections at Berkeley,
Davis, and Riverside.
The original stock consisted of 38 inseminated females, maintained
in separate cultures. Crosses made among the 38 cultures. The holo¬
type, allotype and paratypes are progenies from these crosses.
Drosophila equinoxialis Dobzhansky
This species becomes the nominate subspecies, Drosophila equinoxialis
equinoxialis Dobzhansky.
Drosophila equinoxialis caribbensis Ayala, new subspecies
Morphologically indistinguishable from Drosophila equinoxialis equinoxialis
as described by Dobzhansky (1946), but differing from it by allozyme patterns
in electrophoretic assays for following enzymes. (1) Malate dehydrogenase-2: in
buffer system of pH 9.0, nearly always (more than 99.999 percent of individuals)
exhibiting band with further anodal migration than most common band of D. e.
equinoxialis. (2) Acidphosphate-1: in buffer system of pH 8.65, nearly always
(more than 99.9 percent of individuals) with bands migrating anodally either
less (most often) or more than most common band of D. e. equinoxialis. (3)
Glyceraldehyde-3-phosphate dehydrogenase-, in buffer system of pH 7.0, nearly
always (more than 99.9 percent of individuals) exhibiting bands with less anodal
migration than most common band of D. e. equinoxialis. (4) Hexokinase-1: in
buffer system of pH 7.0, nearly always (more than 99.9 percent of individuals)
exhibiting bands with less anodal migration than most common band of D. e.
equinoxialis. (5) Esterase-4 : in buffer system of pH 8.65, nearly always (more
than 99.8 percent of individuals) exhibiting bands with less anodal migration
than most common band of D. e. equinoxialis.
278
The Pan-Pacific Entomologist
Holotype male, laboratory reared, original stock from PUERTO
RICO: Mayagiiez, collected 24-11-1972 by net sweeping over fruit baits
by F. J. Ayala, reared at Department of Genetics, University of Cali¬
fornia, Davis by F. J. Ayala, killed 6-IV-1973. Allotype and 214
paratypes (103 males, 111 females) same data as holotype, with some
paratypes killed 10-IV-1973, either point mounted or preserved in
alcohol. Holotype and allotype, point mounted, deposited in the Cali¬
fornia Academy of Sciences, Department of Entomology, Type Number
11787. Paratypes deposited in the collections of the California Academy
of Sciences, National Museum of Natural History, Washington, D.C.,
and University of California collections at Berkeley, Davis, and River¬
side.
The original stock consisted of 151 females, maintained in separate
cultures. Crosses were made among the 151 cultures. The holotype,
allotype and paratypes are progenies from these crosses.
Geographic Distribution: Puerto Rico, Hispaniola and Costa Rica
(Limon province).
Summary
Two new subspecies of the Drosophila willistoni group are described.
The subspecies are established on the basis of partial reproductive
isolation (hybrid sterility) between allopatric populations. Members
of each pair of subspecies differ from each other by the patterns of
their enzymes as assayed by gel electrophoresis. Using five different
enzymes, the probability of incorrect identification of the subspecies of
an individual of known genotype is less than 1 X 10 -10 . This is the
first time that allozyme patterns have been used in the formal descrip¬
tion of a taxon.
Literature Cited
Ayala, F. J., and J. R. Powell. 1972. Allozymes as diagnostic characters of
sibling species of Drosophila. Proc. Nat. Acad. Sci., USA, 69: 1094-
1096.
Ayala, F. J., J. R. Powell, M. L. Tracey, C. A. Mourao, and S. Perez-Salas.
1972. Enzyme variability in the Drosophila willistoni group. IV. Genetic
variation in natural populations of Drosophila willistoni. Genet., 70:
113-139.
Bobzhansky, T. 1946. Complete reproductive isolation between two morphologi¬
cally similar species of Drosophila. Ecol., 27: 205-211.
Dobzhansky, T. 1956. What is an adaptive trait? Amer. Nat., 90: 337-347.
Spassky, B., R. C. Richmond, S. Perez-Salas, O. Pavlovsky, C. A. Mourao, A.
S. Hunter, H. Hoenigsberg, T. Dobzhansky, and F. J. Ayala. 1971.
Vol. 49, No. 3, July 1973
279
Geography of the sibling species related to Drosophila willistoni and
of the semispecies of the Drosophila paulistorum complex. Evol., 25:
129-143.
Sturtevant, A. H. 1921. The North American species of Drosophila. Carnegie
Inst. Washington Publ. 301, pp. 1-150.
SCIENTIFIC NOTE
Speculation on the distribution of the Southern California species of
Cafius with a new record from the Salton Sea (Coleoptera: Staphylinidae)
-—-Members of the genus Cafius are found throughout the world along the seashore
and on the banks of rivers near the sea. Of the seven species of Cafius known from
southern California, one (C. canescens Maklin) is abundant, three (C. seminitens
Horn, C. lithocharinus LeConte and C. luteipennis Horn) are sometimes common
and three ( C. sulcicollis LeConte, C. opacus LeConte and C. decipiens LeConte)
are rare. Cafius canescens is known from Alaska to Baja California, C. seminitens
and C. luteipennis from British Columbia to Baja California, C. lithocharinus from
Washington to Baja California, C. sulcicollis from southern California and Baja
California and the very rare C. decipiens has been recorded only twice from San
Diego (LeConte 1863, Smithson. Misc. Coll., 167: 1-92; Casey 1885, Bull. Calif.
Acad. Sci., 1: 285-336). Cafius sulcicollis, recorded from Magdelena Island, Baja
California (Horn 1894, Proc. Calif. Acad. Sci., 4: 302-449), is the only species
known to occur in the southern part of the poorly explored peninsula of Baja
California. On March 3, 1968, Kenneth W. Cooper collected three specimens of
C. sulcicollis at Desert Beach, Salton Sea, Riverside County, California. These
specimens are in the collection of the University of California at Riverside. To our
knowledge, this is the first record of any species of seashore beetle from the shores
of the Salton Sea. Bledius ferratus LeConte, a coastal species which has been re¬
ported from the Salton Sea (Herman 1972, Bull. Arner. Mus. Nat. Hist., 149: 113—
254), lives in salt marshes, not on sea shores, and is widespread in the deserts of
southern California. It probably inhabited those areas long before the formation
of the Salton Sea.
The Salton Sea was formed in 1904, when the Colorado River overflowed its
banks. It is located 235 feet below sea level, in the Colorado Desert about eighty
miles inland from the Pacific Ocean over a mountain range whose lowest pass
is 2600 feet, and about 100 miles north of the Gulf of California. Its present sa¬
linity is slightly greater than that of sea water. The shores of the Salton Sea pro¬
vide a habitat more like that of the large enbayments of southern Baja California
than that of the sea beaches of southern California. The climate of Baja California
is semi-desert in the northwest, becoming progressively more arid to the south
until that of the middle of the peninsula is similar to the climate of the Colorado
Desert except for heavy fogs along the Pacific Coast. It seems likely that Cafius
sulcicollis (along with C. decipiens and C. opacus ) is distributed along the Pacific
shores of southern Baja California, and is uncommon in southern California be¬
cause that region is at the northern extreme of its range.
280
The Pan-Pacific Entomologist
Species of Cafius are known to feed at least in part on larval Diptera in decaying
seaweed (James, Moore and Legner 1971, Trans. San Diego Soc. Nat. Hist., 16:
279-289). Some areas of the collection locality at Desert Beach were covered with
a layer of decaying tamarisk needles containing dipterous larvae which could have
provided suitable food.
How Cafius sulcicollis came to the Salton Sea is a matter of conjecture. This
species is a strong flyer which sometimes swarms along the beaches (Leech and
Moore 1971, Wasman J. Biol., 29: 65-70). Gravid females possibly could have
been carried over the mountains during storms, or they might have been trailered
to the Salton Sea in boats. It is unlikely that the species was introduced from the
Gulf of California as no member of the genus is known from the northern part of
the Gulf.— Ian Moore and E. F. Legner, Division of Biological Control, University
of California, Riverside, 92502.
BOOK REVIEWS
The Insects of Australia. Edited by I. M. Mackerras. Melbourne University
Press. U. S. Distributor: International Scholarly Book Services, Inc. 1029 p.,
704 figs., 8 color plates, Color frontice plate. 1970. $22.50 (U.S.).
The days are past in which a single author can produce a comprehensive text¬
book in general entomology with an up to date coverage of the higher classification
of insects to the family level for any major region. Thus the present book is a
product of the efforts of 30 entomologists with diverse specialties. However, this
type of approach also has its problems, especially in providing a uniform, co¬
ordinated and yet sufficiently concise treatment to fit into a single volume. The
efforts of the editor, I. M. Mackerras, and of D. F. Waterhouse, Chief, Division
of Entomology, CSIRO who coordinated many of the activities involved, have been
highly successful in providing us with a well balanced product. Both men and all
the contributors are to be congratulated for their efforts.
This tome of over 1,000 pages is extremely impressive not only in its size and
weight, but in the nature and quality of its contents. The first nine chapters of
about 200 pages provide a general treatment of the morphology, physiology, cyto¬
genetics, developmental biology, natural history, phylogeny, systematics, and
zoogeography of insects. The next 28 chapters of about 750 pages present the
systematic treatments of each of the classes of hexapods and orders of insects.
The difficulties of one author producing a comprehensive treatment of general
entomology are akin to the difficulties of one reviewer trying to evaluate the efforts
of 30 authors. So I will concentrate on the broader aspects of the volume and
and leave the detailed criticisms of specific chapters to experts in those areas.
The first nine chapters provide an excellent coverage of the basic aspects of
insect structure, function and biology. However, those interested in the applied
aspects of entomology will undoubtedly be disappointed in the lack of coverage
given to agricultural, medical and forest entomology and biological control. One
finds only brief references to the importation of Cactoblastis for the control of
Opuntia and the export of Rodolia to California for the control of Icerya, both
Vol. 49, No. 3, July 1973
281
classical examples of biological control involving Australia. Most chapters deal¬
ing with individual orders contain at least a token section in the introduction in
which the economic importance of the order is mentioned. The book is mainly
directed at an audience interested in the classification and natural history of in¬
sects of the Australian region.
The introductory chapters contain much information of a type not found in
other general entomological textbooks available in English. A considerable amount
of biochemistry (structure and formation of the integument, hormones, metabolism,
and excretion) is discussed and pathways figured in a clear manner in the chapter
by D. Gilmour, et al. Chapters on Cytogenetics by M. J. D. White; Fossil history
by E. F. Riek; and Composition and distribution of the fauna by I. M. Mackerras
provide extremely useful summaries of information not readily available to the
general student or specialists in other areas.
Much of the higher classification differs from that found in other recent text¬
books providing us with some refreshing new views about insects and their relatives
and with plenty of fuel for future polemics over our classificatory schemes. The
Collembola, Diplura and Protura are not included in the Class Insecta but each
is treated as a separate class in a chapter on entognathous hexapods. The ectog-
nathous apterygotes are divided into two orders Archeognatha (Meinertellidae and
Machilidae) and Thysanura (rest). Orthopteroid groups including: Blattodea,
Mantodea, Grylloblattodea, Phasmatodea and Orthoptera (sensu Saltitoria) are
each accorded ordinal rank. The Megaloptera, Raphidioptera, and Neuroptera
also are treated as separate orders. On the other hand a conservative grouping is
taken in the Hemiptera (Homoptera and Heteroptera) and the Phthiraptera (Mal-
lophaga and Anoplura). Although three orders (in the sense of this volume),
Grylloblattodea, Zoraptera, and Raphidioptera, do not occur in the Australian
region, the first two are each discussed in brief chapters.
The systematic chapters each contain discussions of adult anatomy, immatures,
biology, world classification at the family level with notations as to the occurrence
and abundance of families in Australia, keys and brief discussions of each family
in Australia.
This will remain one of the most valuable general references for all entomol¬
ogists for some time to come. Those with a special interest in the zoogeographic
relationships of the Southern Continents also will find this a basic tool in their
work. —Robbin W. Thorp, University of California, Davis, 95616.
American Entomologists. By Arnold Mallis. Rutgers University Press, 30 College
Avenue, New Brunswick, New Jersey 08903. Pp. XVIII -j- 549, 211 -f 8 illlus-
trations. October, 1971. $15.00.
Reference books tend to be dull reading, but here is one which can also be
read from cover to cover or kept as a bedside companion, with equal pleasure. It
is surely a labor of love—and over the years a remarkable amount of work must
have gone into its preparation. As a reference it will be in constant use, yet it
is equally for the reader and browser, highly recommended and a pleasure to use.
It is dedicated to S. W. Frost, and to the memory of E. C. Van Dyke and J. J.
Davis.
Mallis retells for us the lives of 211 entomologists. His studies are based on
282
The Pan-Pacific Entomologist
published obituaries, enriched wherever possible from notes obtained by extended
and widespread correspondence with “old timers.” References and quotations are
cited so that we may verify if we wish. But it is his ability to pick out the activ¬
ities, eccentricities and sayings, which make the personalities come alive and be
remembered.
Many entomologists were intentionally omitted from the present volume. In
the Preface Mallis writes “I can only hope that I have used good judgment in my
selection, and, where I have failed, possibly I can make amends at some future
date.” Students will look forward to a second volume with anticipation.
One is impressed by the number of entomologists who spent their boyhoods on
farms or on acreage near a town or city, and by the number who were from, or
moved to, Kansas. Many a lifelong interest was sparked by seeing another person
collecting, or by the chance reading of one of the early textbooks or agricultural
reports, not surprising in a period when most entertainment had to be home-
produced, while books were scarce and prized. Reading the sketches, one often
wishes for more about certain individuals (for instance, J. W. Folsom), yet the in¬
formation may never have been published; perhaps the writers did not know the
deceased intimately, or thought that the human side of a story did not belong in
“serious” obituaries. It is typical of obituaries of entomologists that little credit
is given to their parents, wives or husbands, who so often encouraged them in their
hobby or career at the expense of needed family items and despite public opinion
of mere bug chasers. Appreciation by the entomologists themselves is found in the
dedications of monographs and books.
Attention may be called to one error. The Pacific Coast Entomological Society
is correctly cited in places, but is also called the Pan Pacific Entomological Society
(p. 169) and the Pacific Entomological Society (p. 215). H. C. Fall is stated to
have been the founder of the society (p. 268), but it was first called together by
Charles Fuchs; Fall acted as temporary president for that meeting of 15 August
1901, at which Fuchs was elected president of the society, then known as the Cal¬
ifornia Entomological Club. But these are small points, and we are greatly indebted
to Arnold Mallis for making so many entomologists live again for us.— Hugh B.
Leech, California Academy of Sciences, San Francisco, 94118.
Monographie der Familie Platypodidae Coleoptera. By Karl E. Schedl. Verlag
Dr. W. Junk N. V., Den Haag, The Netherlands. Pp. VI + 322, 73 figs. 1972.
Price 70 Dutch florins (with postage, just over $25.00 U.S.).
The ambrosia beetles are fungus-carrying wood borers of wide distribution,
especially numerous in tropical and subtropical regions. Although small they may
occur in huge populations, and some are of real economic importance; a few spe¬
cies have spread with the aid of man’s commerce. Dr. Schedl has had a lifelong
interest in the systematics and bionomics of bark and ambrosia beetles, his col¬
lection contains over 80% of the recognized species of Platypodidae. The book
is up to date and includes references into 1971; the lists of species, with citations,
are particularly valuable because the previous world catalogue, by Strohmeyer
in Junk and Schenkling’s Coleopterorum Catalogus, dates to 1912.
The first 80 pp. include a short historical sketch on a geographic basis, and
treat of the morphology and anatomy of larvae and adults, the biology in detail,
Vol. 49, No. 3, July 1973
283
population dynamics, outbreaks, economic importance, control, and relevant
literature, fossils, phylogeny, geographic distribution, and comments on errors in
the literature. The remainder of the book is a taxonomic treatment of the family,
with identification keys to subfamilies and genera, and in large genera to species
groups; for instance there is a key of 62 couplets to the descriptively named sec¬
tions (e.g., Platypi opaci-carinati) of the over 500 listed species of Platypus. The
illustrations are good, some outstandingly so; there is a 44 page index. The bind¬
ing is attractive, but the pages, though of good quality paper, are not sufficiently
opaque, so that text and figures show through.
Despite this very fine treatment, it is clear that the last word has not been said.
F. G. Brown’s major paper “Larvae of the principal old world genera of the
Platypodidae” (1972. Trans. Roy. Entomol. Soc. London, 124(2) : 167-190,
54 figs.) uses a very different classification, which is based also on his extensive
knowledge of the adults.— Hugh B. Leech, California Academy of Sciences, San
Francisco, 94118.
Essays in Evolution and Genetics in Honor of Theodosius Dobzhansky. A
Supplement to Evolutionary Biology. Edited by Max K. Hecht and William C.
Steere. Appleton-Century-Crofts. 594 p., 70 figs., 85 tables. 1970. $16.00.
A volume covering a wide variety of subjects dedicated to one of the most out¬
standing evolutionists of our time on the occasion of his seventieth birthday. The
series of 16 essays written by 27 of Professor Dobzhansky’s colleagues begins with
a chapter on his own life summarizing his many achievements and including a
bibliography of over 400 publications.
Many of the essays deal with insects, especially Drosophila. In Chapter 4
“Competition, Coexistence and Evolution,” Francisco J. Ayala presents a theo¬
retical discussion of these concepts and documents most of his arguments with
data from populations of Drosophila. A considerable portion of Chapter 8 “Hetero¬
zygosity and Genetic Polymorphism in Parthenogenic Animals” by M. J. D.
White deals with various parthenogenic insects. Chapter 9 “Evolutionary Studies
on Maniola jurtina (Lepidoptera Satyridae) : ‘The Boundary Phenomenon’ in
Southern England, 1961 to 1968” by E. R. Creed, W. H. Dowdeswell, E. B. Ford
and K. G. McWhirter presents a review of their published studies through 1960
and a year by year account of the shifts in the boundary between the two female
wing spotting forms from 1961 through 1968. Chapters 10 through 15 deal with
a variety of studies on Drosophila and its relatives. Chapter 10 “The Genetic Basis
of a Cell-Pattern Homology in Drosophila Species” by T. M. and Rose M. Rizki
discusses the use of a specific biochemical phenotype (kynurenine producing) of a
cell to study of speciation in relation to developmental genetics of hybrids between
two closely related Drosophila. Chapter 11 “Ecological Factors and the Variability
of Gene-Pools in Drosophila ” by John Beardmore presents evidence that habitat
heterogeneity strongly influences the amount of variability in gene-pools. Chapter
12 “Mating Propensity and Its Genetic Basis in Drosophila ” by Eliot B. Spiess
points out that females do the selecting among the diversity of courtship patterns
presented by the males and thus the organism plays an active role in its own
selective process. Chapter 13 “Observations on the Microdispersion of Drosophila
melanogaster ” by Bruce Wallace stresses that the slight extent to which flies move
284
The Pan-Pacific Entomologist
coupled with the enhanced success of chance migrants in mating fits well with
the observed mating patterns, allelism, and dispersal of D. melanogaster. Chapter
14 “Studies on the Evolutionary Biology of Chilean Species of Drosophila” by
Danko Brncic analyses chromosomal structure in relation to distribution patterns
in space and time. Chapter 15 “The Evolutionary Biology of the Hawaiian Dro-
sophilidae” by Hampton L. Carson, D. Elmo Hardy, Herman T. Spieth and Wilson
S. Stone analyses what has become one of the best known complexes of drosophilids,
primarily due to the efforts of these researchers.
The hook is a great tribute to a great scientist and a worthy addition to an
entomologist’s library.— Robbin W. Ti-iorp, University of California, Davis, 95616.
INSECT PINS
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ENTOMOLOGICAL EXCHANGES AND SALES
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for $1.50 per line. Ads will be arranged in order of their receipt. Contact the
Advertising Manager, W. H. Lange, Department of Entomology, University of Cali¬
fornia, Davis, California 95616.
FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca,
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FOR SALE. Over 900 species of butterflies from the Malaysian and Indonesian
Jungles, very rare moth (14" wing span), Scorpion, giant sized Indonesian flying
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Sim Yam Seng, 21, Joon Hiang Road, Singapore, 19, Singapore.
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American moths. The color photography has been done with excruciating care and attention
to detail, and must be seen to be believed.
THE MOTHS OF AMERICA, written by authorities in each group treated, constitutes a
thorough review of the North American moths, with descriptions of new genera and species
where appropriate.
In addition to the color photographs of adults, illustrations of genitalia or other structures
are included where necessary to assure accurate identification. The comprehensive text gives
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THE MOTHS OF AMERICA will be published in over 100 parts, over a period of about 20
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Please start your subscription now, or order the parts you desire, before the cost be¬
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ALREADY PUBLISHED:
Fascicle 13, Part la, Pyralidae: Scopariinae & Nymphulinae, by Dr. Eugene G. Munroe.
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Fascicle 13, Part lb, Pyralidae: Odontiinae & Glaphyriinae, by Dr. Eugene G. Munroe.
1972. 116 p. $20.60 ($16.50 to subscribers).
Fascicle 20, Part 2a, Bombycoidea: Saturniidae, in part (Citheroniinae and Hemileucinae, in
part), by Dr. Douglas C. Ferguson. 1972. 153 p., 11 color plates, 19 figs. $39.00
($32.50 to subscribers).
Fascicle 20, Part 2b, Bombycoidea: Saturniidae, in part (Hemileucinae, in part, and Satur-
niinae), by Dr. Douglas C. Ferguson. 1972. xxi + 121 p., 11 color plates, 11 figs.
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color), 8 halftones, 19 figs. $24.00 ($19.60 to subscribers).
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Vol. 49
OCTOBER 1973
No. 4
THE
Pan-Pacific Entomologist
MAYO—A new species of Baetis from Ecuador (Ephemeroptera: Baetidae) .. 285
SHAPIRO—Recurrent aberration in Cynthia annabella; A review with four
new records (Lepidoptera: Nymphalidae) _ 289
PARKER AND POTTER—Biological notes on Lithurgus apicalis Cresson
(Hymenoptera: Megachilidae) _ 294
McCAFFERTY AND EDMUNDS—Subgeneric classification of Ephemera
(Ephemeroptera: Ephemeridae) _ 300
MAYO—Four new species of the genus Baetodes (Ephemeroptera: Baetidae) 308
NELSON—Synopsis of the genus Chilenoperla (Plecoptera: Gripopterygidae) 315
WASBAUER—Some new taxa in the Myrmosinae with keys to the females of
North America (Hymenoptera: Tiphiidae) ___ 325
CHEMSAK AND LINSLEY—New species of Eburia from Mexico (Coleop-
tera: Cerambycidae) _______ 338
GRISSELL—New species of Eurytoma associated with Cynipidae (Hymenop¬
tera: Eurytomidae) ____ 354
ALLEN—New species of Leptohyphes Eaton (Ephemeroptera: Tricorythidae) 363
MATHIS—A review of the genus Borboropsis (Diptera: Heleomyzidae) .... 373
CHANDLER—The Hamotus of Arizona (Coleoptera: Pselaphidae) _ 378
DALY—Lectotype designations of African bees of the genus Ceratina
and allies (Hymenoptera, Apoidea) _ 383
CHRISTIANSEN AND BELLINGER—Six new Nearctic species of the genus
Friesea (Collembola: Poduromorpha) _ 389
WHEELER AND WHEELER—Ant larvae of the subfamily Dolichoderinae:
second supplement (Hymenoptera: Formicidae) _ 396
SCIENTIFIC NOTES _ 288, 324, 337, 353, 377, 395, 402, 403
BOOK REVIEW ________ 299
BOOK NOTICE ____. 372
RECENT LITERATURE ____________ 388
ZOOLOGICAL NOMENCLATURE _____ 404
INDEX TO VOLUME 49 ___ 405
SAN FRANCISCO, CALIFORNIA • 1973
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
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The Pan-Pacific Entomologist
Vol. 49
October 1973
No. 4
A New Species of Baetis From Ecuador
(Ephemeroptera: Baetidae)
Velma Knox Mayo
2702 E. Seneca St., Tucson, Arizona, 85716
Baetis ellenae Mayo is one of a group of large species that occurs
from Mexico to Peru. When mature, nymphs characteristically climb
out of the water onto rocks or logs wet by spray.
Baetis ellenae Mayo, new species
(Figs. 2, 3, 6)
Male imago. —Length: body 10 mm; fore wing 10 mm; caudal filaments 26
mm; body color yellow with rust markings. Head bright yellow; bases of antennae
yellow, bordered with brown distally; distal segments dark smoky or black;
median carina with a blackish brown band; eyes turbinate, orange-red, oval and
moderately convex. Thorax bright yellow; mesonotum olive tinged with rust
colored speckles near wing bases; metanotum yellow, postscutellum bright rust;
pleural sclerites yellow, reddish near coxae, on posterior portion of episternum
and on pleural trochantin; fore wing amber tinged, veins distinctly reddish,
marginal intercalaries paired (Fig. 2) ; hind wings margined with reddish-brown
and with heavier reddish-brown border from base to costal projection (Fig. 3) ;
hind wings with only two longitudinal veins; legs yellow to orange with rust
and blackish brown markings; fore coxae yellow with rust streaks; trochanters
tinged with rust; fore femora bright orange tinged with blackish brown apically;
tibiae slightly longer than femora, yellow with blackish brown apex; tarsi slightly
shorter than tibiae, yellow, brown at joints; fourth tarsomere and claw smoky;
middle and hind legs yellow, penciled with black on posterior and anterior margins;
femora yellow, penciled with black distally; tarsi yellow, narrowly margined with
red; fourth tarsal segment and claw smoky. Abdominal terga bright yellow with
blackish-brown posterior borders on either side of midline and laterally near
pleural fold, borders wider on tergum 1. Genital forceps as in fig. 6. Caudal fila¬
ments pale yellow with reddish-brown joints.
Female imago. —Length: body 8 mm; fore wing: 10 mm. Thorax reddish
olive-brown; parapsidal grooves yellow. Abdominal terga reddish brown, posterior
borders dark brown; abdominal sterna yellow with reddish ganglionic markings.
Nymph: figs. 1, 4, 5, 7-10.—Length: body 11-13 mm; caudal filaments 7.5-8.5
mm; body color pale amber. Head pale amber; antennae pale yellow, first seg¬
ment large; mouthparts as in figs. 4, 5, 8-10. Thorax amber; legs pale yellow;
femora reddish brown at apices and along margins, with short spinules on dorsal
crests; tibiae with short spines at apices; tarsal claws with 8 denticles (fig. 7).
The Pan-Pacific Entomologist 49: 285-288. October 1973
286
The Pan-Pacific Entomologist
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Figs. 1-2. Baetis ellenae Mayo. Fig. 1. Dorsal view of nymph. Fig. 2. Fore
wing of imago.
Posterior borders abdominal terga amber with dark reddish brown medially and
mesad to gills (fig. 1) ; terga 3-7 with reddish-brown streaks along midline in
some specimens; gill trachea pale to dark reddish brown; abdominal sterna and
caudal filaments amber.
Holotype male imago, Rio Amayo, Macuchi, Cotopaxi Province, Ecuador;
altitude about 6000 ft., V. K. Mayo, 30 June 1943. Entomological collection, Uni¬
versity of Utah, Salt Lake City. Female and 1 paratype imago, same data, same
deposition as holotype. Paratypes: 4 nymphs, same data, 7 July 1942; 1 nymph
Vol. 49, No. 4, October 1973
287
Figs. 3-10. Baetis ellenae Mayo, various structures of nymph and imago. Fig.
3. Hind wing of imago. Figs. 4^5. Right and left mandible, respectively, of nymph.
Fig. 6. Rights forceps of adult male. Fig. 7. Tarsus and claw of nymph. Figs.
8-10. Labrum, labium and maxilla, respectively, of nymph.
288
The Pan-Pacific Entomologist
same data, 22 November, 1942 ; 4 nymphs same data, 6 March 1943; 16 nymphs
near Pilalo Road at 5 Km., 7 March 1943. 12 nymphs same data, 30 June 1943.
All nymphs same deposition as holotype. Nymphs and adults were associated
by rearing.
Hie nymphs of B. ellenae are the most common species of the genus
in the Macuchi region. Mature specimens commonly climb out of the
water onto a log or rock where the spray wets them. The only other
nymph described in the group with similar habits was designated Raetis
sp. 1 by Roback (1966). Nymphs of the two species are of the same
size but differ in coloration. B, ellenae is pale amber, while Baetis sp. 1
is brown. The two species also differ in details of the mouthparts (fig-
4, 5, 8, 10) and figs. 77, 80-83 (Roback 1966). This species is named
in honor of the writer’s daughter, Ellen Mayo Harbert.
I he writer wishes to acknowledge the help of Jay R. Traver, Massa¬
chusetts L niversity, Amherst, Massachusetts who read the manuscript.
She is also indebted to Richard K, Allen of California State University
at Los Angeles for help in editing the manuscript, and to W. L. Peters,
Florida A. and M. University, Tallahassee, who contributed valuable
suggestions.
Literature Cited
Kohack, S. S. 1966. Catherwood Foundation Peruvian-Amazon Expedition. VI-
Ephemeroptera nymphs. Monogr. Acad. Sci. Ptiila,, 14: 129 -199, 22 pi.
SCIENTIFIC NOTE
Records for Georyssidae and Cyathoceridae (Coleoptera) in Mexico.
VVhen collecting in Sonora on November 31. 1972 at the Arroyo Cuyuchaba, H-5
milts utst of Alamos, Karl Stephan and I picked up three species of two lesser
known families of Coleoptera, the Georyssidae and Cyathoceridae (= Lepieeridae)-
Hie species collected were Georyssus minor Sharp (3 individuals), Cyathocerus
hnnn Sharp (2 individuals) and Cyathocerus bufo Hinton (2 individuals). The
specimens were identified by comparison with Hinton’s descriptions of the three
speeies (1933, Pan-Pac. Entomol. 9: 160-162).
lhe beetles were obtained by placing river drift, which had been deposited on
the banks of the arroyo after two days of heavy rain, in a pan of water and then
ting tin smuggling Coleoptera. The cyathocerids and georyssids were ob¬
served when they were climbing through the debris just under the surface of the
water. They did not attempt to climb on top of the floating debris as did the
other Coleoptera in the pan. Hinton obtained his larger series of the species from
an identical habitat, damp trash piles left by a river just west of Mexico City at
lejupilco.
collection of the three species were the northernmost records for the
tv\ . tarp species (C. bufo is known only from its type locality of Tejupikol-
it records from Alamos extend their range approximately 700 miles, to the
northern edge of the thorn forest in Sonora.— Donald S. Chandler, University oi
Arizona, Tucson 84721.
Vol. 49, No. 4, October 1973
289
Recurrent Aberration in Cynthia annabella
A Review with Four New Records
(Lepidoptera: Nymphalidae)
Arthur M. Shapiro
Department of Zoology , University of California, Davis, 95616
Three species of Painted Lady buUerflies occur in California: Cyn¬
thia annabella Field = C, carye of authors, C. cardui Linnaeus, and
C. virginiensis Drury. All produce aberrant phenotypes characleiizet
by suppression of the discal wing pattern, fusion of the subapical dark
pattern elements, and development of a series of white submaiginal spots.
These phenotypes were named in all three species, mostly in the 1C 4* u
of aberration naming in the first three decades of this centuty. n>
attracted a great deal of attention because of theii remaikabh
in the three related species and because they occurred with excepUona
frequency considering their drastic nature. When Giinnell c e. cr
“ Vanessa earyae (sic!) var. intermedia ” and “var. letchen iGrinnell,
1918) he had before him three and nine specimens respectively, in at-
dition to four of the similar “var. muelleri'— all hut three nort ern
California, and all collected between 1911 and 1917.
The record of these and similar aberrations in the liter at uic 1
tensive, and may be traced in the bibliographies gi\( n by ie
i i r i, * n rardiLi * elymi Rambur, eniictyrtii
under the following names: in L. caraiu. tuyt ’ . f
Verity, hemielymi Gaede; in C. virginiensis. ah teas it re ox,
annabella : muelleri Letcher, intermedia Crinnell, letc en
schraderi Guilder. The oldest records are of C. card,a aberration
elymi,” which has been known since the early nineteenth tenuity am
was figured clearly from England by Newman (1874). In C. anna e a
this type of variation was first described by Letcher (1898) I var. mue -
leri”). Subsequently Grinnell (1918) named “var. intermedia and vat.
letcheri,” less and more extreme manifestations <>f tht same Y
and Gunder {19291 named yet another member of the series ..
form schraderi." All of these are infrasubspecific names and have no
standing in formal zoological nomenclature. I he \ MI,L 1
tensively discussed and numerous specimens were figuret y un ‘ 1 ^
(1925, 1927a, 1927b, 1929) and Comstock (1927). Most pu i is e.
records-about 35-da.e from this period. The unsavory reputation
of aberration naming led to a decline in published records. One recen
record is by Phillips (1971) of a "muelleri from Salt Lake City, L a ,
The Pan-Pacieic Entomology 49 : 289- 293. October 1973
290
The Pan-Pacific Entomologist
«
Vol. 49, No. 4, October 1973
291
near the eastern edge of the range of C. annabella. There is no evidence
that the actual incidence of these phenotypes has decreased-—only the
level of interest in them. They are, however, no better understood today
than forty years ago.
Speculation on their evolutionary significance diverged. Grinnell
was of the opinion that they had arisen in C. annabella quite recently
and were increasing in frequency in the species. He interpreted their
occurrence in the three species as an example of orthogenesis. Gunder
and Comstock, conversely, interpreted it as a primitive trait derived
from a common ancestor—a case of reversion or “atavism.” The ex¬
treme similarity of the phenotypes leaves little doubt that they are homol¬
ogous in origin, but whether they are parallel developments or persis¬
tent from a common ancestor remains unknown. No living species of
Cynthia or its near relatives has a normal pattern anything like that of
any of them.
Dimock (1968) reported the induction of “ elymi ” in C. cardui by
chilling fresh pupae for 14 days at 36° F. As Grinnell noted, however,
the aberrant phenotypes have been taken in the summer months when
chilling to the requisite degree would be very unlikely in the collection
localities. Either Dimock’s specimens are phenocopies and the wild
ones are produced genetically without the intervention of an abnormal
environment, or else the phenotypes can be produced by more than one
kind of stress. The largest number of captures in C. annabella is in
September and October, but there are records for every month but Jan¬
uary, March, April, and December.
I have taken three fresh aberrant C. annabella in the Sacramento
Valley in nine months, the first such cluster of captures reported in some
forty years. The records are: Willow Slough, 2.5 miles NE Davis, Yolo
County v-10-72 (figs, la, 2a) ; Putah Creek Recreation Area, 3 miles
SW Davis, Yolo County xi-17-72 (figs, lb, 2b) ; American River basin,
North Sacramento, Sacramento County ii-18-73 (figs, lc, 2c). In ad¬
dition, another specimen of slighthly less aberrant phenotype was taken
at Suisun City, Solano County on v-19-72 (figs. 3a, b). The pattern
shown by this individual, while clearly related to the others, is unlike
<e-
Figs. 1-2. Cynthia annabella aberrations discussed in text. Fig. 1. Dorsal
surfaces: a, Yolo County, California, v-10-1972; b, Yolo County, xi-17-1972;
c, Sacramento County, ii-19-1972; d, slightly aberrant individual, Yolo County,
iii-7-1973. Fig. 2. Ventral surface of above specimens.
292
The Pan-Pacific Entomologist
Fig. 3. Dorsal and ventral surfaces of slightly aberrant individual, Suisun City,
Solano County, v-19-1973.
any specimen previously figured. These were among about 3800 wild
C. annabella examined for beak marks during this period in the course
of a predation-frequency study. The only other specimen showing the
slightest variation in this direction is shown in figures Id and 2d. It
was taken at Putah Creek on iii-7-73, fresh from the pupa, and shows
a very minor reduction of the discal markings on the forewings. About
100 other individuals of fresh C. annabella were examined in January,
February, and early March after the very cold 1972-73 winter, but none
was abnormal.
Postulating that the aberrant phenotypes of C. annabella are produced
by a single recessive allele whose genotypic frequencies follow the
Hardy-Weinberg law, the homozygote frequency would be 0.001 and
the gene frequency 0.032. About six per cent of the hypothetical pop¬
ulation would be heterozygous carriers. Such frequencies could not be
maintained by mutation alone and would have to be considered a poly¬
morphism. If the mutant allele were dominant, the Hardy-Weinberg
law would give a gene frequency of 5 X 10 -4 , a figure within the range
of reasonable mutation rates. If dominance is an evolved characteristic,
as is generally believed, then if the aberrant phenotypes of C. annabella
and its relatives were due to a dominant, recurrent mutation this would
be strong evidence for the “atavism” interpretation—reflecting the per¬
sistence of previously selected dominance modifiers, a condition called
by E. B. Ford and his colleagues “paleogenic.”
Literature Cited
Comstock, J. A. 1927. Butterflies of California. Author, Los Angeles. 334 p.
Dimock, T. 1968. An extreme experimental aberration of Vanessa cardui
(Nymphalidae). J. Lepid. Soc. 22: 146.
Vol. 49, No. 4, October 1973
293
Field, W. D. 1971. Butterflies of the genus Vanessa and of the resurrected gen¬
era Bassaris and Cynthia (Lepidoptera:Nymphalidae). Smithson. Con-
trib. Zool. 84: 1-105.
Grinnell, F., Jr. 1918. Some variations in the genus Vanessa (Pyrameis).
Psyche 25: 110-115.
Gunder, J. 1925. Several new aberrant Lepidoptera. Entomol. News 36: 193-200.
1927a. New transition forms or “abs” and their classification (Lepid., Rho-
palocera). Entomol. News 38: 129-138.
1927b. Transition forms (Lepid., Rhopalocera). Entomol. News 38: 263-271.
1929. An addition to Cynthia carye Hubner (Lepidoptera, Nymphalidae).
Pan-Pac. Entomol. 6: 9.
Letcher, B. 1898. Variation of Pyrameis carye Hubner. Entomol. News 9: 38.
Newman, E. 1874. An Illustrated Natural History of British Butterflies and
Moths. David Bogue, London. 486 pp.
Phillips, W. L. 1971. Rare aberrant forms of Utah Cynthia, the Painted Ladj
(Lepidoptera: Nymphalidae). Great Basin Natur. 31: 256-260.
294
The Pan-Pacific Entomologist
Biological Notes on Lithurgus apicalis Cresson
(Hymenoptera: Megachilidae)
F. D. Parker and H. W. Potter
Bee Biology and Systematics Laboratory, Agr. Res. Ser., Logan, Utah 84322
The tribe Lithurgini contains only the genus Lithurgus Latreille, a
group that is considered to be among the least specialized of the Mega¬
chilidae (Mitchell, 1962). Nothing has been published concerning the
biologies of the North American species except for floral records (Mich-
ener, 1951), though one species, L. apicalis Cresson, has been pestif¬
erous in constructing its nests in the doorways of homes in Arizona
(G. E. Bohart, personal communication). During June 1972, several
females of this species were observed entering holes in a cottonwood
log near Myton, Utah. In September of that year, pieces of the log
containing nests were collected and studied.
