Vol. 46 JANUARY 1970 No. 1
THE
Pan-Pacific Entomologist
WILLIAMS—Three new species of Vejovis from Death Valley, California (Scor-
pionida: Vejovidae) _ 1
ICISTNER—Two new genera of termitophiles associated with Longipeditermes
longipes (Haviland) (Coleoptera: Staphylinidae; Isoptera: Nasutiter-
mitinae) _ 12
BAKER—Occurrence of Malacosoma incurvum discoloration in Zion Canyon,
Utah (Lepidoptera: Lasiocampidae) '_ 27
GARDINER—Immature stages and habits of Spondylis upiformis Mannerheim
(Coleoptera: Cerambycidae) _ 33
GRIGARICK AND SCHUSTER—A new genus in the tribe Euplectini (Coleop¬
tera: Pselaphidae) _._ 36
NUTTING—Free diurnal foraging by the North American nasutiform termite,
Tenuirostritermes tenuirostris (Isoptera: Termitidae) _ 39
O’BRIEN —Pselactus spadix (Herbst), a European cossonine weevil new to
California (Coleoptera: Cucurlionidae) _ 42
HALSTEAD—A new species of the genus Largus Hahn with a key to the spe¬
cies of the genus in the southwestern United States (Hemiptera: Pyr-
rhocoridae) _ 45
WASBAUER—The type material in the collection of the California Department
of Agriculture_:_.__ 47
SLOBODCHIKOFF—A revision of the genus Grotea (Hymenoptera: Ichneu-
monidae) _ 50
ZOOLOGICAL NOMENCLATURE_ 63
PUBLICATIONS OF THE SOCIETY __ 64
PROCEEDINGS _ 65
SAN FRANCISCO, CALIFORNIA • 1970
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. 46
January 1970
No. 1
Three New Species of Vejovis from Death Valley, California
(Scorpionida: Vejovidae)
Stanley C. Williams
San Francisco State College, California
During April of 1968 a one-week survey of the scorpion fauna of
Death Valley Avas made. Samples were taken along the entire length of
this desert region in as many different ecological situations as was
possible within the available time. Study of the several hundred
scorpion specimens in our samples indicated that Death Valley has a
diverse scorpion fauna. Three distinct and undescribed species be¬
longing to the genus Vejovis were found and are here described.
Vejovis gramenestris Williams, new species
(Figs. 1, 2)
Diagnosis. —Moderate to small sized species of Vejovis. Total body length
probably seldom exceeds 35 millimeters. Coloration light brownish-yellow over
most of body. Pedipalps long and slender, fixed finger longer than palm;
carapace longer than palm; keels developed on palm, medial ones being granular;
large elongate terminal tooth on distal end of fingers. First metasomal segment
broader than long; second metasomal segment as long as wide; fifth segment
longer than either carapace or fixed finger. Pectines with proximal tooth elongate
on female; 16 to 17 teeth in males, 12 to 14 teeth in females. Inferior border of
movable cheliceral finger lacking denticles. Inferior median and lateral keels of
metasoma finely serrate.
Similar to Vejovis wupatkiensis Stahnke but differing in the following
ways: smaller body size; fewer pectinal teeth; light brownish-yellow
in base color and lacking melanic markings; fifth metasomal segment
distinctly longer than movable finger of pedipalp in males and with
fifth metasomal segment approximating movable finger length in fe¬
males (in V. wupatkiensis movable finger slightly longer than metasomal
segment V in males, females with movable finger distinctly longer than
metasomal segment V).
Holotype Male. — Coloration. —Carapace, mesosoma, metasoma and walking legs
pale brownish-yellow with faint suggestion of underlying dusky pigmentation;
telson and metasomal segment V darker and more brownish than other caudal
The Pan-Pacific Entomologist 46: 1-11. January 1970
2
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
1
Figs. 1 and 2. Vejovis gramenestris Williams, holotype male. Fig. 1. Dorsal
view. Fig. 2. Ventral view.
segments; pedipalp fingers slightly more brownish than palm; pedipalps whitish;
pretarsal claws whitish; cheliceral teeth light tan.
Carapace .—Anterior margin with six erect hairs and with median emargination;
median eyes less than % carapace length at that point; carapace surface irregular
and granular.
Mesosoma .—All tergites granular; tergite 7 with two pairs of dentate lateral
keels, posterior granule of each keel largest. Sternites agranular; last sternite
with one pair of smooth to crenulate lateral keels.
Metasoma .—Dorsal keels of segments I to IV serrate, last serration of each keel
largest; dorsolateral keels serrate on I to IV, last serration of each keel largest on
segments I to III, keel composed of more irregular and broad granules on V;
lateral keels serrate and almost complete on I, serrate on posterior % of II,
serrate on posterior % of III, absent on IV, crenulate on anterior % of V; inferior
JANUARY 1970] WILLIAMS-NEW CALIFORNIA SCORPIONS
3
Table 1 . Measurements (in millimeters) of Vejovis gramenestris
Williams, new species, holotype and allotype.
Holotype
(male)
Allotype
(female)
Total length
24.6
32.8
Carapace, length
3.0
3.7
width (at median eyes)
2.2
2.8
Mesosoma, length
7.2
12.3
Metasoma, length
11.4
13.1
segment I (length/width)
1.6/1.8
1.8/2.3
segment II (length/width)
1.8/1.8
2.1/2.2
segment III (length/width)
1.9/1.7
2.3/2.1
segment IV (length/width)
2.6/1.8
2.9/2.1
segment V (length/width)
3.5/1.7
4.0/2.1
Telson, length
3.0
3.7
Vesicle (length/width)
2.1/1.2
2.4/1.6
depth
0.9
1.2
Aculeus, length
0.9
1.3
Pe dip alp
Humerus (length/width)
2.6/0.7
3.4/1.0
Brachium (length/width)
2.9/0.9
3.7/1.3
Chela (length/width)
4.5/1.0
5.9/1.2
depth
1.0
1.3
movable finger, length
2.9
4.0
fixed finger, length
2.5
3.5
Pectines
teeth (left/right)
17/17
14/14
middle lamellae
10
8
lateral keels crenulate on segments I to IV, serrate on V; inferior median keels
smooth to crenulate on I, crenulate on II-IV, serrate on V. Each inferior median
keel on segments I to IV set with three bristles. All intercarinal spaces of segment
V granular.
Telson .—Ventral surface with eight pairs of long reddish hairs, most of these
approximate aculeus in length; vesicle quite smooth, agranular; aculeus with
pronounced curvature; broad subaculear tubercule.
Genital Operculum .—Completely divided longitudinally; set with about eight
reddish hairs; genital papillae visible externally.
Chelicerae .—Inferior border of movable finger smooth, completely lacking
denticles.
Pedipalps .—Palm only slightly swollen; keels distinct. Internal margin of fingers
not scalloped; terminal tooth of each finger distinctively elongate and overhang
each other when fingers closed.
Standard Measurements .—Table 1.
Allotype Female.— Morphologically similar to holotype with the following
exceptions: considerably larger in all body proportions as well as overall length;
4
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
pectines with fewer teeth (14 instead of 17) ; middle lamellae fewer (8 instead of
10) ; proximal pectinal tooth somewhat elongate; no genital papillae; genital
operculum not completely divided longitudinally.
Standard Measurements .—Table 1.
Study of 20 paratopotypes indicated little variation from the de¬
scriptions of holotype and allotype. Two males were juveniles with
carapace lengths of 2.4 millimeters while 18 females varied in carapace
length from 2.7 to 3.8 millimeters (predominantly 3.6 millimeters).
Pectine tooth count varied from 12 to 14 for the females while both
males had 16 pectinal teeth. Several specimens had small “L-shaped”
dark markings at the anterolateral corner of the carapace.
Holotype male , allotype and 20 paratopotypes were collected in
Death Valley at Travertine Spring (% mile east of Furnace
Creek Inn), Inyo County, California, 11 April 1968 by S. C. Wil¬
liams, V. F. Lee, and R. Lewert. Holotype and allotype are permanently
deposited in the California Academy of Sciences.
This species is named “gramenestris” because of its grass dwelling
habits. It was abundantly found living in the grasses adjacent to
springs. At night it was found climbing the grass stems apparently in
search of food.
Eleven additional paratypes were studied from the following locations
in Death Valley, Inyo County, California: Grapevine Spring (4 miles
east of Ubehebe Crater), 12 April 1968 (S. C. Williams, V. F. Lee,
J. Bigelow), 1 female; 20 Mule Team Canyon, 14 April 1968 (M. A.
Cazier, J. Bigelow) ,8 males, 2 females.
This very small species of Vejovis was found only around permanent
springs where it was under travertine rocks during the day and climbing
in the grasses at night. This is a very active species and is capable of
rapid locomotion for extended periods of time. While collecting sam¬
ples two members of our party were stung by this species on the fingers.
The sting was characterized by a sharp burning sensation which lasted
for five to ten minutes. No swelling, no reddening, and no urticaria
accompanied the sting. Most specimens were collected by the ultra¬
violet detection method, but several were found by rock turning.
Vejovis deserticola Williams, new species
(Figs. 3, 4)
Diagnosis.— Moderate sized species of Vejovis. General body color brownish-
amber with legs yellowish, pectines almost white and pedipalp chela reddish.
Inferior median keels of metasoma all present and crenulate or serrate; inferior
lateral keels all serrate. Movable finger of pedipalp distinctly longer than carapace,
palm or metasomal segment V. Male with 21 or 22 pectinal teeth, female with 20
JANUARY 1970] WILLIAMS—NEW CALIFORNIA SCORPIONS
5
Figs. 3 and 4. Vejovis deserticola Williams, holotype male. Fig. 3. Dorsal
view. Fig. 4. Ventral view.
teeth. Proximal tooth of female pectine longest. Movable finger of chelicerae with
inferior border lacking denticles.
Closest known relative is Vejovis wupatkiensis Stahnke. Distinguished
from V. wupatkiensis by the following characters: pectinal teeth greater
in number for both sexes; vesicle densely covered by coarse granules
6
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
on ventral and lateral surfaces; much larger species in all body propor¬
tions.
Holotype Male. — Coloration .—Base color of prosoma and mesosoma brownish-
amber; metasoma similar but more yellowish-amber; walking legs brownish-
yellow; pedipalp hands reddish-amber; pectines almost white; carapace with
faint suggestion of underlying dusky markings. Eyes black, teeth of chelicerae,
and most keel serrations reddish; aculeus reddish-brown; pretarsal claws yel¬
lowish-brown; fingers of pedipalp reddish. Keels on metasoma and pedipalps
appear contrastingly reddish-brown due to heavy development rather than to
underlying pigmentation.
Carapace .—Anterior margin with distinct broad median emargination, set with
6 bristles. Lateral eyes three per group, most anterior distinctly largest, most
posterior one greatly reduced in size; median eye diad distinctly less than 14
carapace width at that point. Carapace surface rough and uneven, covered by large
coarse granules.
Mesosoma. —Tergites densely covered by large granules, tergite 7 with two pair
of well developed conspicuous lateral keels, these serrate. Stemites agranular;
surface texture uneven; one pair serrate keels on last sternite; lateral margin of
sternites with serrate granules.
Metasoma. —Dorsal and dorsolateral keels well developed and regularly serrate
on segments I to IV; dorsolateral keels of segment V distinct, broad and more
granular than serrate. Lateral keels serrate along almost entire length of I,
posterior 14 of II and III, absent on IV, present and serrate on anterior % of V.
Inferior median keels well developed on all segments; crenulate on I to III;
crenulate to serrate on IV; serrate on V. Inferior lateral keels well developed and
regularly serrate on all segments. All inferior intercarinal spaces smooth, agranular.
Telson. —Vesicle globular and densely covered with large granules from ventral
and lateral aspect. Heavy granulation also occurs in lateral regions of dorsal
aspect of vesicle. Subaculear tubercule large, distinct. Aculeus short and well
curved.
Genital Operculum. —Completely divided longitudinally; large distinct genital
papillae visible externally. Genital operculum with six reddish bristles.
Chelicerae .—Inferior border of movable finger smooth, completely lacking den¬
ticles.
Pedipalps. —Palm only slightly swollen, keels well developed, granular. Fixed
finger same length as carapace, movable finger distinctly longer than carapace.
Caudal segment V longer than fixed finger but shorter than movable finger. In¬
ternal margin of fingers not scalloped; distal tooth of each finger greatly elongate
and overlap when fingers closed.
Standard Measurements. —Table 2.
Allotype Female.— Morphologically the same as holotype with the following
exceptions: pectinal teeth fewer (20/20 instead of 22/21) ; pectinal teeth dis¬
tinctly shorter; proximal pectinal tooth distinctly longer than others; no genital
papillae; genital operculum not completely divided longitudinally; slightly longer
total body size.
Standard Measurements. —Table 2.
The holotype male and allotype were collected at Travertine Spring,
V 2 MILE EAST OF FURNACE CREEK INN, DEATH VALLEY NATIONAL MON-
JANUARY 1970] WILLIAMS—NEW CALIFORNIA SCORPIONS
7
Table 2. Measurements (in millimeters) of Vejovis deserlicolci
Williams, new species, holotype and allotype.
Holotype
(male)
Allotype
(female)
Total length
42.6
45.0
Carapace, length
5.2
5.2
width (at median eyes)
3.9
4.2
Mesosoma, length
11.2
13.7
Metasoma, length
20.6
19.8
segment I (length/width)
2.9/3.2
2.8/3.3
segment II (length/width)
3.3/3.2
3.1/3.2
segment III (length/width)
3.5/3.2
3.3/3.2
segment IV (length/width)
4.7/3.1
4.4/3.1
segment V (length/width)
6.2/3.0
6.2/3.1
Telson, length
5.6
6.3
Vesicle (length/width)
3.9/2.8
4.2/2.S
depth
2.2
2.2
Aculeus, length
1.7
2.1
Pedipalp
Humerus (length/width)
5.5/1.3
5.5/1.4
Brachium (length/width)
5.9/1.6
5.9/1.7
Chela (length/width)
9.2/2.3
9.3/2.2
depth
2.7
2.7
movable finger, length
6.5
6.6
fixed finger, length
5.2
5.4
Pectines
teeth (left/right)
22/21
20/20
middle lamellae
14
14
ument, Inyo County, California, 11 April 1968, by S. C. Williams
and V. F. Lee. Both specimens were collected on a rocky slope at night
by means of ultraviolet detection. The holotype is permanently de¬
posited in the California Academy of Sciences.
This species is named “deserticola” because its only known habitat
is Death Valley, one of the harshest deserts in North America.
Vejovis shulovi Williams, new species
(Figs. 5, 6)
Diagnosis.— Moderate sized species of Vejovis in the Paruroctonus subgenus.
Base color pale yellowish with underlying dusky markings on carapace and dorsum
of mesosoma; mature females with contrasting reddish pedipalp fingers. Short
swollen pedipalp hand with fixed finger shorter than palm, carapace or caudal
segment IV; palm heavily keeled. All metasomal keels present, inferior keels
mostly crenate or serrate. Chelicerae with three small denticles on inferior border
8
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Figs. 5 and 6. Vejovis shulovi Williams, holotype female. Fig. 5. Dorsal
view. Fig. 6. Ventral view.
of movable clieliceral finger, the two terminal teeth on this finger subequal.
Pectinal teeth 19 to 20 in males, 13 to 15 in females; teeth borne only on distal %
of comb in females.
Appears related to Vejovis bantai Gertsch and Soleglad but distin¬
guishable by the following characters: pectines with fewer teeth, fe¬
males 13 to 15 (not 17), males 19 to 20 (not 23) ; fewer middle lamel¬
lae; inferior median keels not essentially obsolete on metasomal segments
I and II; inferior median keels of metasomal segments I to IV with
3,4,4,5 pairs of reddish hairs (not 3,3,3,3 pairs) respectively; pedipalp
hand not nearly as wide as long.
JANUARY 1970] WILLIAMS-NEW CALIFORNIA SCORPIONS
9
Holotype Female. — Coloration. —Base color of cuticle pale yellow; carapace
with underlying dusky pigmentation; mesosoma with dusky underlying pigmenta¬
tion; walking legs with faint regional dusky markings; pedipalp fingers con¬
trastingly light reddish; pectines whitish; most of cuticle with polished shiny
appearance.
Carapace. —Anterior margin subtly convex, set with six bristles; median eyes
more than % but less than % carapace width at that point; carapace surface
irregular and set with coarse granules.
Mesosoma. —Tergites with relatively few granules, these small; tergite 7 with
poorly developed median keel and two pair of dentate lateral keels; sternites
agranular.
Metasoma. —Dorsal and dorsolateral keels crenulate to serrate, never ending
in enlarged tooth; lateral keels crenulate to dentate on most of segment I,
dentate on posterior % of II and III, absent on IV; irregularly crenulate on
anterior % of V. Inferior lateral keels smooth to crenulate on segments I to III;
crenulate to serrate on IV; serrate on V. Inferior median keels basically smooth
on segments I to III; crenulate to serrate on IV; serrate on V. Inferior median
keels of segments I to IV with 3, 4, 4, 5 pairs of stout reddish bristles. Inferior
intercarinal space of segment V abundantly granular.
Telson. —Ventral side basically agranular, set with about 10 pairs of moderately
long reddish hairs; aculeus long, slender; subtle broad subaculear tubercule.
Pectines. —Inferior surface densely hirsute; each fulcrum with about six reddish
hairs; each middle lamella with about four reddish hairs; 14 teeth; 12 middle
lamellae; proximal % of pectine without teeth.
Genital Operculum. —Not completely divided longitudinally; with about 15 long
reddish hairs.
Chelicerae. —Inferior border of movable finger with three small denticles;
terminal tooth on superior border smaller but still subequal.
Pedipalps. —Hand swollen, all keels distinct and granular. Both movable and
fixed finger shorter than carapace; palm longer than fixed finger but shorter
than movable finger; fixed finger shorter than metasomal segment IV. Internal
margin of fingers moderately scalloped proximally; proximal teeth do not meet
when fingers closed.
Standard Measurements.- —Table 3.
Allotype Male. —Color and morphology essentially the same as holotype with
the following exceptions: smaller total length; carapace narrower; pectines with
more teeth (19 instead of 14) ; middle lamellae more numerous (14 instead of 12) ;
genital papillae showing externally; pedipalp fingers not reddish; pedipalp keels
not as conspicuous.
Standard Measurements. —Table 3.
Study of 11 paratopotypes indicated little variation from the de¬
scriptions of the holotype and allotype. The two males had carapace
lengths of 2.6 and 3.8 millimeters and pectine tooth counts of 19/20.
Although the genital papillae were visible externally, the smaller body
sizes indicated these specimens may have been sub-adult. The nine
females varied in carapace length from 4.1 to 5.3 millimeters. Pectinal
tooth counts varied from 13 to 15 (predominantly 13 and 14). The
10
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Table 3. Measurements (in millimeters) of Vejovis shulovi Wil¬
liams, new species, holotype and allotype.
Holotype
(female)
Allotype
(male)
Total length
38.3
34.4
Carapace, length
4.9
4.4
width (at median eyes)
3.5
3.2
Mesosoma, length
11.3
11.1
Metasoma, length
16.8
14.5
segment I (length/width)
2.2/2.4
2.0/2.1
segment II (length/width)
2.7/2.3
2.3/2.0
segment II (length/width)
2.8/2.1
2.4/1.9
segment IV (length/width)
3.7/2.0
3.2/1.7
segment V (length/width)
5.4/2.0
4.6/1.7
Telson, length
5.3
4,4
Vesicle (length/width)
3.2/2.0
2.6/1.6
depth
1.7
1.3
Aculeus, length
2.1
1.8
Pedipalp
Humerus (length/width)
3.7/1.3
3.4/1.2
Brachium (length/width)
4.1/1.8
3.5/1.6
Chela (length/width)
7.1/2.4
5.9/1.8
depth
2.6
1.9
movable finger, length
4.4
3.2
fixed finger, length
3.1
2.3
Pectin es
teeth (left/right)
14/14
19/19
middle lamellae
12
14
female with carapace length of 4.1 millimeters was probably a sub-adult
judging by the less swollen pedipalp hands and lack of red coloration
of the fingers.
The holotype female , allotype and 11 paratopotypes were collected in
Death Valley at Grapevine Spring (4 miles east of Ubehebe
Crater) , Inyo County, California, 12 April 1968 by S. C. Williams,
V. F. Lee and J. Bigelow. All specimens were collected by ultraviolet
detection in early evening. The holotype and allotype are permanently
deposited in the California Academy of Sciences.
This species is named “shulovi” in honor of Professor Aaron Shulov
of the Hebrew University in Jerusalem, Israel. Professor Shulov has
made many fine contributions to the understanding of basic scorpion
biology and venoms. His numerous scientific papers have been a source
of great stimulation to students of scorpions all over the world.
JANUARY 1970] WILLIAMS—NEW CALIFORNIA SCORPIONS
11
In addition to the holotype, allotype and 11 paratopotypes, an addi¬
tional 20 paratypes were available for study. These were collected in
the following locality in Death Valley, Inyo County, California: Scot¬
ty’s Ranch, elevation 3000 feet, 13 April 1968 (M. A. Cazier, B. Nevelyn,
J. Bigelow, G. Lytle), 20 females.
This species is known only from the north end of Death Valley,
Inyo County, California. It was found in two localities, and was not
abundant. The females greatly outnumbered the relatively rare males in
our samples. All of the specimens appeared to be mature or nearly so,
younger juveniles were completely absent. A female was selected as
the holotype of this species for several reasons. Males were very rare in
the samples available, and there was some question whether or not
these males were sexually mature. Also, the females appear to have
species characteristics as definitive or more so than the males of this
species.
The habitat in which this species was found was characterized by
fine sedimentary soil, abundantly covered with dark volcanic surface
rock. The area was hot and arid, but a small spring supplied surface
water which formed a small stream which flowed into the desert for
some % of a mile. The vegetation, except adjacent to the stream, was
very sparse.
Acknowledgments. —Much appreciation is due to Vincent F. Lee
for assistance in the field and to Charlene F. Williams for clerical
assistance. This work was partially supported by research grant GB
7679 from the National Science Foundation and by a Faculty Research
Leave from San Francisco State College.
12
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Two New Genera of Termitopliiles Associated with
Longipeditermes longipes (Haviland ) 1
(Coleoptera: Staphylinidae; Isoptera: Nasutitermitinae)
David H. Kistner
Chico State College, California
During his Oriental field trip in 1963, Professor Alfred E. Emerson,
University of Chicago, collected many termitophiles. Among the most
spectacular of these was a series taken in nests of Longipeditermes longi¬
pes (Haviland) from which termite genus, no previous termitophiles
had ever been taken. Needless to say, all of the genera and species col¬
lected are new, some even representing new termitophilous subtribes.
The genera and species are not only remarkable from the point of view
of termitophilous adaptations but are also remarkable because of the
number of species and genera involved. I am therefore beginning a
series of papers in which I shall describe the fauna associated with
this interesting termite. I estimate that at least 3 papers will be devoted
to the description of new forms and then the 4th and last will be devoted
to the general relationships shown by this diverse termitophile fauna.
I wish to thank Dr. Emerson for providing the specimens which form
the basis of this paper, for determining the host termites, and for reading
and commenting on this paper. All types of the species described
herein are in the collection of the author. Specimens of the host termites
are in the Emerson Collection of the American Museum of Natural His¬
tory, New York, and representative series are in the collection of the
author. All measurements are in mm and techniques involved in the
study of the specimens are given by Kistner (1968).
I wish to thank Mr. David Harwood, Miss Lynette Hawver, Mr.
Herbert Jacobson, Mr. Matthew Rees, Miss Lynn Royce, and Miss
Virginia Sleppy, all of Chico State College, for technical assistance.
All of the species herein described belong to the subtribe Termito-
zyrina, tribe Myrmedoniini, subfamily Aleocharinae.
Genus Longipedoxenus Kistner, new genus
Most closely related to Termitosymbia Seevers from which it is dis¬
tinguished by its overall appearance, the presence of only 1 pair of
paratergites on abdominal segments III—VII and the lack of a vertexal
arcade.
1 This study was supported in part by the National Science Foundation (Grant No. GB-6284).
The Pan-Pacific Entomologist 46: 12-27. January 1970
JANUARY 1970] ICISTNER-NEW TERMITOPHILES
13
Figs. 1-2. Longipedoxenus shinneri . Fig. 1. Dorsal view. Fig. 2. Lateral
view.
Overall shape as in Figs. 1, 2, and 3. Head shaped as in Fig. 1. Dorsal sculpture
consists of a depression in middle of vertex. Clypeus membranous. Antennae
inserted in dorsal sockets at inner margin of eyes with no vertexal arcade apparent.
Ventral surface of head capsule without distinction; with no infraorbital carinae.
Gula relatively short; narrowest anteriorly, becoming wider posteriorly. Mentum
almost as long as gula, separately articulated, somewhat membranous. Antennae
11 -segmented, elongate, shaped as in Fig. 13. Labrum shaped as in Fig. 7.
Mandibles nearly symmetrical, shaped as in Figs. 4 and 6. Maxillae shaped as in
Fig. 8 with four-segmented palpi much longer than lacinia and galea. Labium
shaped as in Fig. 5; palpi 3-segmented. Both maxillae and labium somewhat
membranous.
Pronotum shaped as in Fig. 1. Details of pronotal sculpture (partially visible
in Fig. 1) consist of median ridge starting at median point of anterior border and
14
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Fig. 3. Longipedoxenus shinneri. Lateral view showing abdomen in an upright
position.
a longitudinal indentation to each side of this. Prosternum acarinate, relatively
small, leaving a large amount of exposed membrane. Procoxal cavities closed
behind by membrane. Mesothoracic peritremes small, completely surrounded by
membrane, forming a strap-like process from sides of hypomera to median line.
Metasternum about twice length of mesosternum. Mesocoxal cavities in a strict
sense rather widely separated, but indentations of meso- and metasternum into
which the large coxae fit, fall rather closely together at their medial borders.
Mesosternal intercoxal process broad, blunt, acarinate. Elytra shaped as in Fig. 1,
with some median posterior indentations and sinuate lateral borders. Wings
present, of usual staphylinid venation. Legs elongate. Pro-, meso-, and metalegs
shaped as in Figs. 10, 11, and 9 respectively. Tarsal formula 4-5-5.
Abdomen both physogastric and scaphoidal and can be held partially recurved
over back or straight out; shaped as in Figs. 1, 2, and 3. Abdominal segment I
membranous. Abdominal segment II represented by tergite alone. Abdominal
segments III-VII represented by tergite, sternite, and but 1 pair of paratergites
each. Abdominal segment VIII represented by tergite and sternite only. Segment
IX trivalved, shaped as in Fig. 12. Male genitalia bulbous; specific shape variable
by species. Spermatheca present; shape variable by species.
4
Figs. 4-8. Longipedoxenus shinneri. Fig. 4. Right mandible. Fig. 5. La¬
bium. Fig. 6. Left mandible. Fig. 7. Labrum. Fig. 8. Maxilla.
JANUARY 1970]
ICISTNER-NEW TERMITOPHILES
15
16
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Figs. 9-11. Longipedoxenus shinneri. Fig. 9. Metaleg. Fig. 10. Proleg. Fig.
11. Mesoleg.
The abdomen is both physogastric and scapboidal at the same time.
The physogastry is achieved by the enlargement of membranes between
the abdominal segments, particularly those at the anterior of the abdo¬
men. Scaphoidal shape is achieved by broadening the tergites and
JANUARY 1970]
ICISTNER-NEW TERMITOPHILES
17
Figs. 12-13. Longipedoxenus shinneri. Fig. 12. Abdominal segment IX. Fig.
13. Antenna.
sternites, particularly those at the posterior of the abdomen. The accen¬
tuated folds at the anterior borders of tergite IV probably have some¬
thing to do with the glandular system of the abdomen. Evidence from
the sclerotization of abdominal tergites VI and VII indicates 2 small
defense gland reservoirs in the posterior of segment VI.
Type species. — Longipedoxenus shinneri Kistner.
Key to Species
Hairy species, abdominal sternites III-IV with 3^1 rows of long black curly
setae ___ L. entersoni n. sp.
Slightly hirsute species, abdominal sternites III-V with a sparse apical row
of short black setae and an anteapical median group of 2-3 short black
setae -- L. shinneri n. sp.
Longipedoxenus emersoni Kistner, new species
(Figs. 18, 23)
Related to L. shinneri from which it is distinguished by its smaller
size, the differences in chaetotaxy, as well as the shape of the male
genitalia and the spermatheca.
18
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Figs. 14-24. Male genitalia. Figs. 14-16, 22, 23. Median lobes of male geni¬
talia. Fig. 14. Longipedisymbia bouceki. Fig. 15. Longipedoxenus shinneri.
Fig. 16. Longipedisymbia carlislei. Fig. 22. Longipedisymbia bradburni. Fig.
23. Longipedoxenus emersoni. Fig. 24. Lateral lobe of male genitalia, L. shinneri.
Figs. 17-21. Spermatheca. Fig. 17. L. shinneri. Fig. 18. L. emersoni. Fig. 19.
L. carlislei. Fig. 20. L. bouceki. Fig. 21. L. bradburni. Scale represents 0.25
mm and applies to all figures.
Color yellowish-brown throughout except for black eyes, and white membranes
between abdominal sclerites. Chaetotaxy of abdominal tergites II—VIII as follows:
0, 0, 2,2,2,0,2-6. The 2 on tergites IV-VI barely visible; the 2 on tergite VIII are
anterior to row of 6. Chaetotaxy of abdominal sternites III—VIII as follows:
III-V with 3 to 4 rows of long black curly setae; VI with apical row of long
curly setae and anteapical group of 2-3; VII, 4; VIII, 6. Median lobe of male
JANUARY 1970]
KISTNER—NEW TERMITOPHILES
19
genitalia shaped as in Fig. 23. Lateral lobe of male genitalia shaped as in L.
shinneri. Spermatheca shaped as in Fig. 18.
Measurements .—Pronotum length, 0.33-0.35; head length, 0.30-0.32 (measured
from the edge of the vertex to the nuchal ridge). Number measured, 5.
Holotype male No. 13520, Malaya, Sungei Buloh Forest Reserve,
3° 10' N., 101° 34' E., altitude 150 feet, 18 March 1963, Coll. A. and
Eleanor Emerson, C. M. Low, and R. D. Menon, in vicinity of queen.
Paratypes. —4, same data as the holotype.
The species is named for one of the collectors, Dr. A. E. Emerson.
Longipedoxenus shinneri Kistner, new species
(Figs. 1-13, 15, 17, 24)
Related to L. emersoni from which it is distinguished by its larger
size, differences in the chaetotaxy, as well as the shape of the male
genitalia and the spermatheca.
Color yellowish-brown throughout except for black eyes, and white membranes
between abdominal sclerites. Chaetotaxy of abdominal tergites II—VIII as follows:
II-VII, 0; VIII, 6 apical, 2 anteapical. Chaetotaxy of abdominal sternites III—
VIII as follows: III—VIII all with a sparse apical row of short black setae, III-VI with
anteapical median group of 2-3 short black setae additionally. Median lobe of male
genitalia shaped as in Fig. 15. Lateral lobe of male genitalia shaped as in Fig. 24.
Spermatheca shaped as in Fig. 17.
Measurements .—Pronotum length, 0.43-0.45; head length, 0.31-0.32 (measured
from the anterior edge of the vertex to the nuchal ridge). Number measured, 10.
Holotype male No. 13521, Malaya, Sungei Buloh Forest Reserve,
3° 1CK N., 101° 34' E., altitude 150 feet, 18 March 1963, Coll. A. and
Eleanor Emerson, C. M. Low, and R. D. Menon, in the vicinity of the
queen.
Paratypes. —4, same data as the holotype; 6, Sarawak, 1° 38' N., 113°
35' E., altitude 950 feet, 7 February 1963, Coll. A. and Eleanor Emer¬
son and W. King, in cells under log near nest, vial No. 1; 9, Sarawak,
same nest, date, and collectors as previous vial, from nest in hard log,
vial No. 2.
This species is named for the late Mr. Ernest G. Shinner of Chicago in
honor of his long interest in philanthropic causes of the Chicago area.
Genus Longipedisymbia Kistner, new genus
Closely related to both Termitosymbia and Longipedoxenus from
which it is distinguished by the peculiar lateral processes of the pro¬
notum. Also distinguished from Termitosymbia by the approximately
equal-sized paratergites. Also distinguished from Longipedoxenus by
the presence of 2 pairs of paratergites on abdominal segments IV-VI
and the lack of a membranous clypeus.
20
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Figs. 25-27. Fig. 25. Longipedisymbia bouceki, dorsal view. Fig. 26. L.
ccirlislei, dorsal view. Fig. 27. L. ccirlislei, lateral view.
Overall shape as in Figs. 25, 26, and 27. Dorsal head shape somewhat variable
by species. Basically it is subtriangular with a depression in middle of vertex
and 2 smaller depressions on sides of vertex behind antennal fossae. Protuber¬
ances from the head vary between species. Clypeus not membranous. Gula-sub-
JANUARY 1970]
KISTNEK-NEW TERMITOPIIILES
21
Figs. 28-32. Longipedisymbici bradburni. Fig. 28. Right mandible. Fig. 29.
Left mandible. Fig. 30. Labrum. Fig. 31. Maxilla. Fig. 32. Labium.
mentum developed at an angle so that anterior end is lower than posterior
edge at foramen magnum. Mentum separately articulated and about half length of
gula-submentum. Sculpture of head otherwise variable by species. Antennae
22
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
11-segmented, shaped as in Fig. 36, with antennal fossae opening dorsally near
inner comers of eyes; without arcade but with prominence medial to each
antennal insertion. Labrum shaped as in Fig. 30. Maxillae shaped as in Fig. 31,
palpi 4-segmented, with palps much longer than lacinia and galea. Mandibles
symmetrical, shaped as in Figs. 28 and 29. Labium shaped as in Fig. 32, palpi
3- segmented.
Pronotum shaped as in Fig. 25, with lateral projection from each anterior
corner. Other sculpture consists of 2 depressions, each lateral to median keel-like
structure. Prosternum less than % length of pronotum; without carina. Pro-
thoracic coxal cavities closed behind by membrane. Mesothoracic peritremes
weakly sclerotized and not developed from lateral edges of pronotum. Meta-
stemum about twice length of mesosternum. Mesocoxal cavities widely separated
by broad, blunt, acarinate mesothoracic intercoxal process. Elytra shape variable
by species; what varies appears to be only the process at posterior lateral corner
which can be virtually absent or extremely well-developed. Wings present and
with usual staphylinid venation. Legs extremely long with elongate coxae. Pro-,
meso-, and metalegs shaped as in Figs. 37, 33, and 34 respectively; tarsal formula
4- 5-5.
Abdomen physogastric and held in extended position (at least in all of the
dead specimens) ; shaped as in Figs. 26 and 27. Abdominal segment I membra¬
nous. Abdominal segment II represented by tergite alone. Segment III repre¬
sented by tergite and sternite alone (no paratergites). Segments IV-VI
represented by tergite and sternite and 2 pairs of paratergites each. Segment VII
represented by tergite, sternite, and 1 pair of paratergites, shaped as in Fig. 40.
In one species anterior margin of the paratergites notched which may indicate
paratergites fused, and in 2 other species paratergites actually double. Segment
VIII represented by tergite and sternite only; shaped as in Fig. 39. Segment IX
trivalved and shaped as in Fig. 38. Male genitalia bulbous, specific shape variable
by species. Spermatheca present; shape variable by species.
Physogastry of the abdomen is achieved by the expansion of the
membranes. Two grooves are present in the chitin at the anterior
corners of tergite IV. These probably represent egress points for some
of the abdominal glandular system. From the sclerotization of tergites
VI and VII, there would appear to be a small median reservoir for the
defense gland.
Type species. —Longipedisymbia carlislei Kistner.
Key to Species
1. Head with well-developed angular projections from lateral border behind
eyes (Fig. 25) _ 2
Head without well-developed angular projections from lateral border be¬
hind eyes - L. bradburni n. sp.
Figs. 33-36. Longipedisymbia bradburni. Fig. 33. Mesoleg. Fig. 34. Metaleg.
Fig. 35. Lateral lobe of the male genitalia. Fig. 36. Antenna.
JANUARY 1970]
KISTNER—NEW TEKMITOPHILES
23
24
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
2. Elytra with a long process from lateral posterior corner; sternites IV-VI
with but an apical and an anteapical row of short black setae _
_ L. bouceki n. sp.
Elytra with but a small bump at lateral posterior corners; sternites IV-VI
with many long, black, curly setae which are not arranged in definite
rows _ L. carlislei n. sp.
Longipedisymbia bouceki Kistner, new species
(Figs. 14, 20, 25)
Distinguished from L. carlislei to which it is most closely related by
the shape of the elytra and the chaetotaxy of the abdomen as well as the
shape of the male genitalia and spermatheca.
Color light reddish-brown throughout except for the white abdominal membranes.
Head with definite projections from anterior border of vertex median to antennal
fossae and with well-developed angular projections from lateral border behind eyes
(Fig. 25). Elytra with well-developed projections from posterior lateral borders
(Fig. 25). Abdominal segment VII with 2 distinct pairs of paratergites. Chaetotaxy
of abdominal tergites II-VIII as follows: 0, 6, 16, 14, 8, 4-0, 4-4. Sternite III with
apical row of short black setae and anteapical cluster of 4^6 setae in center.
Sternites IV-VI with apical and anteapical row of short black setae. Sternites
VII and VIII with apical and anteapical row of setae slightly longer than those
of preceding segments. Outer paratergites on segments IV-VI with 1 black
seta each. Median lobe of male genitalia shaped as in Fig. 14. Lateral lobes of
male genitalia shaped as L. bradburni (Fig. 35). Spermatheca shaped as in
Fig. 20.
Measurements .—Pronotum length, 0.65-0.67; head length, 0.31-0.32; elytra
length, 0.45-0.47. Number measured, 6.
Holotype male No. 13522, Malaya, Sungei Buloh Forest Reserve,
3° 10' N., 101° 34' E., 18 March 1963, Coll. A. and Eleanor Emerson,
C. M. Low, and R. D. Men on, altitude 150 feet, near queen. In collection
of D. H. Kistner.
Paratypes. —5, same data as holotype, (D.K.).
This species is named for Mr. George Boucek, Secretary of the
Shinner Foundation, Chicago.
Longipedisymbia bradburni Kistner, new species
(Figs. 21, 22, 28-40)
Distinguished from L. bouceki to which it is most closely related by
the lack of projections from the lateral border of the head behind the
eyes, its slenderer and smaller form, the abdominal chaetotaxy, as well
as the shape of the male genitalia and the spermatheca.
Color reddish-brown throughout except for white abdominal membranes. Pro¬
notum and appendages somewhat lighter in color than head, elytra, and abdomen.
JANUARY 1970]
KISTNER-NEW TERMITOPHILES
25
Figs. 37-40. Longipedisymbia bradburni. Fig. 37. Proleg. Fig. 38. Abdom¬
inal segment IX of female. Fig. 39. Abdominal segment VIII. Fig. 40. Abdom¬
inal segment VII.
Head with vertex squared behind eyes; without definite projections median to
antennal fossae. Elytra with well-developed projections from posterior lateral
borders. Abdominal segment VII with 2 distinct, although thin pairs of paratergites.
Chaetotaxy of abdominal tergites II-VIII as follows: 0, 2, 4, 4, 4, 0, 4; those on
tergites III-VI very tiny. Chaetotaxy of abdominal sternites III—VIII: 2-4, 4-6,
2-6, 0-6, 6, 4. Those mentioned first on sternites III-VI are anteapical, those
26
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
mentioned second are apical. Median lobe of male genitalia shaped as in Fig. 22.
Lateral lobe of male genitalia shaped as in Fig. 35. Spermatheca shaped as in
Fig. 21.
Measurements .—Pronotum length, 0.56-0.62; head length, 0.32-0.35; elytra
length, 0.40-0.42. Number measured, 5.
Holotype male No. 13523, Malaya, Sungei Buloit Forest Reserve,
3° 10' N., 101° 34' E., 18 March 1963, Coll. A. and Eleanor Emerson,
C. M. Low, and R. D. Menon, altitude 150 feet, near queen.
Paratypes.- —2, same data as the holotype; 4, Sarawak, 1° 38' N., 113°
35' E., 7 February 1963, altitude 950 feet, Coll. A. and Eleanor Emerson
and W. King, vial No. 2, ex nest in hard log.