Biology
Nesting site. —Ten miles NW Myton, Duchesne County, Utah, in a
weathered, barkless cottonwood log (foundation of an abandoned po¬
tato cellar). The log was oriented east-west, and the bees were entering
holes in the west end.
Nest construction. —The log in which Lithurgus was nesting was
riddled with fingerlike tunnels, but we could not determine whether the
bees had excavated their own burrows. European species of this genus
are reported to gnaw out their tunnels in rotted wood (Malyshev, 1936),
and Torchio (personal communication) has observed L. echinocacti
(Cockerell) nesting in rotted wood in Arizona. In extracting the nests
from the logs, most were extensively damaged, but five completed nests
were located and extracted from lateral galleries leading off an en-
->
Figs. 1-10. Figs. 1-3. Nests of L. apicalis: (1) nest with 2 cells, a) en¬
trance plug (note leaf pieces and other debris), b) cocoons. (2) three-celled
nest with pollen provisions. A) indicates cell partitions. (3) Enlarged view of
bottom cell in fig. 2. A) indicates egg chamber. Figs. 4-6. Cocoons of L. apicalis:
(4) Cocoon with fecal pellets covering outer surface. (5) Cocoon with anteriorly
placed pellets. (6) Cocoon with posteriorly placed pellets. Figs. 7-8. Pre¬
pupae of bombyliid parasite, Anthrax cintalapa: (7) Lateral. (8) Ventral.
Figs. 9-10. Prepupa of L. apicalis: (9) Ventral. (10) Lateral.
The Pan-Pacific Entomologist 49: 294-299. October 1973
Vol. 49, No. 4, October 1973
295
296
The Pan-Pacific Entomologist
larged tunneled-out area. The tunnels ranged from 5.3 to 8.9 cm long
and 6 to 7 mm in diameter.
The bees had initiated cell series in both clean (new?) and old tun¬
nels (with debris at the end) by packing pollen tightly against the basal
end of the burrow and constructing a partition on top of the packed
provisions. (The pollen adhered to the partitions, Fig. 3.) In multi-
celled nests, the cells were placed end to end and were separated by
partitions that served both as cell bases and caps. Each cell was packed
with pollen from partition to partition.
The cell partitions were made primarily from scrapings of wood fi¬
bers, but they also included debris from old Lithurgus nests. The ma¬
terial for the partitions was packed into shallow saucer-shaped discs
ranging in thickness from 2 to 6 mm and thinly coated above with nectar.
This material readily dissolved in water. The entrance plugs, 1.3 to 6.3
cm long, were composed mostly of debris such as wood fibers, feces,
pieces of cocoons, parts of bees, and pollen. Several nests had pieces of
leaves incorporated into the entrance plug (Fig. 1) but these undoubt¬
edly came from old nests of Megachile pacifica (Panzer) [=M. rotun-
data authors] since cells of this leaf cutter were found in other galleries,
and females were observed entering the holes actively used by Lithurgus.
No vestibular cell was made, and the entrance plug was constructed di¬
rectly above the last cell by filling the tunnel with tightly packed debris
(Fig. 1) and by varnishing the entrance with a thin coating that ap¬
peared to be nectar. In vertical nests, the entrance plug was horizontal
across the burrow, but in obliquely-oriented tunnels, the plugs were
slanted.
Nest provisions. —The provisions were exclusively Opuntia pollen,
which was tightly packed into the cells with little or no nectar. The pol¬
len grains were easily dislodged from the cells. An egg chamber was
made in the middle of the provision. We were unable to find an un¬
disturbed egg chamber; but in one cell, a young larva had died shortly
after hatching and this chamber measured 3X3 mm and was cup-shaped
(Fig. 3). An old world species of this genus, L. fuscipennis Lepeletier,
constructs an egg chamber near the bottom and within the provision
(Lieftinchk, 1939).
Figs. 11-19. Figs. 11-13. Pupa of bombyliid parasite, Anthrax cintalapa:
(11) Dorsal. (12) Ventral. (13) Lateral. Figs. 14-16. Female pupa of L. ap-
icalis : (14) Dorsal. (15) Ventral. (16) Lateral. Figs. 17-19. Male pupa of
L. apicalis: (17) Dorsal. (18) Ventral. (19) Lateral.
Vol. 49, No. 4, October 1973
297
298
The Pan-Pacific Entomologist
Larval habits. —The larval feeding habits could not be reconstructed
accurately, but apparently the larva moves about the cell, at least during
cocoon spinning. In some nests, no cell partitions were found between
cocoons; and we concluded that they had been dislodged during cocoon¬
ing (cell partitions easily fell apart when touched). In nests where lar¬
vae had died before cocooning, all cells (22 mm long) were separated
by partitions. However, we did find longer cells (44 mm) in some nests.
These may have resulted when an egg or larva failed to mature and the
larva above or below it dislodged the cell partition and moved the con¬
tents of both cells around.
Most fecal pellets were in short and uniform links (Fig. 4) but some
ribbon like strands were noted. The shape and color of the pellets were
uniform: flattened dorsoventrally, twice as wide as high, round later¬
ally, and yellowish-tan.
The walls of the brood cells were closely lined with silk to make a
cocoon that was smooth and shiny on the inside but had loose strands
and a crinkled texture on the outside. Cocoon shape was influenced by
the configuration of the cell walls, hut generally the bottom was round,
and the top was flat (Figs. 5, 6). In some cells, additional silk was spun
against the cell partition so a cap covered the anterior tip of the cocoon
similar to cocoons of another megachilid, Hoplitis fulgida Cresson. The
cocoons were white when first observed; but after overwintering, they
became light brown (cocoons in frozen nests remained white).
Cocoons made in larger cells had feces at either end (Figs. 5, 6), but
those in shorter cells had excrement surrounding but not incorporated
into the cocoon (Fig. 4). Cocoons spun by female larvae averaged
longer, 14.1 (10-17 mm) than those spun by males, 10.6 (8-12 mm).
Prepupal larvae were pale white, C-shaped with an enlarged posterior
end, spiculate with sparse, scattered setae (0.06-0.08 mm long), (Figs.
9, 10). The larvae of L. apicalis have an aptypical setal pattern; other
megachilid genera have more setae. Both male and female pupae are
illustrated in Figs. 14-19.
Overwintering. —The species entered diapause as a prepapal larvae.
The prepupal larvae were rather flaccid and moved slowly after being
disturbed and overwintered within the cocoon in this stage.
Sex ratio. —Only 24 adults emerged; 13 were females.
Parasites. —Many (41.5%) of the cells contained immature stages
of the bombyliid, Anthrax cintalapa Cole. These beefly larvae were
found within the cocoon of the host, and they overwintered as prepupae.
The prepupa and pupa of this parasite are shown in Figs. 7-8, 11-13.
Vol. 49, No. 4, October 1973
299
Literature Cited
Lieftinchk, M. A. 1939. Uit het leven van Lithurgus atratus, een Indisch hout-
bijtje. De Tropische Natuur 28: 193-201.
Malyshev, S. 1936. The nesting habits of solitary bees. A comparative study.
Eos, Madr. 11: 201-309.
Miciiener, C. D. 1951. Genus Lithurgus Latreille, p. 1136-7. In C. F. W. Muese-
beck et al. Hymenoptera of America North of Mexico. Synoptic Cat¬
alog. USD A Agr. Monogr. 2, 1420 p.
Mitchell, T. B. 1962. Bees of the Eastern United States. Vol. II. Tech. Bull.
152, No. Carolina Agr. Exp. Sta., 557 p.
BOOK REVIEW
An Index to the Described Life Histories, Early Stages and Hosts of the
Macrolepidoptera of the Continental United States and Canada. Harrison
Morton Tietz. Published by A. C. Allyn, Allyn Museum of Entomology, Sara¬
sota, Florida. Two Vols. iv -j- 1042 pp., First Edition, March 1973. Available
from Entomological Reprint Specialists, P. O. Box 77224, Dockweiler Station,
Los Angeles, California 90007. $25.00.
This comprehensive bibliography of the described immature stages and reported
foodplants of the North American Macrolepidoptera will he welcomed by lepi-
dopterists. An extensive literature has been reviewed by Dr. Tietz, covering the
period following Heniy Edwards’ Bibliographical, Catalogue of the Described Trans¬
formations of North American Lepidoptera (1889) through about 1950. The book,
prepared from a manuscript left by Dr. Tietz after his death in 1963, is introduced
by W. D. Field and J. F. Gates Clarke. The first volume begins with a list of
over 350 periodicals and separate works consulted, followed by a list of nearly
1000 common names of butterflies and moths associated with their scientific names.
The main body of the book lists species alphabetically by scientific name. Sub¬
species, lower taxa and many synonyms are included in this list and are cross-
referenced to the main entry under the valid name. Each main entry is followed
by a bibliography of life history references and a list of foodplants reported for
that species. These entries continue into the second volume. A list of common and
scientific names of plants, and a list of foodplants with each entry followed by a list
of Lepidoptera reported to feed on that plant are also included.
As a guide to published foodplant records, life history data, and as an aid in
attempting to identify immatures, this book is extremely useful. It should not
be used as a source of data however. Foodplants are listed without citation of
supporting data (oviposition record, larval rearing, etc.) and cross reference to
literature sources are not given. In a few cases foodplants are given which are
not mentioned in the literature cited for that species.
Although it is regretable that the book was not published some twenty years
ago, anyone investigating nearly any aspect of foodplant ecology of Macrolepidop¬
tera will find it a valuable guide to the literature.— Carll Goodpasture, Uni¬
versity of California, Davis, 95616.
300
The Pan-Pacific Entomologist
Subgeneric Classification of Ephemera 1, 2
(Ephemeroptera :Ephemeridae)
W. P. McCafferty
Department of Entomology, Purclue University, West Lafayette, Indiana 47907
and George F. Edmunds, Jr.
Department of Biology, University of Utah, Salt Lake City, Utah 84112
The distinctive genus, Ephemera Linnaeus, is the oldest recognized,
the largest in number of species, and the most widespread geographically
of all the genera of Ephemeridae. It is primarily of Holarctic and
Oriental distribution, and also known from the Ethiopian Region.
The name Ephemera was first established by Linnaeus (1758) to
include all mayflies. After limiting the genus, Eaton (1868) designated
E. vulgata L. as its type. Since being comprehensively redefined by
Eaton (1883-88) Ephemera has maintained relatively stable taxonomic
limitations. The ephemerid genera Eatonica , Hexagenia, and lchthy-
botus were originally based on species first described in Ephemera.
Navas (1922) erected the genus Nirvius ; subsequently, however, Lestage
(1922) synonymized this name with Ephemera.
Demoulin (1955) erected the genus Afromera for three Ethiopian
species, two of which had been originally described as Ephemera.
McCafferty (1971) described an additional species of Ephemera from
Africa and first presented evidence for doubting the generic status of
Afromera. On the basis of a detailed review of all character distribution
within the family, we herein designate Afromera as a synonym of
Ephemera: Ephemera Linnaeus (—Afromera Demoulin new synon¬
ymy. We therefore presently recognize four described species of Ephem¬
era in Africa including Ephemera congolana (Demoulin) new combi¬
nation. The characters used by Demoulin to distinguish Afromera ,
namely the tarsal claws of the male fore legs, the posterior margin of the
subgenital plate, and the anastomose venation of the wings, have all been
found to be variable to inconsistent degrees on either an individual or
specific level throughout Ephemera. Furthermore, there is no evidence
from the larval stages that would support recognition of a separate genus.
The genus Ephemera can best be distinguished from other genera of
Ephemeridae by the following characteristics. In both sexes of the
1 The research on which this report is based was in part supported by grants from the National
Science Foundation to the University of Utah, G. F. Edmunds, Jr., Principal Investigator.
2 Published with the approval of the Director of the Purdue University Agricultural Experiment
Station as Journal Series No. 5124.
The Pan-Pacific Entomologist 49 : 300 - 307 . October 1973
Vol. 49, No. 4, October 1973
301
Fig. 1-6. Ephemera ( Ephemera ) spp. 1 and 2. Fore and hind wings, E.
supposita Eaton. 3. Female terminal abdominal segments, ventral, E. simulans
Walker. 4. Larval head, E. supposita. 5. Antennal base, Ephemera sp. 6. Gill 1,
E. sp.
adults the median terminal filament is subequal in length to the cerci.
Other characters common to both sexes are found in the wing venation.
In the hind wings (Fig. 2) MP 2 is almost always attached basally to
CuA; and in the fore wings there are 3 or more veinlets extending from
Ai to the anal margin. Additionally, in the males the tarsal claws of
the fore legs are never hooked and the genital forceps are 4-segmented.
Larval Ephemera can easily be differentiated from other larval
ephemerids on the basis of the presence of (1) the non-spuriferous
bifurcate frontal process, (2) long setae whorled over most of the length
302
The Pan-Pacific Entomologist
of the antennal flagellae, (3) labial paraglossae which are not extended
basally, and (4) the distally rounded tibiae of the fore legs.
The recent study of striking and unusual characteristics possessed
by newly discovered larvae from Thailand, India, and South Africa
has warranted a subgeneric classification of Ephemera as follows.
Subgenus Ephemera
(Figs. 1-6)
Imago. Length of male body, 10.0-20.0 mm.; fore wings, 8.0-17.0 mm. Length
of female body, 12.0-25.0 mm.; fore wings, 10.0-22.0 mm. Fore wings almost
always with MP 2 joining CuA at the basal curvature (Fig. 1). Hind wings usually
over one third of length of fore wings; at least 1 longitudinal intercalary between
R .2 and IR 3 (Fig. 2). Subanal plate of female with posterior margin variously
convex (Fig. 3). Terminal abdominal sternum of female as in Figure 3, usually
with posterolateral processes.
Mature larva. Frontal process of head longer than wide, but always less than
twice as long as wide (Fig. 4). Pedicels of antennae usually with at least 1
heavily sclerotized seta ventrally (Fig. 5). Mandibular tusks slender, circular in
cross section, more than twice length of body of mandible; left tusk sometimes
more curved, slightly longer, and usually crossing ventrally to right tusk. Labial
palpi 3-segmented. Tibiae of prothoracic legs usually with somewhat indistinct
comb of stout apical setae of posterodistal margin (similar to Fig. 21). Gill 1
with both forks similar in size (Fig. 6).
Etymology. — ephemeros, Gr.: short lived.
Type species. —Ephemera vulgata Linnaeus, 1758, by subsequent
designation (Eaton, 1868).
Discussion. Ephemera s.s. is by far the largest and most widely
distributed of the subgenera of Ephemera. It is cosmopolitan except
for the Neotropical and Australian Regions. The character states dis¬
cussed above when taken in combination will distinguish it from the
known stages of the following subgenera.
Dicrephemera new subgenus
(Figs. 7-17)
Imago. Fore wings with MP 2 not joining CuA at basal curvature (Fig. 7).
Hind wings less than one third of length of fore wings; no longitudinal intercalaries
between R 2 and IR 3 (Fig. 8). Subanal plate of female with posterior margin
straight (Fig. 9). Terminal abdominal sternum of female as in Figure 9, without
posterolateral processes.
Mature larva. Frontal process of head twice as long as wide (Fig. 10).
Pedicels of antennae with no heavily sclerotized setae ventrally. Mandibular tusks
atrophied (Fig. 11). Labial palpi with second and third segments appearing
fused. Tibiae of prothoracic legs usually lacking comb of stout apical setae at
posterodistal margin in posterior view (Fig. 12). Gill 1 with outer fork much
larger than inner fork (Fig. 13).
Vol. 49, No. 4, October 1973
303
Etymology. — dicros, Gr.: forked; ephemeros, Gr.; short lived.
Type species. —Ephemera siamensis Ueno, 1969. Ephemera siamensis Ueno,
1969, p. 235; institutional custody of male holotype not indicated.
Male imago (in alcohol). Length: body 11-11.5 mm.; fore wings 10-10.5 mm.
Head yellow dorsally, ivory anteriorly, dark brown between bases of ocelli, bases
of lateral ocelli black. Antennae with basal segments ivory, terminal segments
yellowish-brown to brown. Upper portion of compound eyes grayish-black, lower
portion black. Prothorax brown dorsally, white laterally, usually with brown
marking posterior to coxae ventrally. Mesothorax and metathorax cinnamon-brown,
lateral sutures lighter. Femora of prothoracic legs ivory, tinged with brown
apically, tibiae of prothoracic legs dark brown, tarsi light brownish-yellow, tinged
with brown proximally. Meso- and metathoracic legs ivory. Wings hyaline, vena¬
tion reddish-brown, crossveins very lightly margined with same tint. Fore wings
with subcostal area shaded dark brown becoming lighter distally, MA fork
shaded somewhat at origin. Hind wings smoky reddish-brown marginally. Ab¬
dominal terga (Fig. 14) ivory with markings as follows: tergites 3-9 each with
pair of dark brown longitudinal markings, markings in tergites 3-7 progressively
increasing in length and found almost entirely in anterior half of each segment,
markings in tergites 8 and 9 extending nearly entire length of segments, tergite
9 also with thinner submedian longitudinal markings connected posteriorly to
broader lateral markings. Abdominal sterna (Fig. 15) ivory with markings as
follows: sternite 1 cinnamon-brown except for posterior margins, sternites 3-8
each with pair of thin longitudinal dark brown markings, increasing in length
posteriorly, sternite 9 with broad marking covering entire segment posteriorly
and tapering anteriorly. Genitalia (Fig. 16) with posterior margin of subgenital
plate concave, penes broadly divergent and lobes rounded apically, covered ven¬
trally for almost entire length by thin translucent membrane arising from sub¬
genital plate, titillators short and rounded apically, often covered in ventral view
by subgenital plate membrane. Caudal filaments brown.
Female imago (in alcohol). Length: body 10-12 mm.; fore wings 10-10.5
mm. Head dark brown dorsally. Femora of fore legs light brownish-yellow with
smoky brown streak along outside margin, tibiae and tarsi of prothoracic legs
light brownish-yellow, tibiae tinged with dark brown proximally. Wing venation
light brownish-yellow, entire shading occurring in proximal half of subcostal area
of fore wing. Abdominal terga variably marked (Fig. 17) but usually yellow
with markings as follows: tergite 1 light cinnamon-brown, tergites 3-6 each with
pair of broad longitudinal dark brown markings joined posteriorly and becoming
faded at junction; tergites 7-9 marked with dark brown nearly throughout except
for median stripe, pair of short submedian stripes, and sometimes anterior margin.
Abdominal sterna without distinct markings. Caudal filaments pale with brown
annulations at apex of proximal segments. Other characters as in male except
for usual sexual differences.
Mature larva (in alcohol). Length: body 16-19 mm.; caudal filaments
5-6.5 mm. Color patterns generally corresponding to adults, although usually
lighter. Forks of frontal process (Fig. 10) slightly curved dorsally at apices,
lateral margins only slightly rounded, dorsal surface covered with short golden
setae for nearly entire length; eyes black; labrum slightly emarginate in median
third. Pronotum with pair of small dark brown triangular shaped markings, with
304
The Pan-Pacific Entomologist
Figs. 7-13. Ephemera {Diorephemera) siamensis Ueno. 7 and 8. Fore and
hind wings. 9. Female terminal abdominal segments, ventral. 10. Larval head.
11. Right larval mandible. 12. Parts of larval tibia and tarsus of left prothoracic
leg, posterior. 13. Gill 1.
bases meeting at mid line of notum; legs generally ivory, with prothoracic legs
tinged at joints with brown; tibial processes of metathoracic legs tinged with
dark brown setae, appearing projected anteriorly due to revolvement of femora.
Gills light yellowish-gray. Lateral setae along entire length of filaments, becom¬
ing shorter apically.
Specimens examined. 19 male imagos, 14 male subimagos, 17 female imagos,
16 female subimagos, and 223 larvae as follows: Thailand, Chiengmai Province,
Mae Ping, Chiengmai, 1000 feet; 9 November 1964 or 13 November 1964, W. L.
and J. G. Peters, collectors. The majority of specimens housed at the University
of Utah, Salt Lake City, and representatives deposited with the Laboratory of
Aquatic Entomology, Florida A & M University, Tallahassee; Institut Royal des
Sciences Naturelles, Brussels; British Museum (Natural History), London; and
the Laboratory of Insect Diversity, Purdue University, West Lafayette, Indiana.
Discussion. Dicrephemera presently includes only E. siamensis from
Thailand; however, we have examined larval specimens from the
Vol. 49, No. 4, October 1973
305
Figs. 14-17. Ephemera ( Dicrephemera ) siamensis Ueno. 14. Male abdominal
tergites. 15. Male abdominal sternites. 16. Male genitalia, ventral. 17. Female
abdominal tergites.
Bhavani R., in India, and the Wilge R. in the eastern Transvaal, Re¬
public of South Africa, which most assuredly represent undescribed
species of Dicrephemera. The subgenus is based primarily on the very
distinct larval stage, with the absence of well developed mandibular
tusks being most diagnostic. The adults are rather weakly differentiated
from those of Ephemera s.s., and all of the characters discussed must
be used with caution. The possibility remains that as more of the larvae
of little known Ephemera species from the Oriental and Ethiopian Re¬
gions are discovered they may prove to be Dicrephemera.
Many of the adults of E. siamensis were reared from the larvae by
Dr. and Mrs. Peters in Thailand, therefore assuring a definite associa¬
tion. The adults of E. siamensis are redescribed because of the large
series available to us in comparison to the small series upon which
Ueno (1969) based his description. The larvae are described herein
for the first time.
306
The Pan-Pacific Entomologist
Figs. 18-22. Ephemera ( Aethephemera ) nadinae, larva. 18. Head. 19 and 20.
Left and right mandibles, lateral. 21. Parts of tibia and tarsus of left prothoracic
leg, posterior. 22. Dorsal section of right gill 2.
Aethephemera new subgenus
(Figs. 18-22)
Imago. Unknown.
Mature larva. Frontal process of head as long as wide (Fig. 18). Pedicels
of antennae with no heavily sclerotized setae ventrally. Right mandibular tusk
much reduced, less than three fifths of length of left tusk (Figs. 19 and 20).
Labial palpi 3-segmented. Tibiae of prothoracic legs with comb of stout apical
setae at posterodistal margin (Fig. 21). Gill 1 with both forks slender (similar
to Fig. 6).
Etymology. — aethes, Gr.: unusual; ephemeros, Gr.; short lived.
Type species. —Ephemera ( Aethephemera ) nadinae new species.
Mature larva (in alcohol). Length: body 18.5-20.0 mm.; caudal filaments
6.9 mm. General color dark brown dorsally, brown ventrally, with golden setae
and brown spurs. Frontal process of head golden brown, deeply concave anteriorly,
lateral margins distinctly rounded, margin of concavity fringed dorsally with
long golden setae (Fig. 18) ; eyes black; labrum emarginate only in median third.
Vol. 49, No. 4, October 1973
307
Thoracic notum unicolorous brown; legs generally yellowish-brown, with femora
of metathoracic legs darker; dorsal surface of tibiae of prothoracic legs entirely
spuriferous; tibial processes of metathoracic legs with dense covering of golden-
brown setae. Abdominal terga light brown with pale median and pair of sub¬
median longitudinal markings bordered by dark brown; sterna brown with pair
of dark brown submedian longitudinal maculae on segments 7, 8, and 9; gills
grayish-purple; dorsal portion of gill 2 (Fig. 22) with inner margin produced
ventrally at base; lateral setae along entire length of cerci, and along three fourths
of length of terminal filament.
Holotype. —Mature female larva. South India, Kodaikanal grade, 1600 m.;
30 March 1962, E. S. Ross and D. Cavagnaro collectors. Deposited in the California
Academy of Sciences, San Francisco. Paratype .—mature male larva (parts on
slides) same data and deposition as holotype.
Etymology.— E. (A.) nadinae is named in honor of the wife of W. P.
McCafferty.
Discussion. Aethephemera is presently known only from the larval
stage, and is known only from India. The distinct subgenus, however,
may prove to be more widespread as larvae become known.
Acknowledgments
We wish to express our gratitude to Mr. R. G. Noble of the National
Institute for Water Research, Pretoria, South Africa, for providing us
with larval material from Africa. We also wish to acknowledge Mr.
Arwin Provonsha, Laboratory of Insect Diversity, Purdue University,
for assisting in the preparation of figures.
Literature Cited
Demoulin, G. 1955. Afromera gen. nov., Ephemeridae de la faune ethiopienne
(Ephemeroptera). Bull. Ann. Soc. Roy. Entomol., 91: 291-295.
Eaton, A. E. 1868. An outline of a re-arrangement of the genera of Ephemeridae.
Entomol. Mon. Mag., 5: 82-91.
1883-88. A revisional monograph of recent Ephemeridae or mayflies. Trans.
Linn. Soc. Lond., Sec. Ser. Zool., 3: 1-352.
Lestage, J. A. 1922. Notes sur le genre Nirvius Nav. zz Ephemera L. Bull. Ann.
Soc. Entomol. Belg., 16: 253-254.
Linnaeus, C. 1758. Systema naturae per regna tria naturae, secundum classes,
ordines, genera, species cum characteribus, differentiis, synonymis, locis.
Editio decirna, reformata, Tom. I. Laurentii Salvii, Holmiae, 824 pp.
McCafferty, W. P. 1971. New burrowing mayflies from Africa (Ephemeroptera:
Ephemeridae). J. Entomol. Soc. S. Afr., 34: 57-62.
Navas, P. L. 1922. Efemeropteros nuevos o poco conocidos. Bol. Soc. Entomol.
Espan., 7: 54-63.
Ueno, M. 1969. Mayflies (Ephemeroptera) from various regions of southeast
Asia. Oriental Ins., 3 : 221-238.
308
The Pan-Pacific Entomologist
Four New Species of the Genus Baetodes
(Ephemeroptera: Baetidae)
Velma Knox Mayo
2702 E. Seneca St., Tucson, Arizona 85716
A detailed description and discussion of the generic characteristics
of the nymphs of the genus Baetodes is presented by Mayo (1972).
Herein are described four new species of Baetodes from Mexico, Bolivia,
and Venezuela. The genus has not previously been described from
Bolivia and Venezuela. Koss (1972) described two new species of
Baetodes from southwestern United States, the first recorded from
North America. He proposed some new terms for structures which
have previously been described as spines. Those on the dorsal crests
of the femora he called clavate setae. Those along the ventral edge of
the tibiae and tarsi he designated as spurs and those on the labrum as
spine-like setae. These new terms are used in the following descriptions.
Baetodes proiectus Mayo, new species
(Figs. 1-8, 10)
Immature female nymph. —Body length 6 mm. Caudal filaments broken.
Head: as in fig. 6; vertex planar between eyes; sides of head planar between eyes
and bases of antennae; eyes narrowly rimmed with pale yellow with pale area ex¬
tending to genae; dark brown between eyes and in fork, on either side of epicranial
suture and between antennae; pale spot anterior to small ocellus; labrum dark
brown, darker than clypeus; mouthparts as in figs. 1-3, 5, 7, 8; sclerotization
along midline behind apical cleft of labrum pronounced; one bladelike seta on
tip of each glossa and on tip of each paraglossa, respectively (figs. 8, 5) ; basal
segment of labial palpi longer than other segments combined (fig. 7) ; two blade¬
like setae among the slender setae on crowns of maxillae; antennae pale brown,
white around bases; intersegmental membranes white. Thorax : pronotum dark
brown, with few white setae; posterior border elevated at midline forming fairly
prominent tubercle; mesonotum dark brown, mottled with yellow laterally and
at bases of wing pads; midline white; wing pads brown with pale veins; meta-
Figs. 1-8. Baetodes proiectus. Fig. 1. Right maxilla. Fig. 2. Labrum. Fig.
3. Left mandible. Fig. 4. Claw. Fig. 5. Left paraglossa. Fig. 6. Head. Fig. 7.
Labial palpus. Fig. 8. Left glossa. Fig. 9. Labrum of B. peniculus. Fig. 10.
Tubercle of metanotum and abdominal tubercles 1-9 of B. proiectus. Figs. 11-14.
B. peniculus. Fig. 11. Claw. Fig. 12. Left mandible. Fig. 13. Tip of left glossa
showing pinnate seta. Fig. 14. Tubercle of metanotum and abdominal tubercles
1 - 10 .
The Pan-Pacific Entomologist 49: 308-314. October 1973
Vol. 49, No. 4, October 1973
309
310
The Pan-Pacific Entomologist
notum pale, washed with pale brown along anterior border, and with prominent
white tubercle posteromedially (fig. 10) ; pleural sclerites brown, narrowly
rimmed with reddish brown posteriorly and medially above coxal articulations;
coxae and trochanters light brown; dorsal crests of coxae produced; femora light
brown with large pale area proximally, rimmed with dark brown distally; six
dark reddish brown clavate setae and fringe of white setae along dorsal crests;
tibiae light brown with longitudinal white streak, rimmed with dark brown
proximally; tarsi light brown medially, dark brown dorsally at tibial articulations
and near claws; claws as in fig. 4; segment I of sternum pale yellow; apodemes
dark brown; segment II with brown projection from sclerotized rim of apodeme
forming an erect round-tipped lobe. Abdomen: terga 1-10 brown, darker than
mesonotum, paler at bases of gills; prominent tubercles on abdominal terga 1-9
as in fig. 10; all tubercles pale, slightly tinged with pale brown; abdominal gills
white; two elongated gills on each coxa; sternum 1 pale yellow, 2-5 pale brown,
6-9 darker brown; 10 brown laterally, white medially; sternum lighter than terga;
Cerci yellow on basal third, brown distally; terminal filament yellow.
Mature male nymph. —Length: body 4 mm; caudal filaments 6 mm. Similar
to female except anterior thoracic sterna yellow with reddish brown sclerites as
on subimago; sternum III pale brown, finely mottled with light brown; abdominal
sterna 1-5 pale brown, with broad light brown band on posterior borders; sterna
6-8 dark brown; sternum 9 dark brown on anterior half and along pleural fold,
brown area wider at midline, remainder of segment yellowish; sternum 10 yellow
with reddish brown streak half way between midline and lateral border branching
to each side of bases of cerci; cerci yellowish brown, encircled with small dark
brown spines on each segment to two-thirds of length of cerci, distal third brown
with intersegmental membranes white.
Holotype female nymph, L. PI. 24 km. W. Coripata, Bolivia, 2/5 May, 1961,
P. and P. Spangler, University of Utah collection, Salt Lake City. 6 female and
3 male paratypes, same data as holotype.
Discussion. On sternum II is a brown projection from the rim of
the sclerotized area around the apodeme on each side, forming a small,
erect, round-tipped lobe on immature specimens. This is unique to
the species. On mature specimens this projection is not present, and
the sternum is yellow with reddish brown sclerites as in the sub imago.
The species was named for the projections on immature specimens.
On mature male specimens small dark brown spines encircle each seg¬
ment of the cerci. Mature females are not known. Baetodes sanc-
ticatarinae Mayo, 1972, has similar spines on the cerci of mature
specimens. However there are no abdominal tubercles on B. sanc-
ticatarinae, while those of B. proiectus are prominent.
Baetodes peniculus Mayo, new species
(Figs. 9, 11-14)
Male nympii. —Length: body 5 mm; caudal filaments broken. Dorsum brown
with unsclerotized areas pale yellow; venter pale. Head : midline and epicranial
suture pale; dark brown laterally and in fork; turbinate eyes dark red; pale
Vol. 49, No. 4, October 1973
311
yellow laterad to large ocelli; genae brown, pale yellow anterior to small ocellus,
pale around bases of antennae; two basal segments of antennae pale ventrally,
brown dorsally; distal segments brown; mouthparts as in figs. 9, 12, 13; pinnate
bladelike seta apically on each glossa (fig. 13) ; long straight simple seta medially,
slightly curved at tip, on each paraglossa; two pointed bladelike setae among
setae on crowns of maxillae. Thorax : pronotum amber, midline pale, with two
brown stripes on either side of midline and dark brown band along anterior
border; mesonotum dark reddish brown laterad of pale midline, amber laterally
and on scutellum with paired dark reddish brown spots laterad of midline anterior
to scutellum, an elongate dark spot parallel to border of wing pad, and two dark
spots anteriorly near bases of wing pads; wing pads amber brown dorsally, yellow
ventrally; venation pale yellow; metanotum brown, midline pale with brown
tubercle posteriorly (fig. 14) ; pleural sclerites brown with narrow reddish brown
posterior border; coxae and trochanters brown; femora brown with a pale yellow
spot near proximal articulation; pale longitudinal stripe and irregular brown
border along dorsal crests becoming dark reddish brown at tibial articulations;
dorsal crests with six short, barely discernable clavate setae; tibiae brown with
distinct white streaks; tarsi brown, darker distally; claw as in fig. 11; sternum
white with brown apodemes at leg bases. Abdomen: terga 1-3 and 7-10 brown;
terga 4-6 yellow dorsally and brown laterally, dark brown along posterior borders;
each terga with prominent, pointed, brown tubercle directed posteriorly (fig. 14) ;
abdominal gills oval, white with main tracheole brown at proximal half; few
minute setae along margin, especially in basal half; sternum 1 white; sterna 2-9
yellow, anterior and lateral borders washed with brown; sternum 9 darker brown
laterally and delicately shaded with brown medially along posterior border;
sternum 10 brown laterally, pale yellow medially. Cerci and terminal filament
light brown.
Female nymph. —Similar to male except for larger size; body length 6 mm.
Tubercle on metanotum somewhat larger and tubercle on tergum 1 smaller than
in male.
Holotype male nymph, Mer., 13 km. E. Apartaderos, Venezuela, 24 February,
1969, P. and P. Spangler. In University of Utah collection, Salt Lake City. Para-
types, 4 nymphs, 2 males and 2 females same data and deposition as holotype.
Discussion : Baetodes peniculus has a pinnate seta on the tip of each
glossa (fig. 13) and one nonpinnate long straight seta medially on
each paraglossa as on Baetodes sancticatarinae Mayo, 1972. These
species can be distinguished by the absence of abdominal tubercles in
B. sancticatarinae and the absence of spines on the cerci in B. peniculus.
Baetodes longus Mayo, new species
(Figs. 15-23)
Female nymph.— Length: body 4 mm, cerci 6 mm. Color yellow with brown
markings. Head: elongate; light brown between eyes and in fork, pale anterior
to small ocellus; antennae pale yellow. Mouthparts as in figs, 15, 17-21. Labrum
elongate; with 3 long spine-like setae equidistant from cleft and lateral sclerotized
area (fig. 15). Thorax: pronotum margined anteriorly with light brown band,
mottled with light brown laterad of midline; mesonotum margined anteriorly with
312
The Pan-Pacific Entomologist
m0i
'tiiM
Figs. 15-23. Baetodes longus. Fig. 15. Labrum. Fig. 16. Tarsus and claw.
Fig. 17. Right mandible. Fig. 18. Maxillary palpus. Fig. 19. Labial palpus.
Vol. 49, No. 4, October 1973
313
reddish brown, and with light brown stripes laterad of midline, mottled with
light brown laterally; scutellum and wing pads yellow; metanotum brown with
median marginal tubercle posteriorly; pleural sclerites yellow, posterior borders
narrowly rimmed with reddish brown, coxal articulations dark brown; legs yellow,
trochanters reddish brown distally; femora with 10 or more pale brown clavate
setae along dorsal crests; tibiae broadly rimmed with reddish brown proximally,
very narrowly rimmed with reddish brown distally, set with minute brown spurs
ventrally, these longer distally; tarsi yellow, becoming brown dorsally at bases
of claws and with single row of long reddish brown spurs ventrally (fig. 16) ;
claws as in fig. 16. Sternum pale yellow except for brown apodemes at leg bases.
Abdomen: terga 1-8 yellow with reddish brown anteriorly, darker on terga 1-3;
terga 1-7 with prominent yellow tubercles (fig. 22) ; abdominal gills as in fig. 23;
two elongate finger dike gills on each coxa. Sternum 1 pale yellow; sterna 2-10
light yellowish brown. Cerci yellow at basal half, light brown distally. Terminal
filament yellow.
Male nymph. —pronotum paler than mesonotum; brown on thoracic sternum
anterior to leg bases II and III; abdomen as in female but terga 4-6 pale yellow
dorsally half way to pleural fold.
Holotype female, Rio Frio, Tamaulipas, Mexico, 24 December, 1939, L.
Berner, in University of Utah collection, Salt Lake City. 78 paratype nymphs,
same data and deposition as holotype. 153 paratype nymphs, Rio Guayalejo,
Tamaulipas, Mexico, 22 December, 1939, L. Berner. Same deposition as holotype.
Discussion : Baetodes longus can be distinguished from all other
known species of Baetodes by the elongated head and mouthparts (figs.
15, 17, 20) and by the long tarsal spurs (fig. 16). It differs from the
Mexican species Baetodes inermis Cohen and Allen, 1972 by the fol¬
lowing characteristics: the general color of B. inermis is much darker
than that of B. longus ; the abdominal tubercles on terga 1-7 of B.
inermis are poorly developed, those of B. longus are prominent; Coxal
gills are absent on B. inermis , they are present on B. longus. Intra¬
specific variation is as follows; abdominal terga 4-8 may be dark
reddish brown anteromedially; terga 9-10 may have brown tracheations
along the pleural folds.
Baetodes furvus Mayo, new species
(Figs. 24-28)
Mature female nymph. —Length: body 4 mm, cerci about 7 mm. Color dark
brown with white intersegmental membranes. Head: brown with pale yellow
midline, narrowly pale yellow around eyes and over small ocellus; mouthparts as
Fig. 20. Left mandible. Fig. 21. Right glossa. Fig. 22. Abdominal tubercles 1-7.
Fig. 23. Gill. Figs. 24-28. B. furvus. Fig. 24. Left mandible. Fig. 25. Abdominal
tubercles 3-7. Fig. 26. Tip of left glossa. Fig. 27. Crown of maxilla. Fig. 28.
Lab rum.
314
The Pan-Pacific Entomologist
in figs. 24, 26-28; labrum with 2 long, 1 short, 1 long and 2 spine-like setae laterad
at midline (fig. 28) ; crown of one maxillary fringe of hairs bent mesad revealing
two blade-like setae (fig. 27). Thorax : pronotum and mesonotum amber mottled
with reddish brown; anterior margin of mesonotum dark brown, with paired
reddish brown streak laterad of midline terminating in a point followed by an
elongate spot, laterally mottled with reddish brown; scutellum and wing pads
amber; metanotum dark brown with blunt posteromedial tubercle; pleural
sclerites dark brown; femora brown with large yellow area proximally and yellow
stripe medially, set with numerous reddish brown clavate setae along dorsal crests,
narrowly rimmed with dark reddish brown distally; tibiae brown, narrowly
rimmed with reddish brown proximally; tarsi brown, becoming dark brown distally;
claw with 8 denticles. Abdomen: terga dark brown; intersegmental membranes
white; terga 1-6 with prominent tubercles, tubercle on tergum 7 reduced (fig. 25) ;
gills white, oval; coxal gills absent; sterna 2-7 brown with median pale streak
and broad pale band subadjacent to gills; sterna 7-9 dark brown laterally, pale
medially, with pale portion widening posteriorly; sternum 10 pale amber; cerci
light brown, terminal filament pale yellow.