This species is named for Mr. Robert Bradburn of Chicago, Illinois.
Longipedisymbia Carlisle! Kistner, new species
(Figs. 16, 19, 26-27)
Distinguished from L. bouceki to which it is most closely related by
its larger size, the lack of very long processes from the lateral posterior
corners of the elytra, the abdominal chaetotaxy, as well as the shape
of the male genitalia and the spermatheca.
Color reddish-brown throughout except for white abdominal membranes.
Abdominal sternites III-V and appendages are somewhat lighter in color than
rest of body. Head with vertex bearing definite projections from lateral borders
behind eyes. Vertex also with squared-off projections from anterior border of head
medial to antennal fossae. Elytra with scarcely any projections from posterior
lateral corners (Fig. 26). Abdominal segment VII with 2 paratergites fused
together but having indentation at anterior border (Fig. 27). Chaetotaxy of
abdominal tergites II-VIII as follows: 0, 6, 16, 12, 10, 0, 0-8. Setae of tergites
III-VI extremely small and quite probably give inaccurate counts as it is not
always possible to see the socket if a seta is missing. Sternites III-VI have many
setae long, curly, not arranged in definite rows, the setae 4 or more deep from
anterior to posterior. Sternite VII with apical and anteapical row of setae.
Sternite VIII with 4 setae. Median lobe of male genitalia shaped as in Fig. 16.
Lateral lobe shaped as in L. bradburni. Spermatheca shaped as in Fig. 19.
Measurements .—Pronotum length, 0.70-0.75; head length, 0.45-0.47; elytra
length, 0.50-0.55. Number measured, 5.
Holotype male No. 13525, Malaya, Sungei Buloh Forest Reserve,
3° 10' N., 101° 34' E., altitude 150 feet, 18 March 1963, Coll. A. and
Eleanor Emerson, C. M. Low, and R. D. Menon, near queen.
Paratypes. —5, same data as the holotype; 1, Sarawak, 1° 38' N., 113°
35' E., altitude 950 feet, 7 February 1963, Coll. A. and Eleanor Emerson
and W. King, vial No. 1, in cells under log near nest.
This species is named for Dr. W. T. Carlisle of Chicago.
JANUARY 1970 ]
BAKER—MALACOSOMA ECOLOGY
27
Literature Cited
Kistner, D. H. 1968. A taxonomic revision of the termitophilous tribe Termi-
topaedini, with notes on behavior, systematics, and post-imaginal
growth (Coleoptera: Staphylinidae). Misc. Publ. Entomol. Soc. Amer.,
6(3): 141-196.
Occurrence of Malacosomci incurvum discoloratum
in Zion Canyon, Utah
(Lepidoptera: Lasiocampidae)
B. H. Baker
U. S. Forest Service, Intermountain Region, Ogden, Utah 1
The tent caterpillar, Malacosoma incurvum discoloratum (Neumoe-
gen), feeds on a wide variety of plant hosts (Stehr and Cook, 1968).
These authors proposed that the common name of M. incurvum be the
“Southwestern tent caterpillar” due to its range in southwestern North
America. Infestations have been reported on Fremont cottonwood, Popu-
lus fremontii S. Watson in Zion National Park, Utah and in the lower
reaches of the Virgin River as early as 1923. Unlike many tent cater¬
pillar populations, this one has not reached extremely low levels between
population peaks. When endemic levels have been reached in the Zion
population, noticeable centers of insect activity have persisted. Addi¬
tionally, the highly concentrated public use in the Park has caused
interest in population levels that might elsewhere receive little notice.
Egg masses of the insect have been identified on Populus fremontii
S. Wats., P. alba L., P. augustifolia James, Salix sp., and apricot,
(Prunus sp.). Previous synonymy included M. fragilis mus form dis-
colorata (Neumoegen) ; M. fragilis incurva var. discolorata (Neumoe-
gen) ; M. fragilis incurva (Henry Edwards), sensu Dyar; Clisiocampa
fragilis (Stretch), sensu Neumoegen and Dyar; and Clisiocampa mus
var. discolorata Neumoegen (Stehr and Cook, 1968). Local reference
to the population described herein has often been in the form of the
Great Basin tent caterpillar, Malacosoma fragile (Stretch).
The purpose of this paper is to report the insect’s occurrence on
Fremont cottonwood along the Virgin River in the main canyon of Zion
Park. This cottonwood characteristically occupies alluvial stream bot¬
toms and their borders and contributes to their stability (Sudworth,
1 Branch of Insect and Disease Prevention and Control, Division of Timber Management.
The Pan-Pacific Entomologist 46: 27-33. January 1970
28
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
STAGE
JAN-
FEB.
MAR.
APRIL
MAY
JUNE
JULY —
DEC.
EGG
LARVA
PUPA
ADULT
Fig. 1. Seasonal development of Malacosoma incurvum discoloratum on Populus
fremontii in Zion Canyon, Utah.
1934). In Zion Park and in certain other high-use recreation areas in
the arid Southwest, it is aesthetically valuable as well as being the primary
source of shade. Insect collections and measurements and field obser¬
vations of the Zion population were made in recent years to determine
its characteristics and habits.
Life Stages
Although embryological development is yet to he monitored in this
population, it has been recognized that such development in the genus
Malacosoma commences immediately following egg deposition and that
young larvae are fully formed within two or three weeks (Stehr and
Cook, 1968). Nearly 10 months are passed in the egg stage each year.
Eclosion from the egg in early spring is closely associated with initial
leaf expansion of the host tree. Although there is some annual variation
in seasonal development, Figure 1 depicts the life cycle in a typical year.
Larvae, emerging from the eggs, chew through the chorion and the
Table 1 . Head capsule widths of Malacosoma incurvum discolor¬
atum larvae.
Larval instar
Number of specimens
Head capsule width (mm)
Mean Confidence interval*
1
253
0.43
0.42-0.44
2
185
0.66
0.65-0.67
3
301
1.02
1.01-1.03
4
294
1.54
1.53-1.55
5
365
2.22
2.20-2.24
6
480
3.18
3.16-3.20
* 99 % probability level.
JANUARY 1970 ]
BAKER—MALACOSOMA ECOLOGY
29
Figs. 2-5. Malacosoma incurvum discoloratum. Fig. 2. Recently emerged
larvae forming tent adjacent to whitish egg mass. Fig. 3. Well developed tent
resulting from consolidated larval activity. Fig. 4. Male adult. Fig. 5. Late-
instar larva.
spumaline, the protective adhesive layer covering the mass. Most egg
masses in the same general vicinity hatch within a few days. First-instar
larvae then begin building small silken tents on the host plant (Fig. 2).
Tents are frequently abandoned and new ones established elsewhere.
Colonies originating from individual egg masses become consolidated
into larger tents as the larvae increase in size (Fig. 3).
To determine the number of instars, 1,878 larvae were collected at
intervals during two field seasons and their head capsules measured
(Table 1). These data and the resulting frequency histogram (Fig. 6)
show that there are six instars. This is in contrast with five instars of a
related tent caterpillar on P. tremuloides Michx. in New Mexico (Stelzer,
1968). As expected, development is most synchronized in the early
instars with considerable overlap prior to pupation. By 24 April 1968,
larvae were in the fifth and sixth instars (Fig. 5) and had begun the
characteristic prepupational wandering. Pupation occurs in any niche
or crevice, whether it be under bark, beneath rocks, on buildings, or on
parked vehicles. The pupal stage lasts approximately two weeks. Co¬
coons containing living pupae can, however, be found over a more
extended period. Adult emergence (Fig. 4) usually occurs late in May
or early June with oviposition beginning in late May and continuing
30
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
2 3 4 , 5 . 6
0.40 0.80 1.20 1,60 2.00 2.40 2.80 3.20 3.60
HEAD WIDTH IN MILLIMETERS
Fig. 6. Head capsule measurements of 1,878 Malacosoma incurvum discoloration
larvae.
through June. Seasonal development, it must be stressed, may vary con¬
siderably from year to year. Development rates are also quite localized.
While the population in Zion Park (elev. 4,048 ft.) was mostly in the
sixth instar and pupal stage on 16 May 1968, another population of
M. incurvum discoloratum 54 miles away at Littlefield, Arizona (elev.
1,858 ft.) had reached the egg stage.
Biology
Egg Masses .—The white egg masses are deposited on cottonwood
twigs. In the course of annual egg surveys, it has been found that egg
JANUARY 1970 ]
BAKER-MALACOSOMA ECOLOGY
31
masses are usually most abundant in the central portion of the tree
crown. It was also observed that 99.7% of the egg masses are laid
within 21 inches of the branch end. Examination of 42 randomly
selected 1967-1968 egg masses revealed a mean twig diameter at the
site of oviposition of 4.2 mm. The same egg masses were measured
and the mean mass was found to be 14.5 mm long and 10.6 mm wide.
The mean number of eggs was 163.8. Mean laboratory emergence
was 86.8 or about 53% of the mean total number of eggs.
Colonial Behavior .—An effort was made to determine behavioral
patterns of colonies on their cottonwood host. As behavioral patterns
became evident, it was also apparent that separate colonies or portions
of colonies followed different patterns at the same time. Field obser¬
vations were made during both day and night, during both rain and
clear weather, and at temperatures ranging from 46° to 80° F. Under
all conditions and at all examination times, it was possible to find some
of the tagged colonies either quiescent inside the tents or partially
inside the tents with individuals traveling or feeding outside. One
particular tent was observed to be almost entirely occupied by cater¬
pillars but two hours later most of this aggressive colony had vacated
the old tent and traveled 90 cm along adjacent twigs and had constructed
the nucleus of an entirely new tent. During a record low temperature
of 23° F on 20 April 1966, numerous larvae in the Zion population were
found motionless but perfectly healthy on tents and on leaf and branch
surfaces outside their tents. Larval activity resumed with a return to
higher temperatures. The highly variable intercolonial and intra¬
colonial behavior may possibly be explained by the individual larval
differences demonstrated by Wellington (1957). In his studies, he
categorized larvae under two general types of behavior, “sluggish” and
“active.” He found that the presence of active individuals had a
definite influence on the actions of their sluggish followers. The obser¬
vations recorded in Zion Canyon may be, to some measure, a function
of active individuals expressing a tendency to travel or forage out of
the tent under diverse conditions. Individual larval observations would
be necessary to determine this possibility.
The violent anterior “jerking” exhibited by larvae (Stehr and Cook,
1968) has been repeatedly observed on Fremont cottonwood. Various
disturbances have in the past been a possible explanation for this ac¬
tivity. No disturbances such as attacks by parasites were identified
but may well have prevailed. It was impressive to note entire clusters
of larvae exhibiting the jerking behavior simultaneously.
32
THE PAN-PACIFIC ENTOMOLOGIST [vOL. 46, NO. 1
Host Damage
Early-instar larvae consume expanding leaves and catkins. In the last
three instars, before prepupational wandering, damage to leaves increases
considerably. Individual leaves are either partially or entirely con¬
sumed. It is questionable how much Fremont cottonwood mortality
would result if the population went uncontrolled. Like other hardwoods,
the cottonwood has a remarkable propensity to endure defoliation and
refoliate the same season after the cessation of insect feeding. Un¬
doubtedly, certain trees or groups of trees might succumb, but most of
the host type would probably survive from year to year. Scattered
branch mortality indicates some possible evidence of past tent cater¬
pillar feeding.
Insect Mortality
Natural Factors.- —Parasites that have been recovered from larvae
and pupae in Zion Park over the past three years include a tachinid,
Chaetogaedia monticola (Big.) and the chalcid, Brachymeria ovata
(Say). Also recovered was an eulophid egg parasite, Tetrastichus
malacosomae Girault, a common parasite of the genus Malacosoma
throughout the Great Basin. Although a pathogenic virus has been
observed, recent annual biological evaluations have not revealed any
widespread occurrence of virus in the population.
Applied Factors. —Through the years the nuisance of abundant larvae
has been reduced in the heavily used recreational areas of Zion Park.
Records indicate that control was attempted in Zion Park as early as
1923. Applied control of portions of the Zion infestation has evolved
from the use of inorganic and synthetic organic insecticides to the use of
a more selective microbial insecticide. A formulation of Bacillus thurin-
giensis var. thuringiensis Berliner, Thuricide 90TS 2 , was found in 1965
to provide desirable operational characteristics and was claimed by the
manufacturer to have improved biological effectiveness (Klein and
Brueck, 1965). Control results in 1965 were considered acceptable. In
less than 24 hours, infected larvae had ceased feeding and had assumed
a moribund appearance. Applications of B. thuringiensis have achieved
the desired purpose.
Acknowledgments
Credit is due M. Minnoch, L. 0. Sandin, D. J. Curtis, and officials of
Zion National Park for their field and laboratory cooperation. Photo¬
graphic credit is due W. H. Klein. F. W. Stehr was helpful in species
determination and manuscript review.
3 Registered trademark, Bioferm Division, International Minerals Corp., Wasco, California. Does
not necessarily imply endorsement by U.S.D.A.
JANUARY 1970 ] GARDINER-IMMATURES OF SPONDYLIS
33
Literature Cited
Klein, W. H., and F. R. Brueck. 1965. Control of a tent caterpillar, Mctlaco-
soma incurvum discoloration, with a commercial formulation of Bacillus
thuringiensis var. thuringiensis in Zion National Park, 1965. U. S.
Forest Serv., Intermountain Region, Ogden, Utah. Unpub. Report, 7 pp.
Steiir, F. W., and E. F. Cook. 1968. A revision of the genus Malacosoma
Hiibner in North America (Lepidoptera: Lasiocampidae) : system-
atics, biology, immatures, and parasites. U. S. Nat. Mus. Bull., 276.
Smithsonian Institution Press.
Stelzer, M. J. 1968. The Great Basin tent caterpillar in New Mexico: life
history, parasites, disease, and defoliation. U. S. Forest Serv. Res.
Pap., RM-39.
Sudworth, G. B. 1934. Poplars, principal tree willows, and willows of the
Rocky Mountain Region. U. S. Dep. Agr. Tech. Bull., 420.
Wellington, W. G. 1957. Individual differences as a factor in population
dynamics: the development of a problem. Can. J. Zool., 35: 293-323.
Immature Stages and Habits of Spondylis upiformis
Mannerheim
(Coleoptera: Cerambycidae)
L. M. Gardiner
Forest Research Laboratory, Sault Ste. Marie, Ontario, Canada
Three species of Spondylis Fabricius have been recognized, one from
the Palearctic Region, one from western North America, and one from
Mexico (Linsley, 1962). The larva and pupa of the European species, S.
huprestoides L., have been described (Duffy, 1953) but neither the im¬
mature stages nor food habits of the North American species have been
recognized, although the adults are commonly encountered.
In 1968, several larvae and two pupae were collected by the author
from two stumps of white spruce, Picea glauca (Moench) Voss, at the
Kananaskis Forest Experiment Station, Seebe, Alberta. The stumps had
been pulled from the ground for other purposes; otherwise the presence
of the insects would have been unsuspected. One pupa was permitted to
complete development, and the adult was identified as S. upiformis , the
western North American species. The immature stages and habits of this
species are described herein.
Descriptions of Immature Stages
Advanced Larva.-— Form robust, sub cylindrical, length up to 22 mm; in all
respects, typically asemine.
The Pan-Pacific Entomologist 46: 33-36. January 1970
34
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
] 2
Figs. 1-2. Spondylis upiformis. Fig. 1. Ninth abdominal tergum and uro-
gomphi of larva. Fig. 2. Pronotum of pupa.
Head depressed; mouthframe, frons, ventral genal areas, and hypostomal bridge
piceous to dark castaneous; epistoma shallowly declivous, bearing six to eight
setae laterally, midfrontal inflection dark, frontal sutures bisecting antennal
sockets; frons roughly excavated behind antennae, genae moderately densely
clothed with golden setae; antennae minute, retractile, three-segmented, third
segment very small, almost equalled in length by conical supplementary segment;
ocelli absent; hypostoma fused with base of ventral mouthparts, gular sutures
slightly raised.
Clypeus produced at sides; labrum thick, broadly rounded; densely setose,
setae shorter and denser apically; mandible slender, apex depressed, shining;
ventral mouthparts fleshy, cardo well defined, maxillary lobe digitate, last joint
of maxillary palp shorter than second, equal to or slightly longer than last labial.
Pronotum testaceous, posterior half darker, finely asperate, lateral furrows
pronounced, converging slightly anteriorly, pubescence short and sparse between
furrows, lateral setae moderately dense, ferruginous; eusternum finely asperate,
bearing numerous fine, reddish setae; legs three-segmented, unguiculus attenuate,
imbricately spinose.
Dorsal abdominal ampullae finely asperate, very sparsely pubescent, broadly
impressed on midline; spiracles small, ovate to suborbicular, peritreme thin,
with two or three contiguous chambers on posterior rim; pleural tubercles not
evident; ninth tergum with a pair of small, blunt urogomphi separated by less
than basal width (fig. 1).
The last character readily separates this species from S. buprestoides
in which the urogomphi are separated by at least four times their basal
width (Duffy, 1953).
Pupa .—Length 18 mm, single specimen, male.
Head barely visible from above, body widest at fourth abdominal segment.
Tapering rapidly posteriorly.
Vertex of head glabrous, face and mandibles with numerous, very fine, light
setae; pronotum obtusely angled on sides which are thickly clothed with fine
setae (fig. 2), disc with numerous small spines, some with terminal setae;
scutellum with few similar spines; metatergum with broad, V-shaped band of
JANUARY 1970] GARDINER—IMMATURES OF SPONDYLIS
35
Fig. 3. Larval galleries and pupal cells of Spondylis upiformis in spruce root.
small spines and fine setae; femora with few small spines and a fringe of fine
setae at apex, hind femora with large, fleshy basal lobe (actually developing
trochanter).
Abdominal terga with transverse hands of slightly larger spines, sparsely
clothed anteriorly and laterally with fine setae; ninth tergum terminating in a pair
of incurved, heavily sclerotized urogomphi; third to eighth sterna with lateral
groups of small spines; ninth sternum with a pair of short setae; pleura with a
group of setae; functional spiracles on first seven abdominal segments.
The pupa is distinguished from that of S. buprestoides by the lack of
stout, curved, pronotal spines described by Duffy (1953) on the latter
specie's.
The above descriptions are based on ten larvae and one male pupa,
designated as Collection No. SI. 1.3 in the Cerambycid Collection, Forest
Research Laboratory, Sault Ste. Marie, Ontario.
Habits
Linsley (1962) gives the range of this species as “from Alaska south¬
east to the Great Lakes region, south into the Rocky Mountains, and
along the Pacific Coast.” As well as in Alberta, I have collected adults
in mid-June in an area logged for spruce and balsam fir near Sault Ste.
Marie, Ontario.
36
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
The stumps that yielded the immature stages described above were
from trees cut on 15 June and 16 June 1966. Both stumps were pulled
on 17 May 1968 and were examined for insects on 31 July. The larvae
apparently had entered pupal cells by the time of pulling, but were
prevented from achieving normal development and emergence by dry¬
ing of the stump. The length of the larval feeding galleries suggests that
oviposition occurred in 1966, and that this species has a 2-year life
cycle in Alberta.
The method of oviposition is unknown, but the larval galleries began
in the roots, sometimes more than a meter from the stump. Feeding pro¬
ceeded more or less straight along the root towards the stump, the
larva scoring the wood deeply and leaving the gallery filled with fibrous
frass. Pupation occurred in shallow cells excavated in the sapwood of
the stump or in roots near the stump (fig. 3). Feeding and pupation
took place as much as 50 cm below the soil surface. The adults are, how¬
ever, somewhat adapted for digging, having unusually large mandibles
and terminal lamellae on the fore tibiae.
Acknowledgment
I am grateful to W. B. Johnstone, Department of Fisheries and For¬
estry, Calgary, Alberta for information on stump history and location.
Literature Cited
Duffy, E. A. J. 1953. A monograph of the immature stages of British and
imported timber beetles (Cerambycidae). British Museum (Natur.
Hist.), London. Jarrold and Sons, Limited, Norwich. 350 pp. 8 plates.
Linsley, E. G. 1962. The Cerambycidae of North America Part II. Taxonomy
and Classification of the Parandrinae, Prioninae, Spondylinae and
Aseminae. Univ. Calif. Publ. Entomol., 19: 1-102 -)- 1 pi.
A New Genus in the Tribe Euplectini
(Coleoptera: Pselaphidae)
A. A. Grigaricic and R. 0. Schuster
University of California, Davis
This new genus has close affinities with the genus Actium of Casey.
The first visible abdominal tergite of all of the species of Actium is
essentially the same. This segment bears two basal carinae that differ
between species in length and in degree of separation. A species has
been discovered which differs from Actium mainly in that the base of
The Pan-Pacific Entomologist 46: 36-39. January 1970
JANUARY 1970 ] GRIGARICK & SCHUSTER—NEW PSELAPHID GENUS
37
tergite I has a densely pubescent median depression and the carinae are
absent. We consider the addition of this depression and the absence
of the carinate condition fundamental differences that justify placing
the species in a separate genus.
Pilactium Grigarick and Schuster, new genus
Type of Genus. — Pilactium summersi Grigarick and Schuster, new
species.
This genus has those subtribal characteristics typical of the Trimiina
and is defined by the following combination of characters:
1) Ventral surface of head with capitate setae. 2) Two vertexal foveae, connected
by entire U-shaped frontal sulcus. 3) Gular foveae separate. 4) Antennal seg¬
ments IX to XI forming a distinct club, X nearly symmetrical, well separated
from XI. 5) Pronotum with basolateral foveae connected by transverse, slightly
biarcuate sulcus. 6) Elytron with 2 antebasal foveae; sutral stria entire; discal
fovea with sulcoid impression about one-half length of elytron. 7) Prosternum
simple, prosternal foveae large. 8) Mesosternal preplectus medianly carinate;
mesosternum anteriorly with 2 lateral and 1 median foveae. 9) Metasternum
with median paired foveae. 10) Profemur with row of specialized setae on basal
half. 11) Meso- and metacoxae contiguous. 12) Tarsal claws consist of large
primary and smaller secondary structures. 13) First visible tergite with densely
pubescent, median depression; basal carinae absent. 14) Second sternite with
pubescent, median depression.
Pilactium summersi Grigarick and Schuster, new species
(Figs. 1M0
Male.— Teneral specimen, yellow brown. Length 933 1 , width 354. Head 151
long, 181 wide; vertexal foveae 77 between centers; ventral surface with 24
capitate setae; eyes moderate. Antenna 310 long; segment I 43 long, 27 wide; II
34 long, 30 wide; III to VI inclusive 57 long, 20 wide; VII 16 long, 24 wide; VIII
16 long, 27 wide; IX 17 long, 34 wide; X 24 long, 40 wide, nearly symmetrical,
well separated from XI; XI 87 long, 54 wide, as long as preceding 5 segments.
Pronotum 208 long, 225 wide. Elytron 292 long, with 2 antebasal foveae.
Winged. Protrochanter simple; profemur 57 wide; protibia simple. Mesotro-
chanter simple; mesofemur 63 wide; mesotibia with small, blunt apical spine.
Postmesocoxal foveae with apodemes directed obliquely forward. Metafemur
50 wide.
First visible tergite 245 wide at base, without basal carinae but with dense
setae area 100 wide (fig. 4). Sternite II with median, pubescent depression; III
to V unmodified; VI deeply emarginate, with median group of broad setae (fig. 3) ;
penial plate 125 long, setate area nearly round, 93 in diameter. Aedeagus 221
long, 128 deep (fig. 1).
Female.— Segment IX as in figure 2.
1 All measurements in microns.
38
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
s'? 71
^ h S \> V n ? / T / ' « 1
-f v ''~° / V 1 T '
9 > VV,'7 1
f \ f T \ T V-N^X
4
Figs. 1-4. Pilactium summersi Grigarick and Schuster, new species. Fig. 1.
Male genitalia, dorsal. Fig. 2. Female genitalia, mainly ninth segment, orienta¬
tion not established. Fig. 3. Male sternite VI. Fig. 4. Median depression of
first visible tergite.
Distribution.— The holotype male was collected at Quincy, Plumas
County, California on 1 April 1951 by F. M. Summers. A male
paratype was collected at Pine Grove, Amador County, California on
JANUARY 1970] NUTTING—TENUIROSTRITERMES FORAGING
39
12 February 1961 by D. W. Price from pine forest soil. A female
paratype was collected at Dunsmuir, Mt. Bradley, Siskiyou County,
California on 20 September 1961 by R. 0. Schuster. The type is deposited
with the Department of Entomology, University of California, Davis.
Discussion. —This uncommon montane species is known only from
single specimens at three widely separate localities of northern Cali¬
fornia. Expected differences in species that may be collected in the
future, as indicated by related genera, would be in the following struc¬
tures: tibial spines (male) ; number of capitate setae on venter of head;
size relationship of femora (male) ; setation of sternite VI (male) ; and
genitalic differences.
Free Diurnal Foraging by the North American
Nasutiform Termite, Tenuirostritermes tenuirostris 1
(Isoptera : Termitidae)
W. L. Nutting
University of Arizona, Tucson
Tenuirostritermes tenuirostris (Desneux) is one of at least two spe¬
cies of termites possessing nasutiform soldiers which reach the extreme
southern United States. It ranges from southern Mexico and perhaps
Guatemala (Snyder, 1949) into the Upper Sonoran Zone of south¬
eastern Arizona and similar areas in southwestern Texas. The other
species, T. cinereus (Buckley), occurs in southwestern Texas and may
range into south-central Mexico. Two additional species in this genus
are confined to ranges of undetermined extent in Central America and
perhaps the West Indies. All of the species are apparently subterranean
since they are not known to build tree nests or extensive systems of
tubes above ground as do many other genera of nasutiform termites.
Thus, although T. tenuirostris and cinereus are unusual elements in
our fauna, they are not widely known because of their cryptic habits
and limited distribution in the United States.
The following notes were made on T. tenuirostris , incidental to a
field study of termite flight behavior conducted in western Mexico,
from May to October, 1964. This termite was observed foraging on
exposed trails during daylight hours on four separate occasions. The
1 Journal Paper No. 1499 of the Arizona Agricultural Experiment Station. Supported in part by
State Research Project No. 461 and a research grant from the National Pest Control Association
for travel in Mexico.
The Pan-Pacific Entomologist 46: 39-42. January 1970
40
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
first three observations were made in the mountains directly north of
the village of Ajijic, on the north shore of Lake Chapala in Jalisco.
The summer rainy season had begun with traces of rain on May 18, and
showers of more than 0.1 inch on 20 May and 2 June. On 6 June in a
steep canyon (5800 ft.), a group of several hundred workers and sol¬
diers were disturbed in a loose mat of dry leaves and grasses. They
had been operating in an irregular column several feet from galleries
under a small, deeply set stone. Although this was at 1100 hrs, and
very warm, all of the activity was in the deep shade of the canyon wall.
On 29 July after several weeks of showers totalling over 14 inches
of rain, a similar situation was discovered. The colony was located
beneath a stone wall separating pasture from a well-traveled dirt lane
(5200 ft.). A column of termites was moving to and from a hole beside
the wall to a site six feet away where workers were collecting bits of dead
grass and green leaves. Several workers were cutting semicircles, ir¬
regular circles and other shapes (1-2 mm on a side) from the hairy
leaves of a small honeysuckle plant. Some of the leaves, up to 6 inches
above the ground, had been completely removed. Soldiers were spaced
at one-half-inch intervals on either side of, and about one-half inch
from, the two-way traffic of workers. This activity was first seen at
1400 hrs under a complete overcast. When the foragers were disturbed
they quickly moved in to the end of the column and the traffic was
progressively reversed. Small groups of soldiers moved from their
lateral positions to alternate with groups of workers in a remarkably
orderly return to the hole under the wall. Two and one-half hours
later, during a light drizzle, foragers were working about two feet from
the nest in the opposite direction.
At 1045 hrs on 6 August another column was found in dense grass
beside the trail on an open ridge at 5500 ft. A 50% cloud cover pro¬
vided intermittent shade during the entire morning. The last en¬
counter occurred on 21 August in a pass (3850 ft.), 5.2 road miles
(Highway 110) south of Tecalitlan, Jalisco. A short column was found
at 1330 hrs foraging in a stony, short-grass area in an oak-pine wood¬
land. The sky was completely overcast. Each of the colonies north of
Ajijic was checked on several other occasions, and foraging was never
seen during continuous, bright sunlight.
Although these are not the first records of such activity for T.
tenuirostris, they are the first to establish that it will attack living
plants. In connection with her intensive field and laboratory work on
caste development in this species, Weesner (1953) described her ob¬
servations on its foraging behavior in southeastern Arizona. She saw
JANUARY 1970 ] NUTTING—TENUIROSTRITERMES FORAGING
41
workers actually cutting plant material only once. They were accom¬
panied by soldiers and were operating in daylight on a large grass
stalk under the cover of a small board. (1000 hrs, 3 September 1947,
very hot and dry; Weesner, in litt., 27 May 1969.) On another oc¬
casion at night she encountered a short, exposed column of workers
guarded by soldiers. She also noted that stores of plant fragments were
generally found in their chambers and galleries beneath stones. How¬
ever, in all cases this consisted entirely of dry, weathered plant ma¬
terials. The author has made similar observations in colonies of this
species at several localities in southeastern Arizona and in Mexico.
Considerably less is known about the distribution and biology of
T. cinereus. However, Banks and Snyder (1920) cited three early
accounts in Texas where workers, accompanied by soldiers, were found
foraging in the open: once for grass and weed seeds at about sunset,
again for dry leaves (Buckley) and, on the third occasion “destroying
living plants by sucking the juices” (Parman). Most of the observa¬
tions indicated that they probably do not forage in bright sunlight.
Ebeling (1959) mentioned Tenuirostritermes incisus (Snyder) as
having been recorded feeding on the avocado. The record comes from
some locality in the West Indies, although Snyder (1949) listed it only
from Spanish Honduras. The author has collected this species in
Guerrero, and Tenuirostritermes briciae (Snyder) in Nayarit, both in
southern Mexico. Apparently nothing else has been published on these
two species since the original descriptions (Snyder, 1922).
Snyder (1968) has continued to update a growing bibliography on
termites which damage living vegetation. The most notable among
these papers is that by Harris (1962) wherein he provided a world list
of 106 species of termites, from all six families, which feed on living
plants. Many of them are economically important pests of crops and
trees. In all of these records, reports of such damage in the New World
are surprisingly few. The two termitid genera most often mentioned
are Nasutitermes and Syntermes. Syntermes has been observed forag¬
ing from exposed trails (Emerson, 1938), while Nasutitermes generally
works from covered runways.
Although Tenuirostritermes has apparently never been observed to
attack crop plants, its abundance in certain agricultural areas certainly
provides this potential. In foothill and mountain regions of western
Mexico, tenuinostris was found several times in the vicinity of, and
sometimes within, the “milpas” or cultivated plots (chiefly corn). In
southeastern Arizona, where it occurs in areas of lower rainfall, the
land is used almost exclusively as range for cattle.
42
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
I want to thank Frances M. Weesner of Colorado State University,
for reviewing the manuscript and confirming my own identification of
T. tenuirostris from several localities in Mexico.
Literature Cited
Banks, N., and T. E. Snyder. 1920. A revision of the Nearctic termites with
notes on biology and geographic distribution. U. S. Nat. Mus. Bull.,
108: 190-193.
Ebeling, W. 1959. Subtropical fruit pests. Univ. Calif. Press, Los Angeles,
p. 317.
Emerson, A. E. 1938. Termite nests—a study of the phylogeny of behavior.
Ecol. Monogr., 8: 258-259.
Harris, W. V. 1962. Classification of the phytophagous Isoptera. Symposia
Genetica et Biologica Italica. Atti IV Congr. U.I.E.I.S.-Pavia, 9-14
Sett., 1961,11: 193-201.
Snyder, T. E. 1922. New termites from Hawaii, Central and South America,
and the Antilles. Proc. U. S. Nat. Mus., 61: 25-28.
1949. Catalog of the termites (Isoptera) of the world. Smithson. Misc.
Collect., 112: 312-313.
1968. Second supplement to the annotated, subject-heading bibliography
of termites 1961-1965. Smithson. Misc. Collect., 152: 35-40.
Weesner, F. M. 1953. Biology of Tenuirostritermes tenuirostris (Desneux) with
emphasis on caste development. Univ. Calif. Publ. Zool., 57: 263-264.
Pselactus spadix (Herbst), a European Cossonine Weevil
New to California
(Coleoptera: Curculionidae)
Charles W. O’Brien
International Programs in Agriculture , Purdue University
Pselactus spadix (Herbst) is a native of Great Britain and the west
coast of Europe. It has been introduced into the eastern United States,
Australia, and New Zealand, where specimens have been taken in wood
associated with salt water. Published European records are all coastal
and all directly associated with salt water. Specimens have been col¬
lected in stumps regularly covered by tide (Champion, 1871) and in
England a wharf and its piles which were regularly subjected to salt
spray and storm waves were severely damaged (Walker, 1879). How¬
ever, in a more recent work on its presence in Argentina, Viana (1961)
discusses extensive damage in Acer negundo Linne and Salix babylonica
Linne in the field and also in furniture and the flooring of houses.
The Pan-Pacific Entomologist 46 : 42-44. January 1970
JANUARY 1970] O’BRIEN—PSELACTUS IN CALIFORNIA
43
Fig. 1. Pselactus spadix, dorsal and lateral view. Note robust form, long fine
pubescence, and elongate rostrum.
In the United States the species has been collected far from the coast
as well. The economic threat is well demonstrated in the following un¬
published data, courtesy of Mrs. Rose E. Warner-Spilman, U. S. Na¬
tional Museum: Illinois, in pine in house. Maryland, basement, furni¬
ture store. Massachusetts, larvae boring in piles in Boston Harbor;
from timber in Boston and Main R.R. Pier 40 (not submerged) ; in
pile; from cellar timber, with ex. of Hadrobregnus. Missouri, from
44
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
pine board infested with weevils. South Carolina, under bark of dead
Morus sp. twig; in dead Ficus wood; in old pinewood porch (with
Hexarthrum ulkei Horn).
Pselactus spadix was first collected in California by Mr. William
Tyson at the salt flats in Newark, California, near the southeastern
end of San Francisco Bay on 2 January 1966 from beneath pieces
of wood. Subsequent collections of more than 150 specimens were
made in February and the late fall of 1966 at the same locality. The
area is at the south end of the Newark Slough which leads di¬
rectly into San Francisco Bay. The specimens were taken under and
in boards and timbers on levees above salt evaporation ponds. The
weevils were most common on partly buried wood where conditions
were slightly more damp. Evidence of weevil infestation was found on
all boards and timbers. Mr. Tyson indicated that during the wet win¬
ter season the weevils were common under most of the boards. The
timbers used by the salt company to support the pipes which transport
salt water to the evaporation ponds are heavily infested by the weevils
and will eventually have to be replaced.
Several other genera of cossonine weevils are common along the
west coast and in general damage driftwood in a similar manner. These
include the New Zealand Macrancylas littoralis Broun, very common in
the San Francisco Bay area, north at least to Point Reyes. Ellasoptes
marinus Horn and Rhyncholus spp. are common from Oregon to South¬
ern California. In the eastern United States, one may encounter Ma¬
crancylas linearis Leconte, Dryotribus mimeticus Horn, Anchonus spp.,
and Mesites spp. The adults and larvae may be found burrowing
through the wood and the adults are often taken on the under surface of
wood on the ground. Pselactus spadix adults may be separated from the
other cossonine genera occurring in the western United States by the
combination of a robust form and erect long fine dorsal pubescence
(Fig. 1). Anderson (1952) presents a key to and a description of some
cossonine larvae including Pselactus.
Literature Cited
Anderson, W. H. 1952. Larvae of some genera of Cossoninae (Coleoptera:
Curculionidae). Ann. Entomol. Soc. Amer., 45:281-309.
Champion, G. C. 1871. Pselactus spadix occurs in stumps under salt water.
Entomol. Mon. Mag., 8:85.
Viana, M. J. 1961. Presencia en la Argentina de Carcoma de la Madera
(.Pselactus spadix ). Idia, Instituto de Patologia Vegetal INTA (163):
14-16.
Walker, J. J. 1879. Phloeophagus spadix (Herbst), destructive to piles at
Harwich. Entomol. Mon. Mag., 16:40.
JANUARY 1970]
HALSTEAD-A NEW LARGUS
45
A New Species of the Genus Lcirgus Hahn With a Key to the
Species of the Genus in the Southwestern United States
(Hemiptera: Largidae)
Thomas F. Halstead
San Francisco State College, California
During a taxonomic study carried out as part of the requirements for
the Master of Arts degree in biology at San Francisco State College, I
examined a series of nine female and seven male specimens represent¬
ing a different form of the genus Largus Hahn, 1831. The specimens
were found among the collections of several United States museums. A
search of the literature and comparison with many determined speci¬
mens of the various established species of Largus found in North Amer¬
ica led me to consider the specimens as representative of a new species.
I am now publishing the following description so that the name will
be available for biological studies on this genus. The measurements
given below are those of the holotype or allotype followed by the range
of variation exhibited by the paratypes of the appropriate sex.
Largus semipunctatus Halstead, new species
Length of holotype male 13.5 mm (12-13.5 mm), width across pronotum 5 mm
(4-5 mm), width across corium 6 mm (5-6 mm).
Male. —Elongate oval; robust. Coloration: Black. The following all red-
orange: two small medial lines on the collum, posterior margin of pronotum,
lateral margins of posterior half of pronotum, apex of scutellum, margins of
clavus and claval suture, lateral margin of corium next to membi-ane, furrow in
corium, connexivum of abdomen, base of rostrum, base of first and second
antennal segments. Pubescence: Body and femora clothed with dense silver-gray
tomentum giving a frosted appearance. Punctation: Posterior half of pronotum
except lateral margins finely black punctate. Scutellum finely, evenly punctate.
Clavus rather coarsely black punctate. Corium along claval suture and inside of
lateral red-orange margin black punctate; center and posterior of corium im-
punctate. Hemelytra: Not quite reaching tip of abdomen. Length of antennal
segments : I: 2.9 mm (2.6—2.9 mm) ; II: 1.8 mm (1.8-1.9 mm) ; III: 1.0 mm
(0.9-1.0 mm) ; IV: 1.9 mm (1.8-1.9 mm). Total antenna 7.6 mm (7.1-7.6 mm).
Female.— Length 15 mm (15-17 mm) ; width across pronotum 5 mm (4.5-5.5
mm) ; width across corium 7 mm (6-7 mm).
Similar to male in coloration except: smaller black areas of clavus and corium;
first three ventral abdominal segments tinged with rose; posterior margin of
metathorax very slightly tinged rose. Similar to male in punctation except smaller
impunctate area of corium. Pubescence and hemelytra as in male. Length of
antennal segments : I: 2.9 mm (2.7—3.2 mm) ; II: 1.9 mm (1.8-2.2 mm) ; III:
1.1 mm (1.0-1.2 mm); IV: 2.0 mm (1.7-2.1 mm). Total antenna 8.0 mm (7.6-
8.8 mm).
The Pan-Pacific Entomologist 46: 45-46. January 1970
46
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Holotype male , Atascosa Lookout, Atascosa Mt., Santa Cruz
County, Arizona. Collected by Mr. W. L. Nutting, 9 October 1960,
deposited in the University of Arizona collection. Allotype, Same data
as holotype, in University of Arizona collection.
Paratypes. —One male, Santa Catalina Mts., Arizona, 15 July 1924; one male,
Cochise Str., Arizona, Dragoon Mts., 24 March 1956; one male, Molino Basin
Station, Catalina Mts., Arizona, 18 October 1961, on Baccharis flower; one male,
Pena Blanca Spring, Tumacacori Mts., Arizona, 20 June 1940; one female, Pena
Blanca, 10 miles west of Nogales, Arizona, 1 August 1961; one female, Bear Val¬
ley, Arizona, Tumacacori Mts., 28 June 1940; one female, west slope Patagonia
Mts., Santa Cruz Co., Arizona, 9 August 1955; one female, Madera Canyon, Santa
Rita Mts., Pima Co., Arizona, 16 February 1956, all in the University of Arizona
collection. Two males and two females, Baboquivaria Mts., Arizona, with no
date, in the University of Kansas collection. One female, Madera Canyon, Santa
Rita Mts., Arizona, 14 October 1960, in the Oregon State University collection.
One female, San Bernardino County, California, 16 November 1936, in the Cali¬
fornia Academy of Sciences collection.
In general appearance Largus semipunctatus most closely resembles
Largus convivus Stal. However, it is separated from that species by the
absence of punctation on the corium and the very different pubescence,
as well as the coloration of the thorax and legs.
The characteristics of this new species which differentiate it from
the other members of the genus Largus of western North America are
shown in the following key:
Key to the Species of the Genus Largus Hahn
of the southwestern United States
1. Membrane clear to light tan; scutellum, hemelytra, and posterior half
of pronotum reddish to tan; corium with two distinct large black spots
at juncture of claval suture and membrane _ bipustulatus Stal
Membrane and hemelytra color variable; scutellum black; corium with¬
out black spots at juncture of claval suture and membrane _ 2
2. Thoracic sterna around base of coxae red to orange _ 3
Thoracic sterna around base of coxae black _ 4
3. Coxae often red; trochanters and femora black _ convivus Stal
Coxae, trochanters, and proximal end of femora all red to yellow-orange
__ cinctus Herrich-Schaeffer
4. Hemelytra completely covered with dense punctation; pubescence vari¬
able, but not as below; trochanters and proximal ends of femora
orange to orange-red _ succinctus Linne
Hemelytra with corium impunctate; pubescence a dense silver tomentum
over entire body giving the insect a frosted appearance; trochanters
and femora all black _ semipunctatus Halstead, n. sp.
JANUARY 1970] WASBAUER-CALIF. DEP. AGR. TYPES
47
The Type Material in the Collection of the
California Department of Agriculture
Marius S. Wasbauer
California Department of Agriculture, Sacramento
During the past decade, a number of primary types of insects has
accumulated in the Collection of the Insect Taxonomy Laboratory, Cali¬
fornia Department of Agriculture, Sacramento.
Although there are many cogent practical reasons for any institution
which maintains an insect collection to be interested in the retention of
holotypes, we feel that the field of insect taxonomy could best be served
by relinquishing the holotypes now housed in the California Department
of Agriculture.
In the future, it will be the policy of the Insect Taxonomy Laboratory
to require that holotypes based on our material be deposited in the col¬
lection of the California Academy of Sciences, San Francisco, unless it
can be shown that they would be more readily accessible in an in¬
stitution where active work is being done on that particular group of
insects or where a specialized collection of the group is being housed.
A list of type material in the CDA Collection is presented here, ar¬
ranged by order of insect. Following the family name are the initials
of the institution to which the types will be transferred: CAS (Cali¬
fornia Academy of Sciences, San Francisco), UCD (University of
California, Davis), USNM (United States National Museum, Washing¬
ton, D. C.).
ORTHOPTERA
Acrididae (USNM)
Melanoplus buxtoni Strohecker 1963. Pan-Pac. Entomol., 39(3) : 168.
Holotype $, allotype 2, USNM type #70510.
Melanoplus elaphrus Strohecker 1963. Pan-Pac. Entomol., 39(3) : 169.
Holotype $, allotype 2, USNM type #70517.
Melanoplus elater Strohecker, 1963. Pan-Pac. Entomol., 39(3) : 166.
Holotype $, allotype 2, USNM type #70509.
Melanoplus eremitus Strohecker, 1963. Pan-Pac. Entomol., 39(3) : 162.
Holotype $, allotype 2, USNM type #70512.
Melanoplus harperi Gurney and Buxton, 1965. Bur. Entomol., Calif. Dep.
Agr. Occas. Pap. No. 7, p. 4. Holotype $, allotype 2, USNM type
#70513.
Melanoplus siskiyou Strohecker, 1963. Pan-Pac. Entomol., 39(3) : 170.
Holotype $, allotype 2, USNM type #70511.
The Pan-Pacific Entomologist 46: 47-50. January 1970
48
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Melanoplus wintunus Strohecker and Heifer, 1963. Pan-Pac. Entomol.,
39(3): 160. Holotype $, allotype 2, USNM type #70508.
Aerochoreute s stenometopus Strohecker and Buxton, 1963. Pan-Pac. En¬
tomol., 39(4): 261. Holotype $, allotype 2, USNM type #70514.
Hypsalonia merga Gurney and Buxton, 1963. Bull. Brooklyn Entomol.
Soc., 58(2-3): 68. Holotype $, USNM type #70515.
Boonacris ( Karokia ) bland Rehn, 1964. Nolulae Natur., 368: 3. Holotype
$, allotype 2. USNM type #70516.
THYSANOPTERA
Phlaeothripidae (UCD)
Hoplandrothrips irretius Kono, 1964. Bur. Entomol. Calif. Dep. Agr.
Occas. Pap. No. 4, p. 1. Holotype 2, allotype $, UCD type #291.
Liothrips brevitubus Kono, 1964. Bur. Entomol. Calif. Dep. Agr. Occas.
Pap. No. 4, p. 4. Holotype 2, UCD type #292.
Liothrips monoensis Kono, 1964. Bur. Entomol. Calif. Dep. Agr. Occas.
Pap. No. 4, p. 6. Holotype 2, UCD type #293.
Thripidae (UCD)
Chirothrips tuttlei zur Strassen, 1967. Senckenbergiana Biol., 48(5-6):
345. Holotype 2, UCD type #294.
HOMOPTERA
Asterolecaniidae (UCD)
Sclerococcus bromeliae McKenzie 1963. Bull. Calif. Dept. Agr., 52(1) :38.
Holotype 2, slide mount, UCD type #325.
Derbidae (CAS)
Apache californicum Wilkey 1963. Pan-Pac. Entomol., 39(2) : 99. Holo¬
type $, allotype 2, CAS type #10,196.
Diaspididae (UCD)
Diaspis gilloglyi McKenzie 1963. Bull. Calif. Dep. Agr., 52(1): 33. Holo¬
type 2, slide mount, UCD type #326.
Eriococcidae (UCD)
Cornoculus densus Miller 1967. Hilgardia, 38(13): 487. Holotype 2,
slide mount, UCD type #327.
Pseudococcidae (UCD)
Anisococcus abnormalis McKenzie 1964. Hilgardia, 35(10) : 212. Holotype
2 , slide mount, UCD type #328.
Anthelococcus simondsi McKenzie 1964. Hilgardia, 35(10): 215. Holo¬
type 2 , slide mount, UCD type #329.
Chorizococcus californicus McKenzie 1964. Hilgardia, 35(10): 221. Holo¬
type 2 , slide mount, UCD type #330.
JANUARY 1970] WASBAUER—CALIF. DEP. AGR. TYPES
49
Chorizococcus coxindex McKenzie 1967. “Mealybugs of California,” Univ.
Calif. Press, Berkeley, p. 96. Holotype 2, slide mount, UCD type #331.
Chorizococcus fistulosus McKenzie 1967. “Mealybugs of California,” Univ.
Calif. Press, Berkeley, p. 99. Holotype 2, slide mount, UCD type #332.
Chorizococcus interruptus McKenzie 1964. Hilgardia, 35(10): 223. Holo¬
type 2, slide mount, UCD type #333.
Chorizococcus wilsoni McKenzie 1961. Hilgardia, 31(2): 19. Holotype 2,
slide mount, UCD type #334.
Chorizococcus yuccae McKenzie 1961. Hilgardia, 31(2): 21. Holotype 2,
slide mount, UCD type #335.
Cryptoripersia tubulata McKenzie 1964. Hilgardia, 35(10): 228. Holo¬
type 2 , slide mount, UCD type #336.
Dysmicoccus vacuatus McKenzie 1967. “Mealybugs of California,” Univ.
Calif. Press, Berkeley, p. 174. Holotype 2, slide mount, UCD type
#337.
Heliococcus atriplicis McKenzie 1964. Hilgardia, 35(10) : 235. Holotype
2 , slide mount, UCD type #338.
Humococcus ceraricus McKenzie 1964. Hilgardia, 35(10): 236. Holotype
2 , slide mount, UCD type #339.
Phenacoccus cdleni McKenzie 1964. Hilgardia, 35(10): 243. Holotype
2 , slide mount, UCD type #340.
Phenacoccus nonarius McKenzie 1964. Hilgardia, 35(10): 253. Holotype
2 , slide mount, UCD type #341.
Phenacoccus aherrans McKenzie 1962. Hilgardia, 32(14) : 657. Holotype
2 , slide mount, UCD type #342.
Pseudococcus diversus McKenzie 1964. Hilgardia, 35(10) : 256. Holotype
2 , slide mount, UCD type #343.
Pseudococcus malacearum Ferris 1950. “Atlas of the scale insects of North
America,” Vol. V, The Pseudococcidae (Part I), Stanford Univ. Press, p.
185. Lectotype designated by Wilkey and McKenzie 1961, Bull. Calif.
Dep. Agr., 50(4) : 246, UCD type #344.
Pseudococcus prunicolus McKenzie 1964. Hilgardia, 35(10) : 260. Holo¬
type 2 , slide mount, UCD type #345.
Pseudococcus sparsus McKenzie 1962. Hilgardia, 32(14): 667. Holotype
2 , slide mount, UCD type #346.
Puto echinatus McKenzie 1961. Hilgardia, 31(2): 35. Holotype 2, slide
mount, UCD type #347.
Puto latricribellum McKenzie 1961. Hilgardia, 31(2): 37. Holotype 2,
slide mount, UCD type #348.
Rhizoecus bicirculus McKenzie 1967. “Mealybugs of California.” Univ.
Calif. Press, Berkeley, p. 374. Holotype 2, slide mount, UCD type
#349.
Spilococcus haigi McKenzie 1962. Hilgardia, 32(14): 679. Holotype 2,
slide mount, UCD type #350.
50
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Trionymus frontalis McKenzie 1967. “Mealybugs of California.” Univ.
Calif. Press, Berkeley, p. 471. Holotype 9, slide mount, UCD type #351.
Trionymus mocus Ferris 1953. “Atlas of the scale insects of North America,”
Vol. VI, The Pseudococcidae (Part II), Stanford Univ. Press, p. 490.
Slide designated as TYPE, contains 2 adult $ 9—Ferris noted on slide
“Type, the larger spm.” UCD type #352.
LEPIDOPTERA
Gelechiidae (USNM)
Periploca nigra Hodges 1962. Pan-Pac. Entomol., 38(2): 94. Holotype $,
USNM type #70546.
DIPTERA
Tephritidae (CAS)
Aciurina trilitura Blanc and Foote 1961. Pan-Pac. Entomol., 37(2): 75.
Holotype 9, allotype $, CAS type #10,197.
Neaspilota wilsoni Blanc and Foote 1961. Pan-Pac. Entomol., 37 (2) : 78.
Holotype $, CAS type #10,198.
Lonchaeidae (CAS)
Dasiops alveofrons McAlpine 1961. Can. Entomol., 93(7) : 539. Holotype
$, allotype 9 , CAS type # 10,199.
HYMENOPTERA
Andrenidae (CAS)
Perdita salicis tristis Timberlake 1964. Univ. Calif. Publ. Entomol. 28(2):
345. Holotype 9, allotype $, CAS type #10,200.
Pompilidae (CAS)
Calicurgus braziliensis Dreisbach 1961. Amer. Midland Natur., 65(2):
377. Holotype 9 , CAS type #10,201.
A Revision of the Genus Grotea
(Hymenoptera : Ichneumonidae)
C. N. Slobodchikoff
University of California, Berkeley
Grotea is a small genus in the subfamily Xoridinae, tribe Labiini
(Townes et al., 1960). The genus is restricted to the Western Hemi¬
sphere, with specimens recorded from southern Canada to northern
Argentina. The last revision of the genus was by Rohwer (1920), who
listed four North American species. Townes et al. (1960) provide re¬
descriptions and additional information concerning two of the species
The Pan-Pacific Entomologist 46: 50-63. January 1970
JANUARY 1970] SLOBODCHIKOFF—GROTEA REVISION
51
listed by Rohwer, and Townes and Townes (1966) list fives species in
the Neotropical region. The present revision treats eight species, three
previously undescribed.
Biological data on only two species, G. anguina Cresson and G.
californica Cresson, are available at present. Both of these species para¬
sitize the nest of bees of the genus Ceratina. It is probable that the
other six species in the genus parasitize twig-nesting aculeates.
Methods and Materials
All measurements were made at 40X with an American Optical Com¬
pany Spencer microscope and an ocular micrometer. Drawings were
made at the same magnification with an ocular grid. Holotypes of all
the described species were examined at the institutions where they are
deposited. The following abbreviations are used to designate collec¬
tions: AMNH, American Museum of Natural History, New York, N. Y.;
CAS, California Academy of Sciences, San Francisco; CIS, California
Insect Survey, University of California, Berkeley; Cornell, Cornell
University, Ithaca, N. Y.; Davis, University of California, Davis; MSU,
Michigan State University, East Lansing; Ottawa, Canadian National
Collection, Ottawa; Slobodchikoff, C. N. Slobodchikoff collection,
Berkeley, Calif.; Townes, H. K. Townes collection, American En¬
tomological Institute, Ann Arbor, Mich.; USNM, United States Na¬
tional Museum, Washington, D. C. The location of type specimens is
indicated by parentheses in the synonymy.
Genus Grotea Cresson
Grotea Cresson, 1864, Proc. Entomol. Soc. Philadelphia, 3: 397.
Type: Grotea anguina Cresson (Philadephia).
Front wing 6.0 to 14.0 mm long; frons convex; clypeus and labrum small;
upper end of prepectal carina near ventral corner of pronotum; petiole long and
slender, upcurved in its posterior 0.5, spiracle behind middle.
Key to Species of Grotea
1. Distal tip of front wing with conspicuous brown spot _ 2
Distal tip of front wing without, or with only faint brown spot _ 4
2. Antennal flagellum yellow on distal 0.2; mesopleuron in at least posterior
0.6 with horizontal orange-brown band; second pleural area of pro-
podeum 2 or more times longer than wide (Figs. 10, 14) _ 3
Antennal flagellum black on distal 0.2, orange proximally; mesopleuron
lacking horizontal orange-brown band; second pleural area of propodeum
1.5 times as long as wide (Fig. 13) _ G. fulva Cameron
3. Distal end of genal hook an acute angle (Fig. 3) ; pleural carina of pro¬
podeum absent or incomplete (Fig. 10) _
_ G. lokii Slobodchikoff, n. sp.
52
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46, NO. 1
Figs. 1-8. Genal hooks of Grotea spp., lateral view. Fig. 1, G. anguina;
Fig. 2, G. californica; Fig. 3, G. lokii; Fig. 4, G. mexicana ; Fig. 5, G. fulva;
Fig. 6, G. athenae; Fig. 7, G. delicator; Fig. 8, G. perplexa.
Distal end of genal hook spatulate, ellipsoid (Fig. 6) ; pleural carina of
propodeum complete __ G. athenae Slobodchikoff, n. sp.
4. Apical transverse carina of propodeum absent _ 5
Apical transverse carina of propodeum present (Fig. 12) _._
_ G. mexicana Cresson
5. Distal end of genal hook an acute angle (Figs. 1, 3) ; antennal flagellum
yellow with black band at 0.7 its length _ 6
Distal end of genal hook spatulate, ellipsoid (Figs. 2, 7, 8) ; antennal
flagellum otherwise _ 7
6. Pleural carina of propodeum absent or incomplete (Fig. 10) ; distal tip of
front wing with faint brown spot _ G. lokii Slobodchikoff, n. sp.
Pleural carina of propodeum complete (Fig. 11) ; distal tip of front wing
without faint brown spot _____ G. anguina Cresson
Figs. 9-16. Propodia of Grotea spp., dorsal view. Fig. 9, G. californica'. Fig.
10, G. lokii, Fig. 11, G. anguina'. Fig. 12, G. mexicana-, Fig. 13, G. fulva; Fig. 14,
G. athenae'. Fig. 15, G. delicator; Fig. 16, G. perplexa. ate, apical transverse
carina; 11c, lateral longitudinal carina; ma, 1st median area; pa, 2nd pleural
area; pc, pleural carina.
JANUARY 1970] SLOBODCHIKOFF-GROTEA REVISION
53
ma
/
/
15
1 m m
16
54
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
7. Second pleural area of propodeum open posteriorly (Fig. 16) ; propodeum
broadly quadrate in dorsal view; mesoscutum uniformly orange _
_ G. perplexa Slobodchikoff, n. sp.
Second pleural area of propodeum closed posteriorly (Figs. 9, 15) ; pro¬
podeum and mesoscutum otherwise ___ 8
8. Antennal flagellum black; second pleural area of propodeum curved up¬
ward strongly in its anterior 0.5 (Fig. 15) ; mesoscutum yellow with
median complete and pair of submedian incomplete brown-black longi¬
tudinal stripes _ G. delicator (Thunberg)
Antennal flagellum yellow; second pleural area of propodeum not curved
upward strongly in its anterior 0.5 (Fig. 9) ; mesoscutum orange with
position of notauli marked with yellow _ G. calif omica Cresson
Grotea Anguina Cresson
(Figs. 1, 11)
Grotea anguina Cresson, 1864, Proc. Entomol. Soc. Philadelphia, 3: 398; $, 2.
Grotea anguincr, Cresson, 1916, Mem. Amer. Entomol. Soc., 1: 16. Lectotype
designation. Lectotype: $, New Jersey (Philadelphia).
Biology.— Graenicher, 1905, Entomol. News, 16: 43-49.—Rau, 1922, Trans.
Acad. Sci. St. Louis, 24: 43; 1928, Ann. Entomol. Soc. Amer., 21: 383.
Female.— Front wing 9.0 to 11.0 mm long. Clypeus curving-rectangular, upper
and lower margins decurved; genal hook forming an acute angle, its basal part
approximately 2.5 times wider than at 0.66 the distance from base to tip. First
median area of propodeum triangular, distal angle of triangle acute; lateral longi¬
tudinal carina in front of spiracle complete; second pleural area of propodeum
approximately 3.5 as long as wide; pleural carina present; apical transverse
carina absent.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum yellow-orange, with black band at 0.7 its length. Pronotum yellow,
with yoke-like orange band dorsally, posterior margins black; mesopleuron yel¬
low, with broad horizontal orange band; mesoscutum orange, its lateral margins
black, with pair of median yellow stripes that form an elongate V; scutellum
and postscutellum yellow. Propodeum orange, with yellow markings laterally
and apically. Front and middle legs light yellow-orange; hind legs orange.
Petiole brown-black dorsally and orange ventrally, with lateral yellow stripes.
Abdominal sternites yellow; abdominal tergites brown-black medially and yellow
laterally. Distal tip of front wing lacking conspicuous brown spot.
Male.— Front wing 10.0 to 11.0 mm long. Similar to female in structure and
color.
The habits of this widespread eastern Nearctic species have been
studied by Graenicher (1905) and Rau (1922; 1928). Grotea anguina
attacks bees of the genus Ceratina , which construct nests consisting of
several to many cells in hollowed-out twigs with large pithy centers.
The parasite egg is laid on or near the bee egg in a cell. The parasite
larva consumes the bee larva, then consumes the bee-bread in that cell.
Subsequently the parasite larva destroys the partitions and contents
JANUARY 1970] SLOBODCHIKOFF—GROTEA REVISION
55
of one or more cells before it pupates in the twig and emerges the fol¬
lowing spring.
Collection data (4 $, 44 2) .—CONNECTICUT: Hartford, 2, (USNM). IN¬
DIANA: Lafayette, $, 14 April 1960, and 2, 20 July 1960, (USNM). IOWA:
Sioux City, 2, C. N. Ainslie (USNM). KENTUCKY: Golden Pond, 2, June
1965, H. K. Townes (Slobodchikoff). MARYAND: Beltsville, 2, 15 June 1913,
J. R. Malloch (USNM); Glen Echo, 2, 12 June 1921, R. M. Fonts (USNM);
Laurel, 2, 25 June 1933, E. B. Marshall (USNM); Meadowood, near Colesville,
2, 22 July 1961, L. M. Walkley (USNM); Plummers Island, 2 2, 1 June 1913,
H. S. Barber (USNM) and 9 June 1914, R. C. Shannon (USNM) ; Takoma
Park, 2, 28 May 1944, H. and M. Townes (Townes). MASSACHUSETTS:
Martha’s Vineyard, 2, 20 May 1937, F. M. Jones (USNM). MICHIGAN: Ann
Arbor, $, 6 June 1963 and 2 2, 6 June and 8 June 1963, H. and M. Townes
(Townes) ; Cheboygan Co., 2 2,6 July 1947, R. R. Dreisbach (MSU) ; E. Lansing,
2, 3 June 1963, R. B. Willson (MSU) ; Gull Lk. Biological Sta., Kalamazoo Co.,
3 2,23 June 1965, 12 July 1965, and 22 July 1966, R. L. Fischer (MSU) ; Midland
Co., 2, 9 June 1957, R. and K. Dreisbach (MSU), $, 5 to 11 June 1961, and
32, 12 to 21 June 1961, R. R. Dreisbach (MSU) ; Onsted, 2 2, 16 June 1959, G.
C. Eickwort (MSU) and R. A. Scheibner (MSU); Owasso, 2, 29 June 1953,
B. Wilson (MSU) ; Shiawasse Co., 2, 29 May 1959, R. and K. Dreisbach (MSU).
MISSISSIPPI: Stoneville, 2, 15 May 1958, K. Peterson (USNM). MISSOURI:
St. Louis, 2, Rau (USNM). NEW JERSEY: Moorestown, 2 2,4 June 1939 and
24 June 1939, H. and M. Townes (Townes); Ocean Grove, 2, (USNM). NEW
YORK: Buffalo, 2, 30 June 1934, Krombein (USNM) ; Ithaca, $, 10 June 1935,
H. K. Townes (Townes); New York, 2, (USNM). NORTH CAROLINA: Wake
Co., 2, 12 June 1951, H. and M. Townes (Townes). ONTARIO: Cornwall, 2,
29 June 1925, F. Ide (Ottawa); Wakefield, 2, 25 June 1946, G. S. Walley (Ot¬
tawa). PENNSYLVANIA: Dauphin, 2, May 1930, J. N. Knull (USNM); Per-
dix, 2,21 May 1911, W. S. Fisher (USNM) ; Stoverdale, 2, 13 June 1916, W. S.
Fisher (USNM). SOUTH CAROLINA: Greenville, 2, 20 June 1940, H. and G.
Townes (Townes) ; Wattacoo, near Cleveland, ?, 17 May 1959, G. F. Townes
(Townes). TENNESSEE: Knoxville, 2, 1921, C. N. Ainslie (USNM). TEXAS:
Brownsville, 2, C. H. T. Townsend (USNM); Calvert, ?, June 1961, W. W.
Yothers (USNM). VIRGINIA: Arlington, 2, 4 June 1943, R. A. C. (USNM);
Falls Church, 2 2, 30 May and 5 July, N. Banks (USNM). WASHINGTON,
D. C.: Metro area, 2, summer 1966, T. J. Spilman (USNM).
Grotea athenae Slobodchikoff, new species
(Figs. 6,14)
Female. —Front wing 10.0 mm long. Clypeus curving-rectangular, upper mar¬
gin more decurved than lower margin so that clypeus appears faintly pentagonal;
genal hook spatulate, its distal part broadly ellipsoid, its basal part as wide as at
0.66 the distance from base to tip. First median area of propodeum subrectangular;
lateral longitudinal carina in front of spiracle complete; second pleural area of
propodeum approximately 2.5 times as long as wide; pleural carina present;
apical transverse carina absent.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum yellow-orange, with black band at 0.7 its length. Pronotum yellow
56
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
with incomplete yoke-like brown band dorsolaterally; posterior margins of pro-
notum black; mesopleuron yellow, with broad horizontal orange-brown band;
mesoscutum orange-brown, its lateral margins black, with pair of broad median
yellow stripes that form an elongate V and enclose a median brown stripe be¬
tween them; scutellum and postscutellum yellow. Front and middle legs yellow
on anterior face, orange on posterior face. Petiole brown-black on anterior 0.7,
with pair of dorsolateral yellow stripes, and orange on posterior 0.3. Abdominal
sternites and tergites orange. Distal tip of front wing with conspicuous brown
spot.
Male. —Unknown.
Nothing is known about the habits or hosts of this species.
Holotype female , Monteverde, Costa Rica, 9 February 1962, C.
Palmer (Townes).
Paratype female, MEXICO: Veracruz: Orizaba, 12 to 22 August
1961, R. and K. Dreisbach (MSU).
Grotea californica Cresson
(Figs. 2, 9)
Grotea californica Cresson, 1878, Proc. Acad. Nat. Sci. Philadelphia, 1878: 370;
Type: $, California (Philadelphia).
Biology.— Daly et al., 1967, Ann. Entomol. Soc. Amer., 60: 1273-1282.—Slobod-
chikoff, 1967, Pan-Pac. Entomol., 43: 161-168.
Female.— Front wing 8.0 to 11.0 mm long. Clypeus curving-rectangular, upper
margin slightly more decurved than lower margin so that clypeus appears faintly
pentagonal; genal hook spatulate, its distal part ellipsoid, its basal part as wide
as at 0.66 the distance from base to tip. First median area of propodeum rec¬
tangular or subrectangular, distance between the two median longitudinal carinae
in apical part of median area greater than their combined width; lateral longi¬
tudinal carina in front of spiracle complete; second pleural area of propodeum
approximately 3.3 as long as wide; pleural carina present; apical transverse
carina absent.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum orange on proximal 0.7, yellow on distal 0.3. Pronotum yellow, with
yoke-like orange-black band dorsolaterally and posterior margins black; meso¬
pleuron yellow, with broad horizontal orange-black band; mesoscutum orange,
its lateral margins black, with pair of median yellow stripes that form an
elongate V. Propodeum orange, with yellow markings laterally and apically.
Front and middle legs light yellow-orange; hind legs orange. Petiole brown-
black dorsally and orange ventrally, with lateral yellow stripes. Abdominal
sternites yellow-orange; abdominal tergites brown-black medially and yellow
laterally. Distal tip of front wing with faint brown spot.
Male.— Front wing 6.0 to 10.0 mm long. Similar to female in structure and
color.
The parasitic habits of this species, studied by Daly et al. (1967)
and Slobodchikoff (1967), are similar to those of Grotea anguina. The
female oviposits through the wall of a twig into a cell. Usually cells
JANUARY 1970] SLOBODCHIKOFF-—GROTEA REVISION
57
in the lower third of the nest are preferred. An average of two to three
cells, together with their contents, are destroy by the parasite larva.
The pupal cell of the parasite consists of a membranous web and thick
partitions at its upper and lower ends in the twig. Bees in cells below
the parasite pupal cell are unable to penetrate the partitions and cannot
escape from the nest. Often one third of the progeny in a nest may
be destroyed indirectly in this manner.
Collection data (39$, 30$).—BRITISH COLUMBIA: Robson, $, 29 June
1950, H. R. Foxlee (Ottawa) ; Ruskin, $, 1 July 1953, W. R. M. Mason (Ottawa).
CALIFORNIA: Alameda Co.: $, Coquillett (USNM) ; Berkeley, $, 15 June
1933, (Davis), $, 27 June 1933, G. E. and R. M. Bohart (Davis), June 1939,
(Townes), $, 17 June 1933, M. Cazier (USNM), $, 19 May 1966, J. K. Ryan
(Slobodchikoff), ?, 27 May 1934, G. E. and R. M. Bohart (Davis), ?, 3 May
1963, D. C. Rentz (Slobodchikoff); East Oakland, $, 12 June 1948, L. L. Jen¬
sen (Townes); Oakland, $, 12 June 1933, G. E. and R. M. Bohart (Davis), $,
25 May 1937, E. S. Ross (Townes) ; San Leandro, $, June 1948, C. A. Downing
(Townes), ?, 14 April 1948, C. A. Downing (Davis). Amador Co.: Volcano, 3$,
4 June 1961, R. M. Bohart (Davis). Contra Costa Co.: near Orinda, 7$, 11$,
collected over one year period, H. V. Daly (CIS). El Dorado Co.: Camino, $,
29 June 1948, H., M., G., and D. Townes (Townes). Humboldt Co.: Blocksburg,
$, 19 June 1935, E. W. Baker (USNM) ; Blue Lake, $, 20 to 27 June 1907, Brad¬
ley (Townes). Lassen Co.: Hallelujah Jet., $, 4 July 1968, R. M. Bohart
(Davis). Los Angeles Co.: Los Angeles, $, Coquillett (USNM). Marin Co.:
Inverness, $, 24 June 1961, C. A. Toschi (Slobodchikoff); Mill Valley, 4$, 1$,
14 May 1961, D. Q. Cavagnaro (Davis). Napa Co.: Samuel Springs, $, 9 May
1955, R. M. Bohart (Davis), $, 28 April 1956, S. M. Fidel (Davis). Orange Co.:
Green River Camp, $, 8 May 1933, E. P. Van Duzee (CAS). Sacramento Co.:
Carmichael, $, 13 May 1961, T. Gantenbein (Davis). San Bernardino Co.:
Snow Crest Camp, $, 7 July 1952, R. M. Bohart (Davis). San Lrancisco Co.:
San Francisco, $, 1 May 1947, E. S. Ross (CAS). San Luis Obispo Co.:
Atascadero, 2 $ , 1 $ , 31 May 1948, C. H. Dickenson (Townes), $ , 17 June 1948, C.
H. Dickenson (Townes). San Mateo Co.: Daly City, ?, 30 June 1961, A. K.
Lehre (Slobodchikoff). Santa Barbara Co.: Santa Ynez Mts., $, 24 June 1928,
P. M. Marsh (Davis). Santa Clara Co.: Alum Rock Park, $, 25 May 1950, J.
MacSwain (Townes). Santa Cruz Co.: Santa Cruz, ?, 1 June 1936, G. E.
and R. M. Bohart (Davis). Siskiyou Co.: $, 2 June 1911, F. W. Hunon-
rnaches (USNM). Solano Co.: Putah Canyon, $, 20 April 1959, F. D.
Parker (Davis). Trinity Co.: Carrville, 2 $, 15 and 27 May 1934, G. E. and R.
M. Bohart (Davis). Ventura Co.: Ventura, $, 4 May 1961, Gillogley (USNM).
Yolo Co.: Elkhorn Ferry, $, 12 May 1952, R. M. Bohart (Davis). IDAHO:
Banner Co.: Sandpoint, $, 21 June 1955, R. M. Bohart (Davis). NEVADA:
Douglas Co.: Valley Hot Spring, $, $,24 May 1953, R. M. Bohart (Davis).
Washoe Co.: Verdi, S, 26 June 1962, R. M. Bohart (Davis), $, 27 June 1966,
R. L. Brumley (Davis). OREGON: Benton Co.: Corvallis, $, 9 June 1909, J.
C. Bridwell (USNM), $, 23 May 1939, L. G. Hudson (USNM).
58
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Grotea delicator (Thunberg)
(Figs. 7, 15)
Ichneumon delicator Thunberg, 1822, Mem. Acad. Imp. Sci. St. Petersbourg, 8:
259; 1824, Ibid, 9: 307, sex not given. Type: sex ? (abdomen missing),
(French West Indies: Saint) Barthelemy Island (Uppsala).
Atractodes lineatus Brulle, 1846, In Lepeletier: Historic naturelle des insectes,
Hymenopteres, 4: 167. 2- Lectotype: $, “Guiana” (Paris).
Grotea lineata ; Krieger, 1903, Z. syst. Hymen. Dipt., 3: 290. New synonym.
Grotea delicator-, Roman, 1912, Zool. Bidrag Uppsala, 1: 249.
Female. —Front wing 7.0 to 11.5 mm long. Upper margin of clypeus strongly
decurved, clypeus appears hemispheroid; genal book spatulate, its distal part
ellipsoid, its basal part approximately 1.3 times wider than at 0.66 the distance
from base to tip. First median area of propodeum subtriangular; lateral longi¬
tudinal carina in front of spiracle complete; second pleural area of propodeum
curved upward strongly in anterior 0.5, approximately 2.5 times as long as wide;
pleural carina present; apical transverse carina absent.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum black. Pronotum yellow, black along posterior margins; mesopleuron
yellow; mesoscutum yellow, its lateral edges black, with median and pair of
submedian longitudinal orange stripes; scutellum and postscutellum yellow.
Propodeum yellow, with irregularly-shaped black band that includes first median
area, second pleural areas, and portion of propodeum posterior to basal transverse
carina. Front and middle legs yellow; hind legs orange. Petiole yellow, dorsally
with median and pair of sublateral longitudinal black stripes on anterior 0.75 and
orange on posterior 0.25. Abdominal sternites yellow; abdominal tergites orange.
Distal tip of front wing lacking conspicuous brown spot.
Male. —Front wing 7.5 to 10.0 mm long. Similar to female in structure and
color.
Nothing is known about the habits or hosts of this species.
Collection data (5d, 19 2)-—ARGENTINA: Berisso, 2, 8 December 1965,
H. and M. Townes (Townes); Entre Rios: Pronunciamiento, 2, September
1964, and 22, 3d, January 1965 (Ottawa); Jujuy: San Sebastian de Jujuy,
22, 14 January 1966, H. and M. Townes (Townes); Tucuman: S. Pedro d.
Colalao, 2, January 1951, Arnau (Ottawa). BRAZIL: Santa Catarina: Nova
Teutonia, 27°1P S, 52°23' W, 4 2, 16 October 1949, 28 November 1957, 23 De¬
cember 1959, and 9 October 1967, 1 d, 11 January 1957, F. Plaumann (Ottawa) ;
Nova Teutonia, Id, 12, 1?, 24 November 1955, 29 November 1955, and 4
December 1955, F. Plaumann (Townes). SURINAM: Kwatta, 2, 28 February
to 22 March 1964, D. C. Geijskes (Townes) ; Paramaribo, 2 2, 26 September 1963,
Broekhuizen, and 30 November 1963, D. C. Geijskes (Townes). WEST INDIES:
Trinidad: Arima Valley, 2 2, 22 May 1952 and 28 December 1952, (AMNH) ;
Arima, 2? 23 March 1961, N. Gopaul (Ottawa); Piarce, 2, 27 February 1961,
N. Gopaul (Ottawa).
Grotea fulva Cameron
(Figs. 5,13)
Grotea fulva Cameron, 1886, Biologia Centrali-Americana, Insecta, Hymenoptera,
1: 309. Type: 2, Mexico: Presidio (London).
JANUARY 1970] SLOBODCHIKOFF—GROTEA REVISION
59
Female. —Front wing 12.0 to 13.6 mm long. Clypeus curving-rectangular, up¬
per margin only slightly more decurved than lower margin; genal hook spatulate,
its distal part ellipsoid, its basal part approximately 1.3 times wider than at 0.66
the distance from base to tip. First median area of propodeum subrectangular;
lateral longitudinal carina in front of spiracle complete; second pleural area of
propodeum approximately 1.5 times as long as wide; pleural carina present;
apical transverse carina absent.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum orange on proximal 0.8, black on distal 0.2. Pronotum uniformly yellow
anteriorly, with posterior margins black; mesopleuron yellow, with black spot
near posterior margin; mesoscutum yellow, its lateral margins black, with median
longitudinal black stripe and pair of submedian longitudinal orange stripes;
scutellum and postscutellum yellow. Propodeum yellow, with irregularly-shaped
black band that includes first median area, second pleural areas, and small por¬
tion of propodeum posterior to basal transverse carina. Front and middle legs
yellow, hind legs orange. Petiole yellow-orange with dorsal and pair of lateral
black stripes. Abdominal sternites yellow; abdominal tergites orange medially
and yellow laterially. Distal tip of front wing with conspicuous brown spot.
Male.— Unknown.
In the material examined, one specimen from Coca, Ecuador is
morphologically similar to the type but represents a color variant. Its
description is as follows:
Head yellow, with black behind antennal sockets, around ocelli, and posteriorly
between eyes; scape, pedicel, and first flagellar segment black dorsally, orange
ventrally; antennal flagellum orange on proximal 0.8, black on distal 0.2. Prono¬
tum yellow, posterior margins black, with incomplete black band dorsolaterally;
mesopleuron yellow, with median black band on posterior 0.5; mesoscutum black,
with pair of yellow median stripes marking position of notauli and pair of yel¬
low lateral stripes anterior to tegulae; scutellum and postscutellum yellow.
Propodeum yellow, with dorsal mask-like black band extending over first, second
median areas, second pleural areas, second lateral areas, and portions of the first
lateral areas. Front and middle legs yellow anteriorly, black with yellow mark¬
ings posteriorly; hind coxae yellow with mediolateral incomplete black stripe
anteriorly, black posteriorly; first hind trochanter black on proximal 0.8, yellow
distally; second hind trochanter yellow; hind femora yellow with median longi¬
tudinal black stripe anteriorly, black posteriorly; hind tibiae and tarsi black.
Dorsum of petiole yellow on posterior 0.1, black on anterior 0.9 with pair of
dorsolateral yellow stripes. Abdominal sternites yellow; abdominal tergites
black anteriorly, yellow on posterior 0.4. Distal tip of front wing with con¬
spicuous brown spot.
Nothing is known about the habits or hosts of this species.
Collection data (4 2, 1?).—ECUADOR: Coca, 2, May 1965, L. Pena
(Townes); Ecuador or Peru, 2, 1963, L. E. Pena (Townes). MEXICO: Yuca¬
tan: Temax, ?, 1904, Godman-Salvin (London). PANAMA: Rovira, 2, 4
February 1960, V. W. Brown (Townes). PERU: 10 km. S. of Chiclayo, 2, 21
March 1951, Ross and Michelbacher (CAS).
60
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
Grotea lokii Slobodchikoff, new species
(Figs. 3, 10)
Female. —Front wing 7.0 to 10.5 mm long. Clypeus curving-rectangular, upper
margin slightly more decurved than lower margin so that clypeus appears faintly
pentagonal; genal hook forming an acute angle, its basal part approximately 2.0
times wider than at 0.66 the distance from base to tip. First median area of pro-
podeum subtriangular to subrectangular; lateral longitudinal carina in front of
spiracle complete or incomplete; second pleural area of propodeum approximately
4.0 times as long as wide; pleural carina absent or incomplete; apical transverse
carina absent.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum orange, with black band at 0.7 its length. Pronotum yellow, with
ellipsoid black band dorsally, posterior margins black; mesopleuron yellow, with
horizontal orange or orange-black band; mesopleuron yellow, with horizontal
orange or orange-black band; mesoscutum orange, its lateral margins black, with
pair of median yellow stripes that form an elongate V; scutellum and post-
scutellum yellow. Propodeum orange, with yellow markings laterally and apically.
Front and middle legs yellow, hind legs yellow-orange. Petiole brown or black
dorsally and orange ventrally, with lateral yellow stripes. Abdominal sternites
yellow; abdominal tergites brown or black medially and yellow laterally. Distal
tip of front wing with conspicuous brown spot.
Male. —Front wing 6.5 to 11.5 mm long. Similar to female in structure and
color.
The above description is based on the type specimen, but incorporates
the variation found in the paratype series. This species was confused
with Grotea anguina by Townes et al. (1960, pp. 538-540). Nothing
is known about the habits or hosts of the species.
Holotype female , San Pedro Iturbide, 32± icm. west of Linares,
Neuvo Leon, Mexico, 5 October 1962, H. and M. Townes (Townes).
Allotype, same data as type (Townes).
Paratypes (12$, 16$).—ARIZONA: Coconino Co.: 10 mi. W. Jacob Lake,
$, 6 June 1946, R. M. Bobart (Townes) ; Oak Creek Canyon, $, 18 May 1947,
H. and M. Townes (Townes). Gila Co.: Parker Creek, Sierra Ancha, $, 9 May
1947, H. and M. Townes (Townes). Santa Cruz Co.: Sonoita, $, 30 August 1954,
R. M. Bohart (Davis). COLORADO: Boulder Co.: Lyons, $, 25 June 1962, R.
and K. Dreisbach (MSU). Larimer Co.: near Estes Pk., $, 14 June 1948, H.,
M., G., and D. Townes (Townes). Routt Co.: Steamboat Sprs., $, 5 July 1962,
R. and K. Dreisbach (MSU). MEXICO: Distrito Federal: Atzcapolzalco, $, 5
July 1931 (Cornell). Durango: 5 mi. W. Durango, $, 14 May 1962, L. A.