Holotype female, Culinavara, Mexico, 1 January, 1948, collector unknown,
University of Utah collection, Salt Lake City. 9 paratype nymphs same data and
deposition as holotype.
Discussion: Baetodes furvus resembles Baetodes fuscipes Cohen and
Allen (1972) but is distinguished by the following characteristics: B.
fuscipes has a well developed median posterior elevation on the pro¬
notum, there is no posterior elevation on the pronotum of B. furvus;
the abdominal tubercles on terga 1-6 of B. fuscipes are moderate in
size, the tubercles of B. furvus are prominent; abdominal terga in
B. fuscipes vary from dark brown on segments 1-3 to light brown on
9-10, in furvus the terga are uniformly dark brown; in fuscipes the
caudal filaments are light brown, in B. furvus pale yellow.
Acknowledgments
The author wishes to acknowledge the kindness of Dr. Jay R. Traver,
University of Massachusetts, for the loan of the collection of Baetodes
nymphs of Dr. Louis Berner. She is indebted to Dr. George F. Edmunds,
Jr., University of Utah for the loan of Baetodes nymphs from Mexico,
Bolivia, and Venezuela.
Literature Cited
Cohen, S. D. and R. K. Allen. 1972. New species of Baetodes from Mexico
and Central America (Ephemeroptera: Baetidae). Pan-Pac. Entomol.,
48: 123-135.
Koss, Richard W. 1972. Baetodes : New species and new records for North
America (Ephemeroptera: Baetidae). Entomol. News, 83: 93-102.
Mayo, V. K. 1972. New species of the genus Baetodes (Ephemeroptera: Baeti¬
dae) Pan-Pac. Entomol., 48: 226-241.
Vol. 49, No. 4, October 1973
315
Synopsis of the Genus Chilenoperla
(Plecoptera: Gripopterygidae)
Charles H. Nelson 1
Department of Biology, University of Tennessee, Chattanooga 37401
The genus Chilenoperla was first described by lilies (1963) in his
revision of the South American Gripopterygidae and has been repre¬
sented by three species all found only in Chile. Recently, while studying
stonefly material from South America a new species of this genus,
found both in Argentina and Chile, was discovered. Examination of
relevant holotype and paratype material of the known species of
Chilenoperla reveals that lilies mistakenly designated individuals of
this undescribed species as paratypes of Chilenoperla heschi lilies.
Hence, in order to facilitate recognition of this new species descriptions,
figures and notes of the other known species of this genus have been
included in this study. Unless otherwise indicated, the morphological
terms utilized in the present work are those introduced by McLellan
(1971) in his study of the Australian Gripopterygidae. An initial esti¬
mate of the branching relationship of the four species of this genus is
also undertaken and is presented in the form of a Prim Network.
Since the identity of the females of only two species in Chilenoperla
is known, the key below is confined to males.
Key to The Males of Chilenoperla
1. Meso-posterior region of central sclerite slightly produced posteriorly (Figs.
1, 18) ; posterior sclerite demarked from central sclerite (Figs. 3, 19) _ 2
Meso-posterior region of central sclerite moderately produced posteriorly
(Figs. 6, 11) ; posterior sclerite not demarked from central sclerite (Figs.
8, 13) _ 3
2. Sub-anal lobes sharply pointed apically (Fig. 3) ; anterior sclerite with
ventro-lateral projections (Fig. 2) _ C. semitincta lilies
Sub-anal lobes bluntly pointed apically (Fig. 19) ; anterior sclerite lacking
ventro-lateral projections (Fig. 20) _ C. puerilis lilies
3. Epiproct with less than three tooth-like projections on each side (Figs. 8,
9) ; posterior sclerite bifurcated apically (Fig. 6) ; ninth posterior sternal
border broadly rounded (Fig. 7) _ C. heschi lilies
Epiproct with three or more tooth-like projections on each side (Fig. 13, 14) ;
posterior sclerite not bifurcated apically (Fig. 11) ; ninth posterior sternal
border with slight median emargination (Fig. 12) _ C. illiesi Nelson
1 Research reported in this study partially supported by a University of Chattanooga Foundation
grant. Published with the aid of funds from the University of Tennessee, Chattanooga.
The Pan-Pacific Entomologist 49: 315-324. October 1973
316
The Pan-Pacific Entomologist
Chilenoperla semitincta lilies
(Figs. 1-5)
Chilenoperla semitincta lilies, 1963, Mitt. Schweiz. Entomol. Ges. 36(3) : 223-225,
fig. 40; a (fore and hind wings), b (male terminalia, ventral), c (male
terminalia, dorsal), d (male terminalia, lateral), e (male epiproct, lateral),
f (female terminalia, ventral), g (male hind leg, lateral),
lilies, 1966, Das Tierreich, 82: 51.
Zwick, 1972, Studies on Neotrop. Fauna 7: 99.
Male. —Brown in color with dark brown embossings on the occipital region of
head and disc of pronotum. Color of each leg patterned so that proximal region
of femur and most of tibia, except for hasalmost region, yellow, remainder of leg
brown.
Male terminalia. —Ninth sternite (fig. 2) characterized by posteriorly pro¬
duced rounded subgenital plate under tenth segment.
Tenth segment (fig. 1) divided transversely by a narrow strip of membrane
into anteriorly situated anterior sclerite and a posteriorly situated sclerite,
here designated as mesal sclerite (ms). Anterior sclerite becoming greatly nar¬
rowed meso-dorsally and ventrally incomplete. Ventro-laterally on each side of
anterior sclerite a small difficult to discern projection (fig. 2, p) arising from
posteroventral margin. Roughly triangular-shaped mesal sclerite consisting of a
central sclerite (cs) and two lateral sclerites (Is). Lateral sclerites situated one
on each side of but undemarked from central sclerite.
Small, clearly demarlced, triangular-shaped posterior sclerite (figs. 1, 3, ps)
of segment eleven attached to posteroventral surface of mesal sclerite. Epiproct
very broad basally but narrowing considerably apically (figs. 1, 5). When viewed
laterally (figs. 3, 4) margin of each side of antero-dorsal surface bearing four to
five small sharply pointed tooth-like projections. Medially, postero-ventral surface
of epiproct expanded into a keel-like surface.
Cercus of each side consisting of eight segments. Dorso-basal region of basal
cereal segment expanded into a roughly triangular area which extends to lateral
sclerite of that side (figs. 1, 3, a).
Partly membranous and partly sclerotized sub-anal lobes (paraprocts) in lateral
view (fig. 3) hook-shaped in appearance and sharply pointed at apex.
Female terminalia.— The female of this species was not examined during this
study. However, if lilies description and figure of the female is accurate the pos¬
terior margin of the eighth sternite is slightly convex with a median emargination.
Material examined.— Holotype male, Bach bei Niebla (Kustenkordillere),
Meershohe, Prov. Valdivia, Chile.
->
Figs. 1-5 Chilenoperla semitincta lilies. Fig. 1. Male terminalia, dorsal view.
Fig. 2. Male terminalia, ventral view. Fig. 3. Male terminalia, lateral view. Fig. 4.
Male epiproct, lateral view. Fig. 5. Male epiproct, dorsal view, (ms = mesal
sclerite; cs = central sclerite; Is = lateral sclerite; ps = posterior sclerite; a =
dorso-basal projection of basal cereal segment; p = ventro-lateral projection of
anterior sclerite).
Vol. 49, No. 4, October 1973
317
U1U15 3'
318
The Pan-Pacific Entomologist
Chilenoperla beschi lilies
(Figs. 6-10)
Chilenoperla beschi lilies (in part), 1963, Mitt. Schweiz. Ent. Ges. 36(3): 225-
226, fig. 41; a (male terminalia, ventral), b (male terminalia, dorsal), c (male
terminalia, lateral), d (male epiproct, lateral).
Werner, 1965, Beitr. Neotr. Fauna, 4(3) : 231.
lilies, 1966, Das Tierreich, 82: 50.
lilies (1963) described the female of C. beschi in his original descrip¬
tion of this species. The identity of the female was very likely based
on the single instance in the type series where males and females were
simultaneously collected from the same location. Unfortunately, the
males are not members of C. beschi but belong to a species to be
described in this present study. Hence, the female previously described
by lilies as C. beschi is most likely that of a new species of Chilenoperla
and the identity of the female of C. beschi needs to be established.
Werner (1965) has reported the collecting of both males and females
of C. beschi from the same site and examination of his material may
be useful in determining the identity of the female of this species.
Male.— Color pattern similar to that of C. semitincta except that entire femur
of each leg is uniformly brown.
Male terminalia. —Subgenital plate (fig. 7) very similar to that of semitincta.
Anterior sclerite (fig. 6) resembling that of C. semitincta in being characterized
by a small projection arising from the postero-ventral margin of each side. Lateral
sclerites, unlike those of C. semitincta, clearly demarked from central sclerite.
Meso-posterior region of central sclerite appearing to be longer and narrower than
those of C. semitincta (figs. 6, 8). Posterior sclerite of C. beschi not clearly
demarked from central sclerite. When viewed dorsally this sclerite becomes
bifurcated distally with the apical margin of each half bluntly pointed. Epiproct
(figs. 8, 9, 10) bears on each lateral margin of antero-dorsal surface two sharply
pointed tooth-like projections. Dorso-basal region of each cereal segment (figs.
6, 8) apeparing longer and narrower in C. beschi. Sub-anal lobes of C. beschi
similar to those of C. semitincta.
Material examined. —Holotype male, Bach bei Niebla (Kustenkordillere),
Meershohe, Prov. Valdivia, Chile.
Chilenoperla iiliesi Nelson, new species
(Figs. 11-17)
Chilenoperla beschi lilies (in part), 1963, Mitt. Schweiz. Ent. Ges. 36(3): 226,
fig. 41; e (female terminalia, ventral).
The color pattern of this species is nearly identical to that of C. beschi.
Male terminalia.— Posterior margin of subgenital plate (fig. 12) with a slight
median emargination. Anterior sclerite of C. iiliesi, unlike those of C. semitincta
and C. beschi, ventro-laterally lacking a projection on each side. Lateral sclerites
Vol. 49, No. 4, October 1973
319
Figs. 6-10. Chilenoperla beschi lilies. Fig. 6. Male terminalia, dorsal view.
Fig. 7. Male terminalia, ventral view. Fig. 8. Male terminalia, lateral view.
Fig. 9. Male epiproct, lateral view. Fig. 10. Male epiproct, dorsal view.
25imn
320
The Pan-Pacific Entomologist
Figs. 11-16. Chilenoperla illiesi Nelson. Fig. 11. Male terminalia, dorsal view.
Fig. 12. Male terminalia, ventral view. Fig. 13. Male terminalia, lateral view.
Fig. 14. Male epiproct, lateral view. Fig. 15. Male epipeoct, dorsal view. Fig. 16.
Female terminalia, ventral view.
.25mm
Vol. 49, No. 4, October 1973
321
(fig. 11) clearly demarked from central sclerite. Posterior sclerite of C. illiesi
similar to that of C. beschi (figs. 12, 13) in not being clearly demarked from
central sclerite, but not bifurcated. Epiproct (figs. 13, 14, 15) resembling that
of C. semidncta in having four to five small sharply pointed tooth-like projections
on each margin of antero-dorsal surface. Apical portion of epiproct in dorsal view
(fig. 15) broader than those of C. semitincta and C. beschi. Dorsal-basal region
of each basal cereal segment (fig. 11) resembling closely that of C. beschi.
Female terminalia. —Female (fig. 16) of this species differing from that of
C. semitincta in that posterior border of eighth sternite being bluntly rounded and
lacking any median emargination.
Holotype male, Bariloche, Prov. Rio Negro, Argentina, Nov. 1926 (R. and
E. Shannon). Two males and two females, Fundo Malcho, Cordillera Parral, Prov.
Linares, Chile. The holotype will be deposited in the USNM (no. 72595).
This species is named for Dr. Joachim lilies in recognition of his
substantial contribution to the knowledge of the Plecoptera.
Chilenoperla puerilis lilies
(Figs. 18-22)
Chilenoperla puerilis lilies, 1963, Mitt. Schweiz. Ent. Ges. 36(3) : 226-227, fig. 42;
a (male terminalia, lateral), b (male epiproct, lateral). “
lilies, 1966, Das Tierreich, 82: 51.
'j
lilies (1963) description of this species is based on the male of a
late instar nymph from which the genitalia of the adult could be dissected
and examined. However, the determination of the relationship of this
species to others in the genus is somewhat hindered in that these adult
anatomical features are not completely developed.
Male terminalia.— Posterior border of ninth sternite (fig. 19) broadly rounded
and lacking any median emargination. Anterior sclerite of C. puerilis resembling
that of C. illiesi in lacking a ventro-lateral projection on each side. Meso-posterior
region of central sclerite (fig. 18) similar to that of C. semitincta in that it is
broad and only produced slightly posteriorly. Posterior sclerite (fig. 18, 20)
resembling that of C. semitincta in being well demarked from central sclerite and
triangular in shape. Asymmetrical epiproct (figs. 19, 21, 22) with three to four
small sharply pointed tooth-like projections on each margin of antero-dorsal surface.
Apical extremity of sub-anal lobes (fig. 19) differing from those of the other
species of this genus in being bluntly pointed.
Material examined.— Holotype male, Bach hei Laguna Margarita, Puella, Prov.
Llanquihue, Chile.
Relationship of the Species of Chilenoperla
It is premature to construct a phylogenetic history of the species of
Chilenoperla until more information is available concerning the various
stages in the life history of both sexes of all species. However, an
initial approximation, utilizing available characters, can be made in
322
The Pan-Pacific Entomologist
beschi
Figs. 17-23. Fig. 17. Chilenoperla illiesi Nelson, wings. Figs. 18-22.
Chilenoperla puerilis lilies. Fig. 18. Male terminalia, dorsal view. Fig. 19. Male
terminalia, lateral view. Fig. 20. Male terminalia, ventral view. Fig. 21. Male
Vol. 49, No. 4, October 1973
323
Table 1. Data Matrix consisting of the character states for the
species of Chilenoperla. Missing character states are designated by (-).
Species
Characters
1
2
3
4
5
6
7
8
9
10
11
12
C. semitincta
0
0
0
1
1
0
0
0
0
0
1
0
C. beschi
1
-
0
1
0
1
1
1
1
0
0
0
C. illiesi
1
1
1
0
0
1
1
0
0
1
0
0
C. puerilis
-
-
0
0
-
0
0
0
0
-
-
1
the form of a network based on phenetic differences ( sensu Farris,
1967) between species. A network depicts only the branching relation¬
ship of the species included and requires that no judgments be made
concerning the relative primitiveness or advancement of a character.
In this study a Prim Network, utilizing the procedure described by
Farris (1970) was constructed for the known species of Chilenoperla.
Although, in some cases Prim Networks may not yield as reliable an
estimate of branching relationships as do other quantitative cladistic
techniques (e.g., Wagner Networks) they do have advantages in that
they require a minimal amount of computation and do not necessarily
involve the use of a computer.
The Prim Network constructed during this study (Fig. 23) indicates
a close phenetic relationship between C. semitincta and C. puerilis.
Chilenoperla illiesi is nearly equidistant from C. puerilis and C. heschi
but is closer to the former species than to the latter. Analysis of the
data matrix (Table 1) of the four species reveals that C. semitincta and
C. puerilis have identical states for five characters (3, 6, 7, 8 and 9).
Chilenoperla illiesi shares with C. puerilis identical states for three char¬
acters (4, 8 and 9) and with C. heschi identical states for six characters
(1, 5, 6, 7, 11 and 12).
Descriptions of characters. — (1) femur: monocolored = 0; bicolored = 1.
(2) female eighth posterior sternal border: not emarginate = 1; emarginate = 0.
(3) Male ninth posterior sternal border: not emarginate =: 0; emarginate — 1.
(4) Male ventro-lateral projections of anterior sclerite: absent = 0; present rr 1.
(5) Male lateral sclerites: demarked from central sclerite = 0; undemarked = 1.
<-
epiproct, lateral view. Fig. 22. Male epiproct, dorsal view. Fig. 23. Branching
relationship of the four species of Chilenoperla as produced from the Prim Net¬
work. The lengths of the internodes are given in parentheses.
324
The Pan-Pacific Entomologist
(6) Male meso-posterior region of central sclerite: slightly produced posteriorly
= 0; moderately produced posteriorly = 1. (7) Male posterior sclerite: demarked
from central sclerite = 0; not demarked = 1. (8) Male posterior sclerite: not
bifurcated apically = 0; bifurcated apically = 1. (9) Number of tooth-like pro¬
jections on each side of male epiproct: three or greater = 0; less than three m 1„
(10) Apical region of male epiproct: narrow = 0; broad = 1. (11) Dorso-basal
region of male basal cereal segment: long and narrow = 0; broad and triangular¬
shaped = 1. (12) Sub-anal lobes: sharply pointed apically = 0; bluntly pointed
apically 1.
Acknowledgments
The author would like to thank Dr. Oliver S. Flint, Jr., of the United
States National Museum and Dr. Peter Zwick of the Max-Planck
Limnology Institute, Schlitz, Germany for the loan of the material
utilized in this study. The author would also like to thank Dr. Lane P.
Lester and Dr. Gene S. Van Horn for their critical reading of the
manuscript.
Literature Cited
Farris, J. S. 1967. The meaning of relationship and taxonomic procedure. Syst.
Zool., 16: 44-51.
1970. Methods for computing Wagner Trees. Syst. Zool. 19: 83-92.
Illies, J. 1963. Revision der sudamerikanischen Gripopterygidae. Mitt. Schweiz.
Entomol. Ges., 36: 145-248.
1966. Katalog der rezenten Plecoptera. Das Tierreich. No. 82: I-XXX, 1-632.
McLellan, I. D. 1971. A revision of Australian Gripopterygidae (Insecta:
Plecoptera). Australian Jour. Zool., Sup. No. 2: 1-79.
Werner, E. 1965. Beitrag zur kenntnis der Chilenischen Plecopteren. Beitr.
Neotr. Fauna, 4(3) : 227-233.
Zwick, P. 1972. On Neonemura illiesi nov. spec, and some other stoneflies from
Chile (Ins., Plecoptera). Stud. Neotrop. Fauna, 7: 95-100.
SCIENTIFIC NOTE
Synonymy in Pampoconis (Neuroptera, Coniopterygidae). —Meinander,
1973 (Notulae entomologicae 52: 25) described P. dentifera, and summarized the
differences between that species, P. latipennis, “P . punctipennis Adams in litt.,”
and “ P. uncinata Adams in litt.” While an inadvertent error, this appears suf¬
ficient to make the last two names available nomenclatorially. P. punctipennis
Adams, 1973, and P. uncinata Adams, 1973 (Pan-Pac. Entomol. 49: 250-254) are
therefore synonyms of P. punctipennis Meinander, 1973 and P. uncinata Meinander,
1973, and the specimens designated as holotypes are to be regarded as lectotypes.
—Phillip A. Adams, Department of Biological Science, California State University,
Fullerton, California, 92634.
Vol. 49, No. 4, October 1973
325
Some new taxa in the Myrmosinae with keys to
the females in North America
(Hymenoptera: Tiphiidae)
Marius S. Wasbauer
Laboratory Services, California Department of Food and Agriculture,
Sacramento 95814
The Holarctic tiphiid subfamily Myrmosinae consists of moderate
to small sized wasps in which there is a strong sexual dimorphism.
The males are fully winged with a distinctly tripartite mesosoma and
are either entirely black or black with the metasoma reddish. They are
often much larger than the females which are apterous, have a bipartite
mesosoma with a single functional suture and are reddish to brown in
integumental color, often with part or all of the metasoma and sometimes
part of the mesosoma black. In addition, the metasoma of the females
may have cream-colored integumental markings. Both sexes are known
for Myrmosa and Myrmosula. The Nearctic Myrmosina is known from
males only.
The purpose of this paper is to provide the description of a new
genus of Myrmosinae from the Colorado Desert of Southeastern Cali¬
fornia and descriptions of three new species of the Genus Myrmosula.
In addition, keys are presented to the females of the genera of Myrmosi¬
nae and the species of Myrmosula.
For making possible the loan of material used in this study, I wish
to thank the following persons and the institutions they represent: P. H.
Arnaud, Jr., California Academy of Sciences, San Francisco (CAS) ;
R. M. Bohart and R. 0. Schuster, both University of California, Davis
(UCD) ; J. A. Chemsak, California Insect Survey, University of Cali¬
fornia, Berkeley (CIS) ; J. C. Hall and S. Frommer, both University of
California, Riverside (UCR) ; and K. V. Krombein, United States Na¬
tional Museum (USNM). Mrs. Ellen Parker prepared the illustrations.
In the following descriptions, the head is considered prognathous.
Measurement of the propodeal width is between the outer rims of the
spiracles. Explanation of abbreviations used for mensural ratios is
given by Wasbauer (1968).
Leiomyrmosa 1 Wasbauer, new genus
Adult female.— Head, in posterior view, somewhat narrowed behind eyes, in
dorsal view, rounded; antennal insertions covered only by thin, narrow lamellae;
1 Gr. Leios : smooth, and Myrmosa -. a combination of the Gr. mynnex —ant and L. osus —having
the condition or quality of.
The Pan-Pacific Entomologist 49: 325-337. October 1973
326
The Pan-Pacific Entomologist
ocelli absent; clypeus evenly convex, without median proximal spine or tooth;
mandibles tridentate, without lamella on ventral margin; legs slender, not heavily
armed with spines; anterior coxa without tooth at anterior apex; anterior tarsi
with well developed rake, outer apical angle of first and second tarsal segments
with somewhat spatulate spine which is as long as succeeding segment, third and
fourth segments with progressively shorter, less spatulate spines, outer surface of
anterior basitarsus with row of several short spines; hind coxa dorsally at base
with low, rounded elevation; first metasomal sternum with small, low, toothlike
elevation anteriorly; pygidium not differentiated.
Male. —Unknown. Possibly Myrmosina Krombein.
Type of genus. — Leiomyrmosa spilota Wasbauer.
Leiomyrmosa appears to represent a modification of the Myrmosa
line. This modification has been mainly toward loss or reduction of
existing structures as shown by the lack of a clypeal spine, lack of ocelli,
shallow punctation of the head, greatly reduced punctation of the
mesosoma, lack of abundant erect pilosity, especially on the metasomal
terga, the small elevations on the posterior coxae and the presence of
only a very small tooth on the first metasomal sternum. It may be
further distinguished from Myrmosa by the presence of a tarsal rake
and from Myrmosula by the head narrowing immediately behind the
eyes, the tridentate mandibles, without ventral lamellae, the lack of
elevations at the anterior extremities of the hypostomal carinae and
absence of a tooth anteriorly at the apex of the anterior coxa.
At present, the type is the only species known.
Leiomyrmosa spilota Wasbauer, new species
(Figs. 1, 2, 19, 23)
Holotype female.— Head, mesosoma and metasoma uniformly orange-brown
except a pair of enamel white spots anteriorly on sides of second metasomal
tergum; antennae darkened apically, apical third of mandibles reddish-black.
Entire insect shining, head with small, shallow punctures, sparse on vertex, sepa¬
rated by an average of about their diameter on frons, less than their diameter on
clypeus; mesosoma impunctate laterally with faint integumental reticulation;
dorsum of pronotum and mesonotum with small, shallow punctures separated by
average of somewhat more than their diameter, propodeum nearly impunctate.
Metasoma with few small, shallow, scattered punctures before apex of each, seg¬
ment. Vestiture very sparse, consisting entirely of erect, straw-colored hairs, very
short, scarcely noticeable on frons, vertex and dorsum of mesosoma and metasoma
except for a few long hairs on pronotum anterolaterally and numerous shorter
Figs. 1-6. Heads of female Myrmosinae. Dorsal view on left, right lateral
view on right. Figs. 1-2. Leiomyrmosa spilota. Figs. 3-4. Myrmosula parvula.
Figs. 5-6. Myrmosula exaggerata.
Vol. 49, No. 4, October 1973 327
328
The Pan-Pacific Entomologist
hairs on anterior, declivous portion of first metasomal tergum; longest on clypeus,
underside of head, prosternum and coxae; metasomal sterna with scattered long
hairs before apex of each.
Head. —Rounded, slightly broader than long, TFD/FD 1.15, EH/FD 0.49,
EH/MID 0.59, MID/TFD 0.72; vertex extending in an even arc above tops of
compound eyes; antennae short, first four antennal segments in ratio of 3: 1: 1.2:
1.2; median lobe of clypeus convex in profile, apical margin simple, convex, with¬
out median lamella; mandible with apical tooth long, acute, proximal teeth very
small; occipital carina incomplete ventrally; maxillary palpi six segmented, labial
palpi four segmented.
Mesosoma .—Width ratios of mesosomal nota: pronotum 1.3; mesonotum 1.0;
propodeum 1.2; pronotal length (measured from anterior margin of collar to apex
of segment) 0.76 times width, humeral angles rounded, without projections.
Vestiture of legs consisting of sparse, long, erect hairs, without appressed hairs
or pubescence; anterior tibia without preapical spines, tibial spur spatulate, 0.73
times length of basitarsus; middle tibia with three rows of short, slender, slightly
curved spines on anterior surface, dorsal row with six spines; posterior tibia with
two spine rows.
Metasoma .—First tergum smooth, without carinae or ridges.
Length. —3.50 mm.
Variation (n =: 9) ; length 2.68-3.65 mm.; TFD/FD 1.11-1.17; EH/FD 0.45-
0.51; EH/MID 0.55-0.62; MID/TFD 0.71-0.75; color light reddish brown to
nearly black (1 specimen). All paratype material was preserved in 70% alcohol
before mounting and some color change was noted. On some specimens, for
example, the white spots on the second metasomal tergum became very faint.
However, the very dark paratype was nearly black in color when collected, so
apparently considerable color variation occurs in nature.
Holotype (CAS #11896) and six paratypes 1 mile west Blythe, Riverside
County, California, May 23-24, 1970, in cereal bowl pit traps, J. L. Johnson
and M. Wasbauer collectors (CAS, MSW) ; two paratypes, same locality, May 6,
1970, J. L. Johnson and E. L. Paddock collectors (USNM).
The type locality is a somewhat disturbed sand dune area on the
edge of extensive cultivated fields. The first two or three inches of sand
is loose and unconsolidated. Beneath this is packed, consolidated sand.
The maj or plants in the immediate area are Larrea divaricata , Franseria
dumosa and Prosopis juliflora. One hundred cereal bowl pit traps
were deployed in an area roughly 40 by 40 feet. All specimens were
taken in these between 5:00 and 8:00 p.m. Pacific Daylight Time.
Myrmosula Bradley
As currently recognized, the genus Myrmosula includes species re¬
stricted to the Nearctic region. Krombein (1940) recognized four
species, one based on both sexes, one based on the male and two based
on the female. Mickel (1940) described a fifth from the female and
presented a key to the North American females. Krombein (1946)
proposed still another species based on the female, and the descriptions
Vol. 49, No. 4, October 1973
329
presented below bring the total to nine species for North America.
Only one of these species, the widespread M. parvula (Fox) occurs in
the eastern United States. Krombein (1940) gives its distribution as
westward to Texas and Montana. The remainder are of more limited
distribution in the western United States. One species is here described
from Baja California Sur.
Myrmosula nasuta Wasbauer, new species
(Figs. 7, 8, 20)
Holotype female. —Entirely orange-brown; tarsi somewhat darkened; tips of
mandibles reddish black; second, third and fourth metasomal terga with dark
brown apical band, interrupted medially on second and fourth; ultimate tergum
entirely dark brown; second tergum with pair of anterolateral, creamy white,
comma-shaped spots; fifth tergum with medioapical creamy white spot. Entire
insect shining, punctation minute, punctures on vertex separated by average of
less than their diameter; on frons and lower face slightly larger, separated by
somewhat more than their diameter; on mesepisternum, slightly larger, separated
by less than their diameter; sparse, scattered on lateral posterior surfaces of
propodeum. Vestiture consisting of very short, erect to suberect hairs forming
dilute pubescence over most of body, giving distinct sheen to head in certain
lights and loose plush-like appearance to metasomal terga. Longer, slender, erect
hairs numerous on gena, occiput, pronotum, propodeum, first metasomal tergum
and legs, scattered on metasomal sterna.
Head .—Frons strongly produced into rather narrow, nasutiform projection, in
lateral view truncate apically, bearing a pair of short, parallel ridges which are
angulate posteriorly, posterior slope somewhat concave, in dorsal view, not
bifurcate at apex; slightly curved row of three shallow dimple-like depressions
immediately posterior to interantennal projection; medioapical margin of clypeus
subtruncate, distinctly angled at junction with lateroapical margin; mandibles
slender, long, dorsal tooth considerably shorter than ventral tooth, basal mandibular
lamella low, convex; hypostomal carina produced into noticeable recurved tooth
just mesad of posterior mandibular condyle, visible below mandibular lamella.
Mesosoma .—Propodeum in dorsal view wider than remainder of mesosoma.
Width ratios of mesosomal nota: pronotum 1.00, propodeum 1.13; anterior tarsus
with hairs of ventral basitarsal comb equal width of basitarsus at their bases;
propodeum without carina near spiracles, posterolateral surfaces smooth.
Length. —4.45 mm.
Holotype (UCD #794), 18 miles west Blythe, Riverside County, California,
October 15, 1965, F. D. Parker collector (UCD). Paratypes (12): one female,
Seeley, Imperial County, California, March 20, 1968, on ground at sundown, J. L.
Johnson collector (MSW). One female, Imperial County, California, April, 1917,
J. C. Bridwell collector (USNM). One female, Glamis, Imperial County, Cali¬
fornia, April 8, 1964, R. M. Bohart collector (MSW). One female, same locality,
April 12, 1969, sand dunes, A. R. Hardy collector (UCR). One female, Cathedral
City, Riverside County, California, April 11, 1941, ex: kangaroo rat burrow, Ross
and Aarons collectors (CAS). One female, sand dunes, 7 miles southwest of Kelso,
San Bernardino County, California, April 16-17, 1969, M. S. and J. S. Wasbauer
330
The Pan-Pacific Entomologist
ll
Vol. 49, No. 4, October 1973
331
collectors (MSW). One female, Vidal Junction, San Bernardino County, California,
April 3, 1951, P. D. Hurd collector (CIS). Two females, Gila Bend, Arizona,
March 26, 1940, R. H. Crandall collector (USNM). Two females, Tucson, Arizona,
May, 1905, Wm. M. Wheeler collector (USNM).
Variation .—Length 3.46-5.22 mm.; pale integumental spots on second metasomal
tergum comma-shaped to nearly round. Variation in the shape of these markings
is evidently not geographically correlated, as both round and comma-shaped spots
occur in population samples taken at Tucson, Arizona.
Dr. R. M. Bohart informs me that the holotype was collected at a
mat of Euphorbia sp. M. nasuta shares with the following species the
pronounced median nasutiform lobe which projects above the antennal
sockets. It differs primarily in the shape of the lobe which in lateral
view curves more strongly posteriorly and in posterior view is apically
truncate.
Myrmosula boharti Wasbauer, new species
(Figs. 13, 14)
Holotype female.- —Entirely orange-brown; ultimate tarsal segments darkened;
tips of mandibular teeth reddish-black; first through fourth metasomal terga with
dark brown apical band, interrupted medially on second; ultimate tergum entirely
dark brown; second tergum with pair of anterolateral, creamy white, rounded
spots; fifth tergum with medioapical rounded pale spot.
Entire insect shining, punctation minute, punctures on head uniformly small,
separated by an average of less than their diameter; on mesosoma, faint, shallow,
appearing as reticulations, posterior face of propodeum with a number of larger,
shallow, irregular punctures; on metasoma, distinct, separated by slightly more
than their diameters. Vestiture consisting of short, shining hairs mostly decumbent
on head, mesosoma, first metasomal tergum, and second metasomal sternum,
giving these parts distinct dilute sheen in certain lights; slightly longer, erect to
suberect on second and succeeding metasomal terga and third and succeeding
sterna; scattered, slightly longer, erect hairs over entire body, longest on metasomal
sterna, most dense on posterior face of propodeum and first metasomal tergum.
Head .—Frons strongly produced apically into rather narrow nasutiform inter-
antennal elevation, in lateral view wedge-shaped anteriorly, nearly straight dorsally;
in dorsal view with two divergent teeth at apex; frons without depressions posterior
to interantennal elevation; medioapical margin of clypeus subtruncate, set off
from lateroapical margin by small carina; mandibles stout, dorsal tooth subequal
to ventral tooth, basal mandibular lamella low, margin flat for most of its length;
hypostomal carina produced into very strong, recurved tooth just mesad of
posterior mandibular condyle, visible below mandibular lamella.
<r
Figs. 7-12. Heads of female Myrmosinae. Dorsal view on left, right lateral
view on right. Figs. 7-8. Myrmosula nasuta. Figs. 9-10. Myrmosula latericarinata.
Figs. 11-12. Myrmosula peregrinatrix.
332
The Pan-Pacific Entomologist
17
Vol. 49, No. 4, October 1973
333
Mesosoma. —Propodeum in dorsal view narrower than pronotum. Width ratios:
pronotum 1.00; propodeum 0.910; anterior tarsus with hairs of ventral basitarsal
comb equal width of basitarsus at their bases; propodeum without carina near
spiracles, posterolateral surfaces smooth.
Length. —4.18 mm.
Holotype (UCD #795), Thousand Palms, Riverside County, California,
April 9, 1964, R. M. Bohart collector. Paratype female, same data, F. D. Parker
collector (UCD).
Dr. Bohart informs me that both the holotype and paratype were
collected at mats of Euphorbia sp.
Myrmosula latericarinata Wasbauer, new species
(Figs. 9, 10)
Holotype female. —Entirely orange-brown; tips of mandibles reddish-black;
ultimate segment of antenna blackish; second and third metasomal terga with
dark brown apical band, interrupted medially on second; apical tergum entirely
dark brown; second metasomal tergum with pair of poorly defined, anterolateral,
cream-colored spots. Entire insect mat to subshining, punctation minute, punctures
on head separated by less than their diameter except narrow median band on
frons; on disc of pronotum, slightly larger, separated by about their diameter;
on mesonotum, separated by less than their diameter; on mesepisternum, very
small, shallow, scattered, merging with reticulation; on posterior face of propodeum,
sparse with scattered larger punctures; on metasomal segments, small distinct,
separated by about or slightly more than their diameter. Vestiture consisting of
very short, decumbent to longer, erect haris giving distinct sheen in certain lights
to head, mesosoma and parts of metasoma; erect hairs numerous, very short on
head and dorsum of mesosoma, longer on occiput, anterior, declivous face of
pronotum, posterior face of propodeum, first metasomal segment; scattered on
succeeding metasomal sterna; scattered on coxae and femora ventrally.
Head. —Frons produced apically into low, rounded projection, in dorsal view,
broadly bilobed at apex, in lateral view rounded, not strongly projecting dorsally;
without dimple-like depressions posterior to projection; medioapical margin of
clypeus slightly concave, distinctly angled at junction with lateroapical margin;
mandibles stout, dorsal tooth weak, much shorter than ventral tooth; basal
mandibular lamella relatively high, emarginate at broad, shallow notch; occipital
carina present, well developed, not curving toward midline of head anteriorly;
hypostomal carina produced into broad, rounded elevation mesad of posterior
mandibular condyle, visible below mandibular lamella.
Mesosoma.— Pronotum with short dorsal surface and nearly vertical anterior
declivity; mesonotum margined by sharp lateral angulation extending to propodeal
spiracle; propodeum margined by lateral carina terminating in low tooth pos-
<r
Figs. 13-18. Heads of female Myrmosinae. Dorsal view on left, right lateral
view on right. Figs. 13-14. Myrmosula boharti. Figs. 15-16. Myrmosula rutilans.
Figs. 17-18. Myrmosula pacifica.
334
The Pan-Pacific Entomologist
Figs. 19-26. Taxonomic characters of female Myrmosinae. Fig. 19. Left an¬
terior tibia and tarsus, Leiomyrmosa spilota. Figs. 20-22. Lateral view, left
Vol. 49, No. 4, October 1973
335
teriorly; propodeum in dorsal view narrower than pronotum. Width ratios:
pronotum 1.00; propodeum 0.966; anterior tarsus with hairs of ventral basitarsal
comb about half width of basitarsus at their bases.
Length. —4.84 mm.
Holotype (CAS #11897), Lanito, Bahia Concepcion, Territorio Sur de
Baja California, April 16, 1968, M. E. Irwin collector. Paratype female, 25 miles
south Santa Rosalia, April 15, 1968, 200', dry wash, M. E. Irwin collector (UCR).
The holotype has been placed on indefinite loan to the California Academy of Sci¬
ences from the collection of the University of California at Riverside.
Key to Nearctic Genera of Female Myrmosinae
1. Clypeus with median spine or tooth; first metasomal sternum with large,
conspicuous tooth or lamella; ocelli usually present; forms with heavy
punctation and sculpture _ Myrmosa
Clypeus simple; first metasomal sternum simple or with only very small,
inconspicuous tooth; ocelli absent; relatively smooth forms without heavy
punctation or sculpture _ 2
2. Mandibles with large apical tooth and two very small, low teeth on inner
margin (fig. 1) ; ventral mandibular lamella absent; anterior tarsus with
rake consisting of long, spatulate spine at outer apex of each segment (fig.
19) ; anterior coxa without tooth at apex (fig. 23) ___ Leiomyrmosa
Mandibles with two apical teeth (figs. 3, 5) ; ventral mandibular lamella
present (figs. 20, 21, 22) ; anterior tarsus without rake; anterior coxa usually
with at least small tooth at anterior apex (fig. 24) _ Myrmosula
Key to Nearctic Species of Female Myrmosula
1. Antennal tubercles completely separated, frons between their bases not
raised above remainder of frons (figs. 3, 11, 25) _ 2
Antennal tubercles more or less fused into single interantennal elevation
(figs. 5, 15) ; if bilobed, frons between base of lobes raised above remainder
of frons (fig. 26) _ 3
2. Antennal tubercles separated by distance less than their basal width, each
with small tooth posteriorly (figs. 11, 12) _ peregrinatrix Krombein
Antennal tubercles separated by distance equal their basal width, rounded
posteriorly (figs. 3, 4) _ parvula (Fox)
3. Mesonotum margined by sharp lateral angulation; propodeum with raised
dorsolateral carina continuous with spiracle, terminating posteriorly in dis¬
tinct tooth; interantennal elevation low, rounded, without teeth or ridges
(figs. 9, 10) _ latericarinata Wasbauer
Mesonotum not margined; propodeum sometimes with small posterolateral
<-
mandible of Myrmosula females. Fig. 20. M. nasuta. Fig. 21. M. rutilans. Fig.
22. M. exaggerata. Figs. 23-24. Left anterior coxa of female Myrmosinae. Fig. 23.
Leiomyrmosa spilota. Fig. 24. Myrmosula rutilans. Figs. 25-26. Heads of
Myrmosula females, posterior view. Fig. 25. M. parvula. Fig. 26. M. rutilans.
336
The Pan-Pacific Entomologist
tooth hut without raised dorsolateral carina continuous with spiracle; inter-
antennal elevation with low ridges dorsally or teeth at apex (figs. 13, 17) _ 4
4. Interantennal elevation strongly produced into nasutiform projection (figs.
7, 13) ; inferior margin of mandibular lamella evenly convex or nearly
straight (fig. 20) ; anterior tooth of hypostomal carina distinct, recurved,
visible below mandibular lamella when mandibles closed (figs. 8, 14) _ 5
Interantennal elevation not as strongly produced; either a) inferior margin
of mandibular lamella emarginate (or highest near base) (fig. 22) or b)
anterior tooth of hypostomal carina absent or very small, not visible below
mandibular lamella when mandibles closed (fig. 16) _ 6
5. In lateral view (fig. 14), interantennal elevation slender, wedge-shaped,
anterior margin nearly straight; in posterior view, with two divergent apical
teeth; at least indication of occipital carina present _ boharti Wasbauer
In lateral view (.fig. 8), interantennal elevation stout, not wedge-shaped,
anterior margin strongly curved; in posterior view, truncate with two small
apical carinae; no indication of occipital carina present _ nasuta Wasbauer
6. Interantennal elevation with pair of pronounced ridges, convergent pos¬
teriorly, apex of elevation in dorsal view terminating above bases of
antennae (fig. 15), inferior margin of mandibular lamella evenly convex
(fig. 21) _ rutilans (Blake)
Interantennal elevation with pair of low, parallel ridges, apex of elevation
extending between bases of antennae (figs. 5, 17) ; inferior margin of
mandibular lamella broadly, shallowly emarginate or highest near base at
broad angle (fig. 22) _ _ 7
7. Parallel ridges of interantennal elevation widely separated, distance between
them nearly equal their length (fig. 17) _ pacificci Mickel
Parallel ridges of interantennal elevation more closely spaced, distance be¬
tween them equal to about half their length (fig. 5) _ exaggerata Krombein
It seems likely that Myrmosula pacifica Mickel will prove to be a
synonym of M. exaggerata Krombein. I have examined, in addition to
the holotypes of both nominal species, a number of specimens from
southern and central California localities. The only one which will run
satisfactorily to pacifica in MickeTs key is a topotypic specimen col¬
lected by J. W. MacSwain on September 9, 1952. Even this specimen
exhibits a very slight emargination on the inferior margin of the
mandibular lamella. The majority of central California specimens have
the ridges on the interantennal elevation more widely spaced than do
the southern California specimens but the great majority have an
emargination on the inferior margin of the mandibular lamella. Nearly
all specimens examined have a depression on the frons, posterior to
the interantennal elevation, including the holotype of exaggerata.
Despite these considerations, however, the name Myrmosula pacifica
Mickel should be retained until more information is available on
interpopulation variation.
Vol. 49, No. 4, October 1973
DO
37
Literature Cited
Blake, C. A. 1879. Catalog of the Mutillidae of North America with descrip¬
tions of new species. Trans. Amer. Entomol. Soc., 7: 243-254.
Bradley, J. C. 1917. Contributions toward a monograph of the Mutillidae and
their allies of America north of Mexico. IV. A review of the Myrmosi-
dae. Trans. Amer. Entomol. Soc., 43: 247-290, 6 pi.
Fox, W. J. 1899. The North American Mutillidae. Trans. Amer. Entomol. Soc.,
25: 219-292.
Krombein, K. V. 1940. Studies in the Tiphiidae (Hymenoptera: Aculeata).
IV. A revision of the Myrmosinae of the new world with a discussion
of the old world species. Trans. Amer. Entomol. Soc., 65: 415-465.
1946. A new species of Myrmosa (Hymenoptera: Tiphiidae). Proc. Entomol.
Soc. Wash., 48(9) : 247-248.
Mickel, C. E. 1940. The identification of the female of the myrmosid subgenus
Myrmosula (Hymenoptera: Tiphiidae). Pan-Pac. Entomol., 16(3):
132-134.
Wasbauer, M. S. 1968. New genera of male Brachycistidinae with a redescrip¬
tion of Brachycistellus Baker and a key to North American genera
(Hymenoptera: Tiphiidae). Pan-Pac. Entomol., 44(3): 184-197.
SCIENTIFIC NOTE
New distribution and host record for the parasitoid Heteroschema
aeneiventris (Hymenoptera: Pteromalidae) —Intensive collecting during 1971
and 1972 at Pescadero State Beach, San Mateo County, California has resulted
in the discovery of a species of chalcid not previously known from the state. This
is the first report of Heteroschema aeneiventris (Ashmead) in California and its
parasitism of Euarestoides acutangulus (Thomson) (Diptera: Tephritidae), whose
larvae destroy the staminate florets of beach bur, Ambrosia chamissonis (Lessing)
Greene (Compositae), a prominent member of the coastal strand and dune com¬
munity. The discovery of the parasitoid in the state is of particular interest be¬
cause its previously known range in the United States was limited to Kansas (Peck,
1963, Can. Entomol. Suppl. 30: 629), Montana (Novak, 1967, Unpubl. M.S. Thesis,
Kent State Univ., 102 pp.), New Mexico, Texas and Utah (Burks, per. commun.).
The only other recorded host is the tephritid Neotephritis finalis (Loew) (Novak,
loc. cit.).
Laboratory investigation has shown H. aeneiventris to be a primary, internal,
solitary larval-pupal parasitoid. Of 460 field-collected E. acutangulus puparia,
40% were parasitized by this chalcid. The wasp is bivoltine in coastal California,
with peak emergence periods in mid-July and mid-September.
The writer is indebted to Dr. B. D. Burks, U. S. N. M., Washington, D. C., for
the identification of the parasitoid and to the California Department of Parks and
Recreation for permission to collect at the designated area.— Gary L. Piper, De¬
partment of Entomology and Parasitology, University of California, Berkeley, 94720.
333
The Pan-Pacific Entomologist
New Species of Eburia from Mexico
(Coleoptera: Cerambycidae)
John A. Chemsak and E. G. Linsley
Division of Entomology and Parasitology
University of California, Berkeley 94720
Thirty-two species of Eburia , exclusive of those described here, are
presently known from North America, Mexico and Central America.
Of these, eleven occur in the United States (Linsley, 1962) (only four
of these have not yet been found in Mexico) and twenty-eight in Mexico
and Central America (Bates, 1879-1885; Chemsak and Linsley, 1970a)
(including seven of those which extend their range into the United
States). The West Indies, including the Lesser Antilles, contains
twenty-eight known species, one of which, E. stigma (Olivier) is wide¬
spread in Mexico and southeastern United States. The remaining spe¬
cies occur in South America, extending into Argentina.
The object of the present paper is to make known some of the many
undescribed species in Mexico in order that the names may be available
for use in biogeographical and, hopefully, behavioral studies. One such
study (Chemsak and Linsley, 1970b) has revealed a remarkable death-
feigning behavior in two species presently assigned to the genus, E.
ulkei Horn and E. juanitae Chemsak & Linsley.
Appreciation is expressed to the National Science Foundation for
support through Grant GB-4944X and to Celeste Green for preparing
the illustrations.
Eburia affluens, new species
(Fig. 1)
Male: Form moderate sized, slightly tapering posteriorly; integument brownish,
antennal scape and legs orange-brown, mouthparts, pronotal calluses, a narrow
line around eburneous fasciae, apices of antennal segments, apices of femora, and
parts of coxae black or infuscated; pubescence of elytra sericeus, short, appressed,
short erect hairs numerously interspersed; eburneous fasciae cream-colored, con¬
tiguous, basal pairs equal in length, outer median pair longer than inner, inner
pair longer than basal pairs. Head small, median line deep, extending to neck;
palpi subequal, not dilated; genae short, obtuse at apices; antennal tubercles
depressed, not prominently produced, apices acute but blunt; pubescence dense,
appressed; antennae slender, extending about five segments beyond apices of
elytra, scape conical, not impressed, coarsely, not densely punctate, sparsely
pubescent, segments two to six moderately densely pubescent, remainder densely
clothed with short recumbent pubescence, segments to sixth fringed beneath
with long erect hairs, these decreasing in number from basal segments, segments
from sixth carinate along inside edges, third segment much longer than first,
fourth equal to third, fifth longer than third, eleventh longest, slender, third
The Pan-Pacific Entomologist 49: 338-353. October 1973
Vol. 49, No. 4, October 1973
339
segment finely rugosely punctate. Pronotum as long as broad, sides with a
small acute tubercle; lateral calluses behind apex large, dorsal tubercles prominent,
middle with a flat callus toward base; punctures coarse, dense, confluent;
pubescence dense, appressed, obscuring surface; prosternum deeply impressed,
coarsely, deeply punctate over posterior two-thirds, pubescence dense, intercoxal
process abruptly declivous, coxal cavities open behind; meso- and metasternum
finely, densely punctate, densely pubescent, scent glands distinct. Elytra about
two and one-half times as long as broad; eburneous fasciae sparsely pubescent;
each elytron vaguely bicostate between fasciae; punctures moderately coarse,
dense, becoming finer toward apex; pubescence obscuring surface, golden erect
hairs short; apices emarginate, outer angles with a curved spine, sutural angles
with a short straight spine. Legs robust; hind femora reaching almost to apices
of elytra; middle and hind femora with a moderate sized broad spine internally
at apices, outside with a short broad tooth. Abdomen finely punctate, densely
pubescent; last sternite shallowly notched at apex. Length, 14-26 mm.
Female: Form similar. Antennae a little longer than body. Elytra with erect
pubescence much longer. Abdomen with last sternite narrowly rounded at apex,
notched at middle. Length, 17-25 mm.
Holotype male, allotype (California Academy of Sciences) and 60 paratypes
(35 males, 25 females) from X-Can, Quintana Roo, Mexico, various dates from
9 June to 16 July 1967, 10 April to 29 May 1968, 10 March 1969 (E. C. Welling),
5-7 June 1959 (P. & C. Vaurie).
E. affluens is somewhat similar to E. brevispinis Bates but may be
separated by the equal, contiguous basal elytral fasciae, the shorter
contiguous median fasciae, the coarsely punctate and more densely
pubescent pronotum, and the more densely punctate elytra. Variation
in this species, in addition to the sexual differences, primarily involves
size and coloration. A number of specimens have a testaceous rather
than darker brown integument. The eburneous fasciae of the elytra
are well developed in all individuals and are similar in size and shape
throughout.
Eburia brevicornis, new species
(Fig. 2)
Male: Form moderate sized, parallel; integument reddish-brown, appendages
rufo-testaceous; elytral eburneous fasciae yellowish, distinct, each pair contiguous,
each fascia about as broad as apex of third antennal segment, basal pair subequal
in length, shorter than scape, median pair with outer longer than inner, outer
longer than scape; pubescence fine, grayish, appressed, partially obscuring sur¬
face. Head small, front very deeply impressed; median line deep, extending onto
neck; palpi subequal, apical segments dilated; vertex small, median line becoming
a carina; antennal tubercles depressed, obtuse above; pubescence dense ap¬
pressed; antennae extending about two segments beyond elytra, segments five
to ten slightly produced at apices, scape sparsely pubescent, not impressed, coarsely
punctate, segments three and four moderately densely clothed with short ap¬
pressed pubescence, remaining segments subopaque, densely clothed with very
short depressed pubescence, basal segments with numerous long erect hairs be-
340
The Pan-Pacific Entomologist
neath, becoming sparser to sixth segment, absent on apical segments, third segment
longer than first, fourth shorter than third but longer than first, fifth longer than
fourth, eleventh longer than tenth, appendiculate. Pronotum about as long as broad,
sides vaguely tuberculate; disk with two prominent, glabrous, black calluses before
middle, two flat glabrous calluses present at sides behind apical margin; surface
irregular, rugose, punctures transverse; pubescence dense, appressed, with long
erect hairs interspersed; prosternum deeply impressed, coarsely, rugosely punctate
over posterior two-thirds; pubescence dense, prosternal process abruptly declivous,
apex not expanded, coxal cavities open behind, rounded externally; meso- and
metasternum finely densely punctate, densely pubescent, scent glands obvious.
Elytra about two and one-half times as long as broad; eburneous fasciae with a
few erect setae; each elytron usually bicostate between fasciae; punctures around
basal fasciae coarse, contiguous, coarse punctures scattered over surface, especially
near median fasciae; pubescence appressed, dense, except around fasciae, long
erect setae rising out of coarse punctures; apices emarginate truncate, outer angles
with a short tooth, sutural angles with a very acute tooth. Legs slender; posterior
femora not attaining elytral apices; middle and hind femora spinose internally at
apex, spines much longer than those of elytra. Abdomen very finely, densely
punctate, moderately densely pubescent; last sternite broadly rounded at apex.
Length, 14-20 mm.
Female: Form similar. Antennae shorter than body, basal segments with long,
erect hairs beneath and at sides. Abdomen with last sternite rounded at apex.
Length, 20-22 mm.
Holotype male, allotype (California Academy of Sciences) and 25 paratypes
(23 males, 2 females) from X-Can, Quintana Roo, Mexico, 11-16 June 1967,
29-31 May 1967, 5-29 May 1968, 5-26 June 1968 (E. C. Welling) ; 3 male para¬
types from Piste, Yucatan, Mexico, 16 May to 21 June 1968 (E. C. Welling).
This species somewhat resembles E. aegrota Bates but may be readily
separated by the presence of the post-apical marginal calluses of the
pronotum and the externally produced antennal segments. Additionally,
the integument of aegrota is paler.
The only obvious variation within the type series is in size. The
eburneous fasciae are uniform throughout and characteristics of pu¬
bescence and coloration appear similar in all specimens.
Eburia brunneicomis, new species
(Fig. 3)
Female: Form rather large, robust, parallel; integument dark reddish brown,
appendages rufo-testaceous, mouthparts, pronotal calluses, antennal tubercles,
apices of basal antennal segments, apices of femora, and trochanters black;
pubescence very dense, appressed, brownish, obscuring surface, erect hairs nu¬
merous; eburneous fasciae yellowish, ringed with black, basal pair contiguous,
outer slightly longer, both much shorter than antennal scape, median pair con¬
tiguous, outer pair longer than scape, inner pair slightly longer than outer basal
pair. Head rather small, front deeply impressed; median line deep, becoming a
carina on vertex which extends to neck; palpi unequal, apical segments large
Vol. 49, No. 4, October 1973
341
Figs. 1-4. Diagrammatic illustrations (not scaled) of the pronotum, elytra and
middle and hind femora of: Fig. 1. Eburia affluens Chemsak & Linsley. Fig.
2. E. brevicornis Chemsak & Linsley. Fig. 3. E. brunneicomis Chemsak & Linsley.
Fig. 4. E. elegans Chemsak & Linsley.
342
The Pan-Pacific Entomologist
but not dilated; genae very acute at apices when viewed from above; antennal
tubercles depressed, acute above; pubescence very dense, appressed, long erect
hairs interspersed; antennae extending about two segments beyond apices of elytra,
segments from fifth slightly flattened, scape sparsely pubescent, not impressed,
moderately coarsely, densely punctate, second and third segments more densely
pubescent than scape, outer segments very densely clothed with short appressed
pubescence, scape with long erect hairs more numerous beneath, segments to
eighth with numerous long hairs beneath, these becoming less numerous from
basal segments, long, erect hairs also present on outside of basal segments, third
segment longer than scape, fourth shorter than third, longer than first, fifth
slightly shorter than third, eleventh slightly longer than tenth, vaguely appendicu-
late. Pronotum about as broad as long, sides with a large blunt spine at middle
and an obtuse tubercle behind apex; disk with two prominent blunt tubercles
before middle and a vague flattened callus at middle extending back toward base;
surface coarsely rugose; pubescence obscuring surface except on calluses; pro¬
sternum deeply impressed, coarsely, deeply punctate over basal two-thirds,
pubescence dense, intercoxal process abruptly declivous, apex not expanded,
coxal cavities open behind; meso- and metasternum minutely, densely punctate,
very densely clothed wtih short appressed pubescence, scent glands obvious.
Elytra about two and one-half times as long as broad; eburneous fasciae with a
few erect setae; basal punctures coarser especially around fasciae, punctures
becoming finer and sparser toward apex, well separated; short appressed pu¬
bescence obscuring surface, long erect hairs numerous, longer at base and apex;
apices truncate, each bispinose, outer spine longer than inner, shorter than femoral
spines. Legs slender; posterior femora not attaining elytral apices; middle and
hind femora with long spines internally at apices, outer spines very short, as
long as third tarsal segment. Abdomen very finely densely punctate, densely
pubescent; last sternite rounded at apex. Length, 21-30 mm.
Male: Form similar. Antennae extending about three segments beyond elytra.
Abdomen with last sternite broadly rounded at apex. Length, 23 mm.
Holotype female, allotype (California Academy of Sciences) and 6 female
paratypes from X-Can, Quintana Roo, Mexico, 15 June to 26 July 1967 (E. C.
Welling) ; 5 paratypes (1 male, 4 females) from Piste, Yucatan, Mexico, 16 June
to 20 July 1967 (E. C. Welling).
This species differs from E. pedestris White by the thicker brownish
pubescence, the black-ringed elytral fasciae, non-impressed antennal
scape, and by the short outer spines of the femora. From E. championi
it is distinguished by the non-impressed antennal scape, the denser
pubescence, the more numerous erect hairs of the elytra, and the shorter
spines of the elytra. The combination of dense brownish pubescence
and black-ringed eburneous fasciae makes E. brunneicomis easily
recognizable.
Eburia elegans, new species
(Fig. 4)
Male: Form large, robust; integument reddish, appendages orange; eburneous
fasciae yellowish, broad, contiguous, basal pairs subequal in length, about twice
Vol. 49, No. 4, October 1973
343
as long as broad, outer median pair longer than inner, inner longer than basal
pairs; pubescence white, very short, dense, appressed, long erect hairs golden,
not numerous. Head small, front deeply impressed; median line deep, extending
to neck; palpi subequal, apical segments slender; genae acute at apex; antennal
tubercles small, apices bluntly produced; pubescence dense, appressed, yellowish;
antennae slender, elongate, about twice as long as body, scape small, cylindrical,
narrowly impressed above, coarsely, densely punctate, sparsely pubescent, segments
from third densely clothed with short appressed pubescence, long erect hairs dense
beneath on basal segments, becoming sparser toward apex, third segment almost
twice as long as first, fourth equal to third, fifth longer than third, eleventh very
long, slender. Pronotum inflated, broader than long, sides rounded, unarmed;
disk with two small black, glabrous tubercles before middle; middle transversely,
rugosely punctate, sides very deeply, coarsely, subconfluently punctate; pubescence
dense, short, appressed, long erect hairs sparsely interspersed; prosternum not
impressed, coarsely, deeply punctate, densely pubescent, intercoxal process abruptly
declivous, tuberculate, coxal cavities open behind; meso- and metasternum mi¬
nutely, densely punctate, densely clothed, with appressed pubescence, scent glands
obvious. Elytra more than two and one-half times as long as broad; eburneous
fasciae very sparsely pubescent; punctures fine with large seta-bearing punctures
interspersed; pubescence dense, appressed, obscuring surface, long, golden hairs
sparsely interspersed; apices truncate, bispinose, outer spine longer than inner.
Legs very slender; posterior femora reaching to about elytra! apices; middle and
hind femora bispinose at apices, inner spines much longer. Abdomen densely
pubescent; last sternite truncate at apex. Length, 35 mm.
Holotype male (California Academy of Sciences) from X-Can, Quintana
Roo, Mexico, 27 May 1968 (E. C. Welling).
This species is distinctive among the known Mexican and Central
American Eburia by the inflated, non-tuberculate pronotum.
Eburia inarmata, new species
(Fig. 5)
Male: Form rather small, parallel; integument shining, pale reddish brown,
legs reddish, antennae almost black from third segment, area around median
fasciae infuscated; eburneous fasciae creamy, basal pairs narrowly separated,
outer longer than inner, median pairs contiguous, outer longer, both longer than
basal pairs. Head small, front deeply impressed; median line deep, extending
onto vertex; palpi subequal, apical segments slightly dilated; genae acute at
apices; antennal tubercles depressed, not produced apically; pubescence dense,
recumbent; antennae extending about two segments beyond elytra, scape slender,
conical, rather finely, densely punctate, sparsely pubescent, segments from third
flattened, produced at apices, second and third segments sparsely pubescent, re¬
mainder densely clothed with short appressed pubescence, basal segments with
very few erect hairs beneath, third segment a little longer than first, fourth equal
to first, fifth equal to third, eleventh longest, appendiculate. Pronotum as long
as broad, sides broadly angulate, unarmed; post apical lateral calluses absent;
disk with two prominent glabrous, very obtuse tubercles, areas behind tubercles
and toward sides impressed; punctures moderately coarse, dense, except for
344
The Pan-Pacific Entomologist
median glabrous area extending toward base; pubescence dense, recumbent; pro¬
sternum narrowly impressed, deeply punctate, densely pubescent, intercoxal process
abruptly declivous, coxal cavities open behind; meso- and metasternum finely,
densely punctate, densely pubescent, scent pores distinct. Elytra about three
times as long as broad; eburneous fasciae sparsely pubescent; basal punctures
coarse, confluent, becoming very fine and irregular toward apex; pubescence
dense, subrecumbent, obscuring surface partially except at base and around median
fasciae, erect hairs sparse; apices obliquely truncate, inner angles with small
spine. Legs slender; hind femora not extending beyond apices of elytra; middle
and hind femora unarmed at apices. Abdomen finely densely punctate, densely
pubescent; last sternite subtruncate at apex. Length, 16 mm.
Holotype male (California Academy of Sciences) from 16 miles S. Guamuciiil,
Sinaloa, Mexico, 16 June 1961 (F. D. Parker).
This species may be recognized by the very distinctive, flattened
antennae, the impressions of the pronotal disk, and the absence of
pronotal and femoral spines. The grayish, recumbent pubescence is
similar to that of some Elaphidionini rather than the typical pubescence
of most Eburia.
Eburia latispina, new species
(Fig. 6)
Male: Form moderate sized, parallel; integument reddish brown, parts of
head, antennal scape, parts of prothorax, areas around fasciae and parts of femora
infuscated; eburneous fasciae yellowish, inner pair only present at base, median
pairs contiguous, outer slightly longer, both much longer than basal pair; pubes¬
cence fine, very dense, appressed. Head small, front deeply impressed; median
line deep, extending to neck; palpi subequal, apical segments slightly dilated;
genae acute at apices; antennal tubercles depressed, not produced apically;
pubescence dense, appressed; antennae slender, extending about three segments
beyond elytra, scape conical, not impressed, moderately coarsely, densely punctate,
sparsely pubescent, basal segments sparsely pubescent, segments from sixth more
densely clothed with very short appressed pubescence, long, erect hairs numerous
beneath basally, becoming sparse toward apex, third segment longer than scape,
fourth slightly shorter than third, fifth equal to third, eleventh as long as third.
Pronotum broader than long, sides with a large broad acute spine; post-apical
calluses at sides prominent; disk with two glabrous tubercles before middle and
two vague pubescent calluses at sides behind middle, transverse impression be¬
hind apex shallow, one before base narrow; middle somewhat flattened, shallowly
rugose, punctures irregular; pubescence appressed, obscuring surface; prosternum
deeply impressed, sparsely irregularly punctate, densely pubescent, intercoxal
process rounded, coxal cavities open behind; meso-and metasternum minutely,
densely punctate, densely pubescent, scent pores distinct. Elytra more than two
and one-half times as long as broad; eburneous fasciae sparsely pubescent, con¬
nected by vague costae; deep punctures abundant at base, irregularly scattered
over remainder and becoming very fine toward apex; pubescence short, appressed,
obscuring surface except around fasciae; apices obliquely truncate, inner angles
Vol. 49, No. 4, October 1973
345
Figs. 5-8. Fig. 5. Eburia inarmata Chemsak & Linsley. Fig. 6. E. latispina
Chemsak & Linsley. Fig. 7. E. megalops Chemsak & Linsley. Fig. 8. E.
minutivestis Chemsak & Linsley.
346
The Pan-Pacific Entomologist
minutely spined, outer angles vaguely dentate. Legs slender; hind femora not
attaining elytra! apices; middle and hind femora with a short, broad tooth in¬
ternally. Abdomen finely, densely punctate, densely pubescent; last sternite
truncate at apex. Length, 18 mm.
Holotype male (California Academy of Sciences) from Jesus Maria, Nayarit,
Mexico, 6 July 1955 (B. Malkin).
This species is distinctive by the large broad lateral spine of the
pronotum, the configuration of the pronotal disk, the absence of an
outer pair of basal eburneous fasciae, and by the scarcely spinose
elytral and femoral apices.
Eburia megalops, new species
(Fig. 7)
Male: Form moderate sized, rather slender, parallel; integument reddish
brown; eburneous fasciae yellowish, basal pairs separated, outer pair slightly
longer than inner, median pairs contiguous, outer slightly longer, slightly longer
than outer basal pair; pubescence grayish, dense, recumbent. Head rather small,
front not deeply impressed; median line deep, extending onto vertex; palpi un¬
equal, apical segments large but not strongly dilated; genae acute at apices;
eyes large, upper lobes broad, separated on vertex by less than their width; an¬
tennal tubercles depressed, not produced apically; pubescence dense, recumbent;
antennae extending about three segments beyond elytra, segments from third
flattened, apically produced, scape slender, not impressed, rather finely, densely
punctate, sparsely pubescent, remaining segments densely clothed with very short
appressed pubescence, segments three and four with very few long hairs beneath,
third segment longer than scape, fourth slightly longer than scape, fifth longer
than third, eleventh longest. Pronotum about as long as broad, sides irregular,
not tuberculate medially; lateral calluses behind apex small, glabrous; disk with
two small glabrous tubercles and a vague median one, sides vaguely callused
toward base; punctures indistinct, surface uneven; pubescence dense, recumbent,
obscuring surface; prosternum deeply impressed, vaguely punctate, densely pu¬
bescent, intercoxal process abruptly declivous, coxal cavities open behind; meso-
and metasternum minutely, densely punctate, densely pubescent, scent glands
distinct. Elytra almost three times as long as broad; fasciae not pubescent;
punctures irregular, surface finely rugose; pubescence dense, recumbent, erect
hairs lacking; apices obliquely truncate, outer angles with prominent spines,
inner with small spines. Legs slender; hind femora almost reaching apices of
elytra; middle and hind femora spined internally at apices, spines as long as
outer elytral ones, neither spined nor dentate externally. Abdomen finely, densely
punctate, densely pubescent; last sternite broadly rounded at apex. Length, 22 mm.
Holotype male (California Academy of Sciences )from Tequisistlan, Oaxaca,
Mexico, 5 April 1962 (F. D. Parker, L. A. Stange).
The absence of lateral pronotal tubercles, the large dorsal lobes of
the eyes, the flattened antennal segments, and the vague punctation
make this one of the most distinctive species of Eburia.
Vol. 49, No. 4, October 1973
347
Eburia minutivestis, new species
(Fig. 8)
Male: Form small, slender, parallel; integument reddish brown, legs testaceous,
parts of head, prothorax, areas around eburneous fasciae, and parts of sternum
infuscated; pubescence very short, appressed, long, golden, erect hairs numerous
on elytra; eburneous fasciae yellowish, basal pairs contiguous, equal in length,
median pairs contiguous, outer longer, inner longer than basal pairs. Head small,
front deeply impressed; median line deep, extending to neck; palpi unequal,
apical segments not dilated; genae acute at apices; antennal tubercles flat, not
produced apically; pubescence dense, appressed; antennae slender, extending
about two segments beyond elytra, scape slender, conical, not impressed, rather
finely, densely punctate, very sparsely pubescent, segments to fourth sparsely
pubescent, remainder densely clothed with very short, appressed pubescence,
third segment moderately densely fringed beneath, fringe decreasing to sixth
segment, third segment longer than scape, fourth longer than scape but shorter
than third, fifth equal to fourth, eleventh equal to fifth. Pronotum about as
long as broad, sides with small acute tubercles; disk with two prominent glabrous
tubercles; sides behind middle constricted; surface broadly rugose, sides at apex
inflated, very deeply, coarsely, confluently punctate, this area extending partially
onto disk near apex and onto prosternum; pubescence fine, not obscuring surface;
prosternum not impressed, deeply confluently punctate, sparsely pubescent, inter-
coxal process abruptly declivous, coxal cavities open behind; meso- and meta¬
sternum minutely punctate, densely pubescent, scent glands distinct. Elytra less
than three times as long as broad; eburneous fasciae with a few setae; basal
punctures coarse, separated, becoming finer toward apex; pubescence minute,
appressed, long erect hairs numerous; apices truncate, outer angles with a small
recurved spine, inner angles dentate. Legs slender; hind femoral surpassing
apices of elytra; middle and hind femora with a long internal spine at apices,
these much longer than those of elytra. Abdomen finely densely punctate, densely
pubescent; last sternite broadly rounded at apex, notched at middle. Length,
15 mm.
Holotype male (California Academy of Sciences) from Cotaxtla Experiment
Station, Cotaxtla, Veracruz, Mexico, 28 June 1962 (D. H. Janzen).
The basally impressed and very distinctive lateral punctures of the
pronotum and the extremely short pubescence make this species readily
recognizable. The punctation of the pronotum and prosternum is
reminiscent of species of Callidium.
Eburia opaca, new species
(Fig. 9)
Male: Form large, parallel; integument dull, dark reddish brown, front of head,
antennal scape, and legs darker; pubescence very fine, dense, appressed, yellowish;
eburneous fasciae white, small, basal pair separated, very small, median pair sub-
contiguous, outer longer than inner. Head small, median line deep, ending as a
broad carina on vertex; palpi subequal, slender, genae acute at apices from
above; antennal tubercles prominently produced, impressed up middle, blunt at
348
The Pan-Pacific Entomologist
apices; pubescence brownish, dense, appressed; antennae slender, extending about
five segments beyond elytra, scape cylindrical, short, impressed for about two-
thirds its length, rather finely, rugosely punctate, sparsely pubescent, pubescence
denser along impression, segments from third densely clothed with short ap¬
pressed pubescence, segments to sixth with a fringe of rather short hairs
beneath, third segment much longer than scape, fourth equal to third, fifth
equal to third, eleventh longest, slender. Pronotum broader than long, sides with
short, obtuse tubercles, lateral calluses behind apex small; disk with two small
blunt tubercles before middle, a vague flattened callus at middle extending toward
base and two vague calluses at sides near base; surface densely covered with
moderate sized, deep, pock-lilce punctures around calluses; pubescence around
punctures dense, short, recumbent; prosternum impressed, deeply, separately
punctate, intercoxal process abruptly declivous, coxal cavities open behind; meso-
and metasternum finely densely punctate, densely pubescent. Elytra about two
and one-half times as long as broad; eburneous fasciae sparsely pubescent; each
elytron bicostate between fasciae and behind median pair toward apex, inner
costa turning to suture before apex; punctures obsolete; pubescence short, dense
with longer suberect hairs sparsely interspersed; apices narrowly truncate, sutural
angles with very short spines, outer angles dentate. Legs slender; hind femora
not attaining elytral apices; middle and hind femora bidentate at apices. Abdomen
minutely, densely punctate, densely pubescent; last sternite emarginate at apex.
Length, 33 mm.
Female: Form similar. Antennae slightly longer than body. Abdomen with
last sternite rounded at apex. Length, 23-37 mm.
Holotype male, allotype (California Academy of Sciences) and 3 female para-
types from X-Can, Quintana Roo, Mexico, 2 and 4 May 1968 (E. C. Welling).
Two additional female paratypes from Piste, Yucatan, Mexico, 12 June 1967
(E. C. Welling).
This species belongs to the stigmatica group but may be separated
from the other known species by the more densely punctate pronotum,
the very blunt lateral tubercles of the pronotum, the white elytral fasciae
and the nearly contiguous median pair of elytral fasciae. The eburneous
fasciae tend to be reduced and at least the basal pair will probably
prove to be absent in some specimens. The females have minute spines
at the sutural angle of the elytra and no spines or dentation at the
outer angles.
Eburia paraegrota, new species
(Fig. 10)
Male: Form small, parallel; integument testaceous, tips of mandibles, eyes,
pronotal tubercles and small areas around eburneous fasciae black; pubescence
very short, appressed, longer erect hairs sparsely interspersed. Head small, front
not deeply impressed; median line deep, extending onto vertex; palpi subequal,
apical segments broadened; genae blunt at apices; antennal tubercles depressed,
not produced apically; pubescence fine, dense, appressed; antennae slender, ex¬
tending about four segments beyond elytra, scape slender, conical, not impressed,
Vol. 49, No. 4, October 1973
349
Figs. 9-12. Fig. 9. Eburia opaca Chemsak & Linsley. Fig. 10. E. paraegrota
Chemsak & Linsley. Fig. 11. E. poricollis Chemsak & Linsley. Fig. 12. E. submutata
Chemsak & Linsley.
350
The Pan-Pacific Entomologist
moderately coarsely, densely punctate, sparsely pubescent, remaining segments
moderately densely clothed with very short recumbent pubescence, basal segments
with a fringe of long hairs beneath, these becoming less numerous to sixth segment,
third segment longer than first, fourth subequal to third, fifth equal to fourth,
eleventh longest, slender. Pronotum longer than broad, sides with an acute
tubercle behind middle; lateral post-apical calluses absent; disk with two prom¬
inent, glabrous tubercles before middle, middle vaguely callused; punctures coarse,
confluent, rugose; pubescence fine, appressed, dense; prosternum deeply impressed,
coarsely, deeply punctate except for apical one-third, intercoxal process abruptly
declivous, coxal cavities open behind; meso- and metasternum very minutely,
densely punctate, very finely, densely pubescent; scent glands obvious. Elytra
about three times as long as broad; eburneous fasciae white, with a few hairs,
basal pair contiguous, outer slightly longer than inner, median pair contiguous,
elongate, more than twice as long as respective basal fasciae, integument black at
posterior end of basal fasciae and at apex and base of median pairs; basal punc¬
tures moderately coarse, shallow, dense, becoming obsolete toward apex; pubescence
very short, dense, longer, erect hairs sparsely interspersed, more numerous at apex;
apices emarginate-truncate, outer angles with a prominent curved spine, inner
angles dentate. Legs slender; hind femora surpassing elytral apices; middle and
hind femora with a long spine internally, unarmed externally. Abdomen very
finely, densely punctate, densely pubescent; last sternite rounded at apex, shallowly
emarginate at middle. Length, 12-15 mm .
Female: Form similar. Antennae a little longer than body. Abdomen with
last sternite subtruncate at apex. Length, 13-18 mm.
Holotype male, allotype (California Academy of Sciences) and 15 paratypes
(5 males, 10 females) from 5 miles N. Mazatlan, Sinaloa, Mexico, at black and
white lights, 27, 28 July 1964 (J. A. Chemsak, J. A. Powell), 5-7, 24^29 July 1964
(H. F. Howden), 21 July 1972 (J. A. and M. A. Chemsak, A. E. and M. Michel-
bacher), 25, 29 July 1973 (Chemsak, E. G. & J. M. Linsley, A. & M. Michelbacher),
26, 28 July, 1 August 1973 (E. Giesbert).
In his characterization of E. aegrota , Bates stated that his specimen
was a female from Plan del Rio. The only specimen from this unknown
locality in the collections of the British Museum (Natural History) is
a male which we determine as being the type. A female from Jalapa
may be conspecific but another male from Acapulco designated as
aegrota represents another species. Based on the male type of aegrota,
E. paraegrota differs by the long, curved outer spine of the elytra,
longer elytra, longer femoral spines, and by the presence of numerous
erect hairs on the elytra.
Although the type series is uniform in coloration, there is some
variation in the amount of black between the eburneous elytral fasciae.
The lateral pronotal tubercles vary in size but are always present. Also,
the inner apical angles of the elytra vary from a short tooth to a small
spine. One specimen has the inner pair of basal fasciae greatly reduced
and another appears to lack the long spine on the hind femur. How¬
ever, the spine has been broken off and the other leg is absent.
Vol. 49, No. 4, October 1973
351
Eburia poricollis, new species
(Fig. 11)
Male: Form large, robust, subparallel; integument dark reddish black, antennae
usually paler; eburneous fasciae yellowish, small, separated; pubescence beige to
golden, very short and fine on elytra. Head rather small, median line very deep,
bordered by raised carinae which terminate at a deep pit on vertex; palpi sub¬
equal, apical segments prominent but not dilated; genae acute at apices; antennal
tubercles prominently produced, apices blunt; pubescence dense, appressed; an¬
tennae slender, extending more than three segments beyond apex of elytra; scape
robust, broadly impressed for its length, coarsely, rugosely punctate, sparsely
pubescent, segments from third moderately densely clothed with short recumbent
pubescence, third segment densely fringed with long erect hairs beneath, fringe
becoming much sparser to sixth segment, third segemnt longer than first, fourth
equal to third, fifth slightly longer than third, eleventh longest, slender. Pronotum
broader than long, each side with a large, acute, upwardly recurved spine, lateral
callus behind apex prominent; disk with two blunt tubercles before middle and
a median flat glabrous callus toward base; sides near base vaguely callused;
surface with very large pock-like punctures around calluses, punctures absent at
base across disk; pubescence around calluses and punctures dense, appressed;
prosternum impressed, coarsely, irregularly punctate, intercoxal process abruptly
declivous, coxal cavities open behind; meso- and metasternum finely, irregularly
punctate, densely clothed with yellowish appressed pubescence, scent glands
obvious. Elytra about two and one-half times as long as broad; eburneous fasciae
indistinctly pubescent; punctures small, very sparse; pubescence very short,
golden, appressed, not obscuring surface; apices truncate, inner angles with short
spines, outer angles dentate at most. Legs slender; hind femora extending almost
to elytral apices; middle and hind femora broadly bidentate at apices. Abdomen
finely densely punctate, moderately densely pubescent; last sternite shallowly
notched at apex. Length, 23-31 mm.
Female: Form similar. Antennae extending about one segment beyond elytra.
Abdomen with last sternite rounded at apex. Length, 29-30 mm.
Holotype male, allotype (California Academy of Sciences) and 7 paratypes
(4 males, 3 females) from Valerio Trujano, Oaxaca, Mexico, 4500 ft., 28, 29
July 1937 (M. A. Embury). Additional paratypes as follows: 1 male, 11 miles
W. Hidalgo, Michoacan, Mexico, 12 July 1963 (F. D. Parker, L. A. Stange) ;
1 male, 3 miles N. Petalcingo, Puebla, Mexico, 21 August 1963 (Parker and
Stange); 1 male, Jalastoc, Morelos, Mexico, 23 June 1951 (F. Mendoza P.).