Stange (Davis) ; 6 mi. S. Durango, $, 5 July 1964, W. R. M. Mason (Ottawa) ;
18 mi. W. Durango, $, 31 July 1964, H. F. Howden (Ottawa). Hidalgo: Pachuca,
$, 29 July 1954, J. G. Chillcott (Ottawa). Jalisco: Tamozula, $, 10 February
1954, R. and K. Dreisbach (MSU). Mexico: Teotihuacan Pyr., $, 7 July 1951,
H. E. Evans (Townes); Tepexpan, $, $,26 July 1963, F. D. Parker and L. A.
Stange (Davis). Michoacan: 6 mi. N. W. Quiroga, 2$, 11 July 1963, F. D.
JANUARY 1970] SLOBODCHIKOFF—GROTEA REVISION
61
Parker and L. A. Stange (Davis). Morelos: Cuautla, 2, 30 October 1922, E. G.
Smith (USNM) ; Cuernavaca, 2, 30 May 1959, H. E. Evans (Cornell), $, 2, 29
July 1961, R. and K. Dreisbach (Townes). Nuevo Leon: San Pedro Iturbide,
32± km. W. Linares, 2,5 October 1962 and 3 $, 12, 6 October 1962, H. and M.
Townes (Townes). Sinaloa : 6 mi. N. E. Potrerillos, 2, 19 March 1962, L. A.
Stange (Davis). Sonora: Alamos, $, 25 February 1963, P. D. Arnaud, Jr. (CAS).
Vera Cruz: “San Rafael Jilotepec,” 2, (USNM).
Grotea MEXICANA Cresson
(Figs. 4, 12)
Grotea mexicana Cresson, 1874, Proc. Acad. Nat. Sci. Philadelphia 1873: 418.
$, 2- Lectotype: 2, Mexico: Orizaba (Philadelphia).
Grotea mexicana ; Cresson, 1916, Mem. Amer. Entomol. Soc., 1: 42.
Lectotype designation.
Female.— Front wing 7.0 to 9.5 mm long. Upper margin of clypeus strongly
decurved, so that clypeus appears hemispheroid; genal hook forming an acute
angle, its basal part approximately 1.8 times wider than at 0.66 the distance from
base to tip. First median area of propodeum subtriangular; lateral longitudinal
carina in front of spiracle complete; second pleural area of propodeum curved
strongly upward apically, approximately 1.7 as long as wide; pleural carina
present; apical transverse carina present.
Head yellow, suffused with orange around ocelli and behind eyes; antennal
flagellum orange, with black band at 0.7 its length. Pronotum yellow, with
ellipsoid orange-brown band dorsally; mesopleuron yellow, without distinct
horizontal band; mesoscutum uniformly orange medially, its lateral margins
black; scutellum and postscutellum yellow. Propodeum orange, with yellow
lateral and apical markings. Front and middle legs yellow, hind legs orange.
Petiole orange dorsally and ventrally, with lateral yellow stripe. Abdominal
sternites yellow; abdominal tergites orange. Distal tip of front wing lacking
conspicuous brown spot.
Male. —Front wing 7.0 to 8.0 mm. Similar to female in structure and color.
Nothing is known about the habits or the hosts of this species.
Collection data (6$, 42-—MEXICO ?, (USNM). Morelos: 3 mi. N.
Alpuyeca, 2, 8 April 1959, H. E. Evans (Cornell). Sinaloa: 8 mi. S. E. Elota,
2, 19 April 1962, L. A. Stange (Davis). Vera Cruz: Cordoba, 2, 6 July 1962,
J. S. Buckett, M. R. and R. C. Gardner (Davis) ; Jalapa, 5$, 1 to 6 August, R.
R. Dreisbach (MSU 4$, Townes 1 $) ; Orizaba, $, 13 February 1954, R. R.
Dreisbach (MSU), 2, 13 February 1954, R. R. Dreisbach (Townes).
Grotea perplexa Slobodchikoff, new species
(Figs. 8, 16)
Female.— Front wing 11.0 mm long. Clypeus pentagonal; labrum lacking
dorsomedial pit; genal hook quadrate, its basal part approximately as wide as at
0.66 the distance from base to tip. First median area of propodeum subrectangular,
2 times as long as wide at its base; lateral longitudinal carina in front of spiracle
incomplete; second pleural area of propodeum open, extending back to posterior
62
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
margin of propodeum, approximately 2 times as long as wide; pleural carina
present; apical transverse carina absent.
Head white, suffused with brown around ocelli and behind eyes; antennal
flagellum black. Pronotum orange, white on anterior 0.1 and yellow along dorso-
posterior margins; mesopleuron orange with yellow spot along anterodorsal mar¬
gin; mesoscutum orange; metapleuron white; scutellum and postscutellum yel¬
low. Propodeum orange, apical 0.1 white. Front legs white, femora with orange
band on median 0.7 of posterior face; middle coxae white; first trochanter brown
on anterior face, white on posterior; second trochanter white; anterior face of
middle femora white, posterior face white on distal 0.6, orange on proximal 0.4;
middle tibiae white, suffused with orange along inner faces; anterior face of hind
coxae white with orange band on median 0.3, posterior face orange on proximal
0.7 and white on distal 0.3; first trochanter orange, white on distal 0.4; second
trochanter white; hind femora orange, white on distal 0.2; hind tibiae grey,
white on distal 0.2; middle and hind tarsi grey, white on distal 0.2. Petiole
orange. Abdominal sternites white; abdominal tergites brown-black, tergites 2-6
each with median cuneate white spot near posterior margin. Distal tip of front
wing lacking conspicuous brown spot.
Male.— Unknown.
This species is known from only the type specimen. Nothing is
known about its hosts or habits.
Holotype female , Nova Teutonia, 27°11 / S, 52°23' W. Santa
Catarina, Brazil, 31 December 1959, F. Plaumann (Ottawa).
Acknowledgments
The author would like to thank Dr. H. K. Townes, American En¬
tomological Institute, Ann Arbor, Michigan, for providing specimens,
suggestions, and criticism of this manuscript, and Dr. Jerry Powell,
University of California, for critically reading the manuscript. The
author would also like to thank the following people for the loan of
specimens: Dr. R. L. Fischer, Michigan State University; Mr. H. B.
Leech, California Academy of Sciences; Dr. W. R. M. Mason, Cana¬
dian National Collection, Ottawa; Dr. J. G. Rozen, American Museum
of Natural History, New York; Mr. R. O. Schuster, University of
California, Davis; and Dr. Luella Walkley, U. S. National Museum.
Literature Cited
Daly, H. V., G. I. Stage, and T. Brown. 1967. Natural enemies of bees of the
genus Ceratina. Ann. Entomol. Soc. Amer., 60(6) : 1273-1282.
Graenicher, S. 1905. On the habits of two ichneumonid parasites of the bee
Ceratina dupla Say. Entomol. News, 16: 43-49.
Rau, P. 1922. Ecological and behavioral notes on Missouri insects. Trans.
Acad. Sci. St. Louis, 24(7) : 1-71.
1928. The nesting habits of the little carpenter bee, Ceratina calcarata.
Ann. Entomol. Soc. Amer., 21: 380-397.
Rohwer, S. A. 1920. The North American ichneumon-flies of the tribes
JANUARY 1970]
ZOOLOGICAL NOMENCLATURE
63
Labenini, Rhyssini, Xoridini, Odontomerini, and Phytodietini. Proc.
U. S. Nat. Mus., 57: 405-474.
Slobodchikoff, C. N. 1967. Bionomics of Grotea californica Cresson, with a
description of the larva and pupa. Pan-Pac. Entomol., 43: 161-168.
Townes, H. K. and M. Townes. 1966. A catalogue and reclassification of the
Neotropic Ichneumonidae. Mem. Amer. Entomol. Inst., 8: 1-367.
Townes, H. K., M. Townes, and others. 1960. Ichneumon-flies of America
north of Mexico: 2. Subfamilies Ephialtinae, Xoridinae, Acaenitinae.
U. S. Nat. Mus. Bull., 216(2): 1-676.
ZOOLOGICAL NOMENCLATURE: Announcement A. (n.s.) 85
Required six-month’s notice is given on 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. 26, pt. 2, 8 August 1969) :
1864. Suppression of Acarus telarius Linnaeus, 1758, and of neotype designa¬
tion for that species (Acarina).
1650. Type-species for Poekilocerus Audinet-Serville, 1831, and Pamphagus
Thunberg, 1815; Grant of precedence to PYRGOMORPHIDAE Brunner
von Wattenwyl, 1874, over POEKILCERIDAE Burmeister, 1840, and
PHYMATEIDAE Burmeister, 1840 (Insecta, Orthoptera).
1867. Suppression of Huebner’s pamphlet Der Schmetterlinge Lepidoptera Lin-
naei europdisches Heer, circa 1790-1793.
1875. Suppression of Papilio saportae Huebner, 1828/32 (Insecta, Lepidoptera).
1876. Type-species for Agrotiphila Grote, 1875 (Insecta, Lepidoptera).
(see Bull. Zool. Nomencl. 26, pts. 3/4, 24 October 1969) :
1881. Type-species for Platybunus C. L. Koch, 1839 (Arachnida).
1882. Type-species for Rybaxis Saulcy, 1876 (Insecta, Coleoptera).
1885. Suppression of Scaeva arcuata Fallen, 1817 (Insecta, Diptera).
Comments should be sent in duplicate, citing case number, to the Secretary,
International Commission on Zoological Nomenclature, c/o British Museum (Nat¬
ural History), Cromwell Road, London S.W.7, England. Those received early
enough will be published in the Bulletin of Zoological Nomenclature. —W. E.
China, Assistant Secretary to the International Commission on Zoological Nomen¬
clature.
64
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
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Golden Gate Park
San Francisco, California 94118
JANUARY 1970]
PROCEEDINGS
65
PACIFIC COAST ENTOMOLOGICAL SOCIETY
K. S. Hagen R. W. Thorp M. S. Wasbauer P. H. Arnaud, Jr.
President President-elect Secretary Treasurer
Dr. Charles P. Alexander—Honored Member. —By unanimous action of the
Executive Board of the Pacific Coast Entomological Society at a meeting held on
21 November 1969, Dr. Charles Paul Alexander, distinguished student of the
Tipulidae and world renowned entomologist, was elected an Honored Member of
this Society.
Dr. Charles P. Alexander—Honored Member.
This honor in recognition of 60 years of work and continuous publication on the
Tipulidae, also coinciding with Dr. Alexander’s 80th birthday year, is but one of
the many honors that Dr. Alexander has received from the world entomological
fraternity.
Dr. Alexander is one of the world’s most productive systematists, having described
over 1% of the known insect species. As of 1 September 1969, he had published
932 papers on Diptera, totaling over 15,000 pages with over 12,500 figures (all
drawn by Dr. Alexander with the exception of 180 figures). These papers have
appeared in 125 separate publications in more than 30 countries and cover all
regions of the world. The two leading publications containing Alexander papers are
the Philippine Journal of Science, 1922-1969, with 89 papers (including descrip¬
tions of 2,056 new species) and the Annals and Magazine of Natural History, 1920-
1961, with 113 papers (including descriptions of 1,891 new species). A total of
66
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
9,923 new species of Diptera (all but about 100 being in the single family Tipuli-
dae) are described in the 932 papers mentioned above. It is believed that this
figure vastly exceeds the number of new species in a single family, either in zool¬
ogy or botany, ever described by a single person. One of Dr. Alexander’s recent
comprehensive works “The Crane Flies of California” published in the Bulletin of
the California Insect Survey, volume 8, pages 1-269, text figs. 1-524, 1967, is of
great value to western entomologists.
Dr. Alexander was elected to membership in the Pacific Coast Entomological
Society at the meeting of 9 November 1951. He had previously subscribed to the
Pan-Pacific Entomologist with the beginning of volume 3, in July 1926, and con¬
tributed two papers to that volume. Dr. Alexander is the nineteenth person to be
elected an Honored Member of the Society during the past 58 years. The name,
of previously elected Honored Members, listed alphabetically, with year of election,
are as follows: F. E. Blaisdell (1938), Lawrence Bruner (1935), C. D. Duncan
(1966), Alice Eastwood (1912), E. 0. Essig (1948), G. F. Ferris (1948), L. 0.
Howard (1912), 0. B. Johnston (1911), Vernon Kellogg (1912), E. G. Linsley
(1968), E. R. Leach (1948), A. E. Michelbacher (1968), R. C. Miller (1968),
J. J. Rivers (1911), R. L. Usinger (1968), E. P. Van Duzee (1938), E. C. Van
Dyke (1938) and C. W. Woodworth (1912).—P. H. Arnaud, Jr., California Acad¬
emy of Sciences, San Francisco.
Proceedings
Three Hundred and Twenty-Fourth Meeting
The 324th meeting was held Friday, 10 January 1969 at 7:45 p.m. in the Morrison
Auditorium of the California Academy of Sciences, Golden Gate Park, San Fran¬
cisco, with President Hagen presiding.
Members present (49) : R. P. Allen, P. H. Arnaud, Jr., J. Benet, F. L. Blanc,
I. Boussy, T. Briggs, R. G. Brownlee, G. R. Buckingham, D. Burdick, L. E.
Caltagirone, P. Cammer, J. A. Chemsak, J. Cope, H. V. Daly, D. G. Denning, J. G.
Edwards, W. E. Furguson, M. R. Gardner, P. R. Grimstad, J. Guggolz, J. F. Gustaf¬
son, J. D. Haddock, K. S. Hagen, T. E. Hewton, K. Horn, D. S. Horning, W. H.
Lange, H. B. Leech, R. Lem, K. Lorenzen, J. W. MacSwain, A. R. Moldenke, J. A.
Powell, D. Rentz, D. Ribble, E. S. Ross, F. J. Santana, T. A. Schultz, T. A. Sears,
J. E. Slansky, C. N. Slobodchikoff, R. E. Stecker, R. L. Tassan, J. W. Tilden, W. J.
Turner, D. Ubick, M. S. Wasbauer, S. P. Welles, Jr., R. F. Wilkey, S. C. Williams.
Visitors present (32) : C. Adamson, Ana Caltagirone, K. Chen, Renae Cottam,
C. Cushner, Janie Cushner, Mr. and Mrs. P. Dana, Loretta Denning, Alice and
Janie Edwards, R. Fujii, R. Gardner, C. Glaser, Katie Henry, J. Hjelle, Carol
Horning, G. Leung, Shari Lewis, Alison Moldenke, R. Papp, W. Rausden, Kathy
Rentz, T. Rycraft, R. Smith, Anne Suguitan, Sharon Tanner, Cathy Tassan, D. Tie-
mann, B. Tilden, R. M. Vierhus, Judy Weller.
The minutes of the meeting held 20 December 1968 were summarized.
The following names were proposed for membership: Renae Cottam, William A.
Clark, John G. Franclemont; and for student membership, Bob Lem and Darrell
Ubick.
President Hagen called for introductions. Mike Gardner introduced Mr. Darwin
JANUARY 1970]
PROCEEDINGS
67
Tiemann from China Lake, California, who has worked on Phengodidae, especially
the genus Zccrhipis.
President Hagen called for notes and exhibits. Mr. Leech showed slides of a
common lampyrid beetle usually found early in the spring near streams and in
damp places. The immature stages have never been described. Just recently, what
appears to be the fully mature larva was brought in. When disturbed it rolls up
very much like a cutworm. It feeds on snails.
Mr. Leech also showed slides of Mr. Jack Balfour-Browne, the world specialist
on water beetles, who just retired from the British Museum and stopped by the
Academy on his way to Australia. He has been deaf since birth but is an accom¬
plished lip reader.
Dr. J. F. Gustafson showed slides of a fire damaged Yellow Pine tree. The tree
had a fire scar about 15 ft. long and 3 ft. wide at the base. (Buprestid and
Cerambycid beetles had tunneled in the scar.) Woodpeckers then made holes in
the scar in attempting to get the boring beetles and a Peromyscus mouse enlarged
one of the woodpecker holes to make its nest, an interesting case of succession.
Dr. Edwards showed some slides portraying the effects of blackfly bites on the
hand of one of the members of a recent expedition to Yucatan.
The principal speaker of the evening was Dr. Evert I. Schlinger, Chairman,
Department of Entomology, University of California at Riverside. His illustrated
talk was entitled “An Entomologist’s View of Chile—The University of California—
University of Chile Cooperative Project.”
A social hour was held in the entomology rooms following the meeting.—M. S.
Wasbauer, Secretary.
Three Hundred and Twenty-Fifth Meeting
The 325th meeting was held Friday, 21 February 1969 in the Morrison Audi¬
torium of the California Academy of Sciences, Golden Gate Park, San Francisco,
with President Hagen presiding.
Members present (29) : R. P. Allen, P. H. Arnaud, Jr., J. Benet, I. Boussy, T.
Briggs, G. Buckingham, D. Burdick, P. Cammer, J. A. Chemsak, J. G. Edwards,
J. F. Emmel, W. Gagne, F. Gustafson, T. E. Hewton, Jr., R. L. Langston, H. B.
Leech, R. Lem, K. Lorenzen, R. Main, D. Munroe, P. Opler, D. W. Ribble, E. S.
Ross, S. B. Ruth, F. J. Santana, Oakley Shields, Joanne E. Slansky, R. L. Tassan,
J. W. Tilden, W. J. Turner, D. Ubick, M. S. Wasbauer.
Guests present (20) : C. Adamson, C. Baum, E. E. Bedecanax, Alice Edwards,
R. C. Gardner, G. D. Hanna, R. B. Hewitt, R. Huek, M. Marquis, J. Milstead, W.
Pitcher, C. E. Reiner, Bessie D. Ruth, Suriya Sampunth, Phyllis Stecker, Emily
Stobbe, L. E. Stotelmyre, R. P. Swartzell, Cathy Tassan, Betty White.
The minutes of the meeting held 10 January were summarized.
Four new members were elected: Walter G. Goodman, Robert B. Hewitt, David
R. Meissner, Oakley Shields.
Dr. Joel Gustafson reported that the salt marsh investigations are progressing
very well. Two salt marsh investigations are being carried on locally at present.
One of these is being conducted by Guy Cameron, a graduate student at U.C. Davis.
His study area is next to Petaluma Creek on San Pablo Bay. The other is being
done by Robert Lane of San Francisco State College. His area is next to the
Dumbarton Bridge on the south bay. The two investigators have compared notes
63
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
and find that their areas are very different in faunal composition. For example,
Cameron’s area has no tipulids, whereas Lane’s has many. Cameron’s has anthicid
beetles, but Lane’s has none. Cameron’s has many thrips and Lane’s has only one.
Cameron’s has a few staphylinids and Lane has found none. Cameron has found no
dolichopodid flies and Lane’s area has many. As far as possibilities for fruitful
investigation are concerned, the surface has only been scratched in these two areas.
Mr. J. S. Buckett, Program Chairman, announced that in the interest of uni¬
formity, future meetings, at least for the remainder of year, will be held insofar as
possible on the third Friday of each month.
The following notes were presented:
“Flying mouse” identified as Cerogenes auricoma (Burmeister) (Ho-
moptera: Fulgoridae). —Dr. Dennis Breedlove presented to the Academy in early
1967 a series of Fulgoridae that he and an associate collected in Chiapas, Mexico.
They have recently been identified by Dr. Lois O’Brien as Cerogenes auricoma
(Burmeister). This attractive species was described in 1835 (Handbuch der Ento-
mologie 2(1) : 168) from a specimen or specimens collected “Aus Mexilco, zwischen
Villaalla und Oaxaka im Walde von Herrn Deppe entdeckt.” On 1 September 1966,
while D. E. Breedlove and John Emmel were traveling in the state of Chiapas, near
Amates, 13 mi. SE. of Tapanatepec, at an elevation of 3,300 feet, the Cerogenes
auricoma were observed on the trunk and branches of Quercus conspersa Bentham.
To prevent breakage of the long wax tails the collectors injected the specimens with
alcohol to kill them in situ. As can be noted in the display, some specimens of the
series of nine specimens have white wax tails over four inches in length. The
name “flying mouse” seemed to be an appropriate pseudonym to describe the ap¬
pearance of these large bodied and long-tailed fulgorids. They readily took to flight
when disturbed.— Paul H. Arnaud, Jr., California Academy of Sciences , San Fran¬
cisco.
Giant Margarodid Scales from Yucatan. —J. Gordon Edwards exhibited pre¬
served specimens of female scale insects, Llaveia axin (Llave) from Valladolid,
Yucatan. These huge scale insects have been used for centuries by mesoamerican
Indians for various purposes. The bright orange females, each more than an inch
long, are mashed together and kneaded into a wax-like ball which forms the base
for cosmetics, has certain medicinal values, and is the major ingredient of hard
waxy finishes applied to pottery and gourds by the early Maya and Aztec Indians.
Mrs. Katherine D. Jenkins (Berkeley, California) has done more research on these
insects and their uses than anyone else, and has reared both male and female
adults in captivity, feeding them on Erythrina crista-galli (“Coral Tree”). Under
field conditions they usually live on Spondias purpurea (“Hog-plum”) in southern
Mexico and Central America, and are also sometimes found on Acacia trunks. Mrs.
Jenkins cites the name “Ni-in” or “Nije” (used by the ancient Maya) and “Axin”
or “Aji” (used by the Nahua or Aztec Indians), but Edwards questioned dozens
of Indians in Yucatan and Quintana Roo and found none who were familiar with
those names. Instead, they referred to them only as “Tuch Mucuy” (“Tuch” =
“protruding navel”; “Mucuy” = “little dove”). The Indians questioned were unan¬
imous in the opinion that the wax has NO current usage in Yucatan. They once
used it on cuts and tick-bites, but they say it “itches too much,” and they now
prefer commercial medications, instead.—J. G. Edwards, San Jose State College,
San Jose.
JANUARY 1970]
PROCEEDINGS
69
The principal speaker of the evening was Mr. Ron Stecker, San Jose State Col¬
lege. His illustrated talk was entitled “An Illustrated Tour to the Top of a Living
Giant Sequoia.”
A social hour was held in the entomology rooms following the meeting.—M. S.
Wasbauer, Secretary.
Three Hundred and Twenty-Sixth Meeting
The 326th meeting was held Friday, 21 March 1969 in the Morrison Auditorium
of the California Academy of Sciences, Golden Gate Park, San Francisco, with Pres¬
ident Hagen presiding.
Members present (45) : R. J. Adams, C. Annin, P. H. Arnaud, Jr., J. Benet,
F. L. Blanc, R. M. Bohart, I. Boussy, G. Brady, D. Briggs, T. Calvert, P. Cammer,
D. Carlson, R. V. Cottam, C. Dailey, T. W. Davies, R. L. Doutt, J. G. Edwards, J.
Guggolz, T. E. Hewton, Jr., K. Horn, W. H. Lange, R. Langston, R. Lem, K. Lor-
enzen, R. Main, W. W. Middlekauff, R. R. Pinger, Jr., D. W. Ribble, E. S. Ross,
F. Santana, T. A. Schulz, H. Scudder, J. H. Shepard, Joanne E. Slansky, Carolee
Start, R. E. Stecker, L. Stotelmyre, D. Ubick, M. S. Wasbauer, D. L. Wilson.
Visitors present (40) : C. Adamson, Mrs. Betty Armin, H. W. Bourne, Tina Cal¬
vert, Carolyn and Michelle Dailey, Lucinda Doutt, Alice and Janie Edwards, Peggy
Ann Ewart, Barb Glaser, R. Graatman, A. Hamblin, Katie Henry, S. R. Husted,
Katharine D. Jenkins, Ruth M. Jones, P. Jump, T. Lucas, Laurel Main, Ruth J.
Maund, M. Montgomery, M. Ogden, R. W. Pinger, W. W. Pitcher, D. A. Ramsey,
Susan Rickman, Suriya Sampunth, E. A. Schwarz, Muriel R. Sheehy, Mrs. J. H.
Shepard, F. R. Smith, Phyllis Stecker, Anne Suguitan, Richard and Kathy Swartzell,
Sharon D. Tanner, R. M. Vierhus, Betty White, Mrs. David Wilson.
The minutes of the meeting held 21 February were summarized.
Four new members were elected: Dave Carlson, Franklin Ennik, Stan Husted
and Betty White.
President Hagen announced his appointees to the standing committees as follows:
Publication Committee (1971) ; P. D. Hurd, Jr., Chairman and R. F. Wilkey; His¬
torical Committee, R. L. Doutt to replace the late R. L. Usinger.
Dr. Wasbauer announced that because the circulation of the membership book
during the meeting has been considered a somewhat disruptive influence, the book
will be placed on a stand outside the meeting room for members and guests to sign
as they enter.
Dr. Edwards introduced Mrs. Katherine Jenkins of Berkeley. Mrs. Jenkins
showed an excellent series of kodachrome slides taken in Mexico of the large coc-
coid, Llaveia. The fatty body contents of this insect are used for medicinal pur¬
poses, for waterproofing boards and wood products and in the manufacture of the
traditional lacquer ware of Mexico.
The following notes were presented:
Chlaenius tomentosus zunianus (Coleoptera: Carabidae) preying on an
adult Lacinipolia spiculosa (Lepidoptera: Noctuidae). —On 26 September
1966, while I walked across a cut grass field at about 12:30 p.m., west of the dining-
hall of the Southwestern Research Station, 8 km. W. of Portal, Cochise County,
Arizona, a noctuid moth and carabid were observed to be in a fluttering struggle
in the grass. The carabid, as an examination of the moth now indicates, had cap¬
tured the noctuid with its mandibles by the apical third of the right fore and hind
70
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
wings. Both wings show damage across their entire widths. The pair of specimens
were quickly captured in a cyanide bottle to prevent their escape; this however
precluded further observations. The moth has been determined by W. R. Bauer as
a male Lacinipolia spiculosa Grote and the carabid by Dr. R. T. Bell as a male
Chlaenius tomentosus zunianus (Casey). Only fifteen specimens of C. tomentosus
zunianus were available to Dr. Bell at the time of the publication of his revision
of the North American species of Chlaenius (1960, Miscellaneous Publications of
the Entomological Society of America, Vol. 1, No. 3, pp. 97-166, figs. 1-142). This
subspecies was reported as collected at Fort Wingate (type locality) in New Mex¬
ico, and the Catalina Mountains, Santa Rita Mountains, Sonoita, and Fort Hua-
chuca in Arizona. My thanks to these specialists for their determinations. This
pair of specimens is now deposited in the carabid collection of the Academy.—
P. H. Arnaud, Jr., California Academy of Sciences, San Francisco.
The principal speaker of the evening was Dr. J. Gordon Edwards, Department
of Biology, San Jose State College. His illustrated talk was entitled “Entomology
in Yucatan.”
A social hour was held in the entomology rooms following the meeting.—M. S.
Wasbauer, Secretary.
Three Hundred and Twenty-Seventh Meeting
The 327th meeting was held Friday, 18 April 1969, in the Morrison Auditorium
of the California Academy of Sciences, Golden Gate Park, San Francisco, with
President Hagen presiding.
Members present (25) : R. L. Adlakha, P. H. Arnaud, Jr., I. Boussy, D. L.
Briggs, R. Bushnell, D. Carlson, J. A. Chemsak, B. J. Donovan, J. F. Emmel, J. J.
Guggolz, M. R. Gardner, G. A. Gorelick, K. S. Hagen, E. L. Kessel, R. L. Langston,
V. Lee, H. B. Leech, C. D. MacNeill, Kathleen Meehan, P. A. Opler, D. C. Rentz,
D. W. Ribhle, F. Santana, R. W. Thorp, W. J. Turner.
Visitors present (15) : D. Cigston, D. Gunther, Katherine D. Jenkins, Berta Kes¬
sel, K. McClenaghan, Irene McClenaghan, R. Miller, D. M. Muir, Phyllis L. Pas-
qualli, Kathy Rentz, Kathleen Sakai, Paula Schaller, J. Shimizu, Joyce Thorp,
Peggy Ward.
The minutes of the meeting held 21 March were summarized.
The following new members were elected: Brunson P. Bliven, Jean Marie Cadiou,
Kathleen Meehan, John F. Emmel (Student Member).
President Hagen announced that the Society’s Annual Picnic and Field Day will
be held on Saturday, 10 May, at the Recreation Beach at Monticello Dam, ten miles
west of Winters.
President Hagen asked for introductions from the floor. Dr. E. S. Ross intro¬
duced Dr. Dyrce Lacombe of Rio de Janeiro, who is interested in the histology of
barnacles, in kissing bugs of the genus Triatoma and in the internal anatomy of
the Embioptera. He also introduced Dr. E. S. Kessel, a charter member of the
Society, whom some of the newer members may not have met.
Mr. Paul Opler introduced the principal speaker of the evening, Dr. Clyde D.
Willson, Miller Fellow, Division of Entomology, University of California, Berkeley.
His illustrated talk was entitled “Scanning Electron Microscopy: Some Entomo¬
logical Applications.”
A social hour was held in the entomology rooms following the meeting—M. S.
Wasbauer, Secretary.
JANUARY 1970]
PROCEEDINGS
71
Three Hundred and Twenty-Eighth Meeting
The 328th meeting, the annual picnic and field day, was held Saturday, 10 May
1969 at Recreation Beach, below Monticello Dam, Yolo County.
Members present (12) : P. H. Arnaud, Jr., I. Boussy, J. S. Buckett, G. Buxton,
T. W. Davies, M. R. Gardner, K. S. Hagen, W. W. Middlekauff, R. R. Pinger, Jr.,
E. S. Ross, R. W. Thorp, R. F. Wilkey.
Visitors present (17) : Nancy Barker, Davies family, R. Gardner, R. Good, Hagen
family, L. 0. Otero, Harriet V. Reinhard, Suriya Sampunth, Paula Schatter, Taly
Slay, Thorp family, Wilkey family.
The day was quite warm and collecting was good on the chaparral slopes and
along the margins of Putah Creek. Some members took advantage of the trout
fishing in the creek, and some swimming was noted.—M. S. Wasbauer, Secretary.
Three Hundred and Twenty-Ninth Meeting
The 329th meeting was held Friday, 17 October 1969 in the Morrison Auditorium
of the California Academy of Sciences, Golden Gate Park, San Francisco. Presi¬
dent Hagen presided.
Members present (39) : P. H. Arnaud, Jr., F. F. Blanc, I. A. Boussy, R. Bush-
nell, P. Cammer, Helen Court, D. G. Denning, J. G. Edwards, J. F. Emmel, F.
Ennik, W. Gagne, M. R. Gardner, J. F. Gustafson, J. D. Haddock, K. S. Hagen,
T. P. Heck, T. E. Hewton, W. H. Lange, R. L. Langston, H. B. Leech, D. D. Lins-
dale, R. F. Luck, P. Opler, J. A. Powell, J. Prine, D. Rentz, D. W. Ribble, C.
Rogers, R. Schoeppner, T. A. Schultz, H. I. Scudder, T. A. Sears, R. Stecker, L.
Stotelmyre, W. J. Turner, S. C. Williams, Barbara A. Wilson, Joanne Wasbauer, M.
Wasbauer.
Visitors present (38) : Nancy Blair, R. Blair, J. E. Court, Angele Dacosta, Char¬
lotte Davis, Loretta Denning, D. F. and F. L. Dunn, Alice and Janie Edwards, J. R.
Gabel, C. S. Glaser, L. Gormley, L. Green, J. T. Hjelle, Katharine D. Jenkins, Sandy
Jordan, Mr. and Mrs. Peter Jump, Ellen M. Lange, Shari Lewis, D. Merrill, S. R.
Montague, D. D. Munroe, G. Nichols, Sandra Opler, Judy Oppenheim, R. R. Papp,
P. Pasqualli, Roberta Pfadt, J. A. Scott, T. Slery, J. A. Smith, Phyllis Stecker, B.
Tilden, Marilyn Trockman, Jill Vandergrift, Judy Weller.
The minutes of the meetings held 18 April and 10 May 1969 were summarized.
Ten new members were elected: William E. Azevedo, Bruce H. Baker, Michael
M. Benzien, J. Russel Gabel, Thomas Heck, Sandy Jordan, Robert Luck, LeRoy C.
Olson, Fred Punzo, Torolf R. Torgerson.
Mr. Leech announced on behalf of the Historical Committee, a gift from Mrs.
Roxanna Ferris of individual photographs of six entomologists: J. H. Comstock,
V. Kellogg, K. Jordan, J. Waterston, A. D. Imms, F. Sylvestri, prints of H. W.
Bates and G. H. F. Nuttall and a group photograph of Dipterists at the Fourth In¬
ternational Congress of Entomology, Ithaca, New York, 1928. All the pictures are
framed.
The following notes were presented:
Gnophaela latipennis trapped by flowers of milkweed (Lepidoptera:
Pericopidae). —Gnophaela latipennis Boisduval is a common day-flying moth of
the southwestern United States and adjacent Mexico. Constrastingly marked in
black and white, with a wing span of 35 to 55 mm., it attracts attention from a dis¬
tance. On 5 July 1969 I was collecting beetles in Estell Creek, Logan Basin, at the
72
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
southwestern foot of Bearwallow Ridge in the mountains west of Elk Creek, Glenn
County, California, and saw a number of specimens of G. latipennis trapped by the
flowers of milkweed Asclepias sp. Some moths were still in nearly perfect condi¬
tion, constantly fluttering in attempts to escape, others were hanging limply; the
rest were dead, and tattered from being blown by the wind. In each case the moth
was caught by the tarsi of one or more legs in the floral parts of the milkweed, and
I was unable to release them without breaking the legs. —Hugh B. Leech, Califor¬
nia Academy of Sciences, San Francisco.
Status of the Oriental fruit fly in California. —On 11 September 1969, a
single male Oriental fruit fly, Dacus dorsalis Hendel, was caught in one of the
regular statewide multiple-purpose fruit fly traps in a grapefruit tree at El Monte,
Los Angeles County. Additional traps were immediately deployed in the area at a
concentration of 100 per square mile, with peripheral range of 50 and 20 per square
mile over a 64 square mile area. By 1 October a total of 18 male flies were trapped,
all within a radius of 0.92 miles which could be contained in a circle of 2.65 square
mile area. No flies have been taken since 1 October. (Colored slides of adults and
larvae were shown and pinned adult specimens were passed among the members
and guests.) — F. L. Blanc, Program Supervisor, Bureau of Entomology, California
Department of Agriculture.
Flight of Hilara sp. (family Empididae) at Webber Lake, California.—-
Webber Lake, at an altitude of 6774 feet in Sierra County, California (located
northwest of Truckee) is an historical collecting area for Dipterists. The pioneer
Dipterist, Baron Osten Sacken, collected there in the summer of 1876. Webber
Lake was on the early stage coach route over the Sierra Nevada and the hotel built
over 100 years ago is still in existence, though not in use. On 23 and 24 August
1969, I drove to Webber Lake in my camper for a weekend of collecting.
On the evening of 23 August at 1930 hours, daylight saving time, as I walked to
the western shore of the lake it was immediately evident that there was a tremen¬
dous flight of insects over the waters of the lake. The insects were flying close to
the water as though they were just skimming the surface. They were so numerous
that they produced an audible buzzing sound with their flight. They were not con¬
centrated in any one spot, but they were generally dispersed with possibly one or
more specimens flying over every square foot of water. They were flying over the
water as far out as I could see them (about 20 feet from shore). I collected a few
specimens and they were immediately identifiable as specimens of a Hilara sp.
There was some wind, and when the water surface became choppy, they probably
flew a little higher. They were much more difficult to see in the dimming light of
the approaching night. I walked northward along the shore of the lake for 490
paces and they were as abundant there as where first observed. I found a small
floating dock about 35 feet in length and walked to its end. The hilaras were fly¬
ing over the lake at that distance from shore. Returning to my camp at 2010, they
were still in flight, even though it was almost dark and a large moon was rising.
A total of 34 males and 22 females was collected at various points along the shore
of the lake. The hilaras were attracted to an 8 watt safari black light set on the
sand and gravel near the edge of the lake. Hundreds of specimens were attracted
to the black light, including mating pairs. Eighty-eight males and 79 females were
collected as a sample. By 2040 they no longer flew to the black light. In four
insect flight traps operated near the shore of the lake only a few specimens of this
JANUARY 1970]
PROCEEDINGS
73
Hilara were collected during portions of two days and one night of trap operation
(1 male, 2 females on the 23rd; 2 males, 1 female on the 24th). It would be of
interest to discover the resting places of such large numbers of hilaras.
The following morning, at 0745, some hilaras were seen flying over the lake.
Fifty minutes later none were seen. The lake surface was very choppy due to strong
winds.
Trichoptera were exceedingly scarce during the time of the evening flight of
hilaras as only two specimens (a male Dicosmoecus unicolor (Banks) and a female
Hesperophylax sp., both kindly determined by Dr. D. G. Denning) were attracted
to the black light and none were swept from over the water’s surface. The abun¬
dant silver flashes from jumping fish in the late afternoon probably indicated their
feeding on the Hilara since Trichoptera and other conspicuous Diptera were lack¬
ing. The fish could be seen feeding up to several hundred feet from shore, possibly
indicating the distance of the Hilara flight from shore.—P. H. Arnaud, Jr., Cali¬
fornia Academy of Sciences, San Francisco.
The principal speaker of the evening was Dr. Richard Garcia, University of Cali¬
fornia, Berkeley. His illustrated talk was entitled “Entomology in Western Ma¬
laysia.”
A social hour was held in the entomology rooms following the meeting.—M. S.
Wasbauer, Secretary.
Three Hundred and Thirtieth Meeting
The 330th meeting was held at 7:45 p.m. on Friday, 21 November 1969 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park,
San Francisco. President Hagen presided.
Members present (45) : P. H. Arnaud, Jr., W. E. Azevedo, J. Benet, M. M.
Bentzien, R. G. Blair, I. Boussy, R. G. Blair, D. Briggs, T. Briggs, P. Cammer, J. A.
Chemsak, H. V. Daly, T. W. Davies, J. G. Edwards, W. E. Ferguson, J. R. Gabel,
W. Gagne, Lauren Green, P. Grimstad, J. F. Gustafson, K. S. Hagen, J. L. Joos,
P. Jump, R. L. Langston, V. Lee, H. B. Leech, P. Lem, K. Lorenzen, R. Main, P. A.
Opler, W. Pitcher, D. C. Rentz, D. W. Ribble, E. I. Schlinger, T. Schultz, R. 0.
Schuster, R. Stecker, L. Stotelmyre, R. W. Thorp, D. Ubick, Joanne S. Wasbauer,
M. S. Wasbauer, S. P. Welles, S. C. Williams, D. Wilson.
Visitors present (24) : Kathy Beeby, J. G. Brandon, A. Dacosta, W. H. Dana,
R. E. Dietz, Alice and Janie Edwards, L. A. Gormley, Mrs. J. L. Joos, Sandy Jor¬
dan, D. Merrill, R. Montagna, S. Nelson, Gay Nichols, Judy Oppenheim, Gayle Ray,
Avis Reeves, B. Reeves, S. Sims, J. Smith, Joyce Thorp, Mrs. D. Wilson, Kathy
W aeny.
The minutes of the meeting held 17 October 1969 were summarized.
Six new members were elected: Mr. Randall G. Blair, Miss Lauren Green, Mr.
Peter M. Jump, Mr. William W. Pitcher, Mr. John T. Polhemus and Mr. Gary L.
Seghi.
President Hagen called for notes and exhibits. Mr. David Rentz announced the
recent naming of a mountain peak in California after an insect—Acrodectes Peak.
It was named last year by a U. S. Government commission to provide names for
unnamed mountain peaks. The peak is in the southern Sierra Nevada Mountains.
The name Acrodectes is of Greek origin and means “summit climber.”
Mr. Leech announced two books which have just been published. The first, en-
74
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
titled The Kodiak Island Refugium , is an interdisciplinary approach to the flora
and fauna of Kodiak Island. It contains a couple of chapters by entomologists and
includes a complete list of all the insects which were found during the course of a
study of the island. The second is The Distributional History of the Biota of the
Southern Appalachians, Part I, Invertebrates. It contains two entomological papers,
“The Evolution of the Carabidae of the Southern Appalachians” by Thomas Barr
and “The Ecological and Geographical Relations of the Southern Appalachian
Mecoptera” by George Byers.