An additional male, not paratypical, has been seen from Acapulco, Guerrero,
Mexico, 21 June 1935 (M. A. Embury).
This species is apparently related to E. stigmatica Chevrolet but may
be separated by the dark integument, the very long, recurved lateral
spines of the pronotum, the much sparser pubescence of the elytra, and
the lack of apical spines at the outer elytral margin. The eburneous
fasciae are present in all specimens in the type series but there appears
to be a tendency toward reduction of the median pair.
352
The Pan-Pacific Entomologist
Eburia submutata, new species
(Fig. 12)
Male: Form rather large, slightly tapering; integument pale reddish brown,
legs and usually antennae orange, mouthparts, pronotal spines and calluses, apices
of antennal segments narrowly, fasciae of elytra very narrowly, apices of femora,
and coxae in part black or infuscated; eburneous fasciae yellowish, contiguous,
basal pair subequal in length, about twice as long as broad, outer median pair
longer than inner, inner pair slightly longer than basal pairs; pubescence very
fine, grayish, appressed, elytra lacking erect hairs. Head small, median line deep,
extending onto vertex; palpi subequal, apical segments not dilated; genae acute
at apices; antennal tubercles small, blunt at apices; pubescence yellowish, dense,
appressed; antennae slender, extending about four segments beyond apices of
elytra, scape cylindrical, not impressed, finely, densely punctate, rather sparsely
pubescent; segments moderately fringed beneath, hairs becoming sparser toward
apex, third segment longer than scape, fourth equal to third, fifth longer than
third, eleventh longest, slender. Pronotum about as long as broad, lateral spines
prominent, blunt, calluses at sides behind apex prominent; disk with two rather
small tubercles; middle transversely rugose; pubescence dense, appressed, ob¬
scuring surface except for spines and calluses; prosternum deeply impressed,
deeply, separately punctate over basal one-half, pubescence dense, intercoxal
process abruptly declivous, coxal cavities open behind; meso- and metasternum
minutely punctate, very densely pubescent, scent glands obvious. Elytra about
two and one-half times as long as broad; eburneous fasciae with a few incon¬
spicuous hairs; basal punctures coarse, dense, becoming much finer and shallow
toward apex; pubescence very fine, obscuring surface, a few longer hairs present
at apices; apices truncate, bispinose, outer spine as long as bind femoral spine,
inner spines short. Legs slender; hind femora reaching to elytral apices; middle
and hind femora with prominent spines internally at apices, middle pair dentate
externally, hind pair with external spine as long as inner elytral spines. Abdomen
very densely pubescent, first four sternites with a narrow glabrous band at pos¬
terior margins; last sternite truncate at apex. Length, 20-33 mm.
Female: Form similar. Antennae extending about two segments beyond apices
of elytra. Abdomen with last sternite broadly rounded at apex. Length, 24-33 mm.
Holotype male, allotype (California Academy of Sciences) and 7 paratypes
(4 males, 3 females) from San Jeronimo, Volcan Tacana, Chiapas, Mexico,
13 August 1970, 12 & 18 September 1970, 6-15 October 1970, 3-25 November 1970
(E. C. Welling) ; additional paratypes: 3 males, 4 females, Hogar Infantil,
Ocozocoautla, Chiapas, Mexico, 10 August, 14 September, 7 October 1972 (M.
P. Levin).
Variation in the type series is primarily reflected in size differences.
However, the outer spine of the middle femora is sometimes as long
as that of the hind femora and a few of the specimens have vague
costae connecting the two sets of eburneous fasciae.
This species resembles E. mutata Bates but may be separated by the
absence of a median dorsal callus on the pronotum, the more prominent
lateral spines of the pronotum, the smaller eburneous fasciae, and by
Vol. 49, No. 4, October 1973
oco
ooo
the dark annulations of the antennae and femoral apices. E. submutata
is distinguished from E. pedestris White by the paler integument, the
non-impressed antennal scape, and the shorter femoral spines.
Literature Cited
Bates, H. W. 1879-1885. Longicornia. In, Biologia Centrali-Americana, Insecta,
Coleoptera, 5: 1-436, pis. 1-24.
Chemsak, J. A. and E. G. Linsley. 1970a. New Mexican species of Eburia
Serville and Eburodacrys Thomson (Coleoptera: Cerambycidae). Pan-
Pacific Entomologist, 46: 299-305.
1970b. Death-feigning among North American Cerambycidae (Coleoptera).
Pan-Pacific Entomologist, 46: 305-307, 1 fig.
Linsley, E. Gorton. 1962. The Cerambycidae of North America. Part III.
Taxonomy and classification of the subfamily Cerambycinae, tribes
Opsimini through Megaderini. Univ. California Publ. Ent., 20: 1-188,
56 figs.
SCIENTIFIC NOTE
Collection records of the Black Witch Moth, Ascalapha odorata (Lin¬
naeus) in Nevada (Lepidoptera: Noctuidae). —The black witch moth,
Ascalapha odorata (Linnaeus) (Oiticica Filho, 1962. Arq. Mus. Nac. Rio de
Janeiro 52: 137-144), which in the past was most commonly known as Erebus
odora (L.), is predominantly a tropical species commonly found in the southern
United States and occasionally straying into the northern states. The moth has
been reported as far north as Canada, from Colorado (Smith, 1893. U. S. Nat.
Mus. Bull. 44: 366-367; Holland, 1903. The Moth Book, p. 279; Comstock and
Comstock, 1909. A Manual of the Study of Insects, p. 297), Kansas (Howe, 1963.
Our Butterflies and Moths, p. 94) and there are many records from California.
In Hawaii, A. odorata was first observed in 1928 (Swezey and Bryan, 1929.
Hawaiian Entomol. Soc., Proc. 7: 236-237) where it has since increased in
abundance and importance (Bryan, 1972, personal communication). The occur¬
rence of A. odorata in Nevada has not previously been reported.
The following are the known collection records for A. odorata in Nevada:
Washoe Co., Verdi, 1464 m elevation, 16 August 1959, at night light, wings in
ragged condition (Ira La Rivers) Biological Society of Nevada; Churchill Co.,
base of Stillwater Range, near Dixie Valley (town), 1067 m elevation, 17 July
1964 (Thomas S. Briggs) University of California, Berkeley; Esmeralda Co.,
Dyer, 1488 m elevation, 10 July 1968 (Maxine E. Sprouse) Nevada State Depart¬
ment of Agriculture, Reno. (The specimen was originally sent to Robert Lauder¬
dale, University of Nevada Extension Service, then forwarded to Robert Bechtel,
Nevada State Department of Agriculture. No collection data was included in the
letter from M. Sprouse other than the return address and date) ; Washoe Co.,
Reno, 1372 m elevation, 18 July 1972, flying, 1100 hours, daylight saving time,
32°C, wings undamaged.— Thomas Lugaski and William H. Clark, Department
of Biology, University of Nevada, Reno, Nevada 89507.
354
The Pan-Pacific Entomologist
New species of Eurytoma associated with Cynipidae 1
(Hymenoptera: Eurytomidae)
E. E. Grissell
Division of Plant Industry,
Florida Department of Agriculture and Consumer Services, Gainesville, Fla. 32601
The genus Eurytoma Illiger was last revised in 1967 by Dr. R. E.
Bugbee. Since that time a number of additional species have been
described prompting him to update his revision. I am publishing the
following names to make them available for inclusion in his forth¬
coming work as well as for use in my own study of Eurytoma associated
with Cynipidae. In addition, the types of E. californica Ashmead and
E. querci Fullaway have been compared and are considered conspecific.
The latter name has been placed in synonymy with E. californica.
Eurytoma adiacrita Grissell, new species
Holotype female. —Length 4.2 mm. Mostly black with yellow as follows: fore
and midfemora mostly, hindfemur apically, foretibia below entirely and along
margins above, mid and hindtibiae apically, hindtarsus distally. Mandibles and
wing veins reddish brown. Pubescence silvery white. Flagellum filiform with
Fl-4 longer than wide, F5 about as wide as long, pedicel 0.78 times length of FI.
Face not striate, malar space with setigerous pits, post genal lamella present.
Depression on anterior face of forecoxa distinct, moderately deep. Propodeum
with wide median furrow, narrowing ventrally, lateral carinae complete to base,
central carina distinct in upper % of furrow, lateral surfaces wide with irregular
ridges, spaces between ridges with small, uneven punctures. Postmarginal vein
straight, 1.0 times length of marginal vein (Fig. 7). Abdomen oval in lateral
view and somewhat compressed (Fig. 5), petiole directed downwards and with
a distinct projection dorsally, ovipositor tilted about 35° from horizontal axis of
abdomen, lateral surfaces of tergum VI with dense reticulation covering lower %
completely, thinning to anterior margin medially, very sparse reticulation ex¬
tending dorsally as a thin band along anterior margin, terga IV and V laterally
with heavy reticulation covering anterior half of surface, thinning dorsally as
on VI, dorsal surfaces of segments IV-VI appearing highly polished with reticula¬
tions showing only under certain reflections, tergum VI in dorsal view 1.8 times
as long as V and in lateral view % length of abdomen, tergum IX 0.28 mm.
Genitalia as in Figure 2.
Allotype male.— Length 2.3 mm. Color as in female except mid and hind-
femora and tibiae mostly black. Antenna with five pedicellate flagellomeres which
are longer than wide and a terminal unit of two closely fused articles, scape
swollen anteromedially (Fig. 13). Face and coxae as in female. Propodeum
about as in female except central carina distinct in upper half of furrow. Post-
1 Contribution No. 273, Bureau of Entomology, D.P.I., F.D.A.C.S.
The Pan-Pacific Entomologist 49: 354^362. October 1973
Vol. 49, No. 4, October 1973
355
marginal vein 1.0 times length of marginal vein. Abdominal petiole cylindrical,
3.1 times as long as widest point, 1.3 times length of hindcoxa.
Holotype female, Folsom Lake, Placer County, California, emerged May
1967 from Besbicus conspicuus (Kinsey) on Quercus lobata Nee, D. C. Dailey
collector; allotype male, 12 $, and 8 $ paratypes same data as holotype except
em. 2-11 May 1967. Paratypes, 14 2, 12 $ all from California: Contra Costa
Co.: 4 $, 5 $, Mt. Diablo State Park, 4 November 1958 (R. L. Doutt) ex
Antron echinus (Ashmead) on Q. clouglasii Hooker and Arnott. Placer Co.: 8 2,
3 $, Rocklin, em. 12 May to 14 June 1967 (D. C. Dailey) ex Antron echinus on
Q. douglasii. Yolo Co.: 2 2, 4 $, 3 miles sw. Madison, em. 29 October to 15
November 1968 (E. E. Grissell) from Andricus kingi Bassett on Q. lobata.
Holotype, allotype, and paratypes of E. adiacrita are deposited in
the Entomology Museum, University of California at Davis. Paratypes
have also been placed in the Division of Biological Control (University
of California, Albany), in the U.S. National Museum, and in the
Bugbee Collection (Meadville, Pa.).
Variation. —Females of E. adiacrita range in length from 2.7 to 5.3
mm (x = 3.8; N = 36) and males from 1.6 to 3.1 mm (x — 2.3;
N = 29). In females the pedicel averages 0.67 times FI (range 0.47-
0.90), the postmarginal vein averages 0.98 times the marginal vein
(0.86-1.2), tergum VI averages 1.9 times V (1.5-2.3), and tergum IX
averages 0.20 mm (0.11-0.30). In males the postmarginal vein averages
0.98 times the marginal (0.86-1.2) ; the abdominal petiole averages
3.0 times as long as wide (2.4^3.8) and 1.3 times the length of the
hindcoxa (1.1-1.4).
E. adiacrita is predominantly black. The legs, however, may show
some variation in the amount of yellow coloration. Some large speci¬
mens of both sexes have two shallow, submedian depressions just above
the clypeus. In smaller specimens, these depressions are not distinct.
Males, in many instances, have the scape distinctly enlarged antero-
medially (Fig. 13), but this is not always as pronounced as illustrated.
In both sexes the median furrow of the propodeum is complete, hut in
some specimens the lateral carinae become irregular ventrally. While
the central carina is so indistinct ventrally as to appear incomplete, it is
generally present at least in the upper half of the furrow of both sexes.
Hosts.—Eurytoma adiacrita has been reared from galls of the cynipids
Antron douglasii (Ashmead), Andricus kingi Bassett, and Besbicus con-
spicuus on Quercus lobata , and Antron echinus on Q. douglasii. Antron
douglasii and A. echinus also serve as hosts for Eurytoma foligalla
Bugbee, and both species of Eurytoma may occasionally be reared
from the same species of cynipid (though not necessarily the same gall)
occurring in the same locality.
Systematics.- — E. adiacrita may run to one or other of two species
356
The Pan-Pacific Entomologist
in Bugbee’s revision of the genus Eurytoma (1967). Couplet 30 of his
key (Complex IV. Tylodermatis) is based upon the average length
and range of the ninth abdominal tergum. E. adiacrita overlaps either
choice of the couplet depending on whether the average length or range
of the ninth tergum is used. If smaller specimens of E. adiacrita (i.e.
those with shorter ninth terga) are keyed, they run to E. spongiosa
Bugbee. E. adiacrita may be separated from E. spongiosa by its longer
sixth tergum and more dorsally produced stylet arch in the female and
its longer postmarginal vein in both sexes. Larger specimens of E.
adiacrita key to couplet 42. Here, E. adiacrita may be distinguished
from E. spina Bugbee by its dorsally produced stylet arch and its narrow
dorsal valves; it differs from the remaining species by its complete
median furrow.
Among Californian species of Eurytoma attacking cynipids on oak,
E. adiacrita is most easily confused with E. foligalla because they are
often reared from the same hosts. Females of E. adiacrita may be
distinguished by the compressed abdomen, tergum VI almost twice the
length of V, the lack of heavy reticulation dorsally on tergum VI, and
the darker coloration of the scape and legs.
Males of E. adiacrita are not morphologically separable from those
of E. foligalla. On the basis of color, E. adiacrita may generally be
separated by its black trochanters. E. foligalla usually has yellow tro¬
chanters. This character, while useful, is not consistent in all cases.
It is best, where possible, to make associative rearings of both sexes.
Eurytoma bugbeei Grissell, new species
Holotype female.— Length 2.4 mm. Mostly black and yellow, some reddish
brown. Yellow as follows: undersides of scape, pedicel and flagellum, mid and
hindtrochanters, distal third of fore and midfemora, distal tip of hindfemur, all
tibiae and tarsi. Reddish brown as follows: flagellum above, two small spots at
base of antennal sockets, mandibles, wing veins. Pubescence silvery white.
Flagellum filiform with Fl-3 longer than wide, F4-5 about as wide as long,
pedicel 0.80 times length of FI. Striae converging on clypeus from middle of face,
malar space bare, lacking setigerous pits, a thin deep groove present ventrad of
compound eye fading towards mandible (Fig. 10), post genal lamella absent.
Depression on anterior face of forecoxa indistinct, shallow. Propodeum shallowly
depressed, without a median furrow, median and lateral carinae absent, area
covered with evenly spaced, fine punctures. Postmarginal vein 0.50 times length
of marginal vein which is twice as wide as postmarginal (Fig. 9). Abdomen
oval in lateral view (Fig. 4) , ovipositor tilted at about a 35° angle in relation to
horizontal axis of abdomen, lateral surface of tergum VI with heavy reticulations
covering lower half almost completely, narrow band continuing to % tergum
height anteriorly, reticulation on terga IV and V similar to VI but correspond¬
ingly narrower, dorsal and dorsolateral areas of terga IV-VI polished, tergum VI
Vol. 49, No. 4, October 1973
357
Figs. 1-6. Female Eurytoma, lateral view. Figs. 1-3. Genitalia. Figs. 4-6.
Abdomens.
in dorsal view 3.1 times as long as V, and in lateral view about % length of
abdomen, tergum IX 0.15 mm. Genitalia as in Figure 1.
Allotype male.— Length 1.9 mm. Color as in female except as follows: scape,
pedicel, and flagellum entirely black, trochanters all black, fore and midfemora
yellow apically, hindfemur entirely black, hind tibia mostly black. Antenna with
four pedicellate flagellomeres which are longer than wide, F5 separated from the
two-articled terminal unit by a distinct annulation, scape not swollen antero-
medially (Fig. 11). Face, coxae, and propodeum as in female. Postmarginal vein
0.50 times length of marginal vein. Abdominal petiole cylindrical, narrowing
slightly distally, 2.4 times as long as broad, 1.1 times length of hindcoxa.
Holotype female, 1.5 miles northeast Thornton, San Joaquin County,
California, emerged 10 May 1968 from Disholcaspis eldoradensis (Beutenmueller)
on Quercus lobata, S. Rosenthal and E. E. Grissell collectors; allotype male, 2 $,
3 $, same data as holotype. Paratypes, 7 $, 5 $, all from California: Napa Co.:
12,7 miles w. Monticello Dam, em. 1 February 1968 (E. E. Grissell) from
Disholcaspis corallina (Bassett) on Quercus douglasii ; 2 2, Monticello, 11 Sep-
358
The Pan-Pacific Entomologist
tember 1958 (R. L. Doutt) from terminal bud gall on Quercus durata Jepson.
Marin Co.: 1 2, 1 $, Carson Ridge, em. 23 January 1968 (E. E. Grissell) from
Disholcapsis plumbella Kinsey on Q. durata. Sacramento Co.: 2 $, Hood, em.
26 December 1964 and 26 March 1965 (S. Rosenthal) ex Andricus kingi on
Q. lobata. Yolo Co.: 2 5,2 miles e. Monticello Dam, em. 29 January 1968 (E. E.
Grissell) ex Disholcaspis eldoradensis on Q. lobata ; 1 2,2 <5,3 miles sw. Madison,
em. 31 October 1968 (E. E. Grissell) ex Andricus kingi on Q. lobata.
Holotype, allotype, and paratypes are deposited in the Entomology
Museum, University of California at Davis. Paratypes have also been
placed in the Division of Biological Control (University of California,
Albany), the U. S. National Museum, and the Bugbee Collection
(Meadville, Pa.).
Variation. —Females of E. bugbeei range in length from 1.8 to 2.9
mm (x = 2.4; N = 9) and males from 1.3 to 2.2 mm (x = 1.9; N = 10).
In females the pedicel averages 0.83 times FI (range: 0.70-1.0), the
postmarginal vein averages 0.56 times the marginal (0.50-0.67), tergum
VI averages 3.5 times V (3.0^1.3), and tergum IX averages 0.13 mm
(0.09—0.16). In males the postmarginal vein averages 0.54 times the
marginal (0.46-0.67) ; the abdominal petiole averages 2.8 times as
long as wide (2.4-3.3) and 1.1 times the length of the hindcoxa
( 1 . 0 - 1 . 2 ).
E. bugbeei does not appear to vary significantly in structural char¬
acters (e.g. striae on face, smooth malar space, finely punctate pro-
podeum) but may vary slightly in color. In females the red subantennal
spots are present in most cases, but are absent in all but one male (the
allotype). About half the female paratypes show some brownish colora¬
tion of the tegulae correlated with a slight increase in yellowing on
the fore and midfemora. All female paratypes have the mid and hind
trochanters yellow. Male paratypes have both the tegulae and tro¬
chanters black.
Hosts. —All but three specimens of E. bugbeei were reared from galls
caused by members of the genus Disholcaspis. Several species of
Eurytoma from California ( californica , foligalla , furva Bugbee) have
been recorded from the same species of Disholcaspis galls, and it is
apparent that much biological work needs to be done in order to better
understand the exact relationships of these Eurytoma to their host(s).
The species of Disholcaspis galls from which E. bugbeei was reared are
thick-walled, monothalamous galls. It is within the single, central
chamber that the large Disholcaspis larva develops. However, the pithy
walls of the gall may also harbor several larvae of other, smaller species
of Cynipidae. These are inquilines in the genus Synergus. Based pri-
Vol. 49, No. 4, October 1973
359
marily on size, E. bugbeei is probably parasitic on one or more species
of Synergus rather than on the gall-former itself.
Three specimens of E. bugbeei were reared from the monothalamous,
thin-walled gall of Andricus kingi. This gall also harbors members of
the genus Synergus , but the relationships between gall-former, inquiline,
and parasite are not known.
Systematics. —Females of Eurytoma bugbeei key to E. lacunae Bugbee
in Bugbee’s 1967 revision of the genus Eurytoma. These two species
are close morphologically as evidenced by the striae on the face, the
shallow depression of the forecoxae, the distinctive propodeum, the
lengths of the marginal and postmarginal veins, and especially the male
antenna with 4 pedicellate flagellomeres. Females of E. bugbeei may
be separated by the absence of extensive sculpturing dorsally on tergum
VI and the shorter tergum IX even though this species ranges generally
slightly larger than E. lacunae. In addition, both sexes of E. bugbeei
may be separated from E. lacunae by a combination of the following
characters: black coxae, legs mostly black, tegulae dark brown to black,
and wing veins reddish brown with the marginal vein twice the width
of the postmarginal (Fig. 9).
Eurytoma salicigalla Bugbee, described since Bugbee’s 1967 revision
of the genus, also runs to E. lacunae but may be separated from E.
bugbeei as follows: E. bugbeei has the median area of the propodeum
with minute, evenly spaced punctures (E. salicigalla with irregular
carinae), tergum VI 3 to 4 times as long as tergum V (2 to 3 times in
E. salicigalla ), the lower half of tergum VI completely reticulate (lower
% or less in E. salicigalla, and then weakly so), and male with 4
pedicellate flagellomeres (5 in E. salicigalla).
This species is named in honor of Dr. Robert E. Bugbee whose many
years of work constitute our major knowledge of Nearctic Eurytoma.
Eurytoma daileyi Grissell, new species
Holotype female.-— Length 2.5 mm. Mostly black and yellow, some reddish
brown. Yellow as follows: scape, flagellum beneath, coxae and legs entirely,
distal tip of ventral valves. Reddish brown as follows: mandibles, wing veins,
lateral areas of abdominal terga III-VI partly. Pubescence silvery white. Flagellum
moniliform with Fl-3 longer than wide, F4-5 about as wide as long, pedicel
0.86 times length of FI. Face not striate, malar space with setigerous pits, post
genal lamella present. Depression on anterior face of forecoxa distinct, moderately
deep. Propodeum with wide but incomplete median furrow which narrows ven-
trally, lateral carinae distinct in upper %, median carina distinct in upper %,
lateral surfaces wide and unevenly punctate. Postmarginal vein slightly curved,
0.87 times length of marginal vein (Fig. 8). Abdomen oval in lateral view (Fig.
6), ovipositor tilted at about 45° from horizontal axis of abdomen, lateral surface
360
The Pan-Pacific Entomologist
Figs. 7-13. Eurytoma. Figs. 7-9. Wing veins, dorsal view. Fig. 10. Head,
lateral view. Fig. 11. Male antenna, lateral view. Fig. 12. Scape of dciileyi
male, lateral view. Fig. 13. Scape of adiacrita male, lateral view.
of tergum VI with faint reticulation covering lower VL, thinning to lower V 2
anteriorly, terga IV and V laterally with a small area of reticulation antero-
medially, dorsal surfaces of terga IV-VI highly polished, tergum VI in dorsal
view 1.7 times as long as V and in lateral view about % length of abdomen,
tergum IX 0.13 mm. Genitalia as in Figure 3.
Allotype male. —Length 1.9 mm. Color as in female except as follows: scape
yellow on basal %, flagellum entirely black, hindcoxa smoky, hindfemur and
tibia with a black smudge medially, abdomen entirely black. Antenna with five
pedicellate flagellomeres which are longer than wide and a terminal unit of
two closely fused articles, scape distinctly swollen anteromedially (Fig. 12).
Face and coxae as in female. Propodeum as in female except median carina
inconspicuous. Postmarginal vein slightly curved, 1.0 times length of marginal.
Abdominal petiole cylindrical, 2.3 times as long as broad, 0.97 times length
of hindcoxa.
Holotype female, Moronga Ridge, Contra Costa County,
California, emerged 10 March 1968 from galls of Diastrophus
kincaidii Gillette on Rubus parviflorus Nuttall, D. C. Dailey collector,
No. 1051; allotype male, 6 2, and 3 5 paratypes same data except
emergence from February to March. Additional paratypes, 12,3 5,
Vol. 49, No. 4, October 1973
361
Baltic Mine (near Washington), Nevada County, California, em. De¬
cember 1967, same host and collector as holotype.
Holotype, allotype, and paratypes of E. daileyi are deposited in the
Entomology Museum, University of California at Davis. Paratypes
have also been placed in the U. S. National Museum and the Bugbee
Collection (Meadville, Pa.).
Variation. —Females of daileyi range in length from 1.9 to 2.5 mm
(x = 2.3; N = 16) and males from 1.5 to 2.5 mm (x - 2.1; N — 17).
In females the pedicel averages 0.89 times FI (range: 0.73-0.90),
the postmarginal vein averages 0.90 times the marginal (0.76-1.0),
tergum VI averages 1.7 times V (1.6-1.9), and tergum IX averages
0.13 mm (0.09-0.15). In males the postmarginal vein averages 0.89
times the marginal (0.82-1.0) ; the abdominal petiole averages 2.3
times as long as wide (1.9-2.7) and 0.93 times the length of the hind-
coxa (0.82-1.1).
All specimens show at least some yellow on the coxae and legs.
Females vary only slightly in terms of color (i.e. hindcoxa occa¬
sionally with a slight black smudge). Males, however, vary noticeably
in the amount of yellow present on coxae and legs. In some cases all
coxae may have some black infuscation. In other specimens only the
forecoxae are solid yellow while the mid and hindcoxae are entirely
black. In males the legs vary from all yellow to yellow with black
infuscation. Both sexes appear to vary somewhat in the structure of
the propodeum. The lateral carinae are never complete and may be
almost absent, indicated only by short ridges at the extreme top of the
furrow. The central carina may be present but incomplete, or absent.
Punctation varies from fine and dense over the entire surface (when
the lateral carinae are nearly absent) to uneven and limited to lateral
areas and lower third of furrow.
Systematics.- —Females of Eurytoma daileyi key to E. celtigalla
Bugbee in Bugbee’s 1967 revision. These two species appear quite
similar in size, general coloration, relationship of wing veins, the
broad dorsal valves of the genitalia, and the short abdominal petiole
in the males. E. daileyi females, however, may be separated by the
following characters: flagellomeres 4 and 5 as wide as long, pedicel
three-fourths or more length of FI, tegulae black, median furrow in¬
complete, tergum VI in dorsal view usually distinctly less than twice
the length of tergum V, and tergum IX ranging slightly shorter in
length. Males of E. daileyi can best be separated from E. celtigalla
by the incomplete median furrow, the black tegulae, and the long,
slightly curved postmarginal vein.
362
The Pan-Pacific Entomologist
This species is named in honor of Charles Dailey who has con¬
tributed greatly to our knowledge of the parasitoids of gall-forming
Cynipidae.
Eurytoma californica Ashmead
Eurytomci californica Ashmead, 1887: 195.
Eurytoma querci Fullaway, 1912: 278. New Synonymy.
The lectotype female of E. californica is in the United States Na¬
tional Museum. The holotype female of E. querci is housed in the
Los Angeles County Museum, having been moved there from Stanford
University some years ago. I have seen both types as well as several
hundred specimens of E. californica ; the holotype of E. querci clearly
falls within the variation of E. californica. Fullaway associated a male
specimen from a different lot number and host with the female of
E. querci. This specimen is quite similar to males of E. californica
but does not have the flagellomeres pedicellate and lacks the whorls
of long setae on each segment, a condition not known in other North
American species of Eurytoma. It is possible that the single male
represents an aberrant specimen of E. californica or an undescribed
species which has been incorrectly associated with this species. The
possibility also arises that it is the male of an unrecognized genus.
Acknowledgments
I wish to thank R. E. Bugbee (Allegheny College, Meadville, Pa.)
who has given me much-needed advice as well as the gift and loan
of specimens from his collection. I am indebted also to Charles Dailey
(Sierra College, Rocklin, Calif.) for his help and for the loan of his
invaluable collection. Additional material has kindly been provided
by K. S. Hagen (Division of Biological Control, University of Cali¬
fornia, Albany) and C. L. Hogue (Los Angeles County Museum).
I also wish to thank R. M. Bohart and S. L. Clement (both University
of California, Davis) for their comments and suggestions concerning
this manuscript.
Literature Cited
Ashmead, W. H. 1887. Studies on the North American Chalcididae, with
descriptions of new species, chiefly from Florida. Trans. Amer. Ent.
Soc., 14: 183-202.
Bugbee, R. E. 1967. Revision of chalcid wasps of genus Eurytoma in America
north of Mexico. Proc. U. S. Nat. Mus., 118: 433-552.
Fullaway, D. T. 1912. Gall-fly parasites from California. Jour. N. Y. Ent.
Soc., 20: 274-281.
Vol. 49, No. 4, October 1973
363
New Species of Leptohyphes Eaton
(Ephemeroptera: Tricorythidae)
Richard K. Allen
California State University, Los Angeles
Collections of Leptohyphes nymphs from the University of Utah,
Salt Uake City, and Florida A. & M. University, Tallahassee, include
ten undescribed species. Grateful acknowledgment is made to G. F.
Edmunds, Jr. and W. L. Peters for loan of the material and for per¬
mission to publish names and descriptions of the species.
Leptohyphes edmundsi Allen, new species
Nymph. Length: body 4.0-5.0 mm; caudal filaments 3.0-4.0 mm. General
color light brown to brown with dark brown and black markings. Head light
brown with black markings; vertex light brown; head with narrow black stripe
from posterior margin of compound eyes to posterior margin of head then mesad
along posterior margin to near epicranial suture; transverse black band on frons
between lateral ocelli; lateral ocelli small; maxillary palpi 2-segmented. Thoracic
nota light brown with variable black maculae; legs light brown to pale with
black markings; femora with subapical black macula (fig. la-b) ; femora with
moderately long spines (fig. lc) ; .fore femoral band of spines as in fig. la; hind
femora 30 percent longer than fore femora; anterior surface of middle and hind
femora without spines; ventral (leading) margin of middle and hind femora
without spines; dorsal margin of femora with long spines (fig. lb) ; tibiae light
brown with brown subbasal transverse band; tarsi light brown with brown trans¬
verse band near base; tarsal claws with 6-7 marginal denticles, and palisade of
4-5 submarginal denticles near apex (fig. Id). Abdominal terga 1-9 brown with
dark brown markings which are separated by median brown longitudinal stripe;
terga 1-6 with sublateral black maculae mesad to gills (fig. 19) ; operculate gills
brown with dark brown mark at base and pale apically; gills without apical spine
(fig. 11) ; abdominal sterna brown to dark brown, often with dark brown lateral
margins. Caudal filaments light brown.
Holotype, mature nymph, Rio Catiras, Morretes, Parana, Brazil, IV-65,
F. Plaumann, in collection University of Utah, Salt Lake City. Paratopotypes:
20 nymphs, same data as holotype, 2 nymphs each in collections of California
State University, Los Angeles; California Academy of Sciences, San Francisco;
and Canadian National Collection, Ottawa; Smithsonian Institution, Washington
D. C.; and Florida A. & M. University, Tallahassee, others in collection of Uni¬
versity of Utah. Paratypes: 23 nymphs, Rio Warnow, Santa Catarina, Brazil,
XI-65, F. Plaumann; 8 nymphs, Arroio Irapua, Rio Grande do Sul, Brazil, XI-65,
F. Plaumann; 2 nymphs, Rio Cubatao, Santa Catarina, Brazil, XI-65, F. Plaumann;
2 nymphs, Rio das Cobras, Parana, Brazil, III-65, F. Plaumann; 8 nymphs, Rio
Cavallinha, Santa Catarina, Brazil, XI-65, F. Plaumann; 21 nymphs, Arroio do
Hilario, Rio Grande do Sul, Brazil, XI-65, F. Plaumann. All paratypes in collec¬
tion University of Utah.
The Pan-Pacific Entomologist 49: 363-372. October 1973
364
The Pan-Pacific Entomologist
Figs. 1-6. Leptohyphes nymphs, leg parts: (a) fore femur; (b) hind femur;
(c) femoral spine; (d) tarsal claw. fig. 1, L. edmundsi; fig. 2, L. carinus; fig. 3,
L. flinti ; fig. 4, L. pupulus; fig. 5, L. tinctus; fig. 6, L. invictus.
Vol. 49, No. 4, October 1973
365
This species is named in honor of George F. Edmunds, Jr., Univer¬
sity of Utah, in recognition of his many contributions to our knowledge
of mayfly taxonomy. Leptohyphes edmundsi is easily distinguished
from the other described nymphs of South American Leptohyphes by
the distinctive markings on the abdominal terga, the legs, and the
abdominal gills, and by the number and arrangement of the denticles
on the tarsal claws.
Leptoliyplies carinus Allen, new species
Nymph. Length: body 3.5-4.5 mm; caudal filaments broken. General color
brown with black markings. Head brown with narrow black stripe from posterior
margin of compound eyes to posterior margin of head; lateral ocelli small;
maxillary palpi 3-segmented. Thoracic nota brown with irregular black markings;
legs unicolorous brown; femora with short spines (fig. 2c.) ; fore femoral band
of spines as in fig. 2a; hind femora 50 percent longer than fore femora; anterior
surface of middle and bind femora with row of transverse spines at base; dorsal
and ventral margins of middle and hind femora with numerous moderately long
spines in elevated sockets (fig. 2b) ; tibiae and tarsi with long setae; tarsal
claws with 4 blunt marginal denticles near apex (fig. 2d). Abdominal terga 1-3
brown with black transverse bands; terga 4-10 unicolorous brown; terga 4-9
with a round, median tubercle; operculate gills pale, black at base (fig. 12) ; gills
with apical spine; abdominal sterna brown. Caudal filaments broken.
Holotype, mature nymph, Guyumba, 21 km. de Tingo Maria, Huallaga,
Peru, 254X-63, M. Pandura, in collection University of Utah, Salt Lake City.
Leptohyphes carinus is superficially close to L. tuherculatus Allen
as the nymphs of both bear a median row of dorsal abdominal tubercles.
They are distinguished as the former lacks a tubercle at the apex of
the hind femur. These species are further distinguished by the colora¬
tion of the operculate gills, the dentition on the tarsal claws, and the
armature on the legs and body.
Leptoliyplies flinti Allen, new species
Nymph. Length: body 3.0-4.0 mm; caudal filaments 2.5-3.5 mm. General color
pale to brown and suffused with black. Head brown with black stripe on frons
between lateral ocelli; lateral ocelli small; maxillary palpi 3-segmented. Thoracic
nota pale to brown, suffused with black, and with complex black pattern; legs
yellow; femora with moderately long spines (fig. 3c), fore femoral band of spines
as in fig. 3a; hind femora 30 percent longer than fore femora; anterior surface
of middle and hind femora without spines; ventral (leading) margin of middle
and hind femora without spines; dorsal margin of middle and hind femora with
long spines (fig. 3b) ; tarsal claws with 3-5 marginal denticles, and palisade of
4-5 submarginal denticles near apex (fig. 3d). Abdominal terga pale to brown
and suffused with black; operculate gills brown and suffused with black, margins
pale (fig. 13) ; gills without apical spine; abdominal sterna brown. Caudal fila¬
ments brown.
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The Pan-Pacific Entomologist
Holotype, mature nymph, Springfield Est., Dominica, 26-VII-63, 0. S. Flint,
in collection University of Utah, Salt Lake City. Paratopotypes: 6 nymphs, same
data as holotype, 2 in collection California State University, Los Angeles and 2
in collection University of Utah, and 2 in collection Smithsonian Institution.
This species is named in honor of Oliver S. Flint, Jr., Smithsonian
Institution, Washington, D. C., the collector of the type material.
There are two other species of Leptohyphes now known from the West
Indies, and L. flinti is distinguished from these, and all others in the
genus, by the following combination of characters: (1) by the colora¬
tion of the abdomen, legs, and gills; (2) by the length of the femoral
spines; and (3) by the number and arrangement of the denticles on
the tarsal claws.
Leptohyphes populus Allen, new species
Nymph. Length: body 4.0-5.0 mm; caudal filaments 4.0-5.0 mm. General
color brown with dark brown markings. Head brown with dark brown markings
on vertex and frons between lateral ocelli; compound eyes large, divided in male
(fig. 15) ; compound eyes two-colored, outer portion black, inner portion brown;
lateral ocelli large; maxillary palpi 3-segmented. Thoracic nota brown with dark
brown markings; legs brown; femora with short, heavy spines (fig. 4c) ; fore
femoral band of spines as in fig. 4a; marginal spines of hind femora in elevated
sockets (fig. 4b) ; hind femora 45 percent longer than fore femora; anterior
surface of middle and hind femora with spines; dorsal and ventral margins of
middle and hind femora with spines (fig. 4b) ; tibiae and tarsi brown; tarsal
claws with 3-4 marginal denticles (fig. 4d). Abdominal terga brown, suffused
with black and with dark brown spots; abdominal terga 6-8 with dark brown
transverse hand on posterior margin; operculate gills light brown, suffused with
black, gills with apical spine (fig. 14) ; abdominal sterna brown. Caudal fila¬
ments light brown.
Holotype, mature male nymph, Rio Marauia, 3 days above San Antonio
Mission, NW. Taparuquara, Amazonas, Brazil, 26/27-1-63, F. J. Fittkau, in
collection University of Utah, Salt Lake City.
Leptohyphes pupulus is the first species in which the male nymph
is known to possess divided compound eyes. It is distinguished from
all other described South American species by this character, and by
the combination of the dorsal abdominal color pattern, the number,
->
Figs. 7-10. Leptohyphes nymphs, leg parts: (a) fore femur; (b) hind femur;
(c) femoral spine; (d) tarsal claw. fig. 7, L. minimus', fig. 8, L. viriosus; fig. 9,
L. jcimaicanus; fig. 10, L. rolstoni. Figs. 11-18. body parts: fig. 11, gill, L.
edmundsi ; fig. 12, gill, L. carinus; fig. 13, gill, L. flinti', fig. 14, gill, L. pupulus-,
fig. 15, head, L. pupulus-, fig. 16, gill, L. viriosus; fig. 17, gill, L. jamaicanus; fig.
18, gill, L. rolstoni.
Vol. 49, No. 4, October 1973
367
368
The Pan-Pacific Entomologist
shape, and arrangement of the denticles on the tarsal claws, and by
having elevated spines on the hind femora.