The following notes were presented:
Habitat of Atypoides riversi (Araneida: Aiitrodiaetidae) .—A colony of
the turret-building spider Atypoides riversi Pickard-Cambridge was discovered at
Laurel Dell, Mt. Tamalpais, two miles north of Stinson Beach, Marin County, Cali¬
fornia on 8 August 1969. The colony was located on a north-facing slope heavily
forested with Douglas fir ( Pseudotsuga taxifolia ) and California laurel (Umbel-
lularia californica) . The ground was covered with several inches of leaf litter but
lacked low plant cover.
The colony covered an area 100 feet wide by 300 feet long. Tire spiders’ burrows
were located at the base of trees or near fallen logs. The turrets extended as high
as two inches above the surface litter and the burrows were about 8 inches deep.
Only females were present in a sample of 8 spiders collected. One female had sev¬
eral first instar young in her burrow.— Michael M. Bentzien, San Francisco State
College, California.
Philotes enoptes bayensis and tildeni Langston—Four New Localities.—
Both subspecies are found in definite colonies associated with their Eriogonum
foodplants. Since my original 1964 and 1965 publications, additional colonies have
not been generally found by collectors. Therefore, it is of interest to note these
records: 1) P. e. bayensis —Camp Meeker, 2 mi. N., Sonoma Co., 3 July 1965, a
single female among about a dozen Plebejus acmon; Benecia State Park, Solano
Co., 15 June 1969, a single badly worn male with at least 50 P. acmon —farthest
east (and inland) record. 2) P. e. tildeni —Keene, 4 mi. SE., Kern Co., 5 Septem¬
ber 1965, 1 female, yellow-flowered Eriogonum —new county record (since the
pubs.) and farthest south; Polonio Pass, Hwy. 46, S.L.O. Co., 8 September 1969, 5
males, 5 females among more than 100 P. acmon (appears to be a stronger colony
than nearby Cottonwood Pass, both for the blue and for Apodemia mormo intermed.
to A. langei which was present in the hundreds). 3) P. e. smithi —examples ex¬
hibited from the type locality for comparison. 4) P. e. enoptes —examples shown
from the Sierra Nevada for comparison.— Robert L. Langston.
A Naturalist Returns to Tesla Canyon. —I visited Tesla Canyon of 9 November
1951 to collect living specimens for courses in Forest Entomology and beginning
Entomology. This canyon, at that time, was desolate except for the one no tres¬
passing sign where the owner insisted on permission to collect on his land. On
previous trips many groups of us had driven out Tesla Road, at the south edge of
Livermore, and crawled under the fence with its bullet-riddled sign to collect many
kinds of insects which had their most northern representatives in this dry canyon.
Traveling alone in 1951, I collected large numbers of the drywood termite,
Incisitermes ( Kalotermes ) minor (Hagen), from a solitary, wind-felled, digger pine
on the floor of the valley. At the same time, hundreds of the rare hemipterous spe¬
cies Patapius spinosa (Ross) (Leptopodidae) were found around the same tree.
JANUARY 1970]
PROCEEDINGS
75
Usinger had previously recorded one specimen of this species (194111) 1 as a prob¬
able immigrant into California from its only previously known home in the Canary
Islands. Numerous specimens of the California violin spider, Loxosceles unicolor
Keyserling, were also found around or in the sides of the canyon.
This year another solitary trip on 26 October 1969 revealed a vastly changed en¬
vironment. The original sign was obliterated by rust and holes. Other fences on
the south, upper end of the canyon had many new signs, and bullet holes in the
fences and signs while the northwestern side was fenced and has signs in a some¬
what similar condition. Farther down the valley it was open on the south side of
the canyon, and many hundreds of motorcyclists were carrying on their denuding
of the valley floor and north-facing slopes. On the northeast side of the valley, it
is protected by a stout fence and signs of the Atomic Energy Commission (with
fewer bullet holes) but this section is patrolled by federal agents and farther down
is a guard station and many new buildings.
Otherwise (with exceptions) the fauna and flora was the same. The digger pine
had been reduced to a long reddish brown streak of decayed wood but the termite
is surviving, in large numbers, in numerous stumps of cottonwood which had been
downed for firewood but the stumps were too large to move and were almost im¬
possible to break into with my axe and hatchet. Each had a number of termite
colonies.
Leptopodids and the violin spider were still there, but another curious arthropod
can now be added to any faunal list of this unique canyon. After crawling under
an (anonymous) fence I found two large nymphs of the famous kissing bug, Redu-
vius personatus L. This species was recorded as rare in California by Usinger
(1946h) and again by he and Wygodzinsky (from one pinned male, from Kern
County) in (1964b). Nevertheless the species is not included in Essig’s revised
edition of “Insects of Western North America” (1958) nor does Usinger et al.
(1964b) cite Usinger’s previous paper (1946h) in its Bibliography. Further the
1969 edition of Herm’s Medical Entomology lists this species as occurring only as
far west as the Middle West.
The two nymphs are now feeding on Blatella germanica L. nymphs; while the
dirt cave, from which they were removed on the south-facing slope, is heavily popu¬
lated by several species of spiders and a very large number of lepismids. Access to
another intriguing cave could not be investigated.—J. W. MacSwain, University of
California, Berkeley.
Mr. Paul Opler announced that the annual Christmas dance of the Department
of Entomology, University of California, Berkeley, would be held on 13 December
at the Veterans Memorial Hall, Albany. Tickets are $2.00 and will be available at
the door.
President Hagen announced the temporary committees for the year. The nomi¬
nating committee will be Dr. J. A. Powell, Chairman, Mr. D. G. Denning and Dr.
J. W. Tilden. The auditing committee will be Mr. D. C. Rentz, Chairman, Mr. R. L.
Langston and Mr. H. B. Leech. These committees will report at the December
meeting.
The principal speaker of the evening was Dr. Stanley C. Williams, whose il¬
lustrated talk was entitled “Behavior of Scorpions in Relation to the Birth Process.”
1 From: R. L. Usinger’s Bibliography by P. D. Ashlock, 1969, Pan-Pac. Entomol., 45(3): 185-203.
76
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
A social hour was held in the entomology rooms following the meeting.—M. S.
Wasbauer, Secretary.
Three Hundred and Thirty-First Meeting
The 331st meeting was held at 7:45 p.m. on Friday, 19 December 1969 in the
Morrison Auditorium of the California Academy of Sciences, Golden Gate Park,
San Francisco. President Hagen presided.
Members present (23) : P. H. Arnaud, Jr., R. G. Blair, D. E. Bragg, P. Cammer,
T. W. Davies, J. G. Edwards, J. F. Emmel, J. R. Gabel, M. R. Gardner, J. D. Had¬
dock, P. Jump, R. L. Langston, H. B. Leech, W. W. Pitcher, J. A. Powell, D. W.
Ribble, T. A. Schultz, T. A. Sears, R. E. Stecker, L. Stotelmyre, R. W. Thorp, M. S.
Wasbauer, S. C. Williams.
Visitors present (16) : F. G. Andrews, J. L. Bath, G. S. Benham, Jr., Beatrice
Berry, Nancy Blair, E. A. Cameron, Alice Edwards, Marilyn Hamaker, Katharine
D. Jenkins, Harriette Jump, A. D. Moscioni, Judith Oppenheim, J. A. Smith, Joyce
Thorp, R. M. and Sharon Vierhus.
The minutes of the meeting held 21 November were summarized.
Four new members were elected: G. S. Benham, R. L. Dunn, J. E. McPherson,
Jr., K. Smith.
The following notes were presented:
Thrips “biting” man.—The purpose of this note is to present several “biting”
thrips’ records encountered in central California this fall. Bailey (1936, Canad.
ent., 68(5) :95-98) has published a resume of some of the articles reporting thrips
as attacking man in the United States and other world areas. It is of interest to
note that thrips which are ordinarily considered phytophagous and to some extent
predaceous, would attack man. The first two species recorded below appear to be
new records.
While collecting in the area of Frank Raines Park, Del Puerto Canyon, Stanislaus
County, on a field trip on 26 and 27 September 1969, with Drs. Williams’ and
Gustafson’s entomology classes from San Francisco State College, four insect flight
traps were operated in the creek bed. While operating these traps, I was annoyed
by “biting” insects on both my arms and neck which I finally detected to be thrips.
None were collected at that time. On a later field trip, on 1 November, to the exact
locality with Mr. Thomas Davies, in the mid-day, a number of sharp “bites” were
felt on my arms and three culprit thrips were successfully collected in alcohol. The
specimens were determined as presenting two species—one specimen of Franklini-
ella occidentalis (Pergande) (the western flower thrips) and two specimens of
“probfably] Anapliothrips sp., probably undescribed.” It is thought that the latter
species feeds on oaks. More recently, on 28 November 1969, in the Entomology De¬
partment, California Academy of Sciences, San Francisco, LaVeme Trautz, our
Entomology Department technician, felt a “bite” on her arm. It was a thrips and
this specimen was determined as Thrips tabaci Lindeman (the onion thrips). This
species has previously been reported by Bailey (1936:97) in California as “biting.”
My thanks are extended to Mr. Tokuwo Kono, of the California Department of
Agriculture, Sacramento, for his interest and prompt determination of these thrips.
—Paul H. Arnaud, Jr., California Academy of Sciences, San Francisco.
Grooved Entomology.—Glancing through a recent issue of Systematic Zoology,
I was upset to find only two articles which I could read with real understanding.
JANUARY 1970 ]
PROCEEDINGS
77
Doubtless this is a personal failure and problem. Soon after, I chanced to read
“Living nature and the knowledge gap,” by Dr. Paul A. Weiss ( Saturday Review
for 29 November 1969, pp. 19-22, 56). Though written for a much broader view¬
point, it is possible that some of his comments could be applied to entomology.
Having in mind a noticeable trend in articles in Systematic Zoology over the past
few years, I would like to quote a few lines from Dr. Weiss. 1
(p. 19) “One of the drawbacks [of modern science] is that, by its sheer
momentum, a mass of single-tracked workers tends to amplify any trend
once that trend has started rolling. A fashionable course thus becomes
grooved ever more deeply, draining interest, attention, encouragement, and
talent away from solitary prospecting ventures. As a result of these social
dynamics, the research scenery is gradually becoming converted, meta¬
phorically speaking, from a lush mountain meadow evenly irrigated by a
profusion of anastomosing rivulets into a landscape of deep canyons with
raging rivers, separated by wastelands of arid mesas.”
(p. 21) “Education can give him [a young investigator] scope for self-
direction and creativity or it can mold him into a mere cog in a mass pro¬
duction machine. Society needs the latter kind, well-trained for practice,
in increasing numbers, but here we are concerned primarily with those
aiming at the advancement, rather than the application, of knowledge . . .
if the incipient investigator is enticed to keep on riding currents of fashion,
instead of being challenged to chart his own course toward self-chosen
destinations, it is his education that has been at fault. So what is needed
is certainly not ‘planned research,’ but sounder plans for ‘ education for
research ”- —-Hugh B. Leech, California Academy of Sciences.
The effects on man of a natural sting by the scorpion Vejovis coufusus
Stahnke.—The sting of Vejovis confusus Stahnke was experienced accidentally
while conducting field studies on the ecology of this species in the Sonoran desert
near Phoenix, Maricopa County, Arizona, during 1966. The sting occurred at 10:00
AM (MST), some 10 minutes after the capture of the mature specimen. The sting
was inflicted on the middle segment of the third finger of the right hand. An
immediate, intense burning sensation occurred, which lasted at the site of the sting
for some 20 minutes before beginning to lessen in intensity. Fifteen minutes after
the initial sting, swelling became visible around the knuckle above the wound. This
swelling gradually progressed over the entire finger and spread downward into the
other fingers of the same hand within one hour. Within three hours the entire hand
had become very swollen as compared to the left hand. This swelling was sufficient
to prevent effective bending of fingers and wrist. Within five hours, the swelling-
had progressed up the arm to the elbow. The swelling lasted approximately 24
hours before beginning to subside. Within 48 hours the right arm and fingers had
returned to almost normal size. Within 72 hours the hand had fully recovered from
the sting with no apparent after effects. During the period of increased swelling
the intense burning sensation gradually subsided while a general numbness of sen¬
sation developed in the fingers and spread to the elbow of the affected hand. As
the swelling began to disappear, normal sensitivity gradually returned. No indica-
1 Copyright 1969 Saturday Review , Inc. Quoted with the permission of Saturday Review.
78
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
tion of convulsions or other significant systemic actions of the venom were appar¬
ent. No therapy was used to counter the effects of the venom, however, use of the
right hand was essentially discontinued for some 20 hours because of the extreme
swelling. The speed and pattern of travel of the swelling indicated that the lym¬
phatic circulation was primarily responsible for the distribution of the venom.—
Stanley C. Williams, San Francisco State College.
President Hagen called for reports from the chairmen of the standing committees.
Prof. J. G. Edwards, membership committee, reported that 36 new members were
elected and one member, Dr. C. P. Alexander, was elected to honored status during
the year.
Dr. R. W. Thorp reported on behalf of the publication committee that two manu¬
scripts were under consideration for the Memoirs series during 1969, the auto¬
biography of the late Prof. R. L. Usinger, and a revision of andrognathid millipeds
by M. R. Gardner. The publication dates have not been set on these papers, but
they will appear in 1970.
President Hagen then expressed thanks to the program committee and its chair¬
man, Mr. J. S. Buckett, for a successful year of programs.
Dr. P. H. Arnaud, Jr. read the treasurer’s financial report, and Mr. R. L.
Langston of the auditing committee reported that the financial records of the So¬
ciety for 1969 are in good order.
Mr. H. B. Leech, chairman of the historical committee, reported that the file
cases are now upstairs on the main floor of the entomology department, but 2 or
3 more file cases are needed before all the material on hand can be sorted-classified
and filed. Accessions during the year have been noted at earlier meetings. The
most recent acquisition is over 1000 glass lantern slides used in the past at Stan¬
ford University to illustrate lectures. Most are of entomological subjects and some
are of strictly historical interest.
President Hagen announced appointment to the publication committee for 1972
of Dr. Howell V. Daly and Dr. Albert A. Gregarick.
Dr. J. A. Powell, chairman of the nominating committee, presented the following-
slate of nominees for offices in the Society for 1970: Dr. R. W. Thorp, President;
Mr. F. L. Blanc, President-Elect; Dr. M. S. Wasbauer, Secretary; Dr. P. H. Arnaud,
Jr., Treasurer. There were no nominations from the floor. The nominees were
elected to office for 1970 by unanimous vote.
The principal speakers of the evening and their topics are as follows:
G. R. Buckingham-— Searching for a Parasite of the Walnut Husk Fly,
Rhagoletis completa Cresson.-—The walnut husk fly, Rhagoletis completa Cres-
sen, was accidentally introduced into California from the Southwestern United
States in the 1920’s. The maggots eat the walnut husk, which results in staining of
the shell in English walnuts. A native parasite, Opias juglandis Mues. (Hym.
Braconidae), has been reared for several years at the Division of Biological Control,
University of California at Albany. In September 1969 I traveled to Arizona and
New Mexico to obtain more parasite individuals and to make field observations on
the parasite and its hosts. Many parasites were observed on wild black walnut trees
in the Chiricahua Mountains of Southern Arizona. Since Rhagoletis completa does
not occur there, two other husk flies, Rhagoletis juglandis Cresson and Rhagoletis
boycei Cresson, serve as hosts. Ants and a salticid spider were observed preying
upon parasite adults and ants were very often found carrying maggots. This area
JANUARY 1970 ]
PROCEEDINGS
79
of Arizona is very similar to the parasite release areas in California except that it
has frequent summer thunder showers. How much effect the dry summers of Cali¬
fornia will have on the parasite is unknown. In Southern New Mexico, parasites
were found associated with both Rhagoletis juglandis and Rhagoletis complete!.
James D. Haddock — Biological traits of two species of Leptocella. —The
caddis fly genus Leptocella (Leptoceridae) consists of a number of primarily whit¬
ish colored species of insects whose larvae construct elongate, tubular, portable
cases. They are distributed throughout the western hemisphere, occurring from
Alaska to Argentina.
Species limits, based on the morphology of the adults, have been open to con¬
siderable conjecture and debate. A study of the immature stages, however, has
helped to clarify speciation in the genus in North America; two species will be
briefly discussed to illustrate this point.
The adults of Leptocella intervena and L. albida are remarkably similar in regard
to wing pattern, head structures, male genitalia and a number of other structures.
Larval morphology, case-making behavior and ecology, however, clearly separate
the two species. Leptocella intervena has a well developed swimming and feeding
brush that is located on the hind tibia, has a number of tubular tracheal gills lo¬
cated on the anterior segments of the abdomen and has a characteristic color pat¬
tern found on the dorsal portion of the head and thorax. In contrast, L. albida has
no brush, has an absence of tracheal gills and has different color patterns. Lepto¬
cella intervena constructs a larval case composed primarily of minutely cut plant
fragments, while the case of L. albida is composed almost entirely of sand grains.
Leptocella intervena is found in the western U. S. in lowland, highly silted slow
flowing rivers with midsummer water temperatures ranging from 60-102 degrees
Fahrenheit. Leptocella albida appears to be restricted to the cooler waters of the
past-glaciated midwest and east. It is apparently isolated from the western U. S.
by the head waters of the Mississippi River.
It is anticipated that a study of this nature will be useful in the applicability of
insects as indicators of thermal and organic water pollution and erosion.
Katharine D. Jenkins — The fat-yielding coccid, Llaveia, a monophlebine
of the Margarodidae. —This very large soft-scale insect of the American tropics
and subtropics, which has been exploited (sometimes cultivated) for its fat by
peoples of Meso-America since before the Spanish Conquest, has not been system¬
atically collected and studied by entomologists, and the literature on it is meagre
and unsatisfactory. 1
The material here presented was accumulated in the course of ethno-historical
research and field work—directed toward finding the sources of the fatty substance
used in Mexican lacquer-work, gourd-painting and related techniques, and record¬
ing surviving practices in the exploitation of the insect—and personal experience in
rearing progeny of insects collected.
The fat, extracted from the bodies of mature females, has remarkable properties
in forming a tough impermeable film on any surface to which it is applied and has
had many uses, not only in native arts (in waterproofing wood and gourds and
binding pigments to surfaces in a lacquer-like coating, and as a base for face- and
1 Pertinent references are cited in K. D. Jenkins, “Aje or Ni-in (the fat of a scale insect): Paint¬
ing Medium and Unguent,” XXXV Congreso Internacional de Americanisms, Mexico, 1962, Adas y
Memorias 1 (Mexico, 1964) : pp. 625-636.
80
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
body-paint), but also in folk medicine (particularly as applied to wounds, swell¬
ings, skin afflictions, or, combined with herbs or other materials, taken internally
to cure a variety of ailments).
Both the insect and the fatty product are generally called aje, from the Nahua
(Aztec) name axin; but there are other locally used names such as ni-in, nij, or
nije, from areas of Maya influence, or kurron, among the Guaymi of Panama.
Although the substance has been a widely distributed trade item for centuries, it
has never gained economic importance and production has remained an obscure
native industry carried on by a few persons in isolated communities, where the
coccids may be cultivated or gathered from the wild. The occurrence of the insect
has been reported from widely separated localities in “hot country” (i.e., at eleva¬
tions of less than 4,500 feet), from western Mexico to Panama; but the actual dis¬
tribution of the genus Llaveia is not known; and it is not known whether the coc¬
cids of different areas are of one or several species.
The insect was first described in 1830 by Pablo de la Llave, who examined speci¬
mens from Tlacotalpan in southern Veracruz, and named it Coccus axin. In 1875,
the French entomologist Signoret erected a new genus for it, naming it Llaveia,
and he put specimens from Guatemala in a new species, Llaveia bouvari. In 1928
the late Dr. Harold Morrison, of the USDA Bureau of Entomology, suggested that
L. bouvari and L. axin were identical but, lacking adequate material for compara¬
tive study, left the question open.
These Llaveia coccids produce only one generation a year and have a long over¬
wintering egg stage. The adult females are oblong-oval in shape, usually coral-red
in color (sometimes yellow), but are covered with a powdery white wax. In the
fall, when they have attained full size (1.5 to 2.5 cms in length), they produce a
more flocculent wax and, under natural conditions, leave the host plant to go into
the ground, each female encasing herself in a cottony-looking ball in which the
salmon-colored eggs are laid. Crawlers begin to emerge in early spring, hatching-
out in separate batches over a period of about two months, so that, in any colony,
several stages of development may be seen at the same time. Females molt three
times, at intervals of about a month. After the second molt, the males no longer
feed on the host, but find protected places, in the bark of the tree, the undersides
of leaves, the detritus under the trees, where they form “pseudococoons” of a soft
white wax, in which they spend the next two stages of development, to emerge as
winged adults about two months later (July to early September), when the females
have completed their third molt and are ready for mating. The winged males live
only a few days, but the females continue to feed and fatten on the host plant for
several weeks and, by the time they are large enough to be worth gathering and
processing (late September, October or November), all males have disappeared.
Cultivation is a simple matter of setting aside a number of the fattest females at
harvest time, storing them in a gourd or other container during the dry season, and
placing them on a suitable host tree when the crawlers begin to appear in the
spring.
In field trips in the fall seasons of 1966, 1967 and 1968, 2 to places where the
insects had been exploited in the past, colonies of the coccids were found in five
distinct areas, growing under different conditions:
2 Field work made possible by two grants from the American Philosophical Society, in 1966 and
1968.
JANUARY 1970 ]
PROCEEDINGS
81
1. In the wet climate of southern Veracruz, at San Andres Tuxtla, where they
were being cultivated by tradition on Erythrina americana, a species of
Spondias, and Jatropha curcas— all trees which are commonly cultivated.
2. In the drier climate of the Rio Balsas Basin, at Ziritzicuaro (about 20 kms.
south of Huetamo, Michoacan), where they flourished naturally on Acacia
cochliacantha, in a “thorn forest” situation, and were gathered by persons
from Huetamo.
3. In the similarly dry climate of south-central Chiapas, near Venustiano Car¬
ranza, where they were gathered from Acacia angustissima and A. pennatula.
4. In the Department of Baja Verapaz, Guatemala, with semi-dry climate, at
Rabinal, where they were being cultivated on Spondias and Jatropha trees,
and at nearby Salama, where they grew untended on Jatropha planted in
hedgerows around houses.
5. In Yucatan, with its Gulf Coast-Caribbean climate, at Valladolid, where they
covered the branches of a large Spondias tree, growing behind a house—
again, untended, probable survivors of earlier cultivation.
These findings do not fill the gaps in our knowledge of the distribution of
Llaveia, but they raise interesting questions about the history of the insect’s culti¬
vation, which must have affected the natural distribution patterns of the genus.
Colorslides shown depict the Llaveia coccids—-adult male and female, and some
of the earlier instars—and the several habitats in which they were found, as well
as the techniques used to extract the fat and prepare it for market.
J. A. Powell — Moths of the Humboldt Bay area. —Since the insect fauna of
the north coast area of California has been poorly known, we made an effort to
collect thoroughly last summer during the field course in entomology of the Uni¬
versity of California, held at Humboldt State College, Areata. The two principal
habitats worked were the second growth redwood forest back of campus and the
beach dunes south of Samoa. Black light trapping was carried out only at Areata,
in mixed vegetation marginal to the redwoods and in the forest. Weather conditions
were generally better than anticipated; fogs were intermitent and did not prove to
be a strong deterrent. Nightly temperatures ranged in the mid to low 50’s °F.
Moth collecting was not rich in quantity or diversity (50-60 species per night
maximum), with noctuids and larger macros poor, geometrids relatively good,
tortreoids fair, and pyraloids and smaller micros poor. Nor surprisingly, many rep¬
resented range extensions of species previously known further north. The 33 spe¬
cies of Tortricidae (s.l.) and Phaloniidae may shed some light on distributional
affinities of the fauna of the area: only 39% are widespread Nearctic Boreal spe¬
cies (opposed to 55% of Tortricinae in Califoxnia as a whole), while 36% range
along the coast in a Vancouveran distribution pattern (only 10% of the California
Tortricinae).
Illustrated species included: Acleris gloverana (Wlsm.) four polymorphs taken
in late August near Big Lagoon on Picea sitchensis, indicating the California popu¬
lations probably will prove as variable as those to the north; A. hritannia Kft.,
dark forms previously only known to the north prevalent at Areata; A. maccana
(Tr.), taken at Areata in February, 1968, the first record in California for this
Holarctic, boreal species; Cnephasia longana (Haw.), collected at four localities
from beach dunes to 2000' east of Kneeland, an European species previously known
82
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 1
to be established in the San Francisco Bay area and in Oregon and Washington;
Batodes angustiorcma (Haw.), another European species introduced around the
Puget Sound and San Francisco Bay areas, but not recorded in California since
1943, abundant on cultivated Taxus at Areata in late July; Xenotemna pallorana
(Rob.), flying at Samoa dunes during the latter half of June, not previously re¬
corded south of northeastern Oregon; Pandemis limitata (Rob.), at lights during
most of July at Areata, confirming two isolated records along the California coast
ten years ago; Epinotia cruciana (L.), netted in the redwoods on July 4, an Hol-
arctic species not known south of Washington; Zeiraphera ratzeburgiccna (Ratz.)
(= pacifica Free, or canadensis Mut. & Free.), a single male at light in late July
of this Canadian species; Apotomis capreana (Hbn.), at lights in late July, also a
southward extension from Washington; Epimartyria pardella (Wlsm.), at four sites
around Areata and west of Kneeland, the first collected since the type series taken
in southern Oregon in 1872; and Arctia caja L., at light in late July.
T. A. Sears — -A possible explanation for seasonal emergence peaks of
Macrolepidoptera in southwestern Chihuahua. —My recent collecting with
black light traps (funded in part by American Philosophical Society grant #5192,
Penrose fund) in the Barranca del Cobre region of the Sierra Madre Occidental of
southwestern Chihuahua has revealed a distinctive emergence pattern for Macro¬
lepidoptera of the area, probably as a result of adaptive ecological strategies for
survival in a wet-dry climatic regime. There are no specific weather data for this
area as it is so remote, but, in general, there is a wet season of 60 to 80 days ex¬
tending from the first of July to mid-September. During the remainder of the year,
humidity is extremely low. Temperatures, of course, vary with locality and season;
but in all cases the annual variation is considerable.
In 1968 my collecting commenced 4 July, two days before the rains began, at
Choix, Sinaloa, 900 feet elevation. By 10 July, the extremely rapid budbreak and
development of vegetation so characteristic of this region of Mexico had rendered
the area verdant. From 8 July through 25 July, moth collecting was extremely good
both in numbers and diversity. However, shortly thereafter and throughout the
remainder of the summer, collecting dropped off markedly and was, in general, un¬
profitable. The same pattern was also evident at Creel, Chihuahua, elevation 7,600
feet. Around 10 September diversity and numbers of Macrolepidoptera increased
and collecting was again excellent. Since the intensity of collecting was not con¬
sistent throughout the summer, no conclusions were drawn at that time; and I at¬
tributed the poor collecting to a combination of moon phase and weather factors.
With consistent and continuous collecting throughout the summer of 1969 a gen¬
eral emergence pattern was confirmed. We collected almost continuously in all
weather at two localities—Cuiteco, Chihuahua, elevation 5,200 feet and Temoris,
Chihuahua, elevation 4,500 feet. Rough counts of the numbers of species and total
numbers reveal distinctly bimodal distribution with peaks around 20 July and 10
September. Representative counts for diversity are as follows in numbers of spe¬
cies per trap: 20 July, 150-170; 15 August, 40-60; and 10 September, 100-150. The
figure for 15 August was typical for the period from 25 July to 1 September. The
comparison shows that there is roughly a two- to threefold increase in diversity at
the beginning and again at the end of the summer. The total numbers of Macro¬
lepidoptera taken follow a similar distribution with an estimated 2,000 to 3,000 per
trap/night at the beginning of the season, decreasing to approximately 250 to 300
JANUARY 1970 ]
PROCEEDINGS
83
per trap/night during the middle period, and increasing to 1,000 to 2,000 per trap/
night at the end of the season.
The following interpretation is offered for these data. My collecting in the area
during the dry months of March and December indicates little insect activity dur¬
ing the dry period. One would expect the beginning of the rainy season to trigger
emergence, as food plants are only then generally available. It might be assumed
that as the rains begin to decrease at the end of the season, other species would
emerge. However, in both years the precipitation during the month of September
was fully as great as it was during the rest of the summer; so I do not feel that
decreasing humidity is the most important factor leading to this second emergence
peak.
In most areas of the United States emergence of insects and budbreak of plants
is correlated with accumulated day-degrees. There tends to be a relatively constant
turnover of species throughout the spring and summer, with attenuated emergence
peaks. However, in this region of Mexico, temperature is not the limiting factor;
and, therefore, day-degrees are of lesser importance in determining emergence.
With the addition of water to the environment, development of plants and emer¬
gence of insects is instantaneously initiated. Hence, the gradual turnover of species
is not as evident. I speculate that the two emergence peaks noted arise from the
use of two ecological strategies for survival during the dry season.
These two strategies operate as follows. As the stages of the life cycle most
resistant to desiccation are the egg and pupal stages, most moths would be expected
to pass the dry months in these states. With sufficient moisture insects in both
stages emerge. The first peak of adult emergence would then be due to the hatch¬
ing of the pupae. These adults lay their eggs at the beginning of the season, the
resultant larvae develop, and pupation until the following year takes place. The
second emergence peak is composed of the species which overwinter as eggs. The
eggs hatch with the coming of the rains, the larvae develop, there is a short pupal
stage, and the adults emerge late in the season, laying their eggs which then over¬
winter.
In summary it should be emphasized that the data and conclusions apply only to
Macrolepidoptera. However, it might be expected that other obligately phytopha¬
gous insect groups might utilize the same ecological strategies. Secondly, as this
study covered a relatively small region of Mexico subject to the climatic regime de¬
scribed (the most distant collecting localities being separated by 150 miles), it
cannot be said in how large a geographic area one would obtain the same observa¬
tions. However, the same emergence pattern might be found in general throughout
parts of the southwestern United States and northwestern Mexico where such cli¬
matic conditions prevail.
A social hour was held in the entomology rooms following the meeting.—M. S.
Wasbauer, Secretary.
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Vol. 46 APRIL 1970 No. 2
THE
Pan-Pacific Entomologist
SL0B0DCHIK0FF—Ratios: an intuitive vs. a quantitative approach in
Grotea (Hymenoptera: Ichneumonidae) - 85
KINN—A new genus of Celaenopsidae from California with a key to the
genera (Acari: Mesostigmata) _ 91
KELSEY—A new Scenopinus from the 1964 Galapagos International Scientific
Project (Diptera: Scenopinidae) _ 96
CLIFFORD—Variability of linear measurments throughout the life cycle
of the mayfly Leptophlebia cupida (Say) (Ephemeroptera: Lepto-
phlebiidae) _ 98
ROBERTSON—A new species of Pleocoma from southern California
(Coleoptera: Scarabaeidae) _ 106
ERWIN—The Nearctic species of the genus Leistus Frolich (Coleoptera:
Carabidae) _ 111
KUMAR AND LAVIGNE—A new genus and new species of Callipterinae
from India (Homoptera: Aphididae) _ 120
LEHMKUHL—The life cycle of Rhithrogena morrisoni (Banks) in western
Oregon (Ephemeroptera: Heptageniidae) _ 124
LINSLEY—New genera and species in the lepturine complex related to
Euryptera and Choriolaus (Coleoptera: Cerambycidae) _ 128
SMITH—Biology and structure of the dobsonfly, Neohermes californicus
(Walker) (Megaloptera: Corydalidae) _ 142
NEW BIOLOGICAL CONTROL ORGANIZATION _ 150
SCIENTIFIC NOTE _ 151
SOCIETY NOTICE _ 151
BOOK REVIEW _ 152
NEW BULLETIN _ 152
SAN FRANCISCO, CALIFORNIA • 1970
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
THE PAN-PACIFIC ENTOMOLOGIST
EDITORIAL BOARD
R. W. Thorp, Editor
E. G. Linsley R. 0. Schuster, Asst. Editor E. S. Ross
P. D. Hurd, Jr. D. L. Briggs, Editorial Assistant H. B. Leech
P. H. Arnaud, Jr. Treasurer W. H. Lange, Jr. Advertising
Published quarterly in January, April, July, and October with Society Proceed¬
ings appearing in the January number. All communications regarding nonreceipt
of numbers, requests for sample copies, and financial communications should be
addressed to the Treasurer, Dr. Paul H. Arnaud, Jr., California Academy of
Sciences, Golden Gate Park, San Francisco, California 94118.
Application for membership in the Society and changes of address should be
addressed to the Secretary, Dr. Marius S. Wasbauer, Bureau of Entomology, Cali¬
fornia Department of Agriculture, 1220 N. St., Sacramento, California 95814.
The annual dues, paid in advance, are $5.00 for regular members of the Society
or $6.00 for subscriptions only. Single copies are $1.50 each or $6.00 a volume.
Make checks payable to Pan-Pacific Entomologist.
The Pacific Coast Entomological Society
Officers for 1970
R. W. Thorp, President Paul H. Arnaud, Jr., Treasurer
F. L. Blanc, President-elect Marius S. Wasbauer, Secretary
Statement of Ownership
Title of Publication: The Pan-Pacific Entomologist.
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Location of Office of Publication, Business Office of Publisher, Publisher and Owner: Pacific Coast
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94118.
Editor: Dr. Robbin W. Thorp, Department of Entomology, University of California, Davis, California
95616.
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This issue mailed 29 May 1970.
Second Class Postage Paid at Lawrence, Kansas, U.S.A. 66044.
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yoiNTet, LAWRENCE, KANSAS
The Pan-Pacific Entomologist
Vol. 46
April 1970
No. 2
Ratios: An Intuitive vs. A Quantitative Approch in Grotea
(Hymenoptera : Ichneumonidae)
C. N. Slobodchikoff
University of California, Berekeley
The genus Grotea contains eight species, all from the Western Hemi¬
sphere (Slobodchikoff, 1970). Prior to revision, only five species
were described. Descriptions of these five species were based entirely
on color. When a revision was undertaken, it appeared that several
characters other than color would be ‘useful for separating species.
The intuitive taxonomic approach indicated that the shape and dimen¬
sions of the first median area of the propodeum, the second pleural
area of the propodeum, and the genal hook (the process ventral to the
lower end of the occipital carina) were good, definitive characters. It
seemed useful to convert the dimensions into ratios, and to use these
ratios in the descriptions and the keys to species.
Taxonomic keys often present characters in the form of ratios. For
example, a key may state that structure x is 0.6 the length of structure
y, or structure a is 1.5 the width of structure b. Such ratios often do
not have any accompanying statement of the degree of variation that
one may expect to find. It is implicit that some variation does occur,
but a non-specialist working a key often may not know just how much
variation he should compensate for and still correctly identify a given
species. The purpose of this present study was to find the degree of
variation in each of the characters mentioned above, and to test the
validity of using ratios as diagnostic characters.
The ratios were computed on the basis of the following sets of mea¬
surements: the length of the second pleural area of the propodeum vs.
its width at midpoint; the width of the genal hook at its base vs. its
width at 0.66 the distance from base to tip; and the anteroposterior
length of the first median area vs. its anterior width. All measurements
were made at 40 X with an American Optical Company Spencer micro¬
scope and an ocular micrometer. Means, standard deviations, and cor¬
relation coefficients were calculated using the University of California
Computer Center Statpak program and an IBM 7094 computer. The
The Pan-Pacific Entomologist 46: 85-91. April 1970
86
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
Table 1 . Means, standard deviations, and variances for three char¬
acters of Grotea spp.
Species
Pleural area
Median area
Genal hook
G. anguina
x = 3.66
x = 2.06
x = 2.59
N = 25
N = 25
N = 25
s = 0.580
s = 0.539
s = 0.323
s 2 = 0.3388
s 2 — 0.2916
s 2 = 0.1024
G. lokii
x — 4.38
x — 1.79
x = 1.88
N = 21
N = 21
II
to
1—1
s - 0.659
s = 0.471
s -- 0.383
s 2 = 0.4356
s 2 = 0.2209
s 2 = 0.1444
G. californica
x = 3.25
x — 1.28
x = 1.26
N = 25
N = 25
N = 25
s = 0.725
s = 0.254
s =- 0.190
s 2 = 0.5329
s 2 r= 0.0625
s 2 = 0.0361
G. californica
x = 3.04
x — 1.14
x = 1.26
local population
N = 18
N = 18
N = 18
s = 0.424
s - 0.249
s - 0.130
s 2 - 0.1764
s 2 = 0.0623
s 2 = 0.0169
G. mexicana
x = 1.76
x = 1.34
x : 1.80
N = 10
N = 10
N = 10
s — 0.144
s = 0.691
s = 0.391
s 2 - 0.0207
s 2 = 0.4774
s 2 = 0.1529
G. delicator
x = 2.83
X ~ 2.78
x = 1.49
N = 25
N = 25
N = 25
s = 0.272
s = 0.606
s = 0.162
s 2 = 0.74
s 2 = 0.366
s 2 = 0.026
characters of only five species were analyzed. Three species, Grotea
fulva , G. athenae , and G. perplexa were excluded due to the lack of a
sufficient number of specimens.
The pertinent data for the degree of variation for each character
are presented in Table 1. Correlation-coefficient analysis indicated that
each character varies independently of the other two. The actual mea¬
surements (in absolute units) and the resultant ratios are shown in
APRIL 1970]
SLOBODCHIKOFF—GROTEA RATIOS
87
x
Figs. 1-6: solid circle, Grotea anguina-, open circle, G. californica; solid square,
G. californica, Orinda population; open square, G. mexicana ; solid triangle, G.
delicator; open triangle, G. lokii. Measurements are in absolute units. Fig. 1.
Second pleural area of propodeum; y-axis, width at midpoint, x-axis, antero¬
posterior length.
figures 1-8. Figure 1 shows the scatter of ratios for the second pleural
area, figure 2 shows the ratios for the genal hook, and figure 3 shows
the ratios for the first median area.
Inspection of figure 1 shows that three species, Grotea anguina, G.
lokii, and G. californica clearly fall into a band of ratios between 3.06
and 5.46 and cannot be distinguished from one another on this basis.
Grotea mexicana, with a mean ratio of 1.76 and a standard deviation of
0.144, clearly falls ouside this band (x + s = 1.904), while Grotea
delicator, with a mean ratio of 2.83 and a standard deviation of 0.272,
falls mostly, yet with some overlap, outside this band (x + s = 3.102).
It may be concluded that second pleural area ratios are useful for
separating Grotea mexicana from the other four species, less useful
for separating Grotea delicator, and useless for separating Grotea an¬
guina, G. lokii, and G. californica from one another.
Inspection of figure 2 shows that three species, Grotea delicator,
G. lokii, and G. mexicana fall mostly into a band of ratios between 1.37
and 2.16, while Grotea californica (x s = 1.45) and G. anguina
88
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
X
Fig. 2. Genal hook; y-axis, 0.66 the length from base to tip, x-axis, basal
width. (See Fig. 1 for explanation of symbols).
X
Fig. 3. First median area of propodeum; y-axis, posterior width, x-axis, an¬
terior width. (See Fig. 1 for explanation of symbols).
APRIL 1970 ]
SLOBODCHIKOFF—GROTEA RATIOS
89
8 -
7-
Y
6 -
5 -
4 -
3 -
2~
O
o"
o
o ■
o "o o o
o o
o
o o
o
o o
o
o 4
-i-1-1- 1 - 1 -1- 1 -i-1-1-1- r~
4 5 6 7 8 9 10 11 12 13 14 15
X
Fig. 4. Second pleural area of propodeum; y-axis, width at midpoint, x-axis,
anteroposterior length. (See Fig. 1 for explanation of symobls).
5"
Y
4 -
3-
2 -
O
O
■ O
o
1 -
t -r
1 2
5
3
n r
4 5
X
6
7 8
Fig. 5. Genal hook; y-axis, 0.66 the length from base to tip, x-axis, basal
width. (See Fig. 1 for explanation of symbols).
90
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
5
4
Y
3
2 -
1 -
x
Fig. 6. First median area of propodeum; y-axis, posterior width, x-axis, an¬
terior width. (See Fig. 1 for explanation of symbols).
(x + s = 2.267) mostly fall above and below the band, respectively.