Leptohyphes tinctus Allen, new species
Nymph. Length: body 3.0-4.Q mm; caudal filaments broken. General color
pale with brown markings. Head pale with brown markings; head with paired
brown maculae on occiput mesad of compound eyes; frons brown except pale
around median ocellus and bases of antennae; lateral ocelli small; maxillary
palpi 2-segmented, apical segment with terminal seta. Thoracic nota pale with
small black maculae; pronotum pale; mesonotum with pale median macula be¬
tween base of wing pads; legs pale with brown markings; femora pale at base
and with median brown macula (fig. 5a-b) ; femora with long spines (fig. 5c) ;
fore femoral band of spines as in fig. 5a; hind femora 10 per cent longer than
fore femora; anterior surface of middle and hind femora with 3-5 long spines
near base (fig. 5b) ; dorsal and ventral margins of middle and hind femora with
long spines (fig. 5b) ; tibiae pale with subbasal and subapical transverse bands;
tarsal claws with 6-8 marginal denticles and palisade of 5-7 submarginal denticles
near apex (fig. 5d). Abdominal terga pale with brown and black maculae as in
fig. 20; operculate gills pale with brown markings (fig. 20) ; gills without apical
spine; abdominal sterna pale with brown lateral margins and paired sublateral
black maculae. Caudal filaments pale with brown annulations.
Holotype, mature nymph, Rio Ivay, Rio Grande do Sul, Brazil, XI-65, F.
Plaumann, in collection University of Utah, Salt Lake City.
This species is most easily distinguished by the distinctive color
pattern on the abdominal terga and operculate gills.
Leptohyphes invictus Allen, new species
Nympi-i. Length: body 2.5-3.5 mm; caudal filaments 2.0-3.0 mm. General
color light brown with brown markings; head light brown with an irregular, large,
dark brown macula on frons; head with fine spicules; lateral ocelli small;
maxillary palpi 3-segmented. Thoracic nota light brown with irregular dark brown
maculae; nota with fine spicules; legs light brown with apical brown tibial and
tarsal bands; femora with moderately long spines (fig. 6c) ; fore femoral band
of spines as in fig. 6a; hind femora 20 percent longer than fore femora; anterior
surface of middle and hind femora with short spines (fig. 6b) ; ventral (leading)
margin of middle and hind femora with short spines; dorsal margin of middle
and hind femora with long spines (fig. 6b) ; tarsal claws with 2-3 marginal
denticles and 1 submarginal denticle near apex (fig. 6d). Abdominal terga light
brown; posterior margins of terga 2-9 with long spines; operculate gills light
brown, dark near apex as in fig. 12; gills without apical spines; abdominal sterna
pale. Caudal filaments pale.
Holotype, mature nymph, Tulumayo, 29 km. E. Tingo Maria, San Martin
Province, Peru, 23-VI-63, W. L. Peters, in collection University of Utah, Salt Lake
City. Paratopotypes: 2 nymphs, same data as holotype, 1 each in collections
California State University, Los Angeles, and University of Utah.
This species is distinguished from others described from South Amer-
Vol. 49, No. 4, October 1973
369
ica by the arrangement and degree of development of the spines on the
fore femora, and by the arrangement of the denticles on the tarsal claws.
Leptohyphes minimus Allen, new species
Nymph. Length: body 2.0-3.0 mm; caudal filaments broken. General color
brown with dark brown and black markings. Head brown with a pale transverse
band on vertex between compound eyes; lateral ocelli small; maxillary palpi 2-
segmented, apical segment with terminal seta. Thoracic nota brown with pale
and black markings; pronotum brown with irregular black markings and black
posterior margin; mesonotum brown with two sets of paired median pale maculae,
and black lateral margins; legs brown with pale markings; femora light brown
with brown subapical mark and pale apically (fig. 7a-b) ; femora with moderately
long spines (fig. 7c) ; fore femoral band of spines as in fig. 7a; hind femora 10
percent longer than fore femora; anterior surface of middle and hind femora with
long spines (fig. 7b) ; ventral (leading) margin of middle and hind femora with
moderately long spines; dorsal margin of middle and hind femora with long
spines; tibiae brown with subbasal and subapical brown bands; tarsi with median
brown band; tarsal claws with 5-7 marginal denticles and palisade of 5-6 denticles
near apex (fig. 7df. Abdominal terga brown with pale and black maculae; terga
1- 8 with pale submedian and black sublateral maculae; terga 3-8 with black
median macula; tergum 10 pale with paired lateral brown maculae (fig. 21) ;
operculate gills brown with pale markings and pale transverse line near middle
of lamellae (fig. 21) ; gills without apical spine; abdominal sterna brown. Caudal
filaments brown.
Holotype, mature nymph, Arroio Lageado, Rio Grande do Sul, Brazil,
XI-64, F. Plaumann, in collection University of Utah, Salt Lake City. Paratypes:
7 nymphs, same data as holotype, 3 in collection California State University, Los
Angeles, and 4 in collection University of Utah.
Leptohyphes minimus is distinguished from all other nymphs of the
genus by its small size, and distinctive color patterns on the abdominal
terga and tracheal gills.
Leptohyphes viriosus Allen, new species
Nymph. Length: body 3.0-4.0 mm; caudal filaments broken. General color
pale with brown and black markings. Head pale with brown markings; occiput
with paired brown maculae mesad to compound eyes; frons brown except pale
around median ocellus and bases of antennae; lateral ocelli small; maxillary palpi
2- segmented, apical segment with terminal seta. Thoracic nota pale with small black
maculae; legs brown with darker brown markings; femora brown; femora with
long spines (fig. 8c) ; fore femoral band of spines as in fig. 8a; hind femora 10
percent longer than fore femora; anterior surface of middle and hind femora with
spines (fig. 8b) ; ventral (leading) margin of middle and hind femora with
moderately long spines; dorsal margin of middle and hind femora with long
spines (fig. 8b) ; tibiae pale with brown transverse bands; tarsal claws with
6-8 marginal denticles, and palisade of 5-6 submarginal denticles near apex (fig.
8d). Abdominal terga brown with pale maculae and black spots; terga 2-7 with
pale lateral margins; terga 6-9 with median black spot (fig. 22) ; operculate gills
370
The Pan-Pacific Entomologist
Vol. 49, No. 4, October 1973
371
brown, dark brown at apex, and with pale heart-shaped maculae near base; gills
without apical spine (fig. 16) ; abdominal sterna pale with brown lateral margins
and paired sublateral black maculae. Caudal filaments pale with brown an-
nulations.
Holotype, mature nymph, Rio Ivay, Rio Grande do Sul, Brazil, XI-64, F.
Plaumann, in collection University of Utah, Salt Lake City.
This species is most easily recognized by the distinctive heart-shaped
macula on the operculate gills, and it may be distinguished from all
other described species by the color pattern on the abdominal terga, the
length, shape and arrangement of the femoral spines, and the arrange¬
ment of the dentition on the tarsal claws.
Leptohyphes jamaicanus Allen, new species
Nymph. Length: body 4.0-5.0 mm; caudal filaments 3.0-4.0 mm. General
color pale with black markings. Head pale with black dots on vertex and black
marking on frons between compound eyes; lateral ocelli small; maxillary palpi
3-segmented, and with long apical spine. Thoracic nota pale with complex black
pattern; legs pale with black markings; fore femora pale with black subapical
band (fig. 9a) ; middle and hind femora pale, black at base and with black sub-
apical band (fig. 9b) ; femora with long sharp spines (fig. 9c) ; fore femoral
band of spines as in fig. 9a; hind femora 30 percent longer than fore femora;
anterior surface of middle and hind femora with row of spines near base (fig. 9b) ;
dorsal and ventral margins of middle and hind femora with spines; tibiae pale
with black transverse band near base; tarsi pale; tarsal claws with 7-9 marginal
denticles (fig. 9d). Abdominal terga 1-10 each with median black transverse
band and pale lateral margins; operculate gills nearly triangular in shape (fig.
17) ; gills pale with black markings; gills without apical spines; abdominal sterna
pale. Caudal filaments pale.
Holotype, mature nymph, Clarenden, Rio Minho, 1 mi. E. Frankfield,
Jamaica, 6-III-63, T. H. Farr & A. Barrett, in collection Florida A. & M. Uni¬
versity, Tallahassee.
This is only one of three species of Leptohyphes known from the
West Indies and it is distinguished from the others by the distinctive
black markings on the body and legs. It is further distinguished from
these, and all other species, by the shape of the operculate gills, the
shape and degree of development of the femoral spines, and by the
dentition on the tarsal claws.
Leptohyphes rolsloni Allen, new species
Nymph. Length: body 4.0-5.0 mm; caudal filaments 2.0-3.0 mm. General
color pale to light brown with black markings. Head light brown with black
<-
Figs. 19-22. Leptohyphes nymphs, abdomens: fig. 19, L. edmundsi; fig. 20,
L. tinctus ; fig. 21, L. minimus ; fig. 22, L. viriosus.
372
The Pan-Pacific Entomologist
markings; vertex and occiput with penciled black markings; frons with black
marking between compound eyes and lateral black markings from compound
eyes anterior to clypeus; lateral ocelli small; maxillary palpi 3-segmented. Thoracic
nota pale to light brown with variable black markings; legs pale with black
markings; fore femora pale with black apical macula (fig. 10a) ; middle and hind
femora pale with black markings as in fig. 10b; femora with moderately long
spines (fig. 10c) ; fore femoral band of spines as in fig. 10a; hind femora 25 per¬
cent longer than fore femora; anterior surface of middle and hind femora with
row of spines near base (fig. 10b) ; dorsal and ventral margins of middle and
hind femora with spines (fig. 10b) ; tibiae and tarsi pale; tarsal claws with 7-9
small marginal denticles, and palisade of 4^5 submarginal denticles near apex
(fig. lOd). Abdominal terga pale with black transverse band on each segment,
band interrupted medially on segments 9-10; operculate gills pale with large black
macula at base (fig. 18) ; gills without apical spine; abdominal sterna pale.
Caudal filaments pale.
Holotype, mature nymph, San Cristobal, Dominican Republic, 5-IX-66, L.
H. Rolston, in collection Florida A. & M. University, Tallahassee. Paratopotypes:
35 nymphs, same data as holotype, 5 nymphs each in collections California State
University, Los Angeles and University of Utah, Salt Lake City, remainder in
collection Florida A. & M.
This is the first Leptohyphes to be described from the Dominican
Republic, and is distinguished from the other West Indian species by
the color of the legs, the number of denticles and the shape of the
tarsal claw, and by the shape and the markings on the operculate gills.
This species is named in honor of L. H. Rolston, the collector of the
type material.
BOOK NOTICE
Flies and Disease, Volume II. Biology and Disease Transmission —by Bernard
Greenberg. Princeton University Press, Princeton, New Jersey, 1973. XI -f- 447
pp. $ 18 . 00 .
The first volume of this set consisted largely of cross-indexed lists of flies and
disease organisms they transmit, with selected references. In contrast, Volume II
contains extensive review and analysis of the role of flies as vectors of human
(Chapter 4) and animal (Chapter 5) diseases. Together, these chapters comprise
nearly 60% of the text. Chapter 1 provides a summary of prescientific attitudes
and beliefs concerning flies. Chapter 2 is a review of the biology of selected species
of medical importance which have not been treated in the recent literature. In¬
cluded is a brief discussion of methods for estimating dispersal rates and popula¬
tion sizes. Chapter 3 concerns the fly as a vector, including discussions of
morphological, physiological and ecological characteristics pertinent in disease
transmission, as well as short discussions of laboratory rearing methods and tissue
culture of flies.—Editor.
Vol. 49, No. 4, October 1973
OHO
0(0
A review of the genus Borboropsis
(Diptera: Heleomyzidae) 1
Wayne N. Mathis
Department of Entomology, Oregon State University, Corvallis, Oregon 97331
The genus Borboropsis Czerny was originally proposed in 1902 to
include the single Holarctic species B. fulviceps (Strobl). A second
species, B. steyskali n. sp., is described below from Oregon. Notes on
the generic limits, figures, and a key are also included.
Although the genus Borboropsis is readily placed in the Heleomyzinae,
the absence of well defined preapical tibial bristles and prominent costal
spines are anomalies, atypical of most Heleomyzidae. Moreover, the
absence of these structures, or their underdeveloped states, partially
characterize this unique genus and led Collin (1943: 234) to consider
Borboropsis as “the most aberrant Palaearctic genus of this family.”
With the addition of a second new species the generic concept must be
altered somewhat, especially with reference to the development of pre¬
apical bristles and costal spines.
Czerny (1924, 1927) mentioned that B. fulviceps may have indis¬
tinct preapical bristles on the hind tibiae, but that no costal spines are
present. In the most recent revision of the North American Heleomyzi¬
dae, Gill (1962) stated that dorsal preapical bristles are absent in
Borboropsis , and further mentioned that the costal spines are absent,
or that they are present but obscure.
Because of apparent ambiquities concerning the character states of
these features in B. fulviceps , it might well be that the above authors
were dealing with more than one species or that the species is polymor¬
phic or polytypic. To assess these possibilities, I examined specimens
determined as B. fulviceps from most of the major North American
museums. Based on the material before me, B. fulviceps is neither
polytypic nor polymorphic for these characters. The preapical dorsal
spine on the hind tibia, although small and hair-like, is still evident,
but I found no evidence of costal spines. B. steyskali , on the other hand,
has a definite dorsal bristle on the hind tibia and the costal spines are
distinct, although the development of both structures is somewhat less
than typical of heleomyzids in general.
The addition of B. steyskali further complicates other taxonomic
issues of importance. Subsequent to his description of Borboropsis ,
1 The work reported in this paper was supported by NSF grant GB-20963 to the Coniferous Forest
Biome, Ecosystem Analysis Studies, US/IBP; this is contribution no. 82.
The Pan-Pacific Entomologist 49: 373-377. October 1973
374
The Pan-Pacific Entomologist
Czerny (1924) described a second genus, Oldenbergiella , which is
closely related to Borboropsis. According to Czerny’s description,
Oldenbergiella has costal spines and one species, O. callosa Czerny,
has distinct preapical bristles on all tibiae. Furthermore, B. steyskali
runs to Oldenbergiella rather than Borboropsis in Czerny’s key (1927).
Gill (p. 512), gives one additional character for separating the
genera in his key and discussion of Oldenbergiella, “This genus is
likely to be confused only with Borboropsis Czerny, from which it can
be distinguished by the absence of a bristle in the posterior portion of
the mesopleuron.” The validity of Oldenbergiella, in view of apparent
overlaps with B. steyskali, is perhaps questionable. Further study, espe¬
cially with the male postabdomen, will be necessary to resolve this
question.
B. steyskali is placed in the genus Borboropsis because the meso¬
pleuron has a bristle in the posterior portion, because of its close simi¬
larity to B. fulviceps, and because Borboropsis has priority.
Because specimens are rare, a more complete systematic study of the
two genera is not yet possible. It would be desirable to compare Nearctic
specimens with Palaearctic material with special reference to genitalia.
Recognition of B. steyskali as a new species is justified in view of the
differences already noted and in the key. This is particularly true of
characters found on the male postabdomen.
Borboropsis Czerny
Borboropsis Czerny, 1902: 256; 1924: 67; 1927: 21—Collin, 1943: 2—Gill, 1962:
511.
Key to the Species of Borboropsis
Costa with distinct spines, length 3-4 mm., male genitalia as in figure a, dorsal
preapical bristle present on hind tibiae . steyskali n. sp.
Costal spines absent, length 2.4-3.1 mm., male genitalia as in figure b, preapical
dorsal bristle on hind tibiae absent or weakly developed . fulviceps (Strobl)
Borboropsis steyskali Mathis, new species
Diagnosis.— This species is closely related to B. fulviceps, but differs
from it with respect to the characters mentioned in the key.
Male.— Head from lateral view with large cheeks, cheek-eye ratio 1.7; anterior
half of front yellow and haired, becoming black and generally bare posteriorly;
fronto-orbital plates with 2 bristles, covered with grayish pollinosity; ocellar
triangle also covered with grayish pollinosity; eyes round; face and cheeks
whitish-yellow; oral vibrissae distinct, strongly developed; cheeks haired; first
and second antennal segments reddish-yellow; second segment with prominent
dorsomedial bristle; third segment black, subglobose, covered with whitish pu¬
bescence; aristae pubescent; palpi brownish-yellow.
Vol. 49, No. 4, October 1973
375
.5 mm
Fig. 1. Lateral view of epanclrium, cercus, and surstylus. a. Borboropsis
steyskali. b. B. fulviceps.
Thorax black; pollinosity of dorsum sparse, brownish-gray, becoming stronger
and grayer on humerus, noto- and mesopleurae; acrostical hairs in 4 rows an¬
teriorly, becoming stronger posteriorly and reduced to 2 rows; a pair of prescutellar
bristles present; dorsocentrals 2 + 3, but only the posterior 2 pairs distinctly
larger than dorsocentral hairs; 1 presutural bristle; 1 supra-alar bristle; 2 post-
alar bristles; scutellum bare except for 2 pairs of lateral bristles; humerus haired
with one bristle; notopleuron with 2 bristles on ventral margin; mesopleuron
haired on posterior margin, 1 distinct posterodorsal bristle, 1 propleural bristle;
sternopleuron with one posterodorsal bristle, a few hairs anterior to bristle.
Wings with costal spines distinct though small; hyaline.
Legs black to dirty yellow, becoming more yellow apically; fore coxae yellowed;
mesotibiae with several ventral apical bristles; preapical bristle on hind tibiae
present but small.
Abdomen generally black but yellowed marginally; male protandriuni asym¬
metrical, terga of segments 6, 7, and 8 strongly displaced; andrium more sym¬
metrical; epandrium from a lateral view subquadrate, posteroventral margin
slightly drawn out; cerci bacilliform and hairy; surstyli rounded and enlarged
ventrally, tapering and recurved toward apex, ventral and dorsomedial margin
bearing several long bristles.
Female. —Similar to male except for the female postabdomen.
Distribution. —Oregon, March-May.
Holotype male, Oregon, Lane County, H. J. Andrews Experimental
Forest near Blue River, Clearcut area number L107, April 23,
1973, Rotary net, M. Mispagel collector, sample # (43-17), IBP Biome
Survey-W. P. Nagel and G. Daterman. Allotype (female), Oregon,
Lane County, H. J. Andrews Experimental Forest near Blue River.
376
The Pan-Pacific Entomologist
Young age Doug Fir stand, number L105, March 27, 1973, Rotary net,
M. Mispagel collector, sample # (41-13), IBP Biome Survey-W. P.
Nagel and G. Daterman. Paratypes: 12 males, 6 females, same data
as holotype and allotype except for the dates which go from the 27th
of March through the 9th of May, 1973.
The holotype, allotype, and 2 paratypes will be deposited in the
USNMNH (no. 72839) ; a male and female paratype will be deposited
in the California Academy of Sciences, San Francisco; the Canadian
National Collection, Ottawa; Washington State University, Pullman;
and at Oregon State University, Corvallis. The remaining paratypes
will remain in the collection of the author.
Remarks. —The type specimens were all collected in a rotary trap
in an insect survey being conducted by the International Biological
Program, Coniferous Forest Biome.
I am pleased to dedicate this new species to my good friend and
dipterist, George C. Steyskal, whose assistance is gratefully ac¬
knowledged.
Borboropsis fulviceps (Strobl)
Anthomyza fulviceps Strobl, 1898: 269.
Borboropsis fulviceps (Strobl), Czerny, 1902: 256; 1924: 67; 1927: 22.
Diagnosis.— This species is very similar to B. steyskali except as indicated in
the discussion and key. The male andrium and cerci are of special importance
and are illustrated in figure b.
Distribution.— Holarctic.
Specimens examined.— CANADA: Yukon Territory, Herchel I., 22 July 1971,
W. R. M. Mason, 1 female; North West Territories, Muskox, 64°45' N., 108°10' W.,
4 August 1953, J. G. Chillcott, 1 female, 2 males; British Columbia, Robson, 30
September 1949, H. R. Foxlee, 1 female; Saskatchewan, Otosquen, 12 September
1959, J. R. Vockeroth, 1 female, 1 male; Manitoba, Hudson Bay Road, mile 500,
3 August 1952, J. G. Chillcott, 1 female, Churchill, 30 July 1952, J. G. Chillcott,
1 female, Churchill, 25 August 1950, J. G. Chillcott, 1 male, Fort Churchill, 11
August 1952, J. G. Chillcott, 1 male; Quebec, Harrington Lake, Gatineau Park,
26 October 1960, G. E. Shewell, 1 female. UNITED STATES: Alaska, Matanuska,
28 September 1945, Rotary trap collection, J. C. Chamberlin, 1 male, 1 female;
Alaska, Matanuska, 5 October 1945, J. C. Chamberlin, 1 female; Washington,
Longmire, 27 June 1935, A. L. Melander, 1 male.
Remarks. —This species is most readily distinguished from B.
steyskali by differences in the male genitalia, absence of costal spines,
weakly developed dorsal preapical bristles on the hind tibiae, and the
reduced size. The species has only been rarely collected and nothing
is known of its biology. It is interesting and perhaps significant that the
Alaskan specimens were also collected in a rotary trap.
Vol. 49, No. 4, October 1973
377
Acknowledgments
I wish to thank the following institutions and individuals for per¬
mission to examine specimens: Washington State University, Dr. Wil¬
liam J. Turner; The Canadian National Collection, Dr. B. V. Peterson;
U. S. National Museum of Natural History, Mr. George C. Steyskal.
I also wish to thank Drs. John D. Lattin and Paul W. Oman for
reviewing the manuscript and offering valuable comments and Dr. Bill
Nagel for facilitating financial arrangements.
Literature Cited
Collin, J. E. 1943. The British species of Helomyzidae (Diptera). Entomol.
Mon. Mag., 79: 23Y-240; 241-251.
Gill, G. D. 1962. The heleomyzid flies of America north of Mexico (Diptera:
Heleomyzidae). Proc. U. S. Natl. Mus., 113(3465): 495-603.
Czerny, L. 1902. Bemerkungen zu den Arten der Gattungen Anthomyza Fll.
und Ischnomyici Lw. Wien. Entomol. Ztg., 21: 249-256.
1924. Monographic der Helomyziden (Dipteren). Abh. Zool.-Bot. Ges. Wien.,
15(1): 1-166.
1927. Helomyzidae, Trichoscelidae, und Chiromyidae. [Fam.] 53a-53c, pp.
1-56 {— lfg. 22). In Lindner, E. (ed.), Die Fliegen der palaearktisclien
Region. Yol. 5. Stuttgart.
Strobl, G. 1898. Die Dipteren von Steiermark, IV. Theil. Mitt. Naturwiss.
Ver. Steiermark, 34: 192-298.
SCIENTIFIC NOTE
Fumigation of museum specimens in oven-cooking bags. —Insect boxes of
lightweight cardboard and designed for mailing provide no protection against mu¬
seum pests. Damage during storage can be reduced by placing the boxes in poly¬
ethylene hags with crystals of paradichlorobenzene. Unfortunately the fumigant
readily escapes through the bag wall and thereby creates a health hazard. Oven¬
cooking bags of Reynolon® film (heat stabilized polymer resin as used in Reynolds
Brown-in-Bag® oven bags) minimize loss of the fumigant while retaining the de¬
sirable features of transparency and adaptable shapes and sizes. As a demonstra¬
tion, 20 grams of paradichlorobenzene were placed in a 14 X 20 inch bag of each
type, inflated, and closed with rubber bands. After 49 days no crystals or odor
remained in the polyethylene bag, but only 1.8% of the weight of the crystals had
been lost in the Reynolon film bag. This method of protection is also suitable for
anthropological and art objects of wood, leather, fur, feathers, etc. Furthermore,
Reynolon film bags retain the vapors of ethyl acetate, chloroform, and carbon tetra¬
chloride, thus offering an improvement for the sweep-sampling technique described
by Chemsak (1957, J. Econ. Entomol., 50(4) : 523). Reynolon film bags are avail¬
able in several sizes from retail stores and in tube form from the manufacturer.—
Howell V. Daly, University of California, Berkeley.
378
The Pan-Pacific Entomologist
The Hamotus of Arizona
(Coleoptera: Pselaphidae)
Donald S. Chandler
Department of Entomology, University of Arizona, Tucson 85721 1
Hamotus Aube is a large, almost entirely neotropical genus, which
has been little known from the United States. It was first recognized
in Arizona with Raffray’s (1896) correct placement of Tyrus elongatus
Brendel in Hamotus. Fletcher (1932) added H. opimus in 1932. These
two are the currently known species in the United States. Two more
species from Arizona, H. nodicollis Raffray and H. populus , new spe¬
cies, can now be added to the list.
The Arizona species can be placed in Group XII, subgenus Hamo-
toides, by Park’s (1942) key to Raffray’s groups due to the presence
of the antebasal sulcus and the large median fovea of the pronotum.
H. elongatus keys out to nodicollis and is separated by the characters
given in my key to the Arizona species. H. populus cannot be placed
by Park’s key, being distinct by having antennal segments IX and X
similar in form and size, and more than three times as long as segment
VIII.
The species in Arizona inhabit different altitudinal areas and can
be identified by this difference. H. nodicollis is found in rotting cactus
and white-throated woodrat nests in Arizona and Sonora and can be
considered a desert associate. H. elongatus is found under bark of
dead trees (all records are pine) above 6,000 feet in mountainous
areas. H. populus was collected in the intermediate elevations under
bark of dead cottonwood.
An unplaced female specimen, characterized by thick, white setae on
the antennal club, represents either an extreme of nodicollis or a new
species. It is interesting as it was probably collected in association
with Zootermopsis laticeps (Banks) (Isoptera: Hodotermitidae) when
an ultraviolet light was run in a cage containing Zootermopsis- infested
wood from Patagonia, Arizona (W. L. Nutting).
I would like to thank D. C. Rentz of the Academy of Natural Sciences
of Philadelphia for the loan of the lectotype of H. elongatus ; K. W.
Brown of the Peabody Museum of Natural History, Yale University;
M. W. Sanderson of the Illinois Natural History Survey and K. Stephan
of Tucson for the loan of their collections. Dr. F. G. Werner checked
the manuscript, for which I am grateful.
1 Current address: Department of Entomology, The Ohio State University, Columbus 43210.
The Pan-Pacific Entomologist 49: 378-382. October 1973
Vol. 49, No. 4, October 1973
379
Key to Arizona Species of Hamotus
1. Antennal segments IX and X similar in form and size, each more
than three times as long as segment VIII (Fig. 1) _
_ populus Chandler, n. sp.
Antennal segment X larger than segment IX, IX at most barely
more than twice as long as VIII (Figs. 2-3) _ 2
2. Male profemur with tubercle near base (Fig. 4) ; elevations above
5,000 feet _ elongatus (Brendel)
Male profemur without tubercle; elevations below 3,000 feet
_ nodicollis Raffray
Hamotus (Hamotoides) populus Chandler, new species
(Figs. 1, 5)
Castaneous, flavous pubescence moderate in length; head subtruncate at base,
eyes prominent, with posterior emargination across one-third of eye, emargination
with dense, long pubescence; vertexal foveae small, median antennal fovea larger
and oval; third segment of maxillary palpus with moderate groove through entire
length; mandibles each with three teeth; antennae with distinct three-segmented
club. Pronotum with small, prominent pronotal angles anterior to lateral foveae,
anterior edge truncate; median and lateral foveae large, densely pubescent; dis¬
tinct transverse sulcus much narrower than median fovea. Elytron with two basal
foveae; tergites I-IV margined, tergite I with three lateral foveae to a side, II—III
with two lateral foveae, IV-V with one; sternites I-IV with single large, deep
lateral foveae to each side. Male with protrochanter spined, protibia with tubercle
near base. Female without these projections.
Holotype male.—Cochise Stronghold, Arizona. Head 0.53 mm. long, width
across eyes 0.51 mm., width across antennal insertions 0.28 mm.; 0.28 mm. from
head base to median antennal fovea, vertexal foveae 0.09 mm. from median antennal
fovea, 0.08 mm. from eyes; length and width of antennal segments: VIII
0.05 X 0.08 mm., IX 0.17 X 0.10, X 0.17 X 0.11, XI 0.23 X 0.14; third segment
of maxillary palpus 0.30 mm. long, 0.11 mm. wide. Pronotum 0.52 mm. long,
0.54 mm. wide at pronotal angles, angles 0.26 mm. from base, apex 0.28 mm.
wide; median fovea 0.09 mm. from base, 0.22 mm. from lateral foveae, lateral
foveae 0.10 mm. from base. Basal foveae of elytron 0.19 mm. apart, elytron 0.24
mm. wide at base, 0.48 mm. wide near apex, 0.82 mm. long; width between foveae
of abdominal tergites: I 0.41 mm., II 0.50 mm., Ill 0.45 mm., IV 0.40 mm., V
0.27 mm.; tergite V 0.16 mm. long, 0.34 mm. wide; width between lateral foveae
of sternites: I 0.52 mm., II 0.59 mm., Ill 0.54 mm., IV 0.42 mm.; sternite V 0.33
mm. wide. Male profemur 0.59 mm. long, tubercle 0.17 mm. from base.
Holotype male.—Cochise Stronghold, Dragoon Mountains, Arizona, 27/30 July
1970, UV Trap, R. J. Shaw. Paratypes.— Arizona. —Cochise County: Cochise
Stronghold, Dragoon Mountains, 1 female, 26 May/2 June 1970; 1 male, 2
females, 29 June/2 July 1970; 1 female, 2/5 July 1970; 1 female, 5/8 July 1970;
2 males, 18/21 July 1970; 2 females, 21/24 July 1970; 1 female, 24/27 July 1970;
1 male, eutopotypical; 2 males, 1 female, 31 July/2 August 1970; 1 male, 3/6
August 1970; 2 females, 6/9 August 1970; 1 male, 3 females, 9/12 August 1970;
380
The Pan-Pacific Entomologist
Figs. 1-7. Taxonomic characters of Hctmotus. Fig. 1-3. Antennal segments
VIII-XI. 1 , H. populus; 2, H. elongatus; 3, H. noclicollis. Fig. 4. Protrochanter
and profemur of male H. elongatus. Figs. 5-7. Left lateral view of male genitalia
and dorsal view of penis. 5. H. populus-, 6. H. elongatus-, 7. H. noclicollis.
1 female, 12/16 August 1970; 2 females, 16/22 August 1970, UV trap, R. J. Shaw;
1 male, 29 July 1957, light, G. Butler. Pima County: 2 males, 10 mi. S.E.
Mammoth, San Pedro River, 10 November 1971, under hark of dead cottonwood,
D. S. Chandler. 1 female, Tucson, 8 August 1968, K. Stephan. Pinal County:
1 male, Mammoth, 17 November 1969, K. Stephan. Mexico. Sonora .-—1 male,
10 mi. E. Navajoa, 13 August 1969, blk. It. trap, W. L. Nutting & F. G. Werner.
The holotype to be deposited in the California Academy of Sciences. Paratypes
in collections of K. Stephan, the University of Arizona and the author.
Hamotus (Hamotoides) elongatus (Brendel)
(Figs. 2, 4, 6)
Tyrus elongatus Brendel 1890: 239-240. (Type locality: Williams, Arizona).
Hamotus elongatus-. Raffray 1896: 134. Bowman 1934. Park 1942: 308-309.
In BrendeFs description of this species, he mentioned that the male
profemur had a “small, sharp thorn on the basal, and a minute sharp
Vol. 49, No. 4, October 1973
331
tubercle on the second third.” However, this latter tubercle could not
be found after examination of all available specimens and the lectotype.
The femur is ridged somewhat at that point and may possibly be pro¬
duced into a tubercle in some of the specimens Brendel saw. I have
added a few details and measurements to BrendePs description.
Third segment of maxillary palpus with moderate groove through entire length;
mandibles each with three teeth and a small tubercle; tei’gites I-IV margined,
tergites I—III with two lateral foveae to a side, IV with one lateral fovea, V
without foveae; sternites I-IV with single, large, deep lateral fovea to a side.
Redescribed male.—Santa Catalina Mountains, Arizona. Head 0.47 mm. long,
width across eyes 0.43 mm., width across antennal insertions 0.25 mm.; 0.21
mm. from head base to median antennal fovea, vertexal foveae 0.07 mm. from
median antennal fovea, 0.07 mm. from eyes; length and width antennal segments:
VIII 0.04 X 0.07 mm., IX 0.08 X 0.10, X 0.11 X 0.13, XI 0.22 X 0.17; third seg¬
ment of maxillary palpus 0.22 mm. long, 0.10 mm. wide. Pronotum 0.42 mm.
long, 0.46 mm. wide at pronotal angles, angles 0.27 mm. from base, 0.40 mm. wide
at base, apex 0.25 mm. wide; median fovea 0.08 mm. from base, 0.14 mm. from
lateral foveae, lateral foveae 0.07 mm. from base. Basal foveae of elytron 0.06
mm. apart, elytron 0.21 mm. wide at base, 0.49 mm. wide near apex, 0.65 mm.
long; width between lateral foveae of abdominal tergites: I 0.37 mm., II 0.38,
III 0.37, IV 0.35; tergite V 0.15 mm. long, 0.28 mm. wide; width between lateral
foveae of sternites: I 0.36 mm., II 0.47, III 0.46, IV 0.40; sternite V 0.34 mm.
wide. Male profemora 0.40 mm. long, tubercle 0.09 mm. from base.
Distribution.—Through the mountains of southern Arizona and New Mexico.
Arizona.— Cochise County: Rustlers Parle, Chiricahua Mountains, 17 June 1973,
under pine bark, D. S. Chandler. Graham County: Swift Trail, 7,400 ft., Pinaleno
Mountains, 6 July 1973, under pine bark, T. P. Sluss & D. S. Chandler. Pima
County: Santa Catalina Mountains, 7,000 ft., 5 April/23 May 1968, K. Stephan;
8,500 ft., 12 May 1948, A. A. Nichol; Bear Canyon, 25 January/29 March 1970,
under pine bark, K. Stephan; Bear Canyon, 6,000 ft., 16 June 1970, 26 April 1973,
under pine bark, D. S. Chandler. New Mexico. —Lincoln County: Gallinas Moun¬
tain, T1S R11E, 7,000 ft., 1 July 1951, under bark of yellow pine, C. C. Hoff.
Nogal Canyon, nr. Nogal, 7,200 ft., 5 June 1952, under bark, yellow pine log,
Hoff & Parrack.
Hamotus (Hamotoides) nodicollis Raff ray
(Figs. 3, 7)
Hamotus nodicollis Raffray 1883: 240. (Type locality: Mexico).
Hamotus ( Hamotoides) nodicollis : Park 1942: 317; 1945: 413.
Castaneous, moderately dense, long pubescence; head subtruncate at base, eyes
prominent, posterior emargination across one-third of eye, emargination with dense,
long pubescence; vertexal foveae small, median antennal fovea larger and oval;
third segment of maxillary palpus with moderate groove through entire length;
mandibles each with four teeth; antennae with distinct, three-segmented club.
Pronotum with small, prominent pronotal angles anterior to lateral foveae, apex
truncate; median and lateral foveae large, densely pubescent; distinct transverse
sulcus much narrower than median fovea. Elytron with two basal foveae; tergites
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The Pan-Pacific Entomologist
I-IV margined, I—III with two lateral foveae to a side, IV-V with one lateral
fovea; sternites I-IV with single, large, deep, lateral fovea to a side. Male with
protrochanter spined. Female without spine.
Redescribed male.—Tucson Mountains, Arizona. Head 0.45 mm. long, width
across eyes 0.46 mm., width across antennal insertions 0.26 mm.; 0.24 mm. from
head base to median antennal fovea, vertexal foveae 0.09 mm. from median an¬
tennal fovea, 0.07 mm. from eyes; length and width of antennal segments: VIII
0.04 X 0.07 mm., IX 0.10 X 0.10, X 0.12 X 0.13, XI 0.20 X 0.16; third segment
of maxillary palpus 0.21 mm. long, 0.10 mm. wide. Pronotum 0.41 mm. long, 0.49
mm. wide at pronotal angles, angles 0.19 mm. from base, 0.47 mm. wide at base,
apex 0.26 mm. wide; median fovea 0.10 mm. from base, 0.17 mm. from lateral
foveae, lateral foveae 0.10 mm. from base. Basal foveae of elytron 0.15 mm. apart,
elytron 0.22 mm. wide at base, 0.43 mm. wide near apex, 0.68 mm. long; width
between lateral foveae of abdominal tergites: I 0.37 mm., II 0.45, III 0.44, IV
0.41; tergite V 0.20 mm. long, 0.32 mm. wide; width between lateral foveae of
sternites: I 0.50 mm., II 0.56, III 0.51, IV 0.44; sternite V 0.39 mm. wide.
Distribution.—Through Arizonan and Sonoran desert areas into tropical Mexico.
Arizona. —Gila County: Globe, 28 August 1952, light F. H. Parker. Pima County:
Ariz.-Son. Desert Museum, 1/4 August 1962, It. trap, W. L. Nutting & S. Oman.
Colossal Cave Park, 18 April 1968, K. Stephan; 9 October 1971, 27 October 1971,
31 December 1971, in rotting saguaro, D. S. Chandler. 3 mi. S.W. Redington, 10
November 1971, in rotting saguaro, D. S. Chandler. Saguaro National Monument,
30 December 1972, in decayed saguaro, F. Werner. Silverbell, IBP Research Area,
10 mi. W. Marana, 13 October 1971, in rotting saguaro, D. S. Chandler. Tucson
(Catalina Foothills) Den #2, Neotoma sp., 2 August 1962, W. L. Nutting & P.
Mehringer. Tucson Mountains, 15 February 1970, 21 March 1970, saguaro, K.
Stephan. Pinal County: Boyce Thompson S. W. Arboretum, Superior, 14 Sep¬
tember 1949, It. trap, B. W. Benson. Mexico. Sonora.- —5 mi. W. Alamos, 14
August 1959, black light trap, W. L. Nutting & F. G. Werner. 11.5 mi. E. Alamos,
31 November 1972, in rotting Cardon, K. Stephan & D. S. Chandler. 13.5 mi. E.
Alamos, 29 November 1972, under bark, K. Stephan & D. S. Chandler. 10 mi. E.
Navajoa, 13 August 1959, black light trap, W. L. Nutting & F. G. Werner.
Literature Cited
Bowman, J. R. 1934. The Pselaphidae of North America. Privately published,
Pittsburgh, 149 pp. (not seen).
Brendel, E. & H. F. Wickham. 1890. The Pselaphidae of North America. B ull .
Lab. Nat. Hist., State Univ. Iowa, 1: 216-304.
Fletcher, F. C. 1932. Undescribed North American species of Pselaphidae
(Coleoptera), including a synopsis of the genus Rhexidius Casey.
Canad. Entomob, 64: 29-35.
Park, 0. 1942. A study in neotropical Pselaphidae. Northwestern Univ. Studies
Biol. Sci. Med., No. 1, 403 pp.
1945. Further studies in Pselaphidae (Coleoptera) of Mexico and Guatemala.
Bull. Chicago Acad. Sci., 7 : 331-443.
Raffray, A. 1883. Pselaphides nouveaux ou peu connus. Revue D’Entomol.,
2: 229-250.
1896. Notes synonymiques sur les Pselaphides. Ann. Soc. Entomol. France,
65: 131-137.
Vol. 49, No. 4, October 1973
383
Lectotype designations of African bees of the genus Ceratina
and allies
(Hymenoptera, Apoidea)
Howell V. Daly
University of California, Berkeley
Names proposed in the genus Ceratina for African species by C. T.