It may be readily seen that the genal hook ratios are useful for separat¬
ing Grotea californica from G. anguina, but are not useful for separat¬
ing either species from the other three.
Inspection of figure 3 shows that there is no distinct clustering of
ratios, most ratios falling into a broad band between 1.00 and 3.15.
Grotea mexicana seems to have a bimodal distribution of ratios that
does not appear to be correlated with either geographical location or
sexual dimorphism. It may be concluded that first median area ratios
are of no value in separating any of the five species.
A population of Grotea californica from the vicinity of Orinda,
California, was available for sampling, and the scatter of ratios from
that population was superimposed upon the scatter of G. californica
ratios from the entire geographical range of the species to see how
much local variation may be expected. Figures 4-6 show the resulting
scatters. Interestingly enough, there is almost a complete overlap of
the Orinda population ratios and the entire-range ratios for the genal
hook (Fig. 5) and the first median area (Fig. 6). The second pleural
area ratios (Fig. 4) indicate a slight separation into two groups based on
the magnitude of the individual measurements rather than on the
actual ratios.
-T~
2
■ O O o o
■ O o
o ■
o o
o
o o
o
o
1
3
—r
4
T-
5
—i—
6
~r •
8
APRIL 1970 ] KINN—A NEW GENUS OF CELAENOPSIDAE
91
Thus, it may be concluded that the degree of intraspecific variation
is quite large, and that, for one species at least, the degree of local
variation in the above three characters is as great as the degree of
variation found in the entire geographical range of the species. It may
be further concluded that ratios have a considerably more limited
utility, both in the key to the species and in the descriptions of the spe¬
cies, than was first apparent. The intuitive taxonomic approach in¬
dicated that the above ratios could be used for separating each of the
species discussed. A quantitative approach, however, showed that prem¬
ise to be fallacious.
Acknowledgments
The author would like to thank Mr. Andre K. Lehre, University of
California, Berkeley, for his help in computer programming and statis¬
tical techniques, Dr. Howell V. Daly, University of California, Berke¬
ley, for the loan of specimens from the vicinity of Orinda, California,
and the University of California Computer Center at Berkeley for prov¬
ing computer time.
Literature Cited
Slobodchikoff, C. N. 1970. A revision of the genus Grotea. Pan-Pac. Entomol.,
46: 50-63.
A New Genus of Celaenopsidae from California
with a Key to the Genera 1
(Acari : Mesostigmata)
D. N. Kinn
Division of Biological Control, University of California, Berkeley
Mites of the family Celaenopsidae are encountered under the bark of
dead trees, often in association with insects. In the course of a survey
of mite associates of California Scolytidae, a new genus and species be¬
longing to this family was found under the bark of Pinus monophylla
Torr. and Frem. killed by Ips confusus (LeConte). However, this spe¬
cies of mite is probably not associated with this insect.
The Celaenopsidae, until now, has consisted of four genera which
Tragardh (1951) reviewed and keyed. Like Celaenopsis Berlese and
1 This work has been supported in part by the California State Division of Forestry, the T. B.
Walker Foundation, the Surdna Foundation and various forest industries.
The Pan-Pacific Entomologist 46: 91-95. April 1970
92
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
Brachycelaenopsis Tragardh, the new genus, Schizocyrtillus , possesses a
postanal shield, whereas in the other two genera the postanal shield is
coalesced with the ventrianal shield.
Key to the Genera of the Celaenopsidae, Based upon Females
(after Tragardh, 1951).
1. Postanal shield present _ 2
Postanal shield absent _ 4
2. Metasternal shields large; irregularly rectangular ___ 3
Metasternal shields narrow; triangular in shape _-_ Schizocyrtillus
3. Postanal shield well defined_—_ Ceiaenopsis
Postanal shield narrow; weakly sclerotized _ Brachycelaenopsis
4. Median incision on anterior margin of ventrianal shield shorter than half
the width of the anterior margin, obscure, or located between two an¬
teriorly projecting lobes _ Pleuronectocelaeno
Median incision on anterior margin of ventrianal shield as long as half the
width of the anterior margin _ Ceratoceiaenopsis
Schizocyrtillus Kinn, new genus
Diagnosis. —Ventral and anal shields fused; postanal shield present between
ventrianal shield and posterior margin of the body. Female with narrow, slightly
triangular-shaped metasternal shields. Anterior margin of ventrianal shield with
a notched protuberance. Medial margin of ventrimarginal shield with a curved slit¬
like fissure posterior to coxa IV, from which the genus takes its name. Corniculi
of female elongate, bifurcate; bearing a seta-like projection medially. Gnathosomal
chaetotaxy of male altered with only subcapitular setae distinct; distal hypostomal
setae articulated, rod-shaped.
Type species: S. lathrius Kinn, new species.
Schizocyrtillus lathrius Kinn, new species
Female.— Idiosomal dorsum-. Length 514.1/4, mean of five specimens 516.5/4
(490.2-541.6) ; width 340.7/4, mean of three specimens 338.2/4 (336-340.7). Shape
oblong, oval; broadly rounded posteriorly, more pointed anteriorly; widest above
coxae IV; markedly convex; covered by single shield with more than 100 simple
setae, which increase in length toward posterior part of body. Six pairs of slit¬
shaped pores present. Pores I lie anterolaterally on podosoma; pores II to VI lie on
opisthosoma (Fig. 1C). Marginal plate fused with dorsal plate between coxae II
and III. Marginal plate with single row of simple setae; rounded posteriorly. Idio¬
somal venter-. Tritosternal base adjacent to anterior margin of sternal plate; laciniae
free and pilose (Fig. 1A and IB). Sternal shield wider than long; both anterior and
posterior margins concave. Sternal setae simple. Setae I located submarginally on
anterior angles of shield; setae II and III situated between coxae II. Setae III di¬
rectly posterior to setae I and setae II mesad of setae I and III. Pores I posterolateral
to setae I on margin of plate; pores II located laterally and slightly anterior to setae
III. Posterolateral intercoxal projections of sternal plate without points. Metasternal
plates narrow, lying between coxae II and III and bearing setae IV. Setae IV
short, less than half length of setae I to III. Pore III slit-shaped; located lateral
APRIL 1970 ] KINN—A NEW GENUS OF CELAENOPSIDAE
93
Fig. 1. Schizocyrtillus lathrius, paratype. A. Ventral aspect of female. B.
Sterno-genital region of female. C. Dorsal aspect of female.
to setae IV. Posterior margins of metasternal shields sinuous, with heavily
sclerotized indentations to accommodate heads of vaginal sclerites. Ventral, anal,
parapodal, peritremal, and metapodal shields fused. Anterior margin of ventrianal
shield convex, with a notched protuberance medially (Fig. IB). Vaginal sclerites
with ribbed clubs lie internally. Shield with 17 pairs of setae; with numerous
irregularly placed circular pores and two pairs of slit-shaped pores (one posterior
to coxae IV and the other anterolateral to anal orifice). Stigmata lateral to coxae
IV; peritremes sinuous, extending to anterior margin of coxae II. Ventrimarginal
shields border ventrianal shield from level of coxae II to posterior margin; shields
posteriorly separated from one another by postanal plate. Each ventrimarginal
shield with two simple setae, both setae anterior to anal orifice; with five slit¬
shaped pores posterior to coxae IV, and a slit-like fissure immediately posterior to
hind margin of coxae IV. Postanal shield with one pair of setae and two pairs of
slit-shaped pores (Fig. 1A). Legs: Armed with simple setae; chaetotaxy given
in Table 1 (using system of Evans, 1963). Leg I without claws; leg II stouter
than others. Tarsi II to IV with dorsal slit-shaped pores (Fig. 2A). Gnathosoma :
Table 1. Leg chaetotaxy, exclusive of coxa and tarsus, of adult
Schizocyrtillus lathrius.
I
II
III
IV
Trochanter
(1-1/1,1/2-0)
(1-0/1,0/2-1)
(1-1/2,0/1-0)
(1-1/2,0/1-0)
Femur
(1-2/1,2/3-1)
(2-2/1,2/2-1)
(1-2/1,2/1-0)
(1-2/1,3/1-0)
Genu
(1-3/1,2/1-1)
(1-3/1,2/1-1)
(1-2/1,2/1-1)
(1-2/1,3/1-1)
Tibia
(2-3/2,2/2-2)
(1-1/1,2/1-1)
(1-1/1,2/1-1)
(1-1/2,2/1-1)
94
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
300jj
APRIL 1970 ] KINN-A NEW GENUS OF CELAENOPSIDAE
95
Tectum triangular; terminating in an elongated point (Fig. 2B). Digitus fixus
with 7 to 9 small, posteriorly directed teeth. Digitus mobilis with about 10 teeth,
last tooth much larger than others; with two pilose appendages ventrally, anterior
appendage longer, but not extending beyond chela (Fig. 2C). Deutosternum
narrow. Median hypostomal setae longest and lateral shortest. Subcapitular setae
short, very stout. Corniculi bifurcate; hypopharyngeal processes pilose (Fig. 2D).
Chaetotaxy of palpal trochanter, femur and genu: 2, 5, 7. Proximal end of genu
with a slit-shaped pore on dorsal surface; palpal claw 2-tined.
Male. — Idiosomal dorsum : Length 493.0//, mean of two specimens 486.0//; width
320.3//., mean 322.0//. Shape like that of female. Idiosomal venter : Tritosternal
base contiguous with anterior margin of holoventral plate; laciniae free, pilose.
Holoventral plate with simple setae; sternal setae I in anterior angles of plate
and setae II and III situated more medially. Sternal pore I as in female; pore II
located posterolaterally to setae II and pore III near intercoxal projection between
coxae II and III (Fig. 2E). Ventral and anal pores as in female. Ventrimarginal
and postanal plates as in female. Legs: As in female. Gnathosoma: Tectum and
chelicerae like those of female. Deutosternum much wider than in female. Sub¬
capitular setae longer and finer than in female. Hypostomal setae reduced or
modified; paired rods, each bordered by two short setae, probably homologous
with distal hypostomal setae of female. Corniculi not as heavily sclerotized as in
female (Fig. 2F).
Type Material. — Holotype female, Scheideck, Ventura County,
California, 3 January 1969, D. N. Kinn and Y. L. Kinn, from under
the bark of Pinus monophylla Torr. and Frem. killed by Ips confusus
(LeConte). Allotype same collection data. Paratypes: 1 female (24
April 1966), same host and locality, D. N. Kinn, Y. L. Kinn and N. D.
Kinn; 1 female, 1 male (30 April 1966), same host and location, D.
N. Kinn and Y. L. Kinn; 1 female, 1 male (3 January 1969), same host,
location and collectors; 1 female (11 January 1969), same host, loca¬
tion and collectors.
Holotype and allotype are deposited in the United States National
Museum, Washington, D. C. Paratypes are deposited in the author’s
collection.
Literature Cited
Evans, G. 0. 1963. Observations on the chaetotaxy of the legs in the free-living
Gamasina (Acari: Mesostigmata). Bull. British Mus. Natur. Hist.
Zool., 10: 277-303.
Tragardh, I. 1951. Brachycelaenopsis, a new genus of Celaenopsidae (Acarina)
from Tasmania. Entomol. Tidskr., 72: 60-64.
4-
Fig. 2. Schizocyrtillus lathrius , paratype. A. Leg III of female. B. Tectum of
female. C. Chelicera of female. D. Gnathosome of female. E. Venter of male. F.
Gnathosome of male.
96
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
A New Scenopinus from the 1964 Galapagos
International Scientific Project 1
(Diptera : Scenopinidae)
L. P. Kelsey
Department of Entomology and Applied Ecology, University
of Delaware, Newark
As records of Scenopinidae from the Islands of the Pacific are
uncommon, it was of particular interest to the author to be able to
study three specimens collected on the 1964 Galapagos International
Scientific Project.
These proved to be a new species in the Velutinus Group of the genus
Scenopinus , common to North and South America and some of the
Pacific islands. In the keys to the Neotropical species (Kelsey, 1969)
it would terminate at S. schulzei Enderlein. However, it differs from
that species primarily in size, being only half as large; in the shape and
color of the antennae; in the shape of the 8th sternum; though both
species have a bump just proximal to the posterior margin; and in
the shape of the 9th sternum and bursal cavity.
Scenopinus galapagosensis Kelsey, new species
(Fig. 1)
Female.— Head black-brown to red-brown; eyes brown to red-brown, with a
narrow postocular ridge; frons moderately wide, flattened, with sharply sloping
sides, produced above the antennae, transversely striated, and with a median de¬
pression on lower fourth; ocellar tubercle prominent; ocelli brown; mouthparts
well developed, filling oral cavity, and with the rostrum brown; palpi dark
brown, flattened, and nearly as long as the rostrum; antennae black-brown,
second segment lighter distally, third segment pear-shaped, a little over twice as
long as wide. (See figures.)
Thorax dorsum black-brown to red-brown, transversely striated, with a silvery
pilose patch above the humeral callus; humeral callus red-brown, supra-alar
callus tan; pleural areas red-brown; halter stem red-brown, knob red-brown; wing
hyaline, veins brown, R4 branching from beyond the middle of the cell, cell R5
open to the tip of the wing; legs with coxae to tibiae red-brown, tarsi yellow-
brown.
Abdomen red-brown, 8th sternum subequal to tergum, and with a bulge on
midline anterior to distal margin; bursal pocket small, triangular. (See figures
for details.)
Length: 2 body 2.5 mm., wing 1.9-2.0 mm.
Male. —Unknown.
1 Contribution No. 70 Charles Darwin Foundation; Miscellaneous Paper No. 609 with the approval
of the Director of the Delaware Agricultural Experiment Station. Publication No. 393 of the De¬
partment of Entomology and Applied Ecology.
The Pan-Pacific Entomologist 46: 96-97. April 1970
APRIL 1970 ]
KELSEY—A NEW SCENOPINUS
97
Fig. 1. Scenopinus galagpagosensis Kelsey female; a. wing; b. c. lateral and
frontal aspects of head; d. enlarged detail of antennae; e. ventral aspect of 8th
and 9th sternites; f. lateral aspect of 8th and 9th segments; g. 9th sternum
and bursa.
Holotype female , Galapagos Archipeligo, Isla Santa Cruz, Horne-
man Farm, 220 M. 5 April 1964 (D. Q. Cavagnaro).
Holotype 2 Deposited in California Academy of Sciences # 9305.
Paratypes $ $ Same data except 18 March 1964. One in California
Academy of Sciences, one in the U. S. National Museum.
Literature Cited
Kelsey, L. P. 1969. A Revision of the Scenopinidae (Diptera) of the World.
U. S. Nat. Mus. Bull., 277, pp. 336, 208 figs.
98
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
Variability of Linear Measurements Throughout the
Life Cycle of the Mayfly Leptophlebia cupida (Say)
(Ephemeroptera : Leptophlebiidae)
Hugh F. Clifford
Department of Zoology, University of Alberta, Edmonton
Introduction
Mayfly nymphs of most species undergo a variable number of molts;
instars cannot be determined from head capsule measurements; and,
because of environmental factors, increase in size cannot be strictly
related to development, larger nymphs sometimes being physiologically
younger than smaller nymphs. Pleskot (1962) advocated that mayfly
life history data be based on developmental stages instead of or in
addition to total length measurements. Developmental stage analysis
has been used for mayflies as early as the 19th Century by Vayssiere
(1882), utilized successfully in the 1930’s by Ide (1935) and Raw-
linson (1939), and recently employed by several workers. Still, today,
most mayfly life history studies are based on linear measurements. In
this respect total length is the most common linear dimension, although
occasionally other dimensions are used, e.g. head width (Britt, 1962;
Levanidova and Rubanenkova, 1965), head length (Bretschko, 1965),
length of wing pads in combination with abdominal tergite width (Ide,
1935) and total length extrapolated from mandible length (Corbet,
1957).
Britt (1953) concluded that, for Ephemera simulans Walker nymphs
preserved in alcohol, head width was a more reliable index of growth
than total length, especially if the specimens had been injured. The
variability of linear measurements with age would be a useful criterion
for evaluating which measurement gives the best indication of the may¬
fly’s progress through its life cycle. However, one cannot determine
absolute age of nymphs from field samples, and without empirical
evidence age cannot be strictly correlated with total length. In fact
total length should be evaluated along with the other linear measure¬
ments. By utilizing developmental stages (each stage representing a
different physiological age) and measurements of relative variability,
I examined statistically how six linear measurements varied throughout
the life cycle of the mayfly Leptophlebia cupida (Say). With these
procedures it was also possible to evaluate the homogeneity of arbi¬
trarily chosen developmental stages.
The Pan-Pacific Entomologist 46: 98-106. April 1970
APRIL 1970]
CLIFFORD-MAYFLY VARIABILITY
99
Methods
Specimens of L. cupida used for this study were originally collected
as part of a continuing limnological program on the Bigoray River,
a subarctic brown-water stream of west-central Alberta (Clifford,
1969) . The nymphs, initially used for an allometry study (Clifford,
1970) , were first grouped into mm size classes (between 10 and
20 nymphs per size class) based on total length excluding the
cerci; nymphs 5.0 mm and longer were further separated into males
and females. Each nymph was then placed into one of four arbitrarily
chosen developmental stages by the appearance and development of the
mesothoracic wing pads: Stage I nymphs had no wing pads; Stage II
nymphs had small wing pads, their length being shorter than the dis¬
tance between the two wing pads; Stage III nymphs had a wing pad
length greater than the distance separating the two wing pads; Stage
IV nymphs had darkened wing pads indicating impending emergence.
Stage IV nymphs were in the last nymphal instar; the other stages each
represented several instars. Using an ocular eyepiece and dissecting
microscope, the following measurements were made to the nearest 0.01
mm on the nymphs of each stage: total length, head length, pronotum
width, mesonotum width, mesonotum length, and width of the fourth
abdominal tergite. Subimagos and imagos were also measured as
above.
The relative variability of each of the six linear measurements
throughout L. cupida ’s life cycle was determined by calculating the co¬
efficient of variation (CV) for each dimension of each developmental
stage, where CV is 100 times the standard deviation(s) divided by the
mean (x). The CVs of Stage IV nymphs, subimagos and imagos are
comparable to those of other animals; they are homogeneous samples
in respect to age and sex. The CV values of the other nymphal stages
are only meaningful for comparing the relative variability of the di¬
mensions within a particular stage. However, since each stage repre¬
sents a progressively older developmental age and collectively all stages
represent the entire life span, it is felt this sort of analysis is a valid
way to evaluate the variability of the dimensional measurements
throughout L. cupida’s life cycle.
Results and Conclusions
Homogeneity of developmental stages .—Figure 1 shows the seasonal
developmental cycle of L. cupida when large numbers of nymphs were
gathered throughout the year and separated into the four develop¬
mental stages. How precise are the arbitrarily chosen stages for inter-
100
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
STAGE
m
STAGE
1C
STAGE
ic
STAGE
I
1966 1967
Fig. 1. Seasonal developmental cycle of L. cupida nymphs from the Bigoray
River; total number of specimens per sampling date (except July 1967) ranged
from 94 to 1,302.
preting L. cupida?& developmental cycle? The CVs of subimagos and
imagos (and Stage IV nymphs except for the small sample size) can
be used as a yardstick for judging the homogeneity of the other stages
(Table 1). Stages III CVs agree well with those of the winged stages;
the CVs of Stage I and especially Stage II are somewhat higher than
those of winged stages. For Stage I it was necessary to determine
CVs without regard to sex; Stage II included unsexed nymphs (juve¬
niles) as well as nymphs whose sex could be determined, and for this
reason the same Stage II juveniles were used to calculate Stage II male
CVs as were used to calculate the Stage II female CVs. Even consider¬
ing the heterogeneity due to the mixing of juveniles with males and
females, the high CV values of Stage II, when compared to the other
stages, suggest that more accurate information on L. cupida ?s develop¬
mental cycle would be obtained if Stage II nymphs were further sub¬
divided by additional developmental criteria. Although it is obvious
APRIL 1970]
CLIFFORD-MAYFLY VARIABILITY
101
from Figure 1 that development is much slower during the long winter
than in other seasons, one should, without further subdividing Stage II,
use caution in interpreting the precise nature of winter development.
Variability of linear measurements .—Certain nymphal dimensions
did have consistently lower CY values (e.g. head length) than other
dimensions (e.g. mesonotum length). A clearer picture is obtained by
ranking the CV values of each dimension from 1 to 6 within each nymphal
stage and each sex. Rank 1 represents the lowest CV value for a given
dimension and 6 the highest. Then for a particular body part the
rank numbers by sex for the four nymphal stages are added up (in¬
cluding Stage I in summations for both males and females). There was
a slightly different sequence for the two sexes (Table 2). For females,
both head length and pronotum width had much lower total rank values
than the other measurements; for males, abdomen width and head
length would be the best linear measurement throughout the entire
nymphal life span. For both sexes, mesonotum length was the most
variable measurement. Total length was near the middle of each range
when the sexes were considered separately.
When the CV of each body part is calculated for each stage without
regards to sex (actual tabulated CV values not shown) and then ranked
as above, total length has a much higher total rank position, the se¬
quence being head length (8 total rank numbers), pronotum width
(12), abdomen width (13), mesonotum width (13), total length (19),
and mesonotum length (19).
It would be instructive to know if any of the CVs of Table 1 in¬
dicate dimensional measurements statistically more variable than others
within a particular stage. Lewontin (1966) shows that the variance of
the logarithms (common or natural) of measurements gives a measure
of relative variability that is invariant under a multiplicative change,
and hence can be used for statistical tests; he also points out for CVs
of about 30 or less that the square of the CV (in percentage form)
closely approximates the variances of natural logarithms. For L.
cupida each of the dimensional CVs of Table 1 was squared, and then
two null hypotheses were tested. (1) Within a particular stage none
of the squared CVs regardless of sex is significantly larger (one-sided
“F” test, 95% level) than the smallest squared CV of that stage. (2)
Within a particular stage none of the squared CVs of a particular sex
is significantly larger (same test) than the smallest squared CV of
that sex.
Certain dimensional measurements of female nymphs and subimagos
were significantly more variable than those of males (Hypothesis 1,
Table 1 . Essential statistics (in mm) and coefficients of variation (CV) of the six dimensional measurements of
juvenile (J), male and female nymphs, subimagos and imagos.
Stage I
J
Stagi
J + 5
e II
J+ $
Stage
$
III
2
Stage IV
$ 2
Subimagos
$ 2
Imagos
2
Number
17 1
45
47
19
29
4
4
17
17
19
Total Length
X
2.40
5.81
5.80
9.95
10.44
9.71
13.00
10.89
11.79
12.21
s
0.50
1.64
1.58
1.06
1.74
0.50
0.91
1.36
1.32
1.33
CV
20.89
28.17
27.18
10.68
16.67
5.15
6.98
12.52
11.16
10.90
Head length
X
0.44
0.97
0.95
1.58
1.58
1.45
1.62
1.03
1.02
1.02
s
0.09
0.24
0.23
0.18
0.17
0.06
0.12
0.09
0.13
0.15
CV
20.45
24.96
23.80
11.35
10.57
3.96
7.12
9.11
12.97
14.96
Pronotum width
X
0.65
1.38
1.41
2.24
2.32
1.96
2.35
1.83
1.85
1.82
s
0.13
0.34
0.36
0.26
0.28
0.16
0.17
0.20
0.26
0.27
CV
20.35
24.50
25.31
11.72
11.91
8.34
7.23
11.10
13.78
14.71
1 Except mesonotum length where there were 13 specimens for Stage I.
bO
102 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO.
Table 1 . Cont.
Stage I Stage II Stage III Stage IV Subimagos Imagos
J J + c? J + 2 $ 2 $ $ $ $ 2
Mesonotum width
X
0.66
1.38
1.40
2.36
2.41
2.38
2.67
2.11
2.04
2.05
s
0.15
0.32
0.35
0.33
0.32
0.22
0.24
0.24
0.30
0.26
CV
23.38
23.10
24.67
13.91
13.10
9.08
8.92
11.18
14.50
12.49
Mesonotum length
X
0.34
0.92
0.93
2.18
2.10
2.60
2.60
2.78
2.79
2.94
s
0.34
0.32
0.33
0.44
0.43
0.14
0.15
0.28
0.46
0.34
CV
99.38
34.46
35.27
20.06
20.42
5.44
5.87
9.89
16.38
11.39
Abdomen width
X
0.58
1.41
1.42
2.37
2.62
2.40
2.79
1.97
2.18
2.15
s
0.12
0.36
0.43
0.22
0.32
0.10
0.15
0.18
0.35
0.30
CV
20.26
25.46
29.97
9.43
12.24
4.17
14.03
9.07
16.14
13.73
APRIL 1970] CLIFFORD-MAYFLY VARIABILITY 103
104
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
Table 2. Total rank number variability of male and female nymphs
for the four developmental stages
Body part
Females
Males
Head length
8
10
Pronotum width
11
13
Abdomen width
15
8
Total length
15
14
Mesonotum width
16
17
Mesonotum length
19
22
Table 3). The variability was especially striking for female adbomen
width, even in the discrete instars, and possibly reflects egg develop¬
ment in the abdominal region. Considering the sexes separately, meso-
notum length (for both males and females) and total length (for fe¬
males of Stage III) were the only measurements significantly larger
than the dimension having the smallest relative variability (Hypothesis
2, Table 4).
In brief, and considering both the rank tabulation and the above
statistical tests, one can conclude that no single linear dimension is
Table 3. Body parts of a particular stage that had squared CVs
significantly larger 1 than the body part 2 with the smallest squared CV
of that stage and regardless of sex
Body part F D.F.
Stage I (Abdomen width)
Mesonotum length
4.87
16,16
Stage II ( $ Mesonotum width)
2 Mesonotum length
2.33
44,46
$ Mesonotum length
2.23
44,44
2 Abdomen width
1.69
44,46
Stage III ( S Abdomen width)
2 Mesonotum length
4.68
18,28
S Mesonotum length
4.54
18,18
2 Total length
3.17
18,28
Stage IV ( $ Head length)
2 Abdomen width
12.25
3,3
Subimagos ( $ Abdomen width)
2 Mesonotum length
3.28
16,16
2 Abdomen width
3.16
16,16
2 Mesonotum width
2.56
16,16
1 One-sided “F” test, 95% level
2 Tn parentheses
APRIL 1970 ]
CLIFFORD—MAYFLY VARIABILITY
105
Table 4. Body parts of a particular stage that had squared CVs
significantly larger 1 than the body part 2 with the smallest squared CV
of that stage and of the same sex
Body part F D.F.
Stage II
Males (Mesonotum width)
Mesonotum length
2.23
44,44
Females (head length)
Mesonotum length
2.20
46,46
Stage III
Males (Abdomen width)
Mesonotum length
4.54
18,18
Females (head length)
Mesonotum length
3.71
28,28
Total length
2.52
28,28
1 One-sided “F” test, 95% level
2 In parentheses
both least variable for males and least variable for females. Consider¬
ing the L. cupida nymphal population without regards to sex, the
usual procedure for interpreting mayfly life history phenomena, head
length (or in all likelihood, head width) and pronotum width would
be most desirable linear measurements, and mesonotum length would
be the least desirable measurement. When the sexes are treated sepa¬
rately, total length is, for most purposes, a seemingly justifiable mea¬
surement throughout most of the nymphal life span. But for the larger
female nymphs or when the nymphs are treated without regards to
sex, several of the other dimensional measurements exhibit much less
variability. Finally it is suggested that, because certain females dimen¬
sions are significantly more variable than those of males, the most
accurate interpretations of size-frequency data would be made when
nymphs are separated into males and females, regardless of the linear
dimension used. .
Acknowledgments
I am grateful for the technical assistance of Mr. Chi-hsiang Lei. This
study was supported by a grant from the National Research Council
of Canada. T ~
Literature Cited
Bretschko, G. 1965. Zur Larvalentwicklung von Cloeon dipterum, Cloeon
simile, Centroptilum luteolum und Baetis rhodani. Z. Wiss. Zool.,
172: 17-36.
Britt, N. W. 1953. Differences between measurements of living and preserved
aquatic nymphs caused by injury and preservation. Ecology, 34: 802-
803.
1962. Biology of two species of Lake Erie mayflies, Ephoron album (Say),
and Ephemera simulans Walker. Bull. Ohio Biol. Surv. 1: 70 pp.
106
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
Clifford, H. F. 1969. Limnological features of a northern brown-water stream,
with special reference to the life histories of the aquatic insects. Amer.
Midland Natur., 82: 578-597.
1970. Analysis of a northern mayfly (Ephemeroptera) population, with
special reference to allometry of size. Can. J. Zool., (in press).
Corbet, P. S. 1957. Duration of the aquatic stages of Povilla adusta Navas
(Ephemeroptera : Polymitarcidae). Bull. Entomol. Res., 48: 243-250.
Ide, F. P. 1935. The effect of temperature on the distribution of the mayfly
fauna of a stream. Univ. Toronto Stud., Biol. Ser. 39. Publ. Ont. Fish
Res. Lab., 50: 3-76.
Levanidova, I. M., AND L. S. Rubanenkova. 1965. On the methods of study¬
ing the life cycles of amphibiotic insects. Zool. Zh., 44: 35-45. (In
Russian).
Lewontin, R. C. 1966. On the measurement of relative variabliity. Syst.
Zool., 15: 141-142.
Pleskot, G. 1962. Einleitung: Symposium XII (Ephemeroptera). XI. Int.
Congr. Entomol. Proc. 1960, 3: 240-241.
Rawlinson, R. 1939. Studies on the life history and breeding of Ecdyonurus
venosus (Ephemeroptera). J. Zool., Proc. Zool. Soc. London (B), 109:
377-450.
Vayssiere, A. 1882. Recherches sur l’organisation des larves des Ephemerines.
Ann. Sci. Natur., (6) Zool. 13: 1-137. (cited from Ide, 1935).
A New Species of Pleocoma from Southern California
(Coleoptera : Scarabaeidae)
James Robertson
Culver City, California
Literature has been sparse and little is known of the habits and dis¬
tribution of the elusive beetles of the genus Pleocoma. However, with
the encroachment of civilization into our more primitive areas, more
locations of occurrence are being found and more material is available
for study. The fact that the adult beetles are active during rains re¬
quires greater than average perseverity on the collector’s part, especially
in regards to those species inhabiting remote mountainous areas. Severe
flooding, frequent landslides, and roads blocked by trees, make access
to these areas impossible at times. The author on several occasions has
been forced to dig through a landslide in order to return home after
a collecting excursion. Female Pleocoma must be diligently sought
after, for being flightless they remain on or in the ground. Searching
The Pan-Pacific Entomologist 46: 106-111. April 1970
APRIL 1970 ]
ROBERTSON-A NEW PLEOCOMA
107
for burrows, or following males to the abode of the female is the nor¬
mal method of collecting.
The habit of the male beetle of flying to lights and to the surfaces of
ponds (Davis, 1934), facilitates their collection. Material taken re¬
cently by myself and Noel McFarland in this manner, represents a new
species. It has the unique characteristic of possessing eight lamellae
in the antennal club. This feature excludes it from either of the “four
to five” or “six to seven” lamellate antennal club groups which are
presently utilized in classifying Pleocoma species (Linsley, 1946, Ellert-
son & Ritcher, 1959).
Pleocoma octopagina Robertson, new species
(Fig. 1-4)
Male (holotype).—Form large, robust, broadly oval, (Fig. 1), piceous-brown,
golden pubescent beneath. Head castaneous, edged with black; clypeal horn
very pronounced, deeply notched, moderately punctured, slightly fimbriate;
horn of vertex long, slightly notched, moderately clothed with long hairs, clypeal
and vertex horns subparallel, (Fig. 2) ; ocular canthi radiused on apex; eyes
prominent; antennae with scape moderately stout, subconical, second segment
moniliform, oblate; third segment four times as long as broad, longer than first
and second segments combined, slightly reflexed with flattened extended process
anteroventrad on outer end; fourth segment lamellate, longer than second and
third combined, fourth-fifths as long as fifth segment, markedly laciniate on
ventral edge, fifth segment lamellate, nine-tenths as long as sixth, less laciniate
than fourth, segments six to eleven lamellate, extremely long, longer than length
from antennal socket to eleventh segment, with pronounced curve outward at
tip, less and less laciniate on ventral edges, all segments containing setae to
varying degrees, especially on periphery, all lamellate segments covered with
minute recumbent hairs, (Figs. 3 & 4). Pronotum tending to blotchey brown
and piceous, approximately two and one-half times as broad as long; sides
subparallel, anterior corners broadly rounded, posterior corner narrowly rounded;
moderate anterior median depression only slightly crinite, rest of surface
finely, rather sparsely punctate, shiny, cirrose on periphery. Scutellum finely,
sparsely punctate, hardly fimbriate. Elytra piceous, somewhat shining; finely,
very sparsely, irregularly punctate, opaque; striae only faintly visible, cir¬
rose around outer margins. Body beneath castaneous, densely clothed in reddish
golden pubescence. Legs castaneous, stout, fossorial, pubescent. Length 27mm,
width 15mm (ranges from total type series, 25-31mm by 14-17mm).
Female and Immatures. —Unknown.
This species is immediately distinguishable from all others of the
genus Pleocoma by having a long lamellate fourth antennal segment,
thereby having eight lamellate plates in the club. All other species fall
into one of two groups with either four to five or six to seven plates.
The configuration of the third antennal segment also differs from that
of previously known Pleocoma. In the others it is cylindrical and
108
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
I mm
Figs. 1 and 2. Pleocoma octopagina Robertson. Fig. 1 (Upper.)
pect, adult male, holotype. Fig. 2 (Lower). Lateral view of head.
Dorsal as-
APRIL 1970 ]
ROBERTSON-A NEW PLEOCOMA
109
Figs. 3 and 4. Pleocoma octopagina Robertson. Fig. 3 (Upper). Dorsal view
right antenna. Fig. 4 (Lower). Ventrolateral view (beneath head) of an
tenna.
110
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
simple, while in octopagina the segment bears a flattened, extended
process.
Holotype male, 7 miles south of Pearblossom (2% miles ssw of
Valyermo), Los Angeles County, California (34°25' N., 117°52'
W.) elevation 4800 ft., 30 November 1965 (Jim Robertson) (Deposited
in the collection of the Los Angeles County Museum of Natural His¬
tory). Paratypes: 36 males, same locality as holotype, dates and col¬
lectors as follows: mid-December 1960 (Noel McFarland-flying at
night) ; 6, 19, 21, 26, 30 December 1963, January, 2 March 1964
(McFarland, in reservoir) ; 30 November, 9 December 1965, 8 Novem¬
ber, 8 December 1969, 10 January 1970 (Jim Robertson, blacklight and
in reservoir) (author’s collection; California Insect Survey Collection,
Berkeley; California Academy of Sciences; Los Angeles County Mu¬
seum; U. S. National Museum).
Distribution. —Thus far, this species has been found only at the
type locality, on the alluvial slopes of the north side of the San Gabriel
Mountains at elevations of 4800 to 4900 feet. Collecting attempts at
nearby points have proven negative, even though dates have been
the same.
Biology. —The type locality is situated in a mixed Chaparral-Pinyon
Woodland ecotone. Common perennial plants in the immediate vicinity
are Arctostaphylos glauca Lindl., Artemisia tridentata Nutt., Ceanothus
greggii var. vestitus (Greene) McMinn., Cercocarpus betuloides Nutt.,
Eriastrum sp., Eriodictyon trichocalyx Heller., Eriogonum fasciculatum
ssp. polifolium (Benth.) S. Stokes., Fremontia californica Torr., Lupi-
nus sp., Malacothamnus sp., Penstemon centranthifolius Benth., Quercus
chrysolepis Liebm., Rhamnus crocea ssp. ilicifolia Greene, Rhus tri-
lohata var. anisophylla (Greene) Jeps. Nearby, but at somewhat lower
elevations, Pinyon ( Pinus monophylla Torr. & Frem.), Juniper ( Juni-
perus calif ornica Carr), Scrub Oak ( Quercus dumosa Nutt.), and Yucca
(Yucca whipplei ssp. caespitosa (Jones) Haines.) become dominant,
along with Salvia dorrii (Kell.) Abrams, and Purshia glandulosa Cur¬
ran. Nearby, at somewhat higher elevations, Quercus chrysolepis
Liebm., becomes more abundant, along with Ceanothus cuneatus
(Hook.) Nutt., and Pinus coulteri D. Don. The thick duff which covers
the ground makes seeking of burrows difficult. Being on the south
edge of the Mojave Desert, summer temperatures (93—95° F.) can be
quite high, in spite of the elevation. In winter, temperatures drop low
enough for occasional snows. At the time of the author’s collecting,
the temperature was in the neighborhood of 40-50° F. Winter rainfall
averages 10.4 to 10.8 inches as recorded at two points, 2% miles on
APRIL 1970]
ERWIN-NEARCTIC LEISTUS
111
either side of the collecting site. Occasional summer rains (cloud¬
bursts) occur in July, August, and September, but it is essentially an
area of summer drought. Two nearby creeks (really dry arroyos)
flow after rains and in the early spring. Primary soil composition is
decomposed granite with occasional outcroppings of clay. The soil
make-up is loosely consolidated and well drained, and contains a black
powdery loam to some degree.
Acknowledgments
The author is indeed grateful to Dr. Charles Hogue of the Los An¬
geles County Museum of Natural History, Noel McFarland of The
South Australian Museum, and Dr. John Chemsak of the University of
California, Berkeley, for their advice and suggestions in the preparation
of this paper.
Literature Cited
Davis, A. C. 1934. A revision of the genus Pleocoma. Bull. S. Calif. Acad. Sci.,
33: 123-130.
Ellertson, F. E. and Ritcher, P. 0. 1959. Biology of rain beetles, Pleocoma
spp. Oreg. Agr. Exp. Sta. Tech. Bull., 44: 3-41.
Linsley, E. G. 1946. A preliminary key to the species of Pleocoma. Pan-Pac.
Entomol., 22: 61-65.
The Nearctic Species of the Genus
Leistus Frolich
(Coleoptera : Carabidae)
Terry L. Erwin
Harvard University, Cambridge, Massachusetts
Introduction
The nebriine genus Leistus Frolich contains many species in the
Palearctic region, but heretofore only two North American species have
been recognized, and one of these from only the female type in the
collection of Thomas Lincoln Casey (USNM). Lindroth mentions these
species in his “Ground-Beetles of Canada and Alaska” (1961). In
1965, while visiting the California Academy of Sciences, I came across
88 specimens of Leistus longipenrtis Casey that Van Dyke had col¬
lected, but never reported. During a visit previous to my own, Lind¬
roth had missed these specimens because they were not with the cara-
bids, and I discovered them by accident only. While studying this
The Pan-Pacific Entomologist 46: 111-119. April 1970
112
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
series, I noticed two types of male genitalia occurred. Further in¬
vestigation satisfied me that this series consisted of specimens of two
species, one of which was undescribed. In this paper the recognized
species are redescribed, one new species is described, the male genitalia
of all three species are figured, and a key is given to separate the
species.
The male genitalia of Nearctic and Palearctic forms are grossly dif¬
ferent in form. On the basis of this fact a new subgenus of Leistus is
proposed to contain the North American forms. In the past, other
authors (Reitter 1885, 1905, Daniel 1903, and Banninger 1925) have
erected various subgenera on the basis of characters such as presence
or absence of wings, and presence or absence of a posterolateral seta on
the pronotum. In a forthcoming paper I will discuss the classification
of Leistus on a world basis.
Methods and Materials
The methods and species criteria used here are those which I have
described in considerable detail elsewhere (Erwin 1965, and Erwin
1970).
The materials used here are specimens borrowed from, or seen in,
the California Academy of Sciences, San Francisco (CAS), the Museum
of Comparative Zoology, Cambridge (MCZ), and the United States
National Collection (USNM). One specimen, the holotype of L.
madmeridianus Erwin, was loaned to me by my colleague at San Jose
State College in 1965, Richard D. Spadoni.
Leistus Frolich
The type species is the European Carabus ferrugineus Linne, 1758: 415. The
following combination of characteristics is diagnostic of the genus: mandibles
moderately to broadly explanate; scrobes unisetose; ligula trifid at apex; cardines,
stipites, base of mentum at sides, and submentum strongly spinose, spines ar¬
ranged as circular cage; palpi extremely slender and elongate; head constricted
behind eyes; lateral thoracic setae present; venter at sides punctate from head to
first or second abdominal sternum.