Bingham, P. Cameron, H. Friese, M. F. Meunier, E. Strand, and J.
Vachal lack designated types. The purpose of this paper is to validate
33 lectotype designations in anticipation of my monographic treat¬
ment of small carpenter bees of the genus Ceratina and its allies in
Africa south of the Sahara and Madagascar. In each case the original
author or subsequent reviser failed to designate a holotype, failed to
indicate whether more than one specimen was examined, or referred
ambiguously to a type among the syntypes. A list of 18 names based
on single specimens is also included to document the existence of
holotype specimens and their locations. The 51 designations below are
intended to establish unequivocal primary type specimens for the
names concerned. All species are discussed below as members of the
genus Ceratina in which they were originally assigned. Wing length
in millimeters was measured from the apex of the costal sclerite to
the maximum curvature of the wing.
In each case the specimen, collection data, and original determina¬
tion labels are consistent with the original description, type locality
or localities, dates of collection, general character of the author’s
labels, and known repository for the type material. The choice of
specimens is also consistent with the published indications of previous
revisers, and their comments are noted in each case. Where both
sexes were represented among the syntypes, males were favored as
lectotypes because their taxonomic features are critical in the separation
of species. The location of each lectotype is indicated by abbreviations
given in the acknowledgments. Each type also has a red label with the
indications: Ceratina , species, author, sex, lectotype or holotype, Howell
V. Daly, 1973.
Twelve names from specimens of African Ceratina in the Brauns col¬
lection of the Transvaal Museum are included in the list of types pub¬
lished by the South African Museum’s Association (Anonymous, 1958:
30-33). Of the twelve, seven are Brauns’ unpublished manuscript
names. Three of these were later described by Cockerell (1932: 272,
277, 278) from other specimens in the British Museum (C. liliputana,
The Pan-Pacific Entomologist 49: 383-388. October 1973
384
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male type No. 17B-255; C. subulata, male type No. 17B-314; C. aloes
female lectotype designated below). Specimens bearing the remaining
five names have red, handwritten labels stating that they are types of
Friese’s species. Dr. Brauns is known to have sent specimens to Friese
for identification and description. However, not one of the alleged
types of Ceratina in his collection has the characteristic determination
label of Friese and the data on specimens bearing three of the names
fail to match Friese’s descriptions. In my judgment no evidence exists
that the specimens in question were parts of the syntype series examined
by Friese and in view of Recommendation 74D, I have chosen all the
lectotypes for the species of Friese from the authenticated syntypes at
the Zoological Museum in Berlin. Each of the designated specimens
in Berlin has Friese’s handwritten determination label, each usually has
a red label of unknown origin with the imprint “Type” which often
identifies the favored specimen, and each agrees in every detail with
the original description.
I am greatly indebted to the following curators for their kind coop¬
eration and assistance in pursuing this task: C. Jacot-Guillarmod and
F. Gess, Albany Museum, Grahamstown (AM) ; R. W. Crosskey, I. H.
H. Yarrow and G. Else, British Museum (Natural History), London
(BMNH) ; J. Oehlke, Institut fur Pflanzenschutzforschung Kleinmach-
now (formerly Deutsches Entomologisches Institut), Eberswalde (DEI) ;
C. O’Toole, University Museum, Hope Department of Entomology,
Oxford (UM) ; P. Dessart, Institut Royal des Sciences Naturelles de
Belgique (IRSNB) ; S. Kelner-Pillault, Museum National d’Histoire
Naturelle, Paris (MNHN) ; L. Vari, Transvaal Museum, Pretoria (TM) ;
E. Konigsmann, Zoologisches Museum der Humboldt-Universitat, Berlin
(MNHU).
This research was supported in part by research grants from the
National Science Foundation (GB-7933, GB-34089).
Lectotypes are designated for the following:
1. aereola Vachal, 1903: 383. Lectotype male; wing length, 4.9 mm. Labels:
1) pale green, printed “Bouyssoui, N’Doro, X-XI-98”; 2) white, written
“aereola $ Vach.”; 3) pale green, printed “Museum Paris, Coll. J. Vachal
1911”; 4) salmon, written “Holotype, Ceratina aereola Vachal, det. Y.
Hirashima.” Hirashima (1969: 651) comments, “In addition to the type
series (2 $ $, 2 $ $ ) which is in the Paris Museum . . . The type of this
species is the male.” (MNHN)
2. allodapoides Strand, 1912: 282. Lectotype female; wing length, 5.3 mm.
Labels: 1) blue, printed “Span. Guinea, Uelleburg, G. Tessman S. G.”;
2) white, written “Ceratina allodapoides m $,” printed “Strand det.”
(MNHU)
Vol. 49, No. 4, October 1973
385
3. aloes Cockerell, 1932: 272. Lectotype female; wing length, 6.5 mm. Labels:
1) white, folded, written “Blaukrans, nr Calvinia, Nov. 17, L. 0.”; 2) white,
Cockerell handwriting “Ceratina aloes Br 3) white, printed “B.M.
1932-324, T. D. A. Cockerell.” (BMNH)
4. bouyssoui Vachal, 1903: 383. Lectotype female; wing length, 9.3 mm. Labels:
1) white, printed “Lambaren, X97”; 2) white, written “Ceratina bouyssoui
$ Vach.”; 3) green, printed “Museum Paris, Coll. J. Vachal 1911”; 4)
salmon, written “Type”; 5) white, written “Megaceratina,” printed “det. Y.
Hirashima.” This specimen commented on by Hirashima (1971: 254) :
“. . . I think, therefore, it should be regarded as the type.” Labels 4 and 5
were placed on the specimen by him. (MNHN)
5. braunsiana Friese, 1905: 12. Lectotype female; wing length, 4.9 mm. Labels:
1) white, printed “Algoa Bay., Capland., Dr. H. Brauns,” written “24 7 96”;
2) white, dashed border, written “Ceratina braunsiana $ Fr,” printed “1904
Friese det.” (MNHU)
6. congoensis Meunier, 1890: cci. Lectotype male; wing length, 4.0 mm.
Labels: 1) white, written “Matadi, 21 Mars 1889”; 2) white, written “Ceratina
congoensis Meunier 3) white, written “Ceratina congoensis Meun. $,
Type,” printed “det. Vachal 1911”; 4) white, black line border, printed in
red “Type.” (IRSNB)
7. daressalamica Strand, 1911b: 18. Lectotype male; wing length, 3.7 mm.
Labels: 1) blue, printed “D. O. Afrika, Daressalam, Pangani u. Hinterld,
Regner S. G.”; 2) white, written “Ceratina daressalamica m printed
“Strand det.” (MNHU)
8. defeminata Strand, 1911b: 20. Lectotype male; wing length, 4.9 mm. Labels:
1) blue, printed “Tanganyika-S. P. Reichard S.”; 2) white, written “Ceratina
defeminata m $ ,” printed “Strand det.” (MNHU)
9. dentipes Friese, 1914: 32. Lectotype male; wing length, 2.9 mm. Labels:
1) white, printed “Java, Buitzorg, Schmiedk.”; 2) white, dashed border,
written “Ceratina dentipes $ m Fr,” printed “1900 Friese det.” (MNHU)
10. elongata Friese, 1905: 14. Lectotype male; wing length, ca. 4.3 mm. Labels:
1) white, printed “D. 0. Africa, Kigonsera, 1903”; 2) white, dashed border,
written “Ceratina elongata $ Fr.,” printed “1904 Friese det.” (MNHU)
11. emarginata Friese, 1905: 12. Lectotype female; wing length, 5.1 mm. Labels:
1) white, “Capland, Willowmor, Dr. Brauns,” written “15-6-1904”; 2) white,
dashed border, written “Ceratina emarginata $ Fr.,” printed “1904 Friese
det.” (MNHU)
12. ericia Vachal, 1903: 384. Lectotype female; wing length, 4.2 mm. Labels:
1) white, printed “Bata, 5 IX 98”; 2) blue, printed “Bouyssou, Congo”;
3) white, written “ericia 9 Vach.”; 4) pale green, printed “Museum Paris,
Coll. J. Vachal 1911”; 5) red, printed “Type.” (MNHN)
13. harrarensis Vachal, 1910: 532. Lectotype male; wing length, 4.4 mm. Labels:
1) blue, printed “Museum Paris, Harrar, Maurice de Rothschild, 1905”; 2)
white, written, “Mars”; 3) white, written “Carat. Harrarandi Mars. No. 9”;
4) salmon, written, “Holotype, Ceratina harrarensis Vachal, 1909, det. Y.
Hirashima.” (MNHN)
14. inermis Friese, 1905: 8. Lectotype male; wing length, ca. 4.8 mm. Labels:
1) white, printed “Shilouvane, N. Transvaal, (Junod),” written “10.”;
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The Pan-Pacific Entomologist
2) white, dashed border, written “Ceratina inermis $ Fr.,” printed “1904
Friese det.” (MNHU)
15. labrosa Friese, 1905: 11. Lectotype female; wing length, 5.9 mm. Labels:
1) white, printed “Capland, Willowmor, Dr. Brauns,” written “9.6 1904”;
2) white, dashed border, written “Ceratina labrosa 2 Fr.,” printed “1904
Friese det.” (MNHU)
16. lativentris Friese, 1905: 10. Lectotype male; wing length, 5.9 mm. Labels:
1) white, written “Madagask., Tolia, 03”; 2) white, dashed border, written
“Ceratina lativentris $ Fr.,” printed “1904 Friese det.” (MNHU)
17. lineola Vachal, 1903: 385. Lectotype female; wing length, 4.6 mm. Labels:
1) gray, printed “Monteiro, Delagoa-Bay”; 2) quadrile, written “lineola
Vach. $.” (IRSNB)
18. ludwigsi Strand, 1914: 174. Lectotype female; wing length, ca. 3.1 mm.
Labels: 1) red, printed “Typus”; 2) white, written “Victoria, Kamerun,
Liidwigs”; 3) white, written “Ceratina ludwigsi m 2,” printed “Strand
det.” (DEI)
19. lunata Friese, 1905: 10. Lectotype male; wing length, 4.8 mm. Labels:
1) white, printed “Shilouvane, N. Transvaal, (Junod),” written “12”;
2) white, dashed border, written “Ceratina lunata Fr.,” printed “1904 Friese
det.” (MNHU)
20. madecassa Friese, 1900: 262. Lectotype male; wing length, 3.3 mm. Labels:
1) white, written “Madagask. Nossi-be, 17-XI-95, Voeltzkow”; 2) white,
dashed border, written “Ceratina madecassa, $ Fr.,” printed “1900 Friese
det.” (MNHU)
21. minuta Friese, 1905: 14. Lectotype male; wing length, 2.7 mm. Labels:
1) white, printed “D. O-Africa, Kigonsera, 1903”; 2) white, dashed border,
written “Ceratina minuta $ Fr.,” printed “1904 Friese det.” (MNHU)
22. moerenhouti Vachal, 1903: 385. Lectotype female; wing length, 4.5 mm.
Labels: 1) blue, printed “Vista (Congo), V. Moerenhout”; 2) white, black
line border, printed in red “Type”; 3) white, written “moerenhouti 2 Vach.”
(IRSNB)
23. mucronata Friese, 1905: 9. Lectotype male; wing length, 5.5 mm . Labels:
1) white, printed “Shilouvane, N. Transvaal (Junod),” written “12”; 2) white,
dashed border, written “Ceratina mucronata $ Fr.,” printed “1904 Friese
det.” (MNHU)
24. nasalis Friese, 1905: 8. Lectotype male; wing length, 5.9 mm. Labels: 1)
white, written “Algoa Bay, Capland, 5 7 96,” printed “Dr. Brauns”; 2) white,
dashed border, written “Ceratina nasalis $ Fr.,” printed “1904 Friese det.”
(MNHU)
25. nigriceps Friese, 1905: 11. Lectotype female, wing length, 4.3 mm. Labels:
1) white, written “Cradock, Capland”; 2) white, dashed border, written
“Ceratina nigriceps 2 Fr.,” printed “1904 Friese det.” (MNHU)
26. opaca Friese, 1905: 13. Lectotype male; wing length, 5.5 mm. Labels:
1) white, printed “Algoa Bay, Capland.”; 2) white, dashed border, written
“Ceratina opaca $ Fr.,” printed “1904 Friese det.” (MNHU)
27. penicillata Friese, 1905: 9. Lectotype male; wing length, 4.5 mm. Labels:
1) white, printed “Shilouvane, N. Transvaal, (Junod),” written “10”; 2)
white, dashed border, written “Ceratina penicillata $ Fr.,” printed “1904
Friese det.” (MNHU)
Vol. 49, No. 4, October 1973
387
28. personata Friese, 1905: 14. Lectotype female; wing length, 3.9 mm. Labels:
1) white, printed “W. Africa, Old Calabar, 190 ( date incomplete), v. Stefe-
nelli”; 2) white, dashed border, written “Ceratina personata $ Fr.,” printed
“1904 Friese det.” (MNHU)
29. subelongata Strand, 1912: 283. Lectotype male; wing length, 4.6 mm.
Labels: 1) blue, written “Sp. Guinea, Uelleburg, 6-8 1908, G. Tessman”;
2) white, written “Ceratina subelongata m $,” printed “Strand det.” (MNHU)
30. sulcata Friese, 1905: 13. Lectotype male; wing length, ca. 5.1 mm. Labels:
1) white, printed “Willowmore, Capland, Dr. Brauns,” written “10.1 1902”;
2) white, dashed border, written “Ceratina sulcata $ Fr.,” printed “1904
Friese det.” (MNHU)
31. tanganyicensis Strand, 1911a: 137. Lectotype male; wing length, 4.8 mm.
Labels: 1) blue, printed “Tanganyilca-S., P. Reichard S.”; 2) white, written
“Ceratina tanganyicensis m $" printed “Strand det.” (MNHU)
32. tibialis Friese, 1905: 9. Lectotype male; wing length, 4.1 mm. Labels: 1)
white, printed “Algoa-Bay, Capland”; 2) white, dashed border, written “Cera¬
tina tibialis $ Fr.,” printed “1904 Friese det.” (MNHU)
33. truncata Friese, 1905: 11. Lectotype male; wing length, 5.9 mm. Labels:
1) white, printed “Capland, Willowmor, Dr. Brauns,” written “9.6 1904”;
2) white, dashed border, written “Ceratina truncata $ Fr.,” printed “1904
Friese det.” (MNHU)
The following names were based on single specimens. The location
of the holotype is indicated in parenthesis:
1. acutipyga Strand, 1911b: 22, female; wing length, 5.0 mm (MNHU).
2. bicarinata Cameron, 1905: 198, female; wing length, 5.3 mm (AM).
3. caesia Vachal, 1903: 385, female; wing length, 3.5 mm (IRSNB).
4. foveifera Strand, 1911b: 27, female; wing length, 3.3 mm (MNHU).
5. furcilinea Strand, 1911b: 25, female; wing length, 4.4 mm (MNHU).
6. guineae Strand, 1912: 283, male; wing length, 6.0 mm (MNHU).
7. guineana Strand, 1911b: 19, male; wing length, 4.9 mm (MNHU).
8. langenburgiae Strand, 1911b: 15, male; wing length, 4.8 mm (MNHU).
9. loa Strand, 1912: 286, male; wing length, 4.6 mm (MNHU).
10. nassiinsignita Strand, 1912: 285, male; wing length, 4.1 mm (MNHU).
11. nyassensis Strand, 1911b: 26, female; wing length, 3.6 mm (MNHU).
12. penicilligera Strand, 1911b: 17, male; wing length, 5.3 mm (MNHU).
13. rothchildiana Vachal, 1910: 531, female; wing length, 5.3 mm (MNHN).
14. senegalensis Strand, 1911b: 24, female; wing length, 3.6 mm (MNHU).
15. tabofae Strand, 1911b: 21, female; wing length, 5.1 mm (MNHU).
16. togana Strand, 1911b: 16, female; wing length, 4.9 mm (MNHU).
17. viriditincta Strand, 1911b: 23, male; wing length, 4.4 mm (MNHU).
18. vittata Bingham, 1912: 382, male (sex misdetermined, description includes
male-specific characters) ; wing length, 4.6 mm (UM).
Literature Cited
Anonymous. 1958. A list of zoological and botanical types preserved in collec¬
tions in Southern and East Africa. Vol. 1. Zoology, Part 1: 1-147.
Published by the South African Museum’s Association, Pretoria.
388
The Pan-Pacific Entomologist
Bingham, C. T. 1912. South African and Australian Aculeate Hymenoptera in
the Oxford Museum. Trans. Entomol. Soc. Lond., 60(2): 375-383.
Cameron, P. 1905. On the Hymenoptera of the Albany Museum, Grahamstown,
South Africa. Rec. Albany Mus., 1(4): 185-244.
Cockerell, T. D. A. 1932. Descriptions and records of bees. CXXXVI. Ann.
Mag. Nat. Hist., (10)10: 267-278.
Friese, H. 1900. Hymenoptera von Madagaskar. Abh. Senckenb. naturforsch.
Ges., 26(2) : 259-268.
1905. Die Keulhornbienen Afrikas. Genus Ceratina (Hym.). Wien. Entomol.
Ztg., 24(1-2) : 1-18.
1914. Die Bienen fauna von Java. Tijdschr. Entomol., 57: 1-61.
Hirashima, Y. 1969. Synopsis of the genus Pithitis Klug of the world (Hy¬
menoptera: Anthophoridae). Pac. Insects, 11(3-4): 649-669.
1971. Megaceratina, a new genus of bees of Africa (Hymenoptera, Anthophori¬
dae). J. Nat. Hist., 5: 251-256.
Meunier, M. F. 1890. (communication in meeting of 26 November 1890). Bull.
Soc. Enotmol. Fr., pp. cci-ccii.
Strand, E. 1911a. Faunistische und systematische Notizen iiber afrikanische
Bienen. Wien. Entomol. Ztg., 30(6-7) : 135-159.
1911b. Neue afrikanische Bienen der Gattungen Sphecocles und Ceratina.
Arch. Naturgesch., 77(1, 4 suppl.) : 12-27.
1912. Zoologische Ergebnisse der Expedition des Herrn G. Tessmann nach
Siid-Kamerun und Spanisch-Guinea. Bienen. Mitt. Zool. Mus. Berlin,
6(2): 265-312.
1914. Beschreibung je einer neuen Allodape- und Ceratina- Art aus Kamerun,
nebst biologischen Bemerkungen (Hym.). Entomol. Mitt., 3(6): 173-
176.
Vachal, J. 1903. Hymenopteres du Congo Francais. Ann. Soc. Entomol. Fr.,
72: 358-400.
1910. Collections recueilles par M. le Baron Maurice de Rothchild dans
l’Afrique orientale, Insectes Hymenopteres: Melliferes. Bull. Mus.
Natl. Hist. Nat. (Paris), 15: 529-534.
RECENT LITERATURE
Scabies —by Kenneth Mellanby. E. W. Classey Ltd., reprint, VI + 81 pp. $3.50
(paperback). U. S. distributor, Entomological Reprint Specialists, P. O. Box
77971, Dockweiler Station, Los Angeles, California 90007.
Primarily of historical interest, this small volume which was first published
during World War II has been reissued virtually unchanged. The text includes
chapters on the anatomy and bionomics of the itch mite, as well as discussions of
the pathology, control and prevention of scabies.—Editor.
Vol. 49, No. 4, October 1973
389
Six New Nearctic Species of the Genus Friesea
(Collembola: Poduromorpha)
Kenneth Christiansen and Peter Bellinger
Grinnell College, Grinnell, Iowa 50112 and
California State University, Northridge 91324
In connection with a study of the collembolan fauna of North Amer¬
ica we have found a number of new species of Friesea , mostly from the
western third of the country, which has an unusually rich fauna of this
genus. Descriptions of these new species are needed for the publication
of an analysis of chaetotaxy in the North American members of the
genus.
Measurements of types and other representative specimens are given
in Table 1.
Friesea cera Christiansen and Bellinger, new species
(Fig. la-d)
Light blue, paler ventrally, eye patches black; fourth antennal segment with
partly withdrawn unlobed apical bulb and with four apical and two basal setae
dorsally; eyes 8 —{— 8; maxilla (fig. la) with larger lamella simple, smaller lamella
with three very small teeth; mandible (fig. lb) with seven or eight teeth;
tibiotarsus without clavate tenent hairs; unguis (fig. Id) untoothed; tenaculum
bidentate; mucrodens (fig. lc) one and one-half to two times length of longest
anal spine, with distinct mucro fused to dens; three curved anal spines, subequal
in length to inner edge of unguis; integumental granulation uniform, moderately
coarse; body setae smooth, acuminate; maximum length 0.7 mm.
Holotype (sex undetermined) from Conard Ecological Research Area,
Jasper County, Iowa, mixed deciduous forest litter (B. Dubach). Addi¬
tional material examined from Essex County, Massachusetts; Middle-
field, Hartford County, Connecticut; Shingletown, and Bear Meadows,
Pennsylvania; and Yolo, Illinois.
Among similar Nearctic species of Friesea (with 8 + 8 eyes and
three anal spines), F. sublimis Macnamara is distinguished by its clavate
abdominal setae, the mucro being separate from the dens, and striking
differences in chaetotaxy; F. mirabilis (Tullberg) has a relatively
shorter mucrodens with a hook-shaped mucro.
Friesea millsi Christiansen and Bellinger, new species
(Fig. le-j)
Pale blue with pigment in scattered granules; eye patches dark; fourth antennal
segment (fig. li) fused with third dorsally, separated ventrally, with deeply with-
The Pan-Pacific Entomologist 49: 389-395. October 1973
390
The Pan-Pacific Entomologist
Fig. 1. Friesea cera, F. millsi and F. quinta. All structures at 1000X magnifica¬
tion unless noted otherwise. Fig. la-d. F. cera, structures of holotype. (a) maxilla,
(b) mandible, (c) mucrones and dentes, dorsal aspect, (d) hind foot complex
(450x); Fig. le-j. F. millsi, structures of type specimens, (e) mandible, (f)
forefoot complex (450X), (g, h) mucrodens and tenaculum of two individuals,
(i) apex of fourth antennal segment, (j) eyepatch, variable eye shown dotted.
Fig. lk-m. F. quinta, structures of type specimens, (lc) mandible, (1) mucrodens
and tenaculum (500X) (m) eyepatch (500x)-
drawn unlobed apical bulb, and with one basal and five distal blunt setae dorsally;
eyes (fig. lj) 7 + 7 to 8-f-8; maxilla with large lamella unidentate, smaller
lamella tridentate; tibiotarsus (fig. If) with three weakly clavate tenent hairs;
unguis with minute subapical inner tooth, without lateral teeth; tenaculum (fig.
lg, h) unidentate or with rudimentary second tooth; mucrodens (figs, lg, h)
knohlike, fused to manubrium, with three minute setae (absent in some indi¬
viduals) ; furcula two-thirds to three-fourths length of longest anal spine; five
straight anal spines as long as inner edge of unguis set on short papillae;
integumental granulations ranging from about eight micra in diameter anteriorly
to seventeen micra posteriorly; body setae smooth, acuminate except longest setae
on fifth and sixth abdominal segments which are clavate; maximum length 0.9
mm.
Vol. 49, No. 4, October 1973
391
TABLE 1. Measurements of taxonomically important structures of Friesea
species. All measurements were made with an ocular micrometer, and are in
millimeters. Measurements are middorsal lengths; “total length” is measured from
the apex of the labrum to the end of the abdomen, and may not correspond to the
sum of individual segment lengths because of dorsal curvature of the body. Not
all specimens could be measured; blank spaces in the table correspond to features
which could not be measured accurately because of distortion. For each species,
the first set of measurements represents the holotype. Succeeding measurements
for F. cera are of specimens from Shingletown, Pennsylvania; Essex County,
Massachusetts; Bear Meadows, Pennsylvania (2 specimens) ; and Volo, Ill. (3
specimens). Succeeding measurements for other species are of paratypes.
Species
rH 5
4 g>
O 0
& ^
B
C
0
-P
0
Thoracic
Segments
II III
Abdominal Segments
II III IV V
2
^ 0 P
op O
'H crj *H
P -P P W
0 O 0
*P -H -P P
(0 rQ W hO
*t r t O »H O C
VI fp Eh Ph JO
F. cera
F. mills!
F. quinta
F. wilkeyi
F. fara
F.. polla
.574
.07
.147
.451
. 066
.089
.357
.047
.089
.335
.052
.099
.348
.056
.085
.422
.052
.108
.730
.038
.085
.296
.026
.079
.528
.067
.125
.470
.055
.092
.653
.080
.127
.595
.070
.122
.672
.075
.127
.758
.075
.141
.509
.089
-
.442
.113
.075
.432
.490
.080
.099
.394
.061
.085
.480
.075
.094
.672
.089
.141
.691
.061
.141
.672
.075
.136
.643
,080
.122
.480
.075
.127
• 797
.089
.174
.701
.075
.155
.480
.070
.108
.432
.061
.099
.490
.066
.108
.14 .18 -.033 .0044
.047 .052 .047 .042 .038 .038 .028 .024 .02 .0066
.038 .042 .028 -.028 .019 .015 .0066
.042 . 042 .038 .033 .056 -— .019 .0066
.037 .033 .024 .024 .024 .033 .022 .020 .013 .0044
.031 .044 .024 .026 .024 .037 .033 .026 .018 .0066
.029 .026 .022 .024 .024 .029 .022 .020 .018 .0066
.029 .031 .024 .022 .022 .026 .018 .015 .013 .0066
.055 .073 .051 .049 .042 .046 .033 .033 .024 .0088
.070 .061 .066 .061 .047 .038 .028 .0044
.066 —..061 .052 .047 .029 .0066
-- .026 -
--.048 .053-
- .029 .029 .035
- .075 .056 .070 .042 .047
.056 .066 .085 .047 .061 .070 .061 .047 .038
.033 .070-.070 .047 .047
.038 .085 .080 .080 .080 .075 .075 .052 .047
.038 .028 .042 .042 .042 .038 .042 .038 .033
.022 .048 .055 .031 .042 -.046 .051
.042 .080 .094 .080 .075 .056 .052 .052 .047
.040 .044 .044 .037 .033 .040 .042 .024 , 0 ?o
.033 .044 .048 .037 .022 .026 .031 .026 .026
.024 .048 .057 .031 .035 .033 .044 .031 .031
.026 .033 .024 .0088
- .024 .018 .0066
- ,020 .0088
- .022 .0088
.024 .024 .020 .0066
-.- .020 .0088
.029 .011
.033 .013
.031 .011
.035 .0088
.026 .0088
.037 .013
.033 .0084
.033 .0088
.020 .0088
.020 .0088
Holotype (sex undetermined) and three paratypes from Bighorn
National Forest, Sheridan County, Wyoming, elevation 8,000 ft., litter
from open pine woods, August 7 (K. Christiansen).
The furcula in this species is variable but always extremely reduced.
F. millsi is distinguished from other Nearctic species with five ana]
392
The Pan-Pacific Entomologist
spines by its clavate posterior body setae; the generally similar European
species F. mauriesi Cassagnau differs in having eight anal spines.
Friesea quinta Christiansen and Bellinger, new species
(Fig. Ik—m)
White with scattered blue pigment granules dorsally, eye patches darker, partially
subdivided; fourth antennal segment with deeply withdrawn unlobed apical bulb,
and with three distal and one basal blunt seta dorsally; eyes 4 + 4 (fig. lm) or
5 + 5; larger maxillary lamella unidentate, smaller lamella with about seven small
teeth; tibiotarsus with longest dorsal seta truncate, others acuminate; unguis
untoothed; tenaculum (fig. 11) bidentate; furcula (fig. 11) without mucro, and
with dens sharply demarcated from manubrium, subequal in length to anal spines,
and bearing three dorsal setae; five straight, stout anal spines subequal in length
to inner edge of unguis; integument moderately coarsely granulate; body setae
acuminate, smooth; maximum length 0.8 mm .
Holotype (sex undetermined) and four paratypes from Wheeler Peak,
Taos County, New Mexico, under aspen bark, July 2, 1955 (K.
Christiansen).
This species resembles F. nauroisi Cassagnau, which differs in having
6 + 6 eyes, a relatively much shorter dens, and distinct chaetotaxy.
F. pentacantha Mills is also similar, but has 8 + 3 eyes and only a single
dental seta.
Friesea wilkeyi Christiansen and Bellinger, new species
(Fig. 2)
White except for black eyes; fourth antennal segment (fig. 2c) with deeply
withdrawn unlobed apical bulb, and with four distal and two basal blunt setae
dorsally; eyes 2 + 2; maxilla with large lamella untoothed, small blade with about
four obscure teeth (fig. 2b) ; mandible (fig. 2a) with six teeth; tibiotarsus (fig.
2f) with short acuminate tenent hairs; unguis with small subapical inner tooth,
without lateral teeth; tenaculum (fig. 2d) unidentate; furcula (fig. 2d) with
mucro fused to dens, mucrodens about three-fourths as long as anal spines; two
straight anal spines (fig. 2e), slightly shorter than inner edge of unguis; in¬
tegumentary granules varying in diameter from about six micra on second thoracic
tergum to about fifteen micra on sixth abdominal segment; body setae acuminate,
smooth or minutely serrate; maximum length 0.9 mm.
Holotype (sex undetermined) from Coalinga, Fresno County, Cali¬
fornia, juniper litter, March 19, 1955 (H. L. Wilson) ; five paratypes
from Wharton Canyon, near Coalinga, Fresno County, California,
juniper litter, January 22, 1958 (H. L. Wilson).
R. F. Wilkey (unpublished) first recognized and described this spe¬
cies. It is similar to F. polla, new species and F. mistrali Rapoport and
Rubio (with 3 + 3 eyes) from Chile, but differs from both in eye num¬
ber and structure of the furcula.
Vol. 49, No. 4, October 1973
393
Fig. 2. Friesea wilkei. All figures at 1000 X magnification, after drawings of
type specimens by R. F. Wilkey. (a) mandible, (b) maxilla, (c) apex of fourth
antennal segment, acuminate setae omitted, (d) mucrodens and tenaculum, dorsal
aspect, (e) apex of abdomen, showing anal spines, (f) hind foot complex.
Friesea fara Christiansen and Bellinger, new species
(Fig. 3a-e)
Color unknown (only cleared specimens available) ; fourth antennal segment
weakly separated from third on ventral side, with an unlobed, deeply withdrawn
apical bulb, and with one basal and four apical blunt setae dorsally; eyes 8 -f- 8;
maxilla (fig. 3b) with large lamella tridentate, small lamella obscure but appar¬
ently with four teeth; mandible (fig. 3a) with six teeth; tibiotarsus (fig. 3d)
with eight to ten clavate setae, one greatly enlarged apically; unguis untoothed;
tenaculum (fig. 3c) varying from untoothed to weakly bidentate; furcula (fig. 3c)
without mucro, with dens reduced to knob half as long as anal spines and bare or
with up to three minute setae; six straight anal spines 1.2 to 1.5 times as long as
inner edge of unguis (fig. 3e) ; integument moderately coarsely granulate, diameter
of granules from twelve to fourteen micra; body setae smooth, acuminate except for
one pair of long, weakly clavate setae on the fourth, fifth, and sixth abdominal
segments, and a second strongly clavate pair on the sixth abdominal segment;
fourth and fifth abdominal segments strongly plurichaetotic; maximum length
0.8 mm.
Holotype (sex undetermined) and six paratypes from Farallon Islands,
California, September 23, 1951 (R. L. Usinger).
This is the only known plurichaetotic species of Friesea , and the only
Nearctic species with six anal spines. In number of eyes and anal spines
and the structure of the furcula, it resembles the European species F.
394
The Pan-Pacific Entomologist
Fig. 3. Friesea fara and F. polla. Figures at 1000 X magnification unless noted
otherwise. Fig. 3a-e. F. fara, structures of type specimens, (a) mandible, (b)
maxilla, (c) mucrodens and tenaculum (500X), (d) hind foot complex (500xC
(e) sixth abdominal segment, lateral aspect (250x)- Fig. 3f-j. F. polla, struc¬
tures of type specimens, (f) mandible, (g) fourth antennal segment, (h) mucro¬
dens, lateral aspect, (i) mucrodens, dorsal aspect, (j) hind foot complex.
handschini Kseneman, which has no clavate tibiotarsal or abdominal
setae.
Friesea polla Christiansen and Bellinger, new species
(Fig. 3f-j)
Color unknown (only cleared specimens available) ; fourth antennal segment
(fig. 3g) weakly separated from third on ventral surface, with deeply withdrawn,
unlobed apical bulb, with one basal and two or three distal blunt setae dorsally;
eyes 8 + 8; large maxillary lamella with a single inner tooth, small lamella with
three or four teeth; mandible (fig. 3f) with eight teeth; tibiotarsus (fig. 3j)
with tenent hair acuminate; unguis without teeth; tenaculum (fig. 3i) with 2 + 2
teeth; mucrodens (fig. 3h, i) subequal in length to anal spines, with small hook¬
like mucro partially separated from the dens; two straight anal spines, subequal
in length to inner edge of unguis; integument moderately coarsely granulate; all
body setae smooth, acuminate; maximum length 0.7 mm.
Holotype male and three paratypes from Pollock Pines, El Dorado
County, California, oak leaf litter, April 5, 1959 (J. R. Smith).
This species resembles F. wilkeyi in the number and structure of the
anal spines, but is distinguished by the furcal structure and eye number.
F. polla also resembles the Antarctic species F. grisea (Schaffer), which
differs in having clavate tenent hairs.
Vol. 49, No. 4, October 1973
395
Disposition of Types
The holotypes of all species described here will be deposited in the
Museum of Comparative Zoology, Harvard University. Paratypes of
F. wilkeyi , F. fara , and F. polla will be deposited in the collection of
the California Department of Agriculture, Sacramento.
Acknowledgments
Much of the work done on this paper was made possible by a grant
to Grinnell College from the Sloan Foundation. We would like to thank
Dr. R. F. Wilkey, who was responsible for making most of the California
material available to us.
Literature Cited
Cassagnau, P. 1958. Les especes europeennes du genre Friesea. Bull. Soc. Hist,
nat. Toulouse, 93: 17-29.
Macnamara, C. 1920. A new species of Friesea (Collembola). Canad. Ent., 53:
125-129.
Massoud, Z. 1967. Monographic des Neanuridae. Biol. Amer. austr., 3: 7-399.
SCIENTIFIC NOTE
Ceratosmicra campoplegicis Burks, a hyperparasite of Perilitus coc-
cinellae (Shrank) (Hymenoptera: Chalcididae, Braconidae). —A single
adult specimen of Hippodamia convergens Guerin-Meneville was collected from
a white fir branch [ Abies concolor (Gord. & Glend.) Lindl.] near Lake Edson,
El Dorado Co., California on 12 July 1969. A cocoon of Perilitus coccinellae
(Shrank) was found attached to its abdomen. The beetle was placed in rearing
and a single male hyperparasite, Ceratosmicra campoplegicis Burks (determined
by B. D. Burks, U. S. National Museum) emerged on 28 July 1969. C. campo¬
plegicis has been previously reared from cocoons of different Ichneumonidae but
this is the first report of its occurrence from a braconid primary parasite (B. D.
Burks, personal communication). This rearing also represents the first report of
a chalcidid hyperparasite on P. coccinellae (personal communication, K. S. Hagen,
Division of Biological Control, University of California, Berkeley).
Portions of this study have been funded by Environmental Protection Agency
Grant #5 ROLE P-00814-05.—F. M. Stephen, Division of Biological Control, Uni¬
versity of California, Berkeley, 94720.
396
The Pan-Pacific Entomologist
Ant Larvae of the Subfamily Dolichoderinae:
Second Supplement
(Hymenoptera: Formicidae)
George C. Wheeler and Jeanette Wheeler
Laboratory of Desert Biology, Desert Research Institute,
University of Nevada System, Reno 89507
Subsequent to the publication of our first supplement on the ant
larvae of the subfamily Dolichoderinae (1966) we have collected or
received from other myrmecologists so much additional material that
it has become necessary to publish another supplement, increasing the
count of species studied by 20%. The additional larvae described here
agree with our original characterization of the Dolichoderinae (1951:
207), with the exception of Dolichoderus scabridus Roger, which has
paxilliform labial palpi.
Dolichoderus Lund
REVISION: In our generic characterization (1951: 172) the second
sentence should read: A pair of ventrolateral bosses frequently present
on the prothorax.
Dolichoderus (Diceratoclinea) scabridus Roger
Lengths (through spiracles) 2.5 and 6 mm; straight length 1.9 and 4.2 mm.
Similar to D. ( H .) taschenbergi (1951: 173) except as follows. T2 and T3 more
swollen. Body hairs numerous on thorax and few on venter of AI. Of two types:
(1) 0.025-0.03 mm long, simple, slender, flexuous, without alveolus and articular
membrane; (2) about 0.048 mm long, stouter, with alveolus and articular mem¬
brane, a few on each somite. Cranium transversely subelliptical. Head hairs
0.05-0.075 mm long, slender, flexuous and numerous (about 200). Labrum sub-
trapezoidal, narrowed ventrally, width four times the length; anterior surface
with 2 sensilla and a few oblique rows of spinules; ventral border with 6 sensilla;
posterior surface with 8 sensilla. Maxillary palp a skewed peg with 5 sensilla;
galea represented by 2 slightly raised sensilla. Labium small and feebly bilobed;
palp a skewed peg with 4 sensilla; an isolated sensillum between each palp and
opening of sericteries. Hypopharynx with minute spinules in short subtransverse
rows. (Material studied: numerous larvae from New South Wales, Tahmoor,
28-XII-1966, courtesy of Rev. B. B. Lowery.)
Dolichoderus (Hypoclinea) australis E. Andre
Length (through spiracles) about 3.1 mm; straight length about 2.4 mm. Prob¬
ably very similar to D. (H.) taschenbergi (1951: 173) except in the following-
details. Spiracles on AVIII vestigial. Body hairs 0.006-0.018 mm long, numerous
on venter of Tl, longest and most numerous on AX. Head with flattened dorsal
The Pan-Pacific Entomologist 49: 396-401. October 1973
Vol. 49, No. 4, October 1973
397
areas less distinct. Head hairs shorter (0.006-0.013 mm long). Labrum with 2
sensilla on anterior surface and 4 sensilla on ventral surface. Mandibles with apical
tooth straighter. Maxillary palp a raised cluster of 4 sensilla; galea 2 slightly
raised sensilla. Labial palp a raised cluster of 4 sensilla; an isolated sensillum
between each palp and opening of sericteries. (Material studied: 4 semipupae
from South Australia, Lofty Range 1500 ft, Morialta Reserve, 1-III-1969, courtesy
of Rev. B. B. Lowery.)