Neoleistus Erwin, new subgenus
Type species.— Here designated, Leistus jerruginosus Mannerheim, 1843: 187.
The following combination of characteristics is diagnostic of this subgenus: pos¬
terior lateral setae of pronotum absent; sides of pronotum straight before square
hind angles; third antennal article shorter than fifth; mandibles broadly ex¬
planate; ligula with ventral bisetose cone posterior to apex; second abdominal
sternum not punctate; male genitalia with median lobe bipartite, apex extended
considerably beyond apical orifice.
APRIL 1970]
ERWIN-NEARCTIC LEISTUS
113
Key to the Species of Neoleistus
1. Humeri prominent (Fig. 2) ; hind wings fully developed; pronotum nar¬
rower than humeri just behind scutellum __ ferruginosus Mannerheim
Humeri strongly sloped (Figs. 1, 3) ; hind wings reduced outside stigma;
pronotum as wide as or wider than humeri _ 2
2. Pronotum as wide as humeri just behind scutellum; male median lobe as
in Fig. 4 _ longipennis Casey
Pronotum wider than humeri just behind scutellum; male median lobe
as in Fig. 6 _ madmeridianus Erwin, n. sp.
Leistus ferruginosus Mannerheim
(Figs. 2, 5, 7)
Leistus ferruginosus Mannerheim, 1843: 187. Lectotype, designated by Lind-
roth, 1961: 56, in University Museum, Helsinki. Type locality.—Sitka, Alaska,
as originally given by Mannerheim.
Leistus ferrugineus Dejean, 1831: 569. Junior homonym of Carabus ferrugineus
Linne 1758: 415. Lindroth 1961: 56.
Leistus nigropiceus Casey, 1913: 45. Holotype, a female, in USNM, number
46,843. Type locality.—Metlakatla, British Columbia, as originally given by
Casey. Lindroth 1961: 56.
Diagnostic combination. —The almost square humeri, narrow pronotum, and
short elytra separate these beetles from those of the following two species.
Description. —Medium-sized beetles, 8.0 to 9.0 mm. Color : Rufopiceous to
testaceous, elytra often slightly iridescent; legs, antennae and venter usually
paler. Microsculpture : Nearly isodiametric on head, more transversely stretched
on disc of pronotum and elytra. Macrosculpture: Anterior and posterior transverse
impressions of pronotum and elytral striae punctate. The following punctate:
head at sides, prosternum at sides, proepimera, mesepisterna, mesepimera,
metepisterna; metasternum at sides, and abdominal sternum 1 (and in some
specimens abdominal sternum 2) at sides. Head: Frontal furrows suggested,
continuous on clypeus. Antennal scape elongate, cylindrical. Mandibles broadly
explanate, seta present in scrobe. Ligula tripartite at apex with median, ventral,
setiferous keel. Mentum and submentum strongly setiferous, setae arranged as
ring around periphery of venter of head “collembola cage,” see Life History ).
Prothorax: Pronotum (Fig. 2) narrower than elytra across humeri; side margins
narrowly reflexed, straight just before hind angles. Pterothorax: Hind wings
fully developed. Elytral humeri prominent, arcuate. Genitalia: Male (Fig. 5) :
Median lobe with apex extended far beyond apical orifice. Apex broad, tapered
to acutely rounded point. Venter of shaft with median keel, nearly straight in
lateral aspect. Basal bend strongly arcuate, basal keel small. Internal sac with¬
out sclerites, but apically with small membranous papillae. Female stylus as in
Fig. 8. Twenty specimens of each sex investigated.
Variation. —The number of setiferous punctures in the third stria
of the elytra varies from two to five. The Alaskan and British Colum¬
bian specimens I have seen are at the lower end of the size range, while
the Washington and Oregon ones are at the higher end.
Flight. —The flight of these beetles has not been recorded.
114
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46, NO. 2
C
4
APRIL 1970 ]
ERWIN—NEAiRCTIC LEISTUS
115
Etymology. —Latin, ferrugineous = iron red, referring probably to
the overall color of these beetles, although they are more piceous than
ferrugineous.
Life history. —I have seen specimens collected from May to Au¬
gust and one teneral adult from 11 June 1936. The adults collected in
May were not teneral and were fully colored. It is possible that adults
overwinter, but see also Lindroth, 1961: 57. According to Howard
Frank (per. comm.) Palearctic species have members which feed on
Collembola using their peculiar mouth parts as a cage (previously un¬
reported in print).
Distribution.— (Fig. 7). The range of this species extends from
Alaska south to northern California, but apparently does not overlap
the range of the following species. Eastward it extends to Jasper, Al¬
berta and Revelstoke, British Columbia. I have seen 143 from the fol¬
lowing localities: British Columbia: (Garibaldi) MCZ, (Metlakatla)
MCZ, (Mountains between Hope and Okanogan) MCZ, (Nanaimo)
CAS, (Sidney) MCZ, (Stanley) CAS, (Steelhead) CAS, (Vancouver
Island) CAS, (Victoria) MCZ, (Yale) MCZ. Alaska: (Elmendorf,
Anchorage) MCZ, (Seward) CAS, (Wrangle) CAS, (Yakatoga Beach)
CAS. California: MCZ. Oregon: (Canon Beach) CAS, (Dilley) CAS,
(Florence) CAS, (Marshfield) CAS, (Olney) CAS, (Waldport) CAS.
Washington: (Crescent Lake) CAS, (Fairmont) MCZ, (Humptulipa)
CAS, (Longmire) CAS, (Monroe) CAS, (Mt. Bonaparte) MCZ, (North-
bend) CAS, MCZ, (Olympia) MCZ, (Port Angeles) CAS, (Seattle)
CAS, (Tacoma) MCZ, (The Forks), CAS. See Lindroth (1961) for
additional records.
Leistus longipennis Casey
(Figs. 1, 4, 7)
Leistus longipennis Casey , 1920: 148. Holotype, a female, in USNM, number
46,842. Type locality.—Humboldt County, California as originally given by
Casey.
Diagnostic combination.— The sloped humeri, wide pronotum, spatulate elytra,
and truncate wings separate these beetles readily from those of the preceding
species, but only male genitalic differences are reliable for positive separation
<-
Figs. 1-3. Pronotum and basal part of elytron, right side, dorsal aspect. Fig.
1. Leistus longipennis Casey, male, Oriclc, California. Fig. 2. L. jerruginosus
Mannerheim, male, Crescent Lake, Washington. Fig. 3. L. madmeridianus Er¬
win, male, Jacoby Creek, California. Fig. 4. Male genitalia of L. longipennis
Casey, a. parameres, b. left lateral aspect of median lobe, c. ventral aspect of
apex, Prairie Creek, California.
116
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 2
APRIL 1970 ]
ERWIN—NEARCTIC LEISTUS
117
from male members of the following species. After comparing numerous speci¬
mens one recognizes that the pronota of L. longipennis specimens are narrower
than the pronota of specimens of the following species.
Description. —Medium-sized beetles, 8.5 to 10.2 mm. Color, Micro sculpture,
Macroscultpure, and Head: As in L. ferruginosus. Prothorax: Pronotum (Fig.
1) wider than L. ferruginosus, slightly wider or subequal to width across humeri
just behind scutellum; side margins more broadly explanate, straightened for
longer distance just before hind angles than in L. ferruginosus. Pterothorax:
Hind wings truncate posterior to stigma. Elytral humeri sloped; elytra long and
spatulate, widened about apical third. Genitalia: Male (Fig. 4) : Median lobe
more strongly arcuate than L. ferruginosus, with ventral keel more prominent,
and basal keel higher. Female (as in Fig. 8). Ten males and twenty females
investigated.
Variation. —Elytral punctation varies as described for L. ferru¬
ginosus. There does not appear to be any geographic size difference in
the sample before me as in the preceding species.
Flight.— It is doubtful that these beetles can fly.
Etymology.— Latin, longus = long or lengthy, pennis = wing; re¬
ferring to the long spatulate elytra of these beetles.
Life history.— I have seen specimens collected in June and July.
The June specimens average paler than those collected in July, but no
markedly teneral adults have been seen.
Distribution.— (Fig. 7). The known range of this species is limited
to the extreme northwestern tip of Humboldt County, California. I
have seen 88 specimens from the following localities: California:
Humboldt County CAS, MCZ, USNM, (Deer Lodge, near Trinidad)
CAS, (Orick) CAS, (Prairie Creek) CAS, MCZ.
Leistus madmeridianus Erwin, new species
(Figs. 3, 6, 7, 8)
Type locality.—Jacoby Creek, 5.0 miles southeast of Arcata,
Humboldt County, California.
Type specimens.— The holotype male and allotype are in CAS. The
holotype was collected at the type locality by R. D. Spadoni on 23 June
<e
Figs. 5-6. Male genitalia, left lateral aspect. Fig. 6. L. madmeridianus Er¬
win, Eureka, California. Fig. 5. L. ferruginosus Mannerheim, Vancouver Is¬
land, British Columbia, Canada. Fig. 7. Distribution map of Nearctic Leistus spp.
Solid circles represent L. ferruginosus Mannerheim. Half-filled circle repre¬
sents state locality only of L. ferruginosus Mannerheim. Open circles represent
L. madmeridianus Erwin. Circle with a cross in middle represents L. longipennis
Casey. Fig. 8. left female stylus of L. madmeridianus Erwin, ventral aspect,
Eureka, California.
118
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
1965. The allotype was collected 7 July 1937 by E. C. Van Dyke.
Three paratypes, all males, collected on various dates in various local¬
ities are herewith designated. One is in CAS, MCZ, and my personal
collection.
Diagnostic combination.— The wide pronotum, wider than the elytral humeri
just behind the scutellum, separates these beetles from members of the two
preceding species.
Description.— Medium-sized beetles, 8.7 mm to 10.0 mm. Color, Micro sculpture,
Macrosculpture, and Head : As in L. ferruginosus. Prothroax : Pronotum (Fig.
3) broad, wider than elytra across humeri just behind scutellum; side margins
more widely reflexed than L. longipennis', straight just before hind angles as in
L. longipennis. Pterothorax : Hind wings truncate posterior to stigma. Elytral
humeri sloped; elytra long and spatulate, widened about apical third. Genitalia:
Male (Fig. 6) : Median lobe similar to L. ferruginosus, except apex beyond
apical orifice longer. Shaft straighter than in two preceding species. Female
(Fig. 8) : Stylus trigonal; with two ventral setae. Membranes at base densely
setiferous. Three males and two females investigated.
Variation.— Too few specimens are known to evaluate variation.
Flight. —It is doubtful that these beetles can fly.
Etymology.— Latin, meridianus = southward; Mad, from the Mad
River; referring to the present known distribution of these beetles south
of the Mad River.
Life history. —Teneral specimens were collected in June and July.
The other specimens seen were not labelled with date of collection.
Distribution.— (Fig. 7). The known range of this species extends
from the Point Reyes Peninsula of California to the type locality in
Humboldt County, California, just south of the Mad River. I have
seen five specimens from the following localities: California: Hum¬
boldt County CAS, (Jacoby Creek, 5.0 miles southwest of Areata) CAS;
Marin County (Point Reyes) CAS.
Discussion
Because a more detailed discussion on the phylogeny and zoo¬
geography will be forthcoming in a paper on Leistus classification, a
few words here will suffice. The Nearctic Leistus are no doubt a mono-
phyletic group that has entered the New World via “Beringia.” They
probably entered with the widespread arctotertiary forests in the early
Tertiary and have been restricted to the remnants of these forests until
now. Two of these species are confined to the “redwood belt” of
northern California, while the third species is more widespread, but
still confined to the more moist coastal forests of the northwestern
United States, British Columbia, and Alaska. The small local popula-
APRIL 1970 ]
ERWIN-NE ARC TIC LEISTUS
119
tions of L. longipermis and L. madmeridianus could either be all that
is left of more widespread populations (restricted along with the re¬
stricted redwoods) or small peripheral populations of L. ferruginosus
which have become isolated, brachypterous, and have undergone specia-
tion.
Acknowledgments
I would like to thank George E. Ball and Donald R. Whitehead for
reading the manuscript of this paper. Also I would like to thank my
wife La Verne for her assistance in preparation of this paper.
Literature Cited
Banninger, M. 1925. Neunter Beitrag zur Kenntnis der Carabinae: die
Nebriini. Entomol. Mitt., XIV: 180-343.
Casey, T. L. 1913. Cicindelidae and Carabidae. Memoirs on the Coleoptera, IV.
New Era Co., Lancaster, Pennsylvania. 192 pp.
1920. Memoirs on the Coleoptera, IX. New Era Co., Lancaster, Penn¬
sylvania. 529 pp.
Daniel, K. 1903. Bestimmungstabellen der europaischen Koleopteren. LII.
Muenchn. Koleopterol. Z., 155-173.
Dejean, P. F. M. A. 1831. Species general des Coleopteres de la collection de M.
le comte Dejean. V. Paris, viii + 883 pp.
Erwin, T. L. 1965. A revision of Brachinus of North America: Part I. The
California species (Coleoptera: Carabidae). Coleopt. Bull., 19(1):
1-19.
1970. A reclassification of bombardier beetles and a taxonomic revision of
the north and middle American species (Carabidae: Brachinida).
Quaest. Entomol., 6(1): 4-215.
Lindroth, C. H. 1961. The ground-beetles (Carabidae, excl. Cicindelinae)
Alaska, Part 2. Opuscula Entomol., Suppl. XX. pp. 1-200.
Linne, C. 1758. Systema Naturae, 10th edition, 824 pp.
Mannerheim, C. G. 1843. Beitrag zur Kaefer-Fauna der Aleutischen Inseln,
der Insel Sitka und Neu-California. Bull. Soc. Natur. Moscu, XVI,
pp. 175-314.
Reitter, E. 1885. Analytische Uebersicht der bekannten europaischen Arten
der Coleopterengattung Leistus Frohl. Wien. Entomol. Ztg., IV. pp.
213-219.
1905. Zur systematischen Gruppeneint-eilung des Coleopteren-Genus
Leistus Frolich und Ubersicht der mir bekannten Arten. Wien. En¬
tomol. Ztg., XXIV. pp. 209-225.
120
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
A New Genus and New Species of Callipterinae
from India 1
(Homoptera : Aphididae)
Rabinder Kumar and Robert Lavigne
University of Wyoming, Laramie
Among the aphids collected by the senior author while working on
the migration of aphids from potato fields to alternate hosts at the
Central Potato Research Institute, Simla, India, were two new species
one of which was found to represent a previously undescribed genus.
Tuberdefectus Kumar and Lavigne, new genus
This new genus closely resembles the genus Tinocallis Matsumura
(1919), in the presence of setae on the outer side of the cornicles and
in the presence of two centrodorsal rows of spinal setae. The pairs of
spinal setae in the new genus are equidistant from each other and do
not have the characteristic pattern found in the genus Tinocallis. Un¬
like Tinocallis the abdominal tubercles are completely absent in the
new genus. This character is shared with genus Therioaphis Walker
(1870).
The type species is T. eastopi Kumar and Lavigne, new species.
Tuberdefectus eastopi Kumar and Lavigne, new species
(Figs. 1-5)
Alatoid nymph.— Color in life: pale to light yellow. Sclerites on dorsum of
abdomen visible with naked eye. Color when macerated: yellow disappears,
antennae and apical portions of legs brownish.
Morphology .—Head blunt without swelling; frontal tubercles absent; ocelli
absent; triommatidion visible. Length of antennal segments: III 0.31-0.40 mm,
IV 0.14-0.21 mm, V 0.17-0.21 mm, and VI 0.12-0.15 + 0.11-0.14 mm; long¬
est hair on antennal segment III nearly % the basal diameter of that segment.
Rostrum reaches beyond the front coxae. Abdomen with two rows of sclerites on
dorsum of segments I—VII; abdominal setae prominent and strongly capitate as in
Fig. 2; lateral abdominal sclerite not developed; each sclerite with 3-7 setae;
seta of abdominal sclerite VI present on the outer side of the cornicle on either
side; dorsal and lateral abdominal tubercles lacking; longest spinal seta present
dorsally on abdominal segment VIII.
Alate male. —Color in life: pale yellow. Sclerites on dorsum of abdomen dis¬
tinctly visible with naked eye. Color when macerated: yellow disappears, an¬
tennae, legs, head, thorax brownish; abdomen with brown patches dorsally.
Morphology .—Head moderately blunt without swelling; frontal tubercles ab-
1 Published with the approval of the Director, Wyoming Agricultural Experiment Station as
Journal Article No. 398.
The Pan-Pacific Entomologist 46: 120-124. April 1970
APRIL 1970 ]
KUMAR & LAVIGNE-NEW CALLIPTERINE APHIDS
121
Figs. 1-5. Tuberdejectus eccstopi Kumar and Lavigne. Fig. 1. Dorsum of ab¬
domen of alate viviparous female. Fig. 2. Dorsum of abdomen of alatoid nymph.
Fig. 3. Right antennal segment III of alate viviparous female. Fig. 4. Right an¬
tennal segment IV of alate viviparous female. Fig. 5. Right fore wing of alate
viviparous female.
sent; ocelli very prominent; triommatidion visible. Length of antennal segments:
III 0.52-0.58 mm, IV 0.24-0.31 mm, V 0.23-0.28 mm, and VI 0.14-0.16 +
0.12-0.16 mm; secondary rhinaria on antennal segments: III 21-23, IV 1-4,
V 4-5 and VI 2-3; these vary in size, position and number in the same specimen
on both antennae and in different specimens; longest hair on antennal segment
III nearly half the basal diameter of antennal segment III; hair present on
antennal segments I-VI. Rostrum reaches beyond the front coxae. Distal setae
single. Prothoracic setae normal and single. Setae on body parts pointed. Two
rows of sclerites dorsally on abdominal segments I—VII; sometimes sclerites of
tergite VI coalesce; each sclerite with a spinal seta; lateral abdominal sclerites
are well developed in abdominal segments I-V; sclerite VI coalesces with base
of cornicle on either side; sclerites VII and VIII very faintly developed; each
sclerite with a seta except on the outside of the cornicle on either side on ab¬
dominal segment VI; dorsum of abdomen without any tubercles; lateral abdominal
tubercles present on abdominal segments II-V. Cauda knobbed and densely
hairy. Anal plate deeply indented. Wings (Fig. 5) and legs normal.
Alate viviparous female. —Color in life: pale yellow. Sclerites on dorsum
of abdomen distinctly visible with naked eye. Color when macerated: yellow
122
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
Figs. 6-10. Chromaphis hirsutustibis Kumar and Lavigne (alate viviparous
female). Fig. 6. Dorsum of abdomen. Fig. 7. Right fore wing. Fig. 8.
Right antennal segment III. Fig. 9. Right antennal segment IV. Fig. 10. Right
femur.
disappears, antennae, legs, head, thorax brownish; abdomen with brown patches
dorsally.
Morphology .—Head moderately blunt without swelling; frontal tubercles ab¬
sent; ocelli very prominent; triommatidion visible. Length of antennal segments:
III 0.62-0.71 mm, IV 0.30-0.36 mm, V 0.26-0.32 mm, and VI 0.15-0.17 -f-
0.15-0.18 mm; secondary rhinaria on antennal segment: III, 20-23 in number,
upper y? of segment III rarely with rhinaria as in Fig. 3, these vary in size,
position and number on both antennae in the same individual and in different
specimens; longest hair on antennal segment III nearly half the basal diameter
of segment III; hair present on antennal segments I-VI (Fig. 4). Rostrum
reaches beyond front coxae. Distal setae single. Prothoracic setae normal and
single. Setae on body parts pointed. Abdomen with two rows of sclerites on
dorsum of abdominal segments I-VII; sclerites of middle tergites sometime
coalesce with those on anterior and posterior abdominal segments; lateral ab¬
dominal sclerites well developed on segments II-V; sclerite VI coalesces with
base of cornicle on either side; sclerite VII only faintly developed; each sclerite
with a seta, that on abdominal sclerite VI present on outer side of cornicle;
setae on abdominal segment VIII large; dorsum of abdomen without tubercles
as in Fig. 1; lateral abdominal tubercles present on segments II-V. Cauda
knobbed and densely hairy. Anal plate deeply indented.
Types.— Holotype alate male, Simla, India, 7000 ft., 23 August 1966
(R. Kumar) on Primus sp. (wild peach). Allotype: alate viviparous
female, same data. Paratypes: one alatoid nymph, one alate male and
APRIL 1970] KUMAR & LAVIGNE—NEW CALLIPTERINE APHIDS
123
one alate viviparous female. Deposited in British Museum (Nat. Hist.),
London, England. Four paratypes in collection of senior author.
Comments. —The aphids were first observed on wild peach on 23
August 1966. Alate males were present only in the latter part of Jan¬
uary. Viviparous reproduction was at its peak during February. All
stages feed near the midribs of the leaves forming dense colonies.
Leaves become very sticky due to aphid excretion and are often visited
by ants.
The authors take great pleasure in naming this species for the dis¬
tinguished aphidologist, Dr. V. F. Eastop, British Museum (Nat. Hist.)
who kindly examined the specimens and compared them with existing
genera.
Chromaphis hirsutustibis Kumar and Lavigne, new species
(Figs. 6-10)
Alatoid nymph.— Color in life: pale yellow with paired patches black dorsally
on abdominal terga. Color when macerated: becomes colorless except for ab¬
dominal patches blackish.
Morphology .—Frontal tubercles poorly developed. Antennae six segmented;
length of antennal segments; III 0.23-0.24 mm, IV 0.11-0.12 mm, V 0.11 mm,
and VI 0.08-0.09 + 0.04 mm; secondary rhinaria absent; longest seta on seg¬
ment VI, setae of antennae, head, thorax and abdomen capitate; apex of an¬
tennal segments V and VI dark brown; most of antennal setae present on the
inner side. Head and pronotum with one pair of setae. Two pairs of spinals on
mesonotum, metanotum and abdominal tergites I—VII. Abdominal tergite VIII
usually with six spinals; with median pair spinals often longest; mostly setae
arise from dark sclerotized rows placed segmentally; abdominal segments I-VII
with 3-4 lateral capitate setae, those on segment VI situated on the ventral sur¬
face of cornicle on either side.
Alate viviparous female. —Color in life: yellowish with prominent black
patches on dorsum of abdomen. Costal margins of wing dark brown. Abdominal
patches and wing markings absent in early stages but become prominent with
maturity. Color when macerated: yellow disappears, whereas brownish areas
remain.
Morphology .—Frontal tubercles poorly developed; frontal ocellus visible when
head viewed from above. Longest hair on head much longer than basal diameter
of antennal segment III, but shorter than setae on dorsum of abdomen. An¬
tennae six segmented; length of antennal segments: III 0.57-0.58 mm, IV 0.25-
0.33 mm, V 0.18-0.20 mm, and VI 0.08-0.16 + 0.04 mm; secondary rhinaria
on segment III: 8-10 (Fig. 8) IV: 1 (Fig. 9); longest hair on antennae % of
basal diameter of segment III and pointed; seta on segments I and II much
longer than on other segments. Proximal part of antennal segments IV, V and
processes terminalis dark brown. Rostrum reaching beyond front coxae. All
body setae pointed. Spinal setae on dorsum of abdomen segmentally arranged,
very prominent median dark brown bars on dorsum of abdominal segments IV
and V which sometime coalesce at points, marginal sclerites present on abdominal
124
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
segments I-IV, usually with 4-5 setae in Fig. 6. Cornicles small and without
flange. Cauda knobbed. Anal plate weakly indented. Wings normal as in Fig.
7. Hind tibiae densely hairy as in Fig. 10, and blackish on the margins in the
middle region, upper half comparatively darker, dark blackish brown patch
present below the apical region on femur.
Types.— Holotype alate viviparous female, Simla, India, 21 September 1966
(R. Kumar) on Juglans regia L. Paratypes: one alate viviparous female and one
alatoid nymph, same data. Deposited in the British Museum (Nat. Hist.), Lon¬
don, England. Four paratypes in collection of senior author.
Comments.— This new species is easily distinguished from other
species of the genus Chromaphis Walker (1870) by the presence of
dense hair on the hind tibiae.
Literature Cited
Matsumura, S. 1919. New species and genera of Callipterinae (Aphididae)
of Japan. Trans. Sapporo Natur. Hist. Soc., 7: 99-115.
Walker. F. 1870. Notes on Aphids. Zoologist, 5: 1966-2000.
The Life Cycle of Rhithrogena morrisoni (Banks)
in Western Oregon
(Ephemeroptera : Heptageniidae)
D. M. Lehmkuhl 1
Oregon State University, Corvallis
Detailed biological data are lacking for the North American species of
Rhithrogena. Some species in the genus are distinguished only by
minute details of the male genitalia, and the females and nymphs are
frequently undescribed. The species treated here will be referred to as
R. morrisoni. Specimens of all stages are deposited in the Oregon State
University Insect Collection, Corvallis.
Rhithrogena morrisoni was described by Banks from Nevada in 1924.
It also occurs in western Canada (Berner, 1959), California (Day,
1963), Idaho (Jensen, 1966), Oregon (Allen and Edmunds, 1956),
and Utah (Edmunds, 1952). Jensen (op. cit.) described the nymphs
and included both adult males and nymphs in his keys to the Rhithro¬
gena of Idaho.
Methods and Collections
Nymphs of R. morrisoni were collected from a variety of streams in
the Benton County area. The description of nymphal development is
1 Present address : Department of Biology, University of Saskatchewan, Saskatoon, Canada.
The-Pan Pacific Entomologist 46: 124-127. April 1970
APRIL 1970 ] LEHMKUHL—RHITHROGENA LIFE CYCLE
125
based on intensive sampling in Oak Creek, near Corvallis, from July
1967 to June 1968. Adults were reared or collected in 1966, 1967, and
1968.
In the sample area, Oak Creek was 5-10 feet wide, 6—18 inches deep,
and had a dense canopy of deciduous trees. The sample area consisted
of a series of riffles and pools, the former having a rubble and gravel
substrate. Other information on the stream can be found in Anderson
and Lehmkuhl (1968) and Lehmkuhl (1968).
Benthos samples were taken by embedding the end of a one foot
length of stovepipe 6 inches in diameter into the substrate. Contents
of the sampler were removed by hand and with a fine aquarium net.
Surface velocity, water depth, distance from shore, and general bottom
type were recorded for each sample. Three samples were taken each
month from each of the major habitats in the stream: slow deep glide,
still backwater, middle of riffle, edge of riffle (12 samples per month).
Mayfly nymphs of equal maturity may vary much in size, i.e., ulti¬
mate instar nymphs (having black wing pads) may differ in length by
several mm. In addition to length measurements, a system of group¬
ings based on biologically significant characters was used, as described
below:
Group
Biological Significance
I
Gills poorly developed, threadlike or
absent; nymphs scarcely resemble may¬
flies.
Newly hatched.
II
Definite mayfly appearance, gills de¬
veloped beyond I, no wing buds.
Young nymphs.
III
Wing buds present on posterior mar¬
gin of meso- and metathorax; meso-
thoracic wing pads do not completely
cover metathoracic wing pads.
Half grow nymphs.
IV
Mesothorarcic wing pads cover meta¬
thoracic wing pads.
Mature nymphs.
V
Wing pads black.
Ready to emerge to
subimago.
To obtain adults, nymphs collected in the field were placed in small
screen cages which were partly submerged in a trough in which water
was circulated by a paddle wheel. Nymphs could rise to the water sur¬
face or crawl up the side of the screen to emerge.
Life Cycle
Of over 100 samples taken in Oak Creek, about 15% contained large
stones (4— 5 /r dia.) and were in water over 2 ft./sec. surface velocity.
126
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
Table 1. Sizes and natural groups of Rhithrogena morrisoni
(Banks) nymphs from Oak Creek bottom samples, 1967-1968.
SEPT.
OCT.
NOV.
DEC.
JAN.
FEB.
Length
(mm)
1-2
3-5
6-8
8-9
none
8-9
Natural
Groups 1
II
II-III
III
IV
—
V
1 See text.
Nymphs of R. morrisoni were found only in these samples indicating
that the nymphs have strict environmental requirements.
Table 1 illustrates the seasonal development of the nymphs in the
1967-1968 sample period. None was found from March to August but
small nymphs may have been overlooked in August because of the
presence of Cinygmula reticulata McDunnough nymphs which might be
confused with Rhithrogena. Nymphs of R. morrisoni were 1-2 mm
long in September and had increased to 9 mm in December. Severe
spates in the fall and winter caused much disturbance in the stream
(Anderson and Lehmkuhl 1968), and this may explain the absence of
nymphs in the January sample. Nymphs increased greatly in maturity
but only slightly in length from November to February (compare
lengths and natural groups, Table 1). Nymphs had black wing pads
(natural group V) in February, and the first adults appeared at this
time.
Adults were collected between the following dates: 1966—25 March
to 26 April; 1967—11 March to 21 April; 1968—29 February to 24
April. In 1968, the main emergence from Oak Creek was early and of
short duration (first week of March), probably because of several days
of unseasonably warm weather at this time (air temp., over 20° C;
water temp., about 7° C). This explains the absence of nymphs in
March and April samples (Table 1). Adults from other areas were
collected as late as 24 April 1968.
Cast skins from laboratory-emerged specimens were found floating
free on the surface of the water, indicating that the nymphs do not
climb out of the water to transform to the winged stage. The subimago
stage lasted from 4-6 days for laboratory reared specimens kept at
outside temperatures of 5-15° C. Subimagos collected on buildings in
Corvallis and kept indoors (21-23° C) usually required 2 days to
transform to the adult stage.
A mating swarm was observed over a roadside ditch (March 2, 1968)
near a stream at 3 P.M. on a misty day. The swarm consisted of a
dozen males widely spaced 8-15 ft. above the ground. Jensen (1966),
APRIL 1970 ] LEHMKUHL—RHITHROGENA LIFE CYCLE
127
reported that the adults swarm over streams and adjacent ground in
mid-morning, but that the main flight occurs in the evening.
No nymphs were collected in the summer indicating that the eggs
have a resting period of several months. This is similar to R. semi-
colorata in England (Macan, 1960).
Discussion
Jensen (1966) reported that R. morrisoni had an early emergence in
Idaho and Utah (May and June). This species also has an early emer¬
gence in the Corvallis area (March and April). Macan (1960) sug¬
gested that the end of the period of emergence occurs when rising water
temperatures reach a point which kills the nymphs. I have shown that
emergence may be correlated with temperature in some species of
Epeorus (Lehmkuhl, 1968), but that temperature alone will not explain
the emergence period in all cases for a given species. In the present
case it is unlikely that high water temperatures limit the period of adult
emergence. Jensen (1966) found R. morrisoni nymphs in water up to
18° C, which is well above the temperature in Oak Creek when the last
adults emerged.
Literature Cited
Allen, R. K., and G. F. Edmunds, Jr. 1956. A list of the mayflies of Oregon.
Proc. Utah Acad. Sci. Arts, Lett., 33: 85-87.
Anderson, N. H., and D. M. Lehmkuhl. 1968. Catastrophic drift of insects
in a woodland stream. Ecology, 49: 198-206.
Banks, N. 1924. Descriptions of new neuropteroid insects. Bull. Harvard
Univ. Mus. Compar. Zool., 65: 421-455.
Berner, L. 1959. A tabular summary of the biology of North American may¬
fly nymphs (Ephemeroptera). Bull. Fla. State Mus. Biol. Ser., 4(1):
1-58.
Day, W. C. 1963. Ephemeroptera. (P. 79-105) in Usinger, R. L. (ed.) Aquatic
Insects of Calfiornia. Second printing, Univ. Calif. Press, Berkeley.
508 p.
Edmunds, G. F., Jr. 1952. Studies on the Ephemeroptera. Unpublished Ph.D.
Thesis, University of Massachusetts. 1-399 pp.
Jensen, S. L. 1966. The mayflies of Idaho. Unpublished M.S. Thesis. Uni¬
versity of Utah, Salt Lake City. 1-367 pp.
Lehmkuhl, D. M. 1968. Observations on the life histories of four species of
Epeorus in western Oregon (Ephemeroptera : Heptageniidae). Pan-
Pac. Entomol., 44: 129-137.
Macan, T. T. 1960. The effect of temperature on Rhithrogena semicolorata
(Ephem). Int. Rev. Ges. Hydrobiol., 45: 197-201.
128
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
New Genera and Species in the Lepturine Complex Related to
Euryptera and Choriolcius
(Coleoptera: Cerambycidae)
E. Gorton Linsley 1
University of California, Berkeley
The Lepturinae of Mexico and Central America, last reviewed by
Bates (1879-1885), are still very poorly known. The present paper
is designed to make available names for certain genera and species
which have special evolutionary importance in relation to the
lepturine fauna of the United States. It is a companion to that of
Chemsak (1969) in which he characterizes for the first time Mexican
and Central American species of Strangalia Audinet-Serville. Large
numbers of undescribed genera and species from Mexico and Central
America are now at hand. It is our intention to make many of these
known as time permits. However, priority will be given to the naming
of presumably mimetic forms and others of special ecological or
evolutionary significance.
Megachoriolaus Linsley, new genus
Form moderately robust; elytra parallel-sided or slightly expanded posteriorly,
not attenuated. Head elongate; antennae usually relatively short in both sexes,
basal segments shining, clothed with coarse suberect hairs, fourth segment short,
outer segments dull, with or without poriferous areas. Pronotum with punctation
not finer than that of base of elytra, usually very much coarser. Prosternum
with intercoxal process very narrow, often laminiform, coxae prominent, exerted,
approximate; mesosternum with intercoxal process prominent but not elevated
above the coxae, steeply but arcuately declivous in front. Legs with posterior
tarsi not longer than tibiae, first segment not or only slightly longer than remaining
segments combined. Elytra with fine, non-elevated costal lines. Abdomen with
last segment not excavated.
Type of genus: Megachoriolaus chemsaki Linsley
This genus is proposed for a species from Mazatlan with lycid-
like coloration. It also includes a number of others from southwestern
United States, Mexico, and Central America which have been previously
assigned to Euryptera by Linsley (1961) including: M. breviceps
(Linsley) n. comb., M. flammatus (Linsley) n. comb., M. ignitus
1 The writer expresses appreciation to the National Science Foundation through Grant GB-4944X
for support of this and other studies of North American Cerambycidae. Special acknowledgment
is also given to authorities of the California Academy of Sciences, San Francisco, the Canadian
National Collection, Ottawa; Cornell University, Ithaca, The University of Kansas, Lawrence, and
the United States National Museum, Washington, D.C., for the loan of critical specimens. John F.
Lawrence, Museum of Vertebrate Zoology, Harvard University, kindly examined the sternal characters
of one of the types of “Euryptera” in that collection, enabling me to place it generically. To
Celeste Green, I am deeply grateful for the illustrations.
The Pan-Pacific Entomologist 46: 128-141. April 1970
APRIL 1970 ] LINSLEY—NEW LEPTURINE CERAMBYCIDS
129
(Schaeffer) n. comb., M. sabinoensis (Knull) n. comb., M. spiniferus
(Linsley) n. comb., and M. texanus Knull n. comb.
Megachoriolaus chemsaki Linsley, new species
(Fig. 1)
Male. —Form somewhat oblong; elytra gradually but narrowly expanded
posteriorly before rounding to apices; integument yellowish-brown, antennae,
apical two-fifths of elytra, abdomen, except first sternite, most of tibiae and tarsi
black, head, with vertex at middle, genae, neck except medially above pronotum
and sterna at sides wholly or partially black or piceous. Head elongate, ratio
of distances between apices of antennal tubercles and line midway between them
and apex of labrum, 2.3:6.0; vertex finely, densely punctate, impressed on each
side of midline; antennal tubercles prominent, lateral areas below antennal
insertions finely, densely punctate; clypeus and labrum shining, sparsely punctate,
surface minutely tessellate; antennae extending beyond middle of elytra, first
four segments shining, clothed with coarse, suberect black setae, fifth with few
at apex, segments five to eleven dull, minutely, densely punctate and pubescent,
poriferous areas of outer segments small and inconspicuous, third segment sub-
cylindrical, but slightly thickened apically, about as long as scape, fourth segment
distinctly shorter than third, widened apically, fifth about as long as third, also
expanded apically, segments six to ten thickened but subparallel, eleventh dis¬
tinctly longer than tenth, at most very feebly appendiculate. Pronotum densely,
moderately coarsely punctate, punctures larger than those of vertex and elytra,
with vague or evanescent median impunctate line; pubescence short, appressed;
sides moderately rounded in front of middle, posterior angles extending out over
humeri, acute; prosternum finely, sparsely punctate and shining; mesosternum
finely, sparsely punctate medially and shining, densely punctate on lateral sclerites.
Legs with posterior tarsi about as long as tibia, first segment scarcely shorter than
remainder of tarsus. Elytra more or less oblong, ratio of humeral width to sub-
apical width, 6:7; surface more finely, discretely punctate than pronotum, discal
punctures mostly separated by one or two diameters, pubescence moderately short,
depressed; apices broadly transversely rounded to external angle which is dentiform.
Abdomen shining, finely, sparsely punctate medially, finely, densely punctate
laterally, last sternite sinuate-truncate at apex, lateral angles dentiform. Length,
11.5 mm.
Female. —Form very similar to that of male, pronotum and basal abdominal
sternites yellow. Length, 11 mm.
Holotype male and allotype female from 5 miles north of Mazatlan,
Sinaloa, Mexico, 27 July 1964, taken in copulo on flowers of Buddleia
wrightii Robins, by J. A. Chemsak, seven paratypes, same locality, 27-28
July 1964 on B. wrightii (J. A. Chemsak), fourteen, 24-29 July 1964 (H.
F. Howden), six bearing flower labels for Jatropha curcas L., and one
for Buddleia wrightii , and nine dated 5-11 August 1964 (H. F. Howden)
without flower data. One additional specimen, also designated paratype,
is from Venedio, Sinaloa, Mexico, 3 July 1918 (E. C. Van Dyke collec¬
tion). The holotype and allotype are deposited in the California
130
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[vol. 46 , no. 2
Fig. 1. Megachoriolaus chemsaki Linsley 9
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131
Academy of Sciences, San Francisco; paratypes in the Canadian Na¬
tional Collection, Ottawa, and the collections of the British Museum
(Natural History) London, the California Insect Survey, University of
California, Berkeley, and the California Academy of Sciences.
This species is smaller than most Megachoriolaus , ranging in size
from 9-12 mm, and is distinctive in form and coloration. The two sexes
lack obvious secondary sexual characters and are very difficult to dis¬
tinguish without dissection. The last abdominal segment usually extends
a little more conspicuously beyond the elytral apices in the female but
the last sternite is similar in both sexes. Among specimens believed
to be males, most, but not all, have the pronotum and sterna pre¬
dominantly black or piceous; conversely most females have the
pronotum predominantly yellow.
This species is named in honor of my colleague, John A. Chemsak,
with whom I have collected at the type locality but not during the season
when adults of this species were active.
Megachoriolaus imitatrix Linsley, new species
Female. —Form robust; elytra gradually but narrowly expanded from humeri
to middle then gradually narrowed posteriorly before rounding to apices, ratio of
width at middle to humeral width, 4.5:4.0; integument black, head, pronotum and
elytra red, prosternum, pro- and mesocoxae and intercoxal process of mesosternum
brown. Head elongate, ratio of distance between apices of antennal tubercles and
line midway between them and anterior margin of labrum, 2.8:6.0; vertex finely
punctate above, more coarsely below, pubescence very sparse, more evident
between upper lobes of eyes; area above antennal supports triangularly impressed
and impunctate and glabrous, impression deeper at sides, antennal supports finely
punctate, subglabrous, triangular area at base of clypeus shining, impunctate,
glabrous; clypeus with length at sides subequal to width, surface coarsely punctate
basally, impunctate apically; general length below eye greater than basal width
of mandibles; antennae extending to about middle of elytra, basal segments
subcylindrical, clothed with short, appressed, coarse black setae, longer at apices,
outer segments dullish, minutely densely punctate and pubescent, expanded but
not serrate, third segment subequal in length to scape, fourth segment distinctly
shorter than third and fifth, slightly thickened apically, fifth segment not quite
as long as third, expanded apically, six to ten thickened and successively slightly
shorter, poriferous areas not evident, eleventh segment a little longer than tenth,
apex attenuated but scarcely appendiculate. Pronotum with sides obtusely rounded
in front of middle, basal angles extending over humeri; disk coarsely sub-
contiguously punctate except for an ill-defined median longitudinal impunctate
line, pubescence short, coarse, recurved, not obscuring surface. Legs with posterior
tarsi distinctly shorter than tibiae, first segment slightly longer than remainder
of tarsus. Elytra about twice as long as broad; surface finely discretely punctate,
punctures mostly separated by one or two diameters, pubescence short, appressed,
not obscuring surface; apices transversely sinuate-truncate, outer angle dentiform.
132
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[VOL. 46 , NO. 2
Fig. 2. Lycomorphoides simulans Linsley $
APRIL 1970 ] LINSLEY-NEW LEPTURINE CERAMBYCIDS
133
Abdomen shining, sparsely punctate medially, densely punctate laterally, last
sternite truncate at apex, lateral angles prominently dentiform. Length, 11 mm.
Holotype female (University of Kansas), from El Salto, San Luis
Potosi, Mexico, 8 June 1961, 1900 ft. elevation (University of Kansas
Mexican Expedition).
This species superficially resembles M. flammatus (Linsley) from
Cuernavaca, but differs markedly in the form of the head which is
more elongate, differently shaped and subglabrous, rather than densely
clothed with long, golden, recumbent pubescence. In addition, the third
antennal segment is as long as the scape and longer than the fifth (it is
shorter than either in M. flammatus ) ; the pronotum is more coarsely,
and the elytra more distinctly, punctate. In general outline it is distinct
from any of the other known species, being more robust, with the
elytra widest at the middle.
Lycomorphoides Linsley, new genus
Form elongate, flat, lyciform, elytra broadly expanded posteriorly, apices
separately rounded. Head elongate, neck prominent, temples short, genal length
below eyes much greater than basal width of mandibles; antennae short, stout,
inserted adjacent to upper anterior margin of lower lobe of eye outside of
emargination, basal segments densely clothed with coarse erect setae, outer seg¬
ments dull, minutely punctate and pubescent, poriferous areas not evident.
Pronotum campanuliform, posterior angles extending over humeri; prostemum
with intercoxal process narrow, laminiform, anterior coxae exerted; mesosternum
with intercoxal process not prominent, much narrower than coxae, sunken between
them, gradually and more or less flatly declivous in front. Legs slender, tibiae
arcuate, posterior tarsi short, first segment elongate. Elytra elongate, costate.
Type of genus: Lycomorphoides simulans Linsley
This genus is proposed for an anomalous Lycus -like lepturine of
the Choriolaus-Strangalia group, with which it agrees in sternal char¬
acters but differs markedly in the form of the elytra which are broadly
expanded and separately rounded without any trace of an external
angle.
Lycomorphoides simulans Linsley, new species
(Fig. 2)
Female. —Integument dark brown, head, except for mouthparts and median
longitudinal stripe on vertex, pronotum except for broader median stripe which
does not reach base or apex, and elytra except for extreme apices, anterior and
intermediate coxae and intercoxal processes and basal portion of tibiae and tarsi
yellow brown; pubescence black on dark areas, golden on pale areas. Head with
neck shining, punctures much larger and less dense than those of vertex; vertex
prominent, opaque, finely densely punctate, densely clothed with appressed
golden pubescence, area above antennal insertions slightly impressed, ratio of
134
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
distance between apices of antennal tubercles and line midway between them
and anterior margin of labrum approximately 2:3; clypeus coarsely punctate
basally; antennae with only apical half extending beyond elytral humeri, scape
about one and one-half times as long as apical width, third segment a little
longer and stouter than scape, fourth segment less than half as long as third,
equally stout, fifth segment a little shorter than third, six to ten successively a little
shorter and less stout, eleventh segment scarcely longer than tenth, with pale conical
projection at apex. Pronotum opaque, finely densely punctate, densely clothed with
fine appressed golden pubescence; prosternum shining, sparsely punctate; meta¬
sternum densely punctate at sides, metepisterna punctate throughout. Elytra about
three and one-third times as long as basal width, subapical width more than twice
basal width; surface subopaque, with two costae extending from base nearly to apex
and two additional evident externally over most of expanded apical half; punctures
dense, larger than those of pronotum, interspaces minutely tessellate; pubescence
long, fine, golden, appressed not obscuring surface; apices narrowly rounded at
suture, broadly rounded externally. Abdomen shining; surface sparsely punctate,
thinly clothed with suberect pubescence; last sternite medially and subapically im¬
pressed, apex rotundate-truncate. Length: 13 mm, greatest width, 6 mm.
Holotype female (United States National Museum), from Turrialba,
Costa Rica, 800 m (Schild, in collection of F. Nevermann).
Macrochoriolaus Linsley, new genus
Form large, robust, subparallel; elytra gradually and very slightly narrowed
posteriorly before rounding to apices which are transversely truncate and feebly
emarginate. Head large, occiput excavated behind, vertex and postocular area
very prominent, wider than upper lobe of eye and continuous with genae, genae
also prominent giving the head proper a squarish outline; antennal tubercles very
prominent; eyes very large, emargination shallower than lateral width of eye,
antennae inserted adjacent to upper anterior margin of lower lobe of eye outside
of emargination, antennal sockets large, conspicuous; area at base of antennal
tubercles deeply transversely impressed; muzzle short, prognathous; clypeus trans¬
verse, twice as wide as long; labrum short, transverse; antennae short, basal seg¬
ments shining, discretely punctate and setose, outer segments minutely, densely
punctate and pubescent, poriferous areas not evident. Pronotum transverse but less
than twice as wide as long, sides obtusely rounded in front of middle, arcuately nar¬
rowed in front of posterior angles, posterior angles produced but do not extend over
humeri, basal width twice apical width but only seven-tenths as wide as elytra across
humeri; prosternum with intercoxal process narrow, laminiform, coxae prominent,
mesosternum with intercoxal process narrow, arcuately declivous in front. Legs
relatively stout, tibiae not arcuate, tarsi short, first segment of posterior pair
much shorter than remainder of tarsus.
Type of genus: Macrochoriolaus elegans Linsley
The remarkable structure of the head immediately sets this genus
apart. Although somewhat suggestive of Megachoriolaus in the form
of the body, the narrow pronotum and heavy appendages will further
distinguish it from that genus.
APRIL 1970 ] LINSLEY-NEW LEPTURINE CERAMBYCIDS
135
Macrochoriolaus elegans Linsley, new species
(Fig. 3)
Female. —Body black, shining, pronotum and elytra brownish-yellow. Head
with vertex and postocular area finely punctate with scattered larger punctures
superimposed, sparsely clothed with erect black hairs, median line on vertex
impressed, impression above antennal supports somewhat V-shaped, punctures
larger and denser than those on lobes of vertex, antennal supports distinctly
punctate, punctures moderately large, mostly well separated, about the size of
those on antennal scape; clypeus with narrow band of coarse punctures at base,
apical two-thirds glabrous, impunctate; labrum glabrous, shining; genae shining
and almost impunctate at sides, irregularly punctate below eyes; antennae with
segments subcylindrical, segments three and four gradually enlarged toward
apices. Pronotum with disk coarsely, closely punctate basally, more finely and
less discretely toward apex, transversely impressed about one-third distance between
basal lobe and apex; pubescence short, erect, golden, not obscuring surface;
scutellum black, apex rounded. Elytra more finely, shallowly punctate than
pronotum, pubescence short, nearly erect, not obscuring surface. Legs densely
punctate, clothed with short suberect hairs, posterior tarsus with last segment
elongate, distinctly longer than first segment. Abdomen shining, sternites more
densely punctate and pubescent at sides, last sternite rotundate-truncate, feebly
emarginate, apex pubescent. Length, 15.5 mm, breadth 5 mm.
Holotype female (Cornell University) from THE VICINITY OF
Olmicteme, Guerrero, Mexico, 29-30 July 1965, in a humid pine-
oak formation at an elevation of 2300 to 2500 meters (Cornell University
Mexico Field Party).
Platerosida Linsley, new genus
Form elongate, slender, flattened. Head relatively short, deeply impressed above
antennal tubercles which are fused and shallowly longitudinally impressed at
apex, triangular area above clypeus deeply impressed, transverse impression at
base of clypeus also deep so that muzzle prognathous; antennae inserted near
upper anterior margin of lower lobe of eye outside of emargination, those of male
extending beyond middle of elytra, basal segments shining, setose outer segments
dull, poriferous areas not evident, scape robust, second segment bead-like, about
as long as broad, third segment short, about three times as long as second, clavi-
form, fourth segment very short, bead-like, only about twice as long as second,
fifth segment about as long as three preceding segments taken together, gradually
widened externally to apex, sixth segment distinctly longer than fifth, similarly
shaped but more slender, seventh segment subequal in length to sixth, segments
eight to ten successively slightly shorter and more slender, eleventh segment a
little longer than tenth, feebly constricted before apex; antennae of female
reaching to about basal one-fifth or one-sixth of elytra, first four segments pro¬
portioned much as in the male but with segments five to eight broadly triangular,
their apices nearly as wide as their length, segments nine and ten also broad but
a little more parallel-sided, eleventh segment not longer than tenth, apex obtusely
pointed. Pronotum broadly constricted at apex, sides obtusely subangulate at
middle, base with lateral angles produced but not extending over elytral humeri;
136
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 2
Fig. 3. Macrochoriolaus elegans Linsley 2
APRIL 1970] LINSLEY—NEW LEPTURINE CERAMBYCIDS
137
prosternum with intercoxal process narrow, laminiform, coxae exerted, ap¬
proximate; mesosternum with intercoxal process much narrower than coxae, sub¬
merged between them, arcuately declivous in front. Legs long and slender in male
with posterior tarsi longer than tibiae, shorter in female with the posterior tarsi
not as long as tibiae. Elytra subparallel, costate, apices separately rounded.
Abdomen with last sternite broadly excavated in male, narrowly so in female.
Type of genus: Platerosida howdeni Linsley
The species upon which this genus is based bears a remarkable super¬
ficial resemblance to 44 Euryptera ” huachucae Schaeffer from southern
Arizona and chihuahuae Bates from the western Chihuahua (prob¬
ably all three have the same or similar small dark lycid models). How¬
ever, the highly modified antennae, particularly in the female, readily
distinguish it from these and other members of the Choriolaus-Strangalia
complex.
Platerosida howdeni Linsley, new species
(Fig. 4)
Male. —Body black, opaque. Head with neck finely, densely punctate, punctures
larger at sides, pubescence sparse, suberect, golden; vertex minutely, densely
punctate and pubescent, with few scattered long erect hairs; eyes fringed with
long, suberect golden pubescence; genae below eyes finely densely punctate and
pubescent; antennal supports densely punctate, triangular area above clypeus
shining, glabrous; labrum finely punctate; antennae with first four segments
finely, discretely punctate and clothed with coarse suberect black setae, segments
five to eleven minutely densely punctate, densely clothed with minute, erect
pubescence. Pronotum with anterior constriction more or less rugosely punctate,
disk finely and very densely punctate, punctures larger on median basal lobe;
entire surface densely clothed with moderately long erect black pubescence with
few scattered very long hairs at sides; pro- and mesosternum finely densely punctate,
metepisterna punctate throughout. Elytra about three times as long as basal
width; surface very densely punctate, punctures slightly larger than those of
pronotum, interspaces minutely tessellate; pubescence depressed but not recumbent;
apices obliquely rounded. Abdomen with sternites densely punctate, thinly clothed
with rather long, suberect hairs; last sternite coarsely punctured at base and sides
of excavation which is elevated at the sides, deeply concave, shining and involving
most of apex of segment, lateral angles prominent, produced. Length, 11.5 mm,
breadth 3 mm.
Female: Form slightly broader; abdomen with last sternite shallowly, tri¬
angularly impressed, apex subtruncate, lateral angles not prominent. Length 12
mm, breadth 3.5 mm.
Holotype male from 10 miles west of El Salto, Durango, Mexico,
11 July 1964 (H. F. Howden), allotype female from P. Buenos Aires,
37 miles west of El Salto, Durango, Mexico, 1 July 1964 (H. F. Howden)
and one male paratype from 10 miles west of El Salto, Durango, Mexico,
9000 ft. (W. R. Mason) and a female paratype from 3 miles west of
El Salto, Durango, Mexico, 9000 ft. (J. A. Powell). The holotype and
138
THE PAN-PACIFIC ENTOMOLOGIST
[vol. 46 , no. 2
Fig. 4. Platerosida howdeni Linsley 2
APRIL 1970 ] LINSLEY—NEW LEPTURINE CERAMBYCIDS
139
allotype are deposited in the Canadian National Collection, the two
paratypes in the collection of the California Insect Survey, University
of California, Berkeley.
Choriolaus sulcipennis Linsley, new species
Male. —Form elongate; elytra gradually expanded posteriorly before rounding
to apices; integument black, head in part, pronotum at sides, and basal one-half
of elytra reddish-orange, vertex brownish; pubescence golden on reddish-orange
areas, otherwise black. Head with vertex convex, with median longitudinal line
extending toward base of clypeus, surface brownish, very finely densely punctate,
clothed above with fine, short, erect golden pubescence; face densely punctate,
antennal tubercles not prominent, antennal insertions not, or scarcely separated
from upper anterior margin of inner lobe of eye; antennae reaching to middle
of elytra, first four segments clothed with coarse suberect setae, fifth less so,
outer segments minutely densely punctate and pubescent, with large, conspicuous
apical poriferous area, scape subcylindrical, third segment subcylindrical but
slightly thickened toward apex, a little shorter than scape, fourth segment shorter
than third, fifth subequal to third, segments six to ten expanded externally, sub-
serrate, successively shorter, eleventh segment distinctly longer than tenth, ap-
pendiculate, appendix conical, pale. Pronotum scarcely rounded at middle; surface
opaque, finely, densely punctate but punctures much larger than those of vertex;
pubescence fine, appressed; median dark area basally at least twice as wide as
scutellum, equally wide at apex. Legs with posterior tarsi about as long as
tibiae. Elytra about three and one-third times as long as humeral width; each
longitudinally shallowly bisulcate, the sulcae separated by an obtuse, costa-like
ridge; surface finely, densely rugoso-punctate, punctures much larger than those
of pronotum, more or less transversely confluent; pubescence fine, suberect, not
obscuring surface; apices rounded to external angle which is distinct but not
acute. Abdomen shining, very sparsely punctate and pubescent, apex truncate.
Length, 10 mm.
Female.— Form a little shorter and broader than male; antennae surpassing
basal one-third of elytra but not attaining middle, outer segments thickened but
not apically expanded, poriferous areas not evident. Length, 8 mm.
Holotype male , allotype female and five paratypes (California
Academy of Sciences) from Cordoba, Vera Cruz, Mexico (A. Fenyes).
An additional paratype from the same series is deposited in the collection
of the California Insect Survey, University of California, Berkeley.
This species bears a very strong superficial resemblance to
“ Euryptera ’ mimula Bates which also occurs at Cordoba. However, it
differs at once in the elongate, apically expanded, longitudinally sulcate
elytra as well as in the apically expanded outer antennal segments of the
male which bear large, conspicuous poriferous areas.
Choriolaus aurifer Linsley, new species
Female. —Form moderately stout; elytra subparrallel, not expanded posteriorly
before rounding to apices; integument golden yellow, vertex, eyes, antennae,
140
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
longitudinal stripe on pronotum, scutellum, apices of femora, tibiae except at base,
and tarsi in part, black or brownish. Head with vertex convex, with median
longitudinal line extending to base of clypeus, surface brownish, very finely
densely punctate, thinly clothed with fine appressed hairs, short and suberect
above; face shining, antennal tubercles finely punctate, sparsely pubescent, frons
with transverse impression at base of clypeus moderate, clypeus and labrum
shining, moderately coarsely punctate; antennal insertion distinctly hut narrowly
separated from upper anterior margin of lower lobe of eye; antennae rather
stout, extending a little beyond middle of elytra, first four segments with coarse
suberect black setae, fifth with fewer, outer segments minutely densely punctate
and pubescent, without poriferous areas; scape subcylindrical, third segment
slightly enlarged toward apex, not shorter than scape, fourth segment shorter
and thicker than third, fifth longer and thicker than fourth but shorter than third,
segments six to ten stout but not expanded at apices, successively gradually de¬
creasing in length and diameter, eleventh segment a little longer than tenth, ap-
pendiculate, appendage conical, pale. Pronotum moderately rounded at middle;
surface finely, very densely punctate but not opaque, punctures larger than those
of vertex, more or less transversely confluent, pubescence appressed, not dense
but partially obscuring punctation, median stripe narrow, at base about one-half
as wide as scutellum, at apex extending only to anterior constriction. Legs with
posterior tarsi about as long as tibiae. Elytra less than two and one-half times
as long as humeral width; surface densely, granulate-punctate, punctures very
much larger than those of pronotum; pubescence fine, moderately short, de¬
pressed but not recumbent; apices rotundate-truncate, external angle acute.
Abdomen shining, finely punctate and pubescent; last segment rotundate truncate,
disk coarsely punctate, impressed before apical margin. Length 8.5 mm.
Holotype female (Department of Entomology, Cornell University),
from El Limon, Tamaulipas, Mexico, 9 June 1951 (H. E. Evans).
This species was taken at the same time and place as examples of
“ Euryptera ” mimula Bates, and, in spite of the different facies and
coloration, was at first assumed to represent a color phase of that species.
However, in addition to the golden coloration, it differs in the shorter,
stouter form and structure of the antennae, as well as details of puncta¬
tion and pubescence. The posterior tarsi are unusually long for a female
of this genus.
Choriolaus similis Linsley, new species
Female.— Form moderately narrow; elytra subparallel, not expanded posteriorly
before rounding to apices; integument golden brown, eyes, antennae, a longitudinal
stripe on pronotum, apices of femora and apices of tibiae black. Head with
vertex convex, with median longitudinal line extending to base of clypeus, surface
very finely, densely punctate, clothed with fine, short, appressed pubescence; face
shining, antennal tubercles finely punctate, sparsely pubescent; frons with transverse
impression at base of clypeus moderate; clypeus and labrum shining, moderately
coarsely punctate; antennal insertions distinctly but narrowly separated from
upper anterior margin of lower lobe of eye; antennae moderately slender, surpassing
APRIL 1970 ] LINSLEY—NEW LEPTURINE CERAMBYCIDS
141
basal one-third of elytra but not attaining middle, first four segments shining,
with numerous coarse, suberect black setae, fifth with fewer, segments six to ten
opaque, minutely densely punctate and pubescent, with distinct apical poriferous
areas, scape feebly claviform, third segment barely shorter than scape, slightly
enlarged apically and emarginate externally, so as to expose attachment of fourth
segment, fourth segment shorter than third, apex wider than that of third, similarly
emarginate externally, fifth a little longer than fourth, shorter than third, sixth
with apex wider than that of fifth, seventh to tenth successively shorter and
slightly narrower, parallel-sided, eleventh segment appendiculate, appendage
conical, pale. Pronotum obtusely rounded at middle; surface finely densely, some¬
what granulate-punctate, punctures larger than those of vertex, pubescence ap-
pressed, not dense but partially obscuring punctation; median stripe narrow, not
wider than scutellum. Legs with posterior tarsi slightly shorter than tibiae. Elytra
twice as long as humeral width; surface densely punctate, punctures very much
larger than those of pronotum, more or less transversely confluent, pubescence
fine, suberect basally, otherwise depressed; apices rounded to external angle which
is distinct but not acute. Abdomen shining, finely, sparsely punctate and pubescent;
last segment truncate, moderately coarsely but not densely punctate, shallowly
impressed medially at apex. Length, 10.5 mm.
Holotype female (California Academy of Sciences) and one para-
type female from Temescal, Oaxaca, Mexico, 6 July 1965 (G. H.
Nelson and family).
This species bears a strong superficial resemblance to C. aurifer
Linsley in aspect and coloration. However, it is less stout, the ground
color is golden brown rather than golden yellow, the outer antennal
segments have distinct apical poriferous areas, and the external angle
of the elytra is not acute. The female paratype differs from the holotype
by having the median pronotal band wider and extending from base
to apex (in the type it does not attain either), the scutellum black,
thoracic sterna, abdominal sternites brownish black, except anterior
femora, intermediate and posterior tarsi, metasternum at middle and
the two subapical abdominal sternites, which are yellowish. None of
these differences materially affect the superficial aspect of this in¬
dividual as a presumed lycid mimic.
Literature Cited
Bates, H. W. 1879-1885. Biologia Centrali-Americana, Coleoptera, 5:1-436, pis.
1-25.
Chemsak, J. A. 1969. New Mexican and Central American species of Strangalia
Audinet-Serville (Coleoptera: Cerambycidae). Jour. New York Entomol.
Soc., 77:2-9, figs.
Linsley, E. G. 1961. The North and Central American species of Euryptera and
a related new genus (Coleoptera, Cerambycidae). Jour. New York
Entomol. Soc., 69:131-141.
142
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
Biology and Structure of the Dobsonfly, Neohermes
californicus (Walker)
(Megaloptera: Corydalidae)
Edward Laidlaw Smith
Chico State College, California
During a study on the evolution of external insect genitalia and other
appendages (Smith, 1969), 2 species of Corydalidae were reared or
collected as adults for dissections. They were Neohermes californicus
(Walker) and Protochauliodes infuscatus Caudell (= N. nigrinus Van
Dyke, teste Chandler 1956). This paper reports observations on the
behavior, rearing and external structure of these peculiar insects. Their
genitalic anatomy will be covered in a separate article.
Relatively little has appeared on California Megaloptera. One of the
most recent papers is on the biology of 2 species of western Sialidae by
Azam and Anderson (1969). Among the most comprehensive works on
corydalids are 2 master’s theses discovered after completing my initial
investigation: Maddux (1952) and Penland (1953). I have drawn
upon them freely to flesh out the life histories.
Rearing Techniques and General Biology
Larvae of Neohermes californicus (Fig. 1), mostly in the final 3
instars, were collected by hand in shallow, stony creeks in the Chico
(Butte County) area of California in April 1969. The small streams
were drying up rapidly, and hellgrammites were most abundant at the
ends of the creeks where the water sank into the riverbed and dis¬
appeared underground. The larvae were usually in shallow excavations
under stones, but some were away hunting prey on the bottom and
others were out of the water and burrowing into the damp volcanic
sand of the banks to pupate. Collections of larvae were reared easily
and continuously en masse in a plastic breadbox 32 cm L X 26 cm
W X 11 cm D. The container was filled to a depth of 6 cm, the bottom
littered with mud and stones, and open areas filled with aquatic plants.
Aeration was provided by an aquarium air pump. Another breadbox
of identical size was inverted over the water-filled one to maintain
humidity and to prevent the restless insects from crawling out.
Azam and Anderson (1969) noted frequent cannibalism among
larvae of Sialis rotunda Banks and S. californica Banks. Maddux (1952)
and Penland (1953) reported similar behavior by the corydalids
Neohermes californicus and Protochauliodes spp. yet I found no fighting
The Pan-Pacific Entomologist 46: 142-150. April 1970
Fig. 1 . Female last instar larva of Neohermes californica (Walker) (Megaloptera: Corydalidae) exposing spiracles (SP) of
segments 7, 8 above the water (somites numbered in circles). Note alternating position of styli, and the gripping action of the claws
on the cereal vesicles.
APRIL 1970] SMITH—NEOHERMES BIOLOGY AND STRUCTURE 143
144
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
and but a single instance of cannibalism out of nearly 100 N.
californicus, and this one took place in the crowded collection jar at
the time of capture. There was no fighting or cannibalism among
specimens in the laboratory, although more than 30 larvae of different
sizes were kept together in less than a liter of water. A very pronounced
avoidance reaction was always displayed between larvae, regardless of
disparate size. This behavior, coupled with an excess of both refuges
and food, was probably responsible for keeping the peace presumably
normal in the natural environment.
Food consisted of mayfly and caddisfly larvae, small tadpoles, a few
small sialids accidentally collected with the pollywogs, and chunks of
raw beef about 1 cm in diameter. The hellgrammites not only readily
accepted insects (alive or freshly killed) and meat offered them in
forceps, but would actively hunt down such material when it was
dropped into the tank. Food was seized with both jaws and front legs,
then macerated sufficiently to be swallowed whole. This engulfing was
observed in sialids by Azam and Anderson. It was very rapid in N.
californicus, and meat of the size described was devoured in less than
5 min. by mature larvae.
In the middle of the plastic tank, a stainless steel pan 13 cm in
diameter and 7 cm deep was filled to the top with a mixture of damp
gravel, sand, clay and small stones. Mature larvae readily climbed
up over rocks heaped around the pan, and excavated rapidly into the
soil to build pupal chambers at varying depths. Maddux (1952) found
most of his pupae under stones along the creek banks, but no larva
reared in this study made a chamber beneath a rock. All hollowed
out a small cavity, usually at the bottom of the pan. The container was
emptied daily and larvae removed to 5-cm petri dishes containing some
of the pupation medium. The petri dishes were labeled and kept in a
darkened (but not light-tight) box. The soil was kept barely moist.
Out of 29 larvae allowed to pupate, an average of 13 days (range
7-20) elapsed from the time they entered the soil until pupal ecdysis.
About 5 days before shedding, a larva would assume the characteristic
comma-shaped prepupal resting position on its side, and take on an
inflated, dropsical appearance. The styli shriveled. Towards the end of
this period, the pupa could be seen beneath the prepupal integument.
While usually quiescent, prepupae suddenly exposed to bright light
would engage in violent activity, and often would vomit an extremely
foul-smelling, dark greenish liquid. This defense is employed also by
larvae when disturbed. The actual molting of the prepupa is very
rapid (about 5 min.). The pupa emerged through the median dorsal
APRIL 1970 ] SMITH—NEOHERMES BIOLOGY AND STRUCTURE
145
slit in the thorax and head, assisted by thrusts of the anal legs (cerci)
and wrenching telescopic motions of the abdomen. In 2 instances, the
head capsule was incompletely cast, and the insects subsequently died
when the adults attempted to emerge. Several females developed an
intersegmental ventral abdominal hernia as they emerged from the
prepupal integument. These abnormalities may have been rearing
artifacts.
Freshly molted pupae were a light cream color, and quite active
when disturbed. Not only were they capable of biting, but could struggle
about using the cerci to push and steady themselves. About 9 days after
emergence from the prepupa, pupae began to darken. The first rearings
died in the closed petri dishes at this point, apparently from excessive
humidity. Subsequent pupae were allowed to complete development in
open petri dishes once the eyes had begun to darken. The soil upon
which they rested was moistened slightly every few days. Regardless
of treatment, the majority of males in all rearings died from an unde¬
termined malady. The affliction was always heralded about 4 days
before death by all wingpads becoming grotesquely distended with
hemolymph, and taking on a roughly tubular appearance. No females
were so affected.
Of 11 surviving specimens not dissected prior to adult emergence,
the pupal period lasted for an average of 11 days (range 9-12) at about
25° C. There was no significant difference between the sexes. Maddux
(1952) also found the average pupal period for 15 specimens of N.
californicus and Protochauliodes aridus Maddux to be 11 days. Penland
(1953) obtained the same data with rearings of P. aridus. Maddux
managed to prolong pupation up to 40 days by placing specimens col¬
lected as pupae in a refrigerator at 4° C. Azam and Anderson reported
pupal stages of but 4-6 days for their sialids.
Just prior to adult emergence, the pupa would crawl out of the open
petri dish and wander about. The method of ecdysis was the same as
that exhibited by the prepupa, and equally as rapid. One of the first
acts of the still teneral adult was to void white, ill-smelling meconium.
Adult corydalids of indeterminate age caught in nature voided
meconium when handled, and so did reared specimens up to 3 days
after emergence. No adults were kept alive for longer than that, but
Maddux (1952) maintained unfed adults 7-10 days in the laboratory.
He obtained oviposition in 5 instances, all but 1 depositing their eggs
on the exposed surfaces of flat rocks. He found a similar egg mass in
the field, and described one from the laboratory as “roughly rectangular
in outline, approximately 500 mm at its greatest length and 250 mm at
146
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 2
its greatest width; it consisted of more than 3,000 white eggs arranged in
a series of parallel rows, with the micropylar ends all facing the same
direction. The mass was without any protective coating. Individual eggs
were approximately 1 mm in length and 3X longer than wide. . . . the
micropylar projection was similar to that seen in the eggs of various
sialids.” This description was for P. aridus , and closely approximates
the pattern of Sialis californica shown by Azam and Anderson in their
figure 2B. The naked condition of the eggs is interesting. The adults
of at least N. californicus and P. aridus emerge only after the small
creeks they inhabited as larvae have dried up. Maddux (1954) noted
that the adult females of at least the latter species deposited their eggs
on stones in the dry creek bed.
Last instar larvae removed from the water before they had voluntarily
come out to pupate would go into an inactive state. If returned to the
tank within 2 weeks, they developed normally. After that time, however,
they shriveled and eventually died. Given the intermittent nature of
many of the creeks various California corydalids inhabit, I suspect
larvae may bore into the mud when the stream temporarily dries up.
Estivation of newly hatched hellgrammites is the rule with P. aridus ,
and probably with N. calif ornicus. Maddux (1954) reported that the
larvae after eclosion immediately burrowed down into the gravel of the
dry stream bed to await the winter rainy season. It was his fellow
worker Penland (1953) who apparently worked out the details. Penland
found abundant egg masses of corydalids deposited during late May
and early June upon the downstream faces of relatively large, flat stones
in well-sheltered areas of dry creek channels in the foothills around
Chico. The eggs hatched in 7-10 days, and the larvae dropped down
onto the soil and excavated 4-12 inches into it. They made small cells
beneath stones, and remained there until the streams flowed once more
(usually by November in the Chico area). When he removed larvae to
a completely dry environment, they died in 3-4 days, so the estivation
site must retain a minimal humidity even during the most scorching
summer days (ground temperatures in the vicinity of Chico often
surpass 55° C). The larvae emerged from their cells about a week after
water reappeared. The only species mentioned in this context was P.
aridus. Estivation is apparently not obligatory, since hatchlings will
accept food. Penland reared P. aridus in a novel way: he kept them
individually in damp soil (rather than in water), and fed them termites.
I suspect N. calif ornicus has a similar life cycle: adults emerged only
after the streams had dried, and there were only mid- to late-instar larvae
present in early spring of the following year, which larvae reached
APRIL 1970 ] SMITH—NEOHERMES BIOLOGY AND STRUCTURE
147
adulthood by June. Indeed, this life history pattern seems likely for
most or all California Corydalidae. Whether hellgrammites in late
instars can rest for extended periods in an emergency (e.g., premature
drying out of their creeks in spring, requiring estivation) as do
hatchlings (and thus take 2 years to develop) is not known, but does
seem unlikely.
Adults of N. californicus and Protochauliodes infuscatus were netted
a few miles from where larvae were collected. The appearance of adults
in nature corresponded with the emergence of reared material. What
may have been a mating swarm of P. infuscatus was watched from
about 1330-1430 (PST), 23 May 1969. Over 15 males were flitting
about some distance apart 0.5-3 m above the grass. The area was a hill¬
side clearing about 100 m square, immediately downwind to a rivulet,
and there was an almost continuous overstory of valley oak ( Quercus
lobata Nee). Only 1 female was discovered in the vicinity. The insects
vanished as abruptly as they had appeared. Similar flights, but com¬
posed of but 2 or 3 individuals (all male), were noted in adjacent areas
of similar ecology. This diurnal activity of adult P. infuscatus also has
been noted by Chandler (1956).
Use and Structure of Abdominal Styli and Terminal Appendages
Unlike Corydalis cornutus (L.), larvae of the much smaller far-
western corydalids normally lack the vesicles borne ventrally on
abdominal coxites I—VI. These permanently everted sacs, the apparent
coxal endites (the lateral styli on abdominal segments I-VIII are the
telopodites, or legs themselves), have cuticular tufts in C. cornutus,
and serve a respiratory function. On the flank of one female N.
californicus pupa, however, I found a permanently everted vesicle (VS,
Fig. 3) having not only the appearance, but the primitive laterodistal
orientation of both the gonapophyses (endites of the genital segments:
VIII, IX) and eversible vesicles of Apterygota (Fig. 2). The slightly
recurved appendage was filled with fat body.
The lateral abdominal styli of Megaloptera have very thin cuticle,
and contain a large median tracheal trunk (Fig. 4, TR), which gives
rise to numerous smaller branches in no obvious sequence among
Corydalidae. The styli are more or less rigid, their shape being main¬
tained by blood pressure. While gas exchange in larvae undoubtedly
takes place across the entire integument, the principal sites of transfer
are the styli. The retention of vesicles and the still further increase in
exchange surface by hollow cuticular extensions from them in Corydalis
cornutus but not in other corydalids is probably related to the much
148
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
Figs. 2-4. Figs. 2, 3. Comparison of the abdominal coxite, stylus and vesicle of an
adult machilid (2) with the apparent homologs of a female Neohermes californica
pupa (3). The permanently everted vesicle (endite?) found on segment IV of this
particular Megaloptera specimen is atavistic in this genus, and not normally re¬
tained. The 1 or 2 vesicles of the microcoryphian coxite are everted by blood
pressure and retracted by coxal muscles, and employed to sponge up water from
damp surfaces. Fig. 4. Terminal segments (numbered) of moulting male
Neohermes californica prepupa; the pupa within is outlined. The left cereal coxite
bears a ventral vesicle armed with paired claws, and a dorsal, segmented (?)
stylus. At this stage, the pupal appendages have pulled away from the prepupal
integument, and the cereal claws have retracted. Abbreviations.— AN, anus;
CE-CX, cereal coxite; Cl, claw; CX, subcoxal-coxal plate (s. lat: = pleuron) ;
P-CE, pupal cercus; S, sternite; SP, spiracle; ST, stylus (= telopodite) ; T,
tergite; TR, trachea; VS, vesicle (= endite?). The scale in Figs. 3 and 4 is
the same.
APRIL 1970 ] SMITH—NEOHERMES BIOLOGY AND STRUCTURE
149
larger size of the former, and resultant smaller ratio of surface area to
body volume. Larval styli also have a sensory function. As the hell-
grammite creeps about, the pairs of one segment are held at an angle
over the body, while those of the next set are angled below it, and so
on alternately (Fig. 1). The styli also help to orient the insect in
crevices.
While there are no styli on segment IX (here replaced by the gonostyli,
which appear externally only in the imago), the invaginated composite
segment X-XI bears a pair of 2-part appendages. Ventrally, each
terminates in a vesicle armed with paired retractile claws (Fig. 4, CL).
A pseudosegmented stylus (ST) homologous with styli on preceding
abdominal segments recurves about each vesicle from the cereal coxa
(CX-CE). The mobile, clawed vesicles thrust the larva forward much
as do the homologous pair (“pygopods”) of Mecoptera, Hymenoptera
and Lepidoptera caterpillars. The appendages also serve to anchor and
maneuver the larva in the current as do the corresponding anal hooks
of Trichoptera immatures. These terminal larval appendages in the
instances cited above give rise to the pupal and adult cerci (Fig. 4,
P-CE). Both pupal and adult corydalids use the cerci to assist in moving
as do the larvae, and employ the same vigorous telescopic contractions
of the abdomen. Cereal mobility in the pupa and adult, however, is
restricted largely to adduction.
The abdominal styli which precede the cereal styli in the larva are
retained by the pupa as small, conical, musculated coxal rudiments
(Fig. 3, ST). They are capable of slight movements. The homologous
coxal plate of the ephemerid subimago (which marks the site of the
nymphal abdominal styli—also respiratory in function) is capable of
similar twitching. The precercal styli of the adult corydalid are mere
dorsolateral nubbins, although recognizable as limb bases.
While the abdominal styli serve as the primary respiratory devices
in young Megaloptera larvae, the most posterior abdominal spiracles
become functional in later instars of corydalids. The spiracles are
dorsolateral, just above and mesal to either stylus, and present on
abdominal segments I—VIII (and also on the meso- and metathorax).
Under the stimulus of low oxygen tension in laboratory rearings, mature
larvae back up to the surface and expose at least the most caudal set
of abdominal spiracles (Fig. 1). According to Chandler, the still—
stagnant water-inhabiting corydalid, Nigronia , bears the last pair of
spiracles on short, individual extensions. The similar tubes of Chauliodes
actually extend caudally beyond the body. These behavioral and
structural modifications probably represent the respiratory patterns
150
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 2
antecedent to the evolution of the complex terminal breathing tube (de¬
rived from the caudal segments and a posterior migration of the 8th
set of abdominal spiracles) so characteristic of aquatic larvae in the
phylogenetically more advanced Coleoptera and Diptera.
Literature Cited
Azam, K. M., and N. H. Anderson. 1969. Life history and habits of Sialis
rotunda and S. californica in western Oregon. Ann. Entomol. Soc.
Amer., 62: 549-558.
Chandler, H. P. 1956. Megaloptera—Corydalidae, p. 231-33. In R. L. Usinger
[ed.], Aquatic Insects of California. Univ. Calif. Press, Berkeley.
Maddux, D. C. 1952. A study of the dobsonflies (Megaloptera) of the Chico,
California area. Master’s thesis, Chico State College, Chico, Calif., 44 p.
1954. A new species of dobsonfly from California. Pan-Pac. Entomol.,
30: 70-71.
Penland, D. R. 1953. A detailed study of the life cycle and respiratory system
of a new species of western dobsonfly, Neohermes aridus. Master’s
thesis, Chico State College, Chico, Calif., 34 p.
Smith, E. L. 1969. Evolutionary morphology of the insect genitalia. 1. Origins
and relationships to other appendages. Ann. Entomol. Soc. Amer.,
62: 1051-1078.
NEW BIOLOGICAL CONTROL ORGANIZATION
This is a brief report on a meeting convened by IUBS at Amsterdam, November
17-19, 1969 to finalize plans for the proposed new International Organization
for Biological Control (IOBC).
Plans for a new global organization in biological control, building from the
existing “Organization Internationale de Lutte Biologica” (OILB), under the
title given above (IOBC), were consummated at Amsterdam in November, as far
as could be done at the time. New statutes were proposed and approved by
delegates, and other agreements reached. It is hoped that the widest possible
interest and support can be made evident before February, 1971, at which time
approval by the current OILB’s General Assembly will presumably occur.
The following slate of candidates for the Executive Committee of the proposed
new Council of IOBC was recommended by unanimous vote of the delegates in
Amsterdam:
Paul DeBach, President
E. Biliotti, Vice President
Frank Wilson, Vice President
Vittorio Delucchi, Secretary General
Fred J. Simmonds, Treasurer
A fuller statement of the meeting in Amsterdam and of the aims and functions
of this new organization, and the progress of developments in other respects is
being published in The Bulletin of Entomological Society of America, to which
interested parties are referred. Such interested parties should contact any of the
above slate of candidates, or for the United States, Dr. Reece Sailer.—C. B.
Huffaker, University of California, Albany.
APRIL 1970 ]
SCIENTIFIC NOTE
151
SCIENTIFIC NOTE
Examples of convergence between blister beetles of California and oth¬
ers from Chile (Coleoptera: Meloidae). —Collections of meloid beetles from
desert areas of western Chile have been made available by their collectors (E. I.
Schlinger, M. E. Irwin, and L. A. Stange). Their close morphological similarity
to Californian genera and even individual species is remarkable, and it is hoped
that further knowledge of the adult and larval morphology and foods will permit
a more satisfactory determination of their phylogenetic affinities.
As suggested in Selander’s (1960, Ill. Biol. Monogr., No. 28) monograph of the
genus Lytta, the South American species (currently assigned to that genus) may
be new genera and may only be named, as at present, due to convergence rather
than close relationship. Certainly, all of them fall within the current definition
of the genus Lytta. One (new and undescribed wingless species) might be com¬
parable to the wingless genus Poreospasta in North America which was placed
in the genus Lytta by MacSwain (1956, Univ. Calif. Publ. Entomol., 12). In the
same paper, Poreospasta polita Horn [=z Lytta ( Poreospasta ) sublaevis (Horn)]
was placed near L. auriculata Horn on the basis of comparative larval morphology.
Edwards (1951, Pan-Pac. Entomol., 27:44) in a short note recorded one pair of
these two species in copulation.
Other genera and species (undescribed, probably) are even more remarkable
in that they superficially resemble members of the North American Tribe
Eupomphini [= Calospastini according to MacSwain (1956)] but when viewed
under the highest magnification are clearly members of the Lyttini