Dolichoderus (Hypoclinea) germaini Emery
Length (through spiracles) about 8.2 mm; straight length about 5 mm. Similar
to D. ( H .) taschenbergi (1951: 173) except as follows. Very short and stout;
no lateral welt on Tl. Body hairs 0.005-0.03 mm long; very few (mostly on
venter of Tl). Maxillae with a few minute spinules in encircling rows; palp a
slightly raised cluster of 3 sensilla. Labium with minute spinules, isolated or in
short rows; palp a slightly raised cluster of 4 sensilla. Hypopharynx with a few
minute spinules. (Material studied: 4 larvae from Brazil, Utiariti, 325 m, Rio
Papagalo, MT., VIII-1961, courtesy of Dr. K. Lenko.)
Iridomyrmex Mayr
REVISION: Our generic characterization (1951: 184) should be
changed to read: Body slightly curved. Anus ventral or posteroventral.
Body and head hairs usually minute. Head moderately large. Antennae
small.
Iridomyrmex melleus Wheeler
(Fig. 3)
Length (through spiracles) about 1.8 mm, straight length about 1.4 mm.
Stout; dorsal profile long and C-shaped; ventral feebly sigmoid; a middorsal
boss on AI; AX with a small projection posterior to the anus. Anus postero¬
ventral. Leg and wing vestiges present. Spiracles small, those on AI largest.
Integument with minute spinules in transverse rows, most numerous and prominent
on venter of anterior somites and dorsum of posterior somites. Body hairs sparse,
short and generally distributed. Of 2 types: (1) 0.025-0.036 mm long, on dorsal
and lateral surfaces, bifid, the branches short to long; (2) 0.008-0.025 mm long,
simple, most numerous on ventral surface of Tl. Cranium about a third broader
than long, broadest dorsally, suboctagonal in anterior view. Antennae large, each
with 3 minute sensilla. Head hairs few, 0.013-0.025 mm long, simple. Labrum
about a third broader than long, bilobed, erectile; each lobe with 1 sensillum on
anterior surface, 1 on ventral border and with 1 plus a few short rows of minute
spinules on posterior surface. Mandibles feebly sclerotized, slightly longer than
broad, subtriangular in anterior view, apical tooth small and blunt, with one
small subapical tooth, posterior surface with coarse denticles; mandibles erectile.
Maxillae adnate; palp a slight elevation with 5 sensilla; galea a short cone with
2 sensilla. Labium small; anterior surface with minute spinules in short trans¬
verse rows; palp a slight elevation with 5 sensilla; opening of sericteries a short
transverse slit. Hypophayrnx densely spinulose, spinules arranged in rows, rows
398
The Pan-Pacific Entomologist
grouped in 2 subtriangles which have their bases near the middle. (Material
studied: numerous larvae, courtesy of Dr. R. Lavigne; in pith chambers of
Cecropia peltata on forest floor, 2-VII-1970, El Verdi Field Sta., Loquilla Forest,
Puerto Rico.)
This is the only species we have seen with erectile mandibles and
labrum.
The remaining species of Iridomyrmex are compared with I. pruinosus
(1951: 185) ; only differences are given here.
Iridomyrmex conifer Forel
Length (through spiracles) about 3 mm; straight length about 2.8 mm. Spiracles
largest on T3 and AI, remainder small. Integument with minute spinules in
numerous short arcuate rows. Body hairs 0.005-0.025 mm long. Cranium sub-
circular. Head hairs few and minute (about 0.001 mm long). Labrum breadth
3 times length; bilobed; each lobe with 2 sensilla on anterior surface and one on
ventral border; entire posterior surface with a few sensilla and with minute
spinules. Mandibles with apical tooth more prominent, straighter and longer.
Maxillary and labial palps represented by clusters of 5 sensilla each. (Material
studied: 2 larvae from Western Australia, Blackwood River near Nannup, 18-
XII-1969, courtesy of Rev. B. B. Lowery.)
Iridomyrmex detectus F. Smith
Length (through spiracles) about 2.9 mm; straight length about 2.7 mm. Body
stouter; dorsal profile longer; ventral shorter and more nearly straight. Head and
anus ventral. Integument with minute spinules, isolated or in short rows on
posterior somites and ventral surface of anterior somites. Body hairs 0.008-0.016
mm long, mostly on Tl and AX. Dorsal cranial outline a smooth curve; integu¬
ment spinulose, spinules of moderate size and isolated or in short subparallel
transverse rows. Head hairs few, 0.013-0.025 mm long. Labrum narrow; distinctly
bilobed; each lobe with minute spinules in short rows and 3 sensilla ventrally
on anterior surface, with 2 sensilla and several minute spinules on ventral border,
and with a few short rows of minute spinules medially and 3 sensilla on the
posterior surface. Mandibles with apical tooth more prominent and sharp-pointed.
Maxillae lobose. Labial palp a rounded knob. (Material studied: 10 larvae from
Queensland, St. George, 6-1-1966, courtesy of Rev. B. B. Lowery.)
Iridomyrmex glaber Mayr
Length (through spiracles) about 3.9 mm; straight length about 3.0 mm. Body
short, stout and subovoidal; head applied to ventral surface below anterior end;
anus posteroventral. Entire integument spinulose, spinules minute and in short
transverse rows ventrally and posteriorly, some isolated spinules laterally. Body
hairs moderately numerous and generally distributed, minute (about 0.003 mm
long). Dorsal outline of cranium broadly curved; integument with short rows
of minute spinules. Head hairs more numerous but still few. Mandibles with
larger apical tooth. Maxillary and labial palps represented by clusters of 3
sensilla each.
Young larva: Length (through spiracles) about 1.4 mm. Body stouter and
Vol. 49, No. 4, October 1973
399
Figs. 1-3. Fig. 1. Forelius foetidus : a, head in anterior view, X107; b, left
mandible in anterior view, X303; c, body hair, X677; d, very young larva in
side view, X35; e, young larva in side view, X35; f, mature larva in side view,
X35. Fig. 2. Iridomyrmex nitidus: head in anterior view, X72. Fig. 3. Irulo-
myrmex melleus: a, head in anterior view, Xlll; b, mouth parts in anterior
view with mandibles and labrum erected, Xlll; c, head in side view with
mandibles and labrum erected, X133; d, larva in side view, X47; e, dorsal
portion of first abdominal somite showing boss in lateral view, Xl33; f and g,
two types of body hairs, X533; h, left mandible in anterior view, X370.
400
The Pan-Pacific Entomologist
outlines straighter, but with 6 distinct middorsal bosses, one each on T2 and
AI-AV. Thirteen feebly differentiated somites. Spinules more prominent on
posterior end. Body hairs 0.002-0.015 mm long, longest ventrally. Otherwise simi¬
lar to mature larva.
Material studied: numerous larvae from New South Wales, DeBurgh’s Bridge,
Lava Cove, Sydney, 26-V-1966, courtesy of Rev. B. B. Lowery.)
Iridomyrmex nitidus Mayr
(Fig. 2)
Length (through spiracles) about 4.9 mm; straight length about 3.4 mm. Body
stouter. Spiracles on AVIII vestigial. Integument of venter of anterior somites
and dorsum of posterior somites with minute spinules, isolated or in short rows.
Body hairs 0.001-0.025 mm long, with a slight concentration around AI spiracle.
Cranium subquadrangular and with a bilobed flattened dorsal portion. Antennae
slightly elevated, each with 3 small sensilla, bearing a spinule each. Head hairs
shorter (0.006-0.012 mm long) and more numerous. Labrum width twice length,
bilobed, each lobe with 2 hairs on anterior surface, with 2 median sensilla on
ventral border and with 3 or 4 sensilla on posterior surface; entire posterior
surface spinulose, spinules minute and in subtransverse rows. Mandibles with
breadth equal to length; apical tooth moderately sclerotized and base feebly
sclerotized; anterior surface of base spinulose, spinules minute and in short rows;
apical portion narrowed, tapering only slightly; apex round-pointed or with
rounded tip and a small subapical medial tooth. Maxillae adnate; palp a slightly
raised, heavily sclerotized base with 5 sensilla; galea a boss with 2 sensilla. Labial
palp a heavily sclerotized knob with 5 sensilla; an isolated sensillum between
each palp and opening of the sericteries. (Material studied: numerous larvae
from New South Wales, Mt. Mullens, Murwillumbah, 2-IX-1966, courtesy of Rev.
B. B. Lowery.)
Iridomyrmex viridiaeneus Viehmeyer
Length (through spiracles) about 6.2 mm; straight length about 5 mm. Body
with dorsal profile long and C-shaped ventral short and nearly straight; head and
anus ventral. Entire integument spinulose, spinules minute and in transverse
rows. Body hairs sparse, more numerous ventrally. Of two types: (1) 0.013-
0.056 mm long, simple, with fine tip; (2) 0.025-0.05 mm long, with a few fine
denticles near tip. Cranium subtriangular in anterior view, a fourth broader than
long. Head hairs twice as numerous and shorter (0.006-0.013 mm long). Labrum
1% times as broad as long; bilobed, each lobe with 2 sensilla and 1 minute
hair on the anterior surface, with minute spinules on ventral border, and with
4 or 5 sensilla in a medial cluster posteriorly; entire posterior surface with
minute spinules in transverse subparallel rows. Mandibles nearly as broad as
long and with the medial border erose. Maxillary palp a heavily sclerotized
cluster of 5 sensilla; galea a sclerotized knob with 2 sensilla. Labial palp a
slightly raised cluster of 5 sensilla; an isolated sensillum between each palp and
opening of sericteries, the latter a wide transverse slit. (Material studied: 5 larvae
from South Australia, 8 miles SW Morgan, courtesy of Rev. B. B. Lowery.)
Vol. 49, No. 4, October 1973
401
Forelius Emery
ADDITION to our generic characterization (1966: 729) : A mid¬
dorsal boss on AIL
Forelius foetidus (Buckley)
(Fig. 1)
Mature larva: Length (through spiracles) about 2.3 mm; straight length
about 1.6 mm. Plump, chunky and turgid; dorsal profile long and C-shaped;
ventral profile feebly sigmoid; a low middorsal boss on All; both ends broadly
rounded; head and anus ventral. Leg, wing and gonopod vestiges present. Spiracles
small; diameter of AI greatest, diminishing posteriorly. Entire integument spinu-
lose, spinules minute and isolated or in short transverse rows. Body hairs very
few, minute (0.007-0.013 mm long), simple, most numerous on venter of Tl.
Cranium subhexagonal, with a depressed transverse dorsal band. Antennae large,
slightly raised, each with 3 sensilla, each hearing a spinule. Head hairs few,
simple and minute (about 0.013 mm long). Labrum short, broad, feebly bilobed:
each lobe with 2 sensilla on anterior surface and 2 on posterior surface; entire
posterior surface sparsely spinulose, spinules minute and isolated or in short
transverse rows. Mandibles very feebly sclerotized, subtriangular in anterior view,
base broad, apex sharp-pointed. Maxillae appearing adnate, with apex rather
broadly round-pointed; palp a cluster of 5 sensilla; galea represented by 2
sensilla. Labium with ventral border paraboloidal, a few minute spinules near
opening of sericteries; palp a slightly elevated cluster of 5 sensilla; an isolated
sensillum between each palp and opening of sericteries, the latter a short trans¬
verse slit. Hypopharynx densely spinulose, spinules in rows, rows grouped in 2
subtriangles which have their bases near middle.
Young larva: Length (through spiracles) about 1.7 mm. Similar to mature
larva except as follows. Body more slender; segmentation more distinct. Diameter
of spiracles on Al much greater than remainder which are minute. Body hairs
about 0.001 mm long.
Very young larva: Length (through spiracles) about 0.9 mm. Similar to
young larva except as follows. Body much more slender; head nearly diameter
of body and on anterior end; 10 distinct somites; boss on dorsum of All very
distinct; posterior end narrowly rounded. Anus ventral and with a distinct pos¬
terior lip. Body hairs shorter (about 0.008 mm).
Material studied: numerous larvae from Arizona, Organpipe Cactus National
Monument, 1600 ft, 10-IV-1968, G. C. and J. Wheeler collection # 108.
Literature Cited
Wheeler, G. C., and J. Wheeler. 1951. The ant larvae of the subfamily
Dolichoderinae. Proc. Entomol. Soc. Wash., 53: 169-210.
1966. The ant larvae of the subfamily Dolichoderinae: supplement. Ann.
Entomol. Soc. Amer., 59: 726-732.
402
The Pan-Pacific Entomologist
SCIENTIFIC NOTES
Parasitization of Malacosonia larvae by Tachinidae in coastal Cali¬
fornia (Lepidoptera: Lasiocampidae; Diptera). —During studies of insects
associated with coast live oak, Quercus agrifolia Nee, I collected large larvae of
the Pacific tent caterpillar, Malacosonia constrictum (Hy. Edwards) and western
tent caterpillar, M. californicum (Packard), from and around oaks on the campus
of California State University, Hayward. I fed them fresh oak leaves which were
first thoroughly scrutinized to eliminate parasite eggs. When cocoons were spun,
each was placed within a 4 oz glass jar with a loosely-attached lid to permit air
circulation. When a parasitoid larva emerged, the resulting puparium was trans¬
ferred to a vial in which was placed a slightly moistened piece of paper towel.
Flies emerged from 45% of these puparia.
No parasitoids resulted from 90 larvae collected in 1970 or 60 collected in 1971.
Results for 1972 appear in Table 1; 51% of 139 larvae were parasitized. High
proportions of parasitization by Tachinidae are somewhat unusual in Malacosoma
larvae. Bess (1936, Ann. Entomol. Soc. Amer., 29: 593-613) reported 23% para¬
sitization of M. disstria Hubner larvae and Kulman (1965, J. Econ. Entomol., 58:
66-70) found up to 39% of M. americanum (F.) parasitized, both in areas of high
host density. In my study area, Malacosoma larvae increased from moderate to
high density during the 3 years, so it is possible that the rise in parasitism rep¬
resents a delayed numerical response to increasing host numbers. Alternate hosts
are doubtless important in maintaining such a high density of parasitoids.
Two species of Tachinidae were recovered: Compsilura concinnata (Meigen),
a parasite of European origin that has been recorded from over 200 hosts includ¬
ing several Malacosoma (Witter and Kulman, 1972, Univ. Minnesota Agr. Expt.
Sta. Tech. Bull., 289) and Chaetogaedia monticola (Bigot), which likewise has
been reared from several Malacosoma (Stehr and Cook, 1968, Bull. Smithson. Inst.,
276; Witter and Kulman, 1972). Both flies parasitized M. constrictum and cali¬
fornicum in approximately equal numbers, but all 18 Chaetogaedia were reared
from host larvae collected away from foliage, presumably searching for cocooning
sites on tree trunks or surrounding litter. Compsilura emerged from host larvae
collected both on and off foliage, and it reached peak numbers somewhat earlier
than did Chaetogaedia. Turnock (1961, Can. Dept. Forest. Bi-M. Prog. Rep., 17:
Table 1. Parasitism of Malacosoma larvae by Tachinidae, Hayward, California
1972.
Date
Host
larvae
Compsilura
Chaetogaedia
8 Apr
35
13 (37%)
1 (3%)
15 Apr
20
8 (40%)
3 (15%)
22 Apr
62
30 (48%)
4 (6%)
26 Apr
15
2 (13%)
6 (40%)
10 May
7
0
4 (57%)
Vol. 49, No. 4, October 1973
403
2) noted spatial differences in parasitism by Sarcophagidae on M. disstria pupae.
It is possible that such spatial and temporal differences as I report here may per¬
mit coexistence of both species of Tachinidae on the same host species. —David J.
Horn, Department of Entomology, The Ohio State University, Columbus, 43210.
Apocephalus borealis Brues parasitic upon Vespula spp. (Diptera:
Phoridae; Hymenoptera: Yespidae).— During 1971 and 1972, larvae, pupae
and adults of a phorid fly, Apocephalus borealis Brues, were recovered from three
species of yellowjackets in northern California. Specimens were kindly identified
by Dr. W. W. Wirth (U. S. National Museum). The parasites were obtained from
two ground nesting species, Vespula pensylvanica (Saussure) and V. vulgaris (L.),
and the aerial nesting species, V. arenaria (F.).
In his recent revision of North American phorids, Borgmeier (1963, Stud. En-
tomol. (n.s.), 6: 1-256) reported that A. borealis had been reared from “Vespula?”
in Minnesota and from the spider, Latrodectus mactans (F.) in British Columbia.
Other New World species of this widespread genus are parasitic on various species
of ants and one parasitizes the cantharid beetle, Chauliognathus fallax Germar,
in South America.
Large numbers of wasps were collected in cone traps placed directly over the
ground nest openings. When the nest appeared vacated, it was dug up and ex¬
amined. Lesser numbers of yellowjackets were trapped in cages baited with tuna
cat food. Samples of V. arenaria were taken from an aerial nest.
Parasitized yellowjackets were observed in a very lethargic state, slowly crawl¬
ing about on the floor of the cage or lying on their back or side with legs quivering.
These affected wasps were closely pursued by other wasps which attacked and
often dismembered them. Social wasps instinctively remove dead or dying members
from their nests. However, I could not determine if the antagonistic behavior to¬
ward the weakened individuals was due to their parasitized condition. Last instar
larvae emerged from affected wasps through the thoracic membranes, mainly in the
cervical region. The maximum number observed to emerge from a single wasp was
eight. Dissection of lethargic wasps revealed that most larvae had developed in the
thoracic haemocoel, but fly larvae were occasionally found in the abdominal cavity.
Upon pupation, the larvae turned light brown and attached themselves to the sub¬
strate. Adult flies emerged 2-4 days later.
Apocephalus borealis is apparently active throughout the yellowjacket season.
Reared specimens were recovered from V. pensylvanica workers and from excavated
nests from July to December and from V. vulgaris workers and a V. arenaria nest
in July and August. In four V. pensylvanica nests, all collected near Martinez,
California, approximately 5% of the yellowjackets were parasitized. Only one or
two parasitized yellowjackets were caught in the baited cone traps even though
large numbers of individuals were trapped. This observation suggests that A. bore¬
alis females confine their activities to the yellowjacket nest. The oviposition habits
of other phorids suggests that A. borealis females lay their eggs primarily on yel¬
lowjacket larvae or teneral adults. Females of A. borealis are equipped with a
flat, heavily sclerotized, lanceolate ovipositor which might be used to puncture
the integumental membranes of the yellowjacket.
Borgmeier (1963) lists Muir Wood, Marin County and Gasquet, Del Norte
404
The Pan-Pacific Entomologist
County in the California distribution of A. borealis. I have recovered material
from San Rafael, Mill Valley (Marin County) and El Cerrito, Point Molate and
Martinez (Contra Costa County). —Franklin Ennik, State Department of Public
Health, Berkeley, 94704.
ZOOLOGICAL NOMENCLATURE
Required six-months’ notice is given of the possible use of plenary powers by
the International Commission on Zoological Nomenclature in connection with the
following names listed by case number:
Announcement A. (n. s.) 90
(see Bull. Zool. Nomencl. 30, part 1, 6th July 1973)
1958. Correction of homonymy of DREPANIDAE in Insecta and Pisces.
2008. Suppression of Macgillivraya Grote, 1894 (Insecta, Collembola).
2009. Validation of Haematopinus eurysternus Denny, 1842 (Insecta, Anoplura).
2017. Suppression of Rhopalidia Lepeltier, 1836 (Insecta, Hymenoptera).
2023. Suppression of Formica maxima Moore, 1842 (Insecta, Hymenoptera).
2025. Suppression of Daristane Walker, 1859 (Insecta, Lepidoptera).
2026. Type-species for Crinocerus Burmeister, 1839 (Insecta, Hemiptera).
2028. Type-species for Onycholyda Takeuchi, 1938 (Insecta, Hymenoptera).
Announcement A. (n. s.) 91
(see Bull. Zool. Nomencl. 30, part 2, 10th October, 1973)
2027. Designation of a neotype for Pieris virginiensis Edwards, 1870 (Insecta,
Lepidoptera).
2028. Designation of a typespecies for Onycholyda Takeuchi, 1938 (Insecta,
Hymenoptera).
2029. Designation of lectotype for Psednura longicornis Sjbstedt, 1920 (Insecta,
Orthoptera).
2033. Suppression of Sceptrophorus Foerster, 1856 (Insecta, Hymenoptera).
2035. Suppression of Drosophila carinata Grimshaw, 1901 (Insecta, Diptera).
2036. Suppression of Hydrophorus jaculus Fallen as type-species of Hydrophorus
Fallen, 1823, in favour of H. nebulosus Fallen. (Insecta, Diptera.)
2037. Designation of a type-species for Oscinella Becker, 1909 (Insecta, Diptera).
2038. Suppression of Leucospis gallica, Villers, 1789 and Cynips lugdunaea Gmelin,
1790 (Insecta, Hymenoptera).
2040. Designation of a type-species for Madiza Fallen, 1810 (Insecta, Diptera).
Comments should be sent in duplicate, citing case numbers, to the Secretary,
International Commission of Zoological Nomenclature, c/o British Museum (Nat¬
ural History), Cromwell Road, London SW7 5BD, England. Those received early
enough will be published in the Bulletin of Zoological Nomenclature. — Margaret
Green, Scientific Assistant.
Vol. 49, No. 4, October 1973
405
INDEX TO VOLUME 49
Adams, new Coniopterygidae, 250
synonymy, 324
Aethephemera, 306
Alcock, Behavior of Microbembex, 144
Allen, new Ephemerellidae, 160
new Leptohyphes, 363
Allen, new Comadia, 240
Antonelli, larva and notes on Antho-
comus, 56
Araneida
Diguetidae, 110
Arnaud, Parrish collection, 48
Cook obituary, 177
Ayala, new Drosophila, 273
Baetis ellenae, 285
Baetodes projectus, 308
peiiiculus, 310
longus, 311
furvus, 313
Bellinger, new Friesea, 389
Benedetti, notes on Neomachilis, 246
Bentzien, Biology of Diguetia imperiosa,
110
Book reviews, 55, 101, 123, 143, 176,
184, 228, 239, 280, 281, 282, 283,
299
Borboropsis steyskali, 374
Brown, new Comadia, 240
Campbell, new Diplolepsis, 174
Chaetocnema repens, 62
Chandler, revision of Formicilla, 31
new Hamotus, 255
new records, 288
Hamotus of Arizona, 378
Chemsak, new Eburia, 338
Chilenoperla illiesi, 318
Chilcott, new Symphoromyia, 5
Christian, new Friesea, 389
Clark, records of black witch moth, 353
Coleoptera
Anobiidae, 102
Anthicidae, 31, 67
Carabidae, 80, 126
Cerambycidae, 338
Chrysomelidae, 61
Cyathoceridae, 288
Georyssidae, 285
Lagriidae, 1
Pselaphidae, 255, 378
Scarabaeidae, 127
Staphylinidae, 42, 279
Collembola, 389
Comadia suaedivora, 240
Dailey, unisexual generation of Andricus,
171
new Diplolepis, 174
synonymy of Andricus, 188
Daly, fumigation of specimens, 377
lectotype designations, 383
Denning, new Trichoptera, 132
Dicrephemera, 302
Diplolepis inconspicuis, 174
Diptera
Drosophilidae, 273
Heleomyzidae, 373
Phoridae, 403
Rhagionidae, 5
Doutt, genus Paranaphoidea, 185
genus Paratrichogramma, 192
genus Polynemoidea, 215
fossil Mymaridae, 221
Drosophila willistoni quechua, 276
ecjuinoxialis caribbensis, 277
Eburia affluens, 338
brevicornis, 339
brunneicomis, 340
elegans, 342
inarmata, 343
latispina, 344
megalops, 346
minutivestis, 347
opaca, 347
paraegrota, 348
poricollis, 351
submutata, 352
Editorial notice, 25, 184
Eighme, new Pulverro, 49
Emmel, correction, 4
variability in Philotes, 74
Ennik, Vespula parasite, 403
Ephemerella (Drunella) kabwlensis,
163
Ephemerella (Serratella) subsolana,
164
Ephemeroptera
Baetidae, 385, 308
Ephemerellidae, 160
Ephemeridae, 300
Tricorythidae, 363
Eurytoma adiacrita, 354
bugbeei, 356
daileyi, 359
Farula honeyi, 137
petersoni, 139
Formicilla longipilosa, 35
alta, 39
vitrea, 40
Foster, larva and notes on Anthocomus,
56
Frankie, habits of Ernobius conicola,
102
Friesea cera, 389
fara, 394
millsi, 389
polla, 934
quinta, 392
wilkeyi, 392
Gardner, new milliped, 270
406
The Pan-Pacific Entomologist
Gilbert, behavior of Trigona, 21
Glossosoma sequoia, 132
Goodpasture, new Plebejus, 149
book review, 299
Gordh, new Trjapitzinellus, 51
Grigarick, Macrobiotus, 299
eggs of Macrobiotus, 258
Grissell, new Torymidae, 232
new Eurytoma, 354
Hammotus cavus, 255
populus, 379
Hardy, new Phobetus, 127
Hemiptera
Reduviidae, 183
Rhopalidae, 196
Homoptera
Coccidae, 264
Horn, Leptocoris, 196
parasitization of Malacosoma, 462
Hydropsyche cora, 135
Hymenoptera
Apidae, 21
Cynipidae, 171, 174, 188
Encyrtidae, 51
Eurytomidae, 354
Formicidae, 207, 396
Ichneumonidae, 124, 197
Megachilidae, 294
Mymaridae, 185, 214, 221
Pompilidae, 182
Pteromalidae, 337
Sphecidae, 26, 49, 144
Tenthredinidae, 93
Tiphiidae, 325
Trichogrammatidae, 192
Incasemidalis meinanderi, 253
Larochelle, notes on Scaphinotus, 80
wing dimorphism, 126
Leech, note on Ageniella, 182
book reviews, 101, 281, 282
Legner, distribution of Cafius, 279
Leiomyrmosa, 325
spilota, 326
Lepidoptera, 149
Lasiocampidae, 402
Lycaenidae, 74
Noctuidae, 353
Nymphalidae, 289
Leptohyphes carinus, 365
edmundsi, 363
flinti, 365
invictus, 368
jamaicanus, 371
minimus, 369
populus, 366
rolstoni, 371
tinctus, 368
viriosus, 369
Linsley, new Eburia, 338
Litus mexicanus, 223
Lugaski, records of black witch moth,
353
Lysiopetalidea, 270
Manohyphella, 160
Manohyphella keiseri, 163
Mathis, review of Borboropsis, 373
Mayo, new Baetis, 385
new Baetodes, 308
McCafferty, classification of Ephemera,
300
McCrea, new flea beetle, 61
Menke, lectotype designation, 257
Miller, mealybugs, 264
Monodontomerus saltuosus, 232
dementi, 235
Moore, note on Dacnochilus, 42
distribution of Cafius, 279
Muchmore, new pseudoscorpions, 43
Murochernes, 43
Myrmosula boharti, 331
latericarinata, 333
nasuta, 329
Nelson, synopsis of Chilenoperla, 315
Neuroptera
Coniopterygidae, 250, 324
Nutting, new Hamotus, 255
Paetzel, behavior of Trypoxylon, 26
Pampoconis punctipennis, 250
uncinatus, 251
Paratrichogramma pretoriensis, 193
californica, 195
Parker, notes on Lithurgus, 294
Parsons, Lagriidae of California, 1
Parthina vierra, 141
Peterman, behavior in Tanarthrus, 67
Phobetus chearyi, 128
Piper, new records for Heteroschema,
337
Plebejus acmon texanus, 149
Plecoptera, 315
Polynemoidea mexicana, 219
Potter, notes on Lithurgus, 294
Proceedings, Pacific Coast Entomologi¬
cal Society, 81
Pseudoscorpionida, 43
Pulverro californica, 49
Rust, ectoparasites of Thomomys, 59
Ryan, behavior of Microbembex, 144
Schuster, Macrobiotus, 229
eggs of Macrobiotus, 258
Scorpionida
Chactidae, 60
Vejovidae, 165
Shapiro, aberration in Cynthia, 289
Shields, variability in Philotes, 74
Slobodchikoff, studies of Therion, 197
Smith, sawfly tribe Lycaotini, 93
Soleglad, synonymy, 60
redescription of Vejovis, 165
Spinolochus disolatus, 124
Sprenger, unisexual generation of An-
dricus, 171
synonymy of Andricus, 188
Vol. 49, No. 4, October 1973
407
Statira dumalis, 1
latitator, 3
Stephen, hyperparasite, 395
Symphoromyia cervivora, 6
inconspicua, 12
nana, 15
truncata, 17
Synaptoneura vopida, 96
Tardigrada, 229, 258
Thorp, book reviews, 55, 123, 143, 176,
184, 228, 239, 280, 283
Thysanura, 246
Tinodes gabriella, 133
Toftner, Macrobiotus, 229
eggs of Macrobiotus, 258
Torgersen, new Spinolochus, 124
Torymus frankiei, 236
memnonius, 237
Trew, variability in Philotes, 74
Trichoptera, 132
Trjapitzinellus microrphanos, 51
Turner, new Symphoromyia, 5
Tynomma schlingeri, 270
Wale, Cook obituary, 177
Wasbauer, note on Ageniella, 182
new Myrmosinae, 325
Wheeler, ant larvae, 207
ant larvae, 396
Wood, Triatoma protracta, 183
Zoological Nomenclature, 73, 404
MAILING DATES FOR VOLUME 49
No. 1—January _ mailed 18 May 1973
No. 2—April _ mailed 11 July 1973
No. 3—July _ mailed 19 October 1973
No. 4—October _ mailed 16 April 1974
Published by the
Pacific Coast Entomological Society
in cooperation with
The California Academy of Sciences
VOLUME FORTY-NINE
1973
EDITORIAL BOARD
J. T. DOYEN, Editor
J. A. CHEMSAK, Assistant Editor
E. G. LINSLEY
HUGH B. LEECH
E. S. ROSS
H. V. DALY
P. H. ARNAUD, JR., Treasurer
PUBLICATION COMMITTEE
1973
1974
1975
D. D. Linsdale
J. A. Powell, Chairman
M. S. Wasbauer
E. I. Schlinger
M. A. Gardner
E. G. Linsley
San Francisco, California
1973
11
CONTENTS FOR VOLUME 49
Adams, P. A.
Coniopterygidae from Chile, with three new species . 250
Synonymy in Pampoconis .,. 324
Alcock, J. and A. F. Ryan
The behavior of Microbembex nigrifrons .... 144
Allen, R. K.
New Ephemerellidae from Madagascar and Afghanistan . 160
New species of Leptohyphes Eaton . 363
Arnaud, P. H., Jr.
Chris Parrish scorpion collection donated to California Academy
of Sciences . 48
Arnaud, P. H., Jr. and M. M. Wale
Thomas Wrentmore Cook (1884^1962) ..... 177
Ayala, F. J.
Two new subspecies of the Drosophila willistoni group . 273
Benedetti, R.
Notes on the biology of Neomachilis halophila on a California
sandy beach . 246
Bentzien, M. M.
Biology of the spider Diguetia imperiosa .. 110
Book Notices... 245, 342
Book Reviews .... 55, 101, 123, 143, 176, 184, 228, 239, 280-283, 299
Brown, R. M. and R. P. Allen
A new Comadia from the San Joaquin Valley of California . 240
Chandler, D. S.
A revision of the new world genus Formicilla . 31
New records for Georyssidae and Cyathoceridae (Coleoptera)
in Mexico . 288
The Hamotus of Arizona . 378
Chandler, D. S. and W. L. Nutting
A new species of Hamotus from galleries of Microtermes septen-
trionalis in Western Mexico . 255
Ill
Chemsak, J. A. and E. G. Linsley
New species of Eburia from Mexico . 388
Christiansen, K. and P. Bellinger
Six new Nearctic species of the genus Friesea ....... 389
Dailey, D. C. and L. Campbell
A new species of Diplolepis from California... 174
Dailey, D. C. and C. M. Sprenger
Unisexual generation of Andricus atrimentus __ 171
Synonymy of Andricus gigas and the bisexual generation of
Andricus crenatus . 188
Daly, H. V.
Fumigation of museum specimens in oven-cooking bags .. 377
Lectotype designations of African bees of the genus Ceratina
and allies ........ 383
Denning, D. G.
New species of Tricboptera . 132
Doutt, R. L.
The genus Paranaphoidea Girault _ 185
The genus Paratrichogramma Girault ..... 192
The genus Polynemoidea Girault..... 215
The fossil Mymaridae . 221
Eighme, L. E.
A new Pulverro from California ... 49
Emmel, T. C. and J. F. Emmel
Correction . 4
Emmel, T. C., H. R. Trew, and 0. Shields
Chromosomal variability in a Nearctic Lycaenid butterfly,
Philotes sonorensis _ 74
Ennik, F.
Apocephalus borealis Brues parasitic upon Vespula spp. 403
Foster, D. E. and A. L. Antonelli
Larval description and notes on the biology of Anthocomus horni 56
Frankie, G. W.
Feeding habits and seasonal history of Ernobius conicola in cones
of Monterey Cypress with notes on cohabiting insects . 102
IV
Gardner, M. R.
A new milliped of the genus Tynomma from California. 270
Gilbert, W. M.
Foraging behavior of Trigona fulviventris in Costa Rica .. 21
Goodpasture, C.
A new subspecies of Plebejus acmon ____ 149
Book Review: An Index to the Described Life Histories, Early
Stages and Hosts of the Macrolepidoptera of the Continental
United States and Canada _____ 299
Gordh, G.
Description of a new species of Trjapitzinellus parasitic on
Coniopterygidae in California _______ 51
Grigarick, A. A., R. 0. Schuster, and E. C. Toftner
Macrobiotus montanus from California ...... 229
Descriptive morphology of eggs of some species in the Macro¬
biotus hufelandii group ... 258
Grissell, E. E.
New species of North American Torymidae .... 232
New species of Eurytoma associated with Cynipidae .. 354
Hardy, A. R.
A new species of Phobetus ... 127
Horn, D. J.
Leptocoris rubrolineatus, an occasional predator of the California
oakworm, Phryganidia californica _ 196
Parasitization of Malacosoma larvae by Tachinidae in coastal
California . 42
Larochelle, A.
Notes on the distribution and ecology of Scaphinotus bilobus .... 80
Wing dimorphism in Carabus meander ... 126
Leech, H. B.
Book Review: Monografia da tribo Ibidionini . 101
Lugaski, T. and W. H. Clark
Collection records of the Black Witch Moth, Ascalapha odorata
(Linnaeus) in Nevada . 353
Mathis, W. N.
A review of the genus Borboropsis .. 373
V
Mayo, V. K.
A new species of Baetis from Ecuador _ 285
Four new species of the genus Baetodes _ 308
McCafferty, W. P. and G. F. Edmunds, Jr.
Subgeneric classification of Ephemera _ 300
McCrea, R. J.
A new species of the flea beetles genus Chaetocnema found on
dichondra in California _ 61
Menke, A. S. and J. T. Polhemus
Lectotype designation for Gerris buenoi Kirkaldy_ 257
Miller, D. R.
Mealybugs of San Miguel Island, California _ 264
Moore, I.
A note on Dacnochilus fresnoensis _ 42
Moore, I. and E. F. Legner
Speculation on the distribution of the Southern California species
of Cafius with a new record from the Salton Sea_ 279
Muchmore, W. B.
A new genus of pseudoscorpions based upon Atemnus hirsutus .... 43
Nelson, C. H.
Synopsis of the genus Chilenoperla _ 315
Paetzel, M. M.
Behavior of the male Trypozylon rubrocinctum _ 26
Parker, F. D. and H. W. Potter
Biological notes on Lithurgus apicalis Cresson _ 294
Parson, C. T.
The Lagriidae of California _ 1
Peterman, R. M.
Possible behavioral thermoregulation in Tanarthrus salinus and
T. inyo _ 67
Piper, G. L.
New distribution and host record for the parasitoid Heteroschema
aeneiventris _ 337
Proceedings, Pacific Coast Entomological Society _ 81
VI
Rust, R. W.
Ectoparasites and nidicolous Acari of the pocket gopher,
Thomomys bottae ___ 59
Shapiro, A. M.
Recurrent aberration in Cynthia annabella a review with four
new records _ 289
Slobodchikoff, C. N.
Behavioral studies of three morphotypes of Therion circum-
flexum _ 197
Smith, D. R.
The sawfly tribe Lycaotini in South America _ 93
Soleglad, M. E.
Uroctonoides fractus a synonymy _ 60
A redescription of the scorpion Vejovis flavus _ 165
Stephen, F. M.
Ceratosmicra campoplegicis Burks, a hyperparasite of Perilitus
coccinellae (Shrank) _ 395
Thorp, R. W.
Book Review: Principles of Insect Chemosterilization _ 55
Book Review: The Moths of America North of Mexico. Fasc.
21, Sphingoidea _ 143
Book Review: Biological Control _ 176
Book Review: Insect Vision _ 184
Book Review: Australian Butterflies _ 228
Book Review: Annual Review of Ecology and Systematics_ 239
Book Review: The Insects of Australia_ 280
Book Review: American Entomologists _ 281
Book Review: Monographie der Familie Platypodidae Coleop-
tera _ 282
Book Review: Essays in Evolution and Genetics in Honor of
Theodosius Dobzhansky _ 283
Torgersen, T. R.
A new species of Spinolochus from North America_ 124
Turner, W. J. and J. G. Chillcott
Four new species of the Symphoromyia pachyceras complex
from California _ 5
Vll
Wasbauer, M. S.
Some new taxa in the Myrmosinae with keys to the females in
North America _ 325
Wasbauer, M. S. and H. B. Leech
A biological note on two species of Ageniella from California .... 182
Wheeler, G. C. and J. Wheeler
The ant larvae of the tribes Basicerotini and Dacetini: second
supplement _ 207
Ant larvae of the subfamily Dolichoderinae: second supplement396
Wood, S. F.
Recent annoyance to man in Utah by Triatoma protracta _ 183
Zoological Nomeclature _ 73, 404
ENTOMOLOGICAL EXCHANGES AND SALES
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FOR SALE. Extensive collection of mayfly reprints. Write R. Brusca,
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THE MOTHS OF AMERICA
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THE MOTHS OF AMERICA, written by authorities in each group treated, constitutes a
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In addition to the color photographs of adults, illustrations of genitalia or other structures
are included where necessary to assure accurate identification. The comprehensive text gives
a full taxonomic treatment, including synonymy, plus information on distribution, foodplants,
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THE MOTHS OF AMERICA will be published in over 100 parts, over a period of about 20
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ALREADY PUBLISHED:
Fascicle 13, Part la, Pyralidae: Scopariinae & Nymphulinae, by Dr. Eugene G. Munroe.
1972. 134 p. $20.60 ($16.50 to subscribers).
Fascicle 13, Part lb, Pyralidae: Odontiinae & Glaphyriinae, by Dr. Eugene G. Munroe.
1972. 116 p. $20.60 ($16.50 to subscribers).
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IN PREPARATION:
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Fascicle 13, Part 1c, Pyralidae: Evergestiinae, by Dr. Eugene G. Munroe (including 14
color plates for all 3 parts of Fascicle 13. 1).
Fascicle 6, Part 2, Gelechioidea: Oecophoridae, by Dr. Ronald W. Hodges.
Fascicle 22, Part 2, Lymantriidae, by Dr. Douglas C. Ferguson.
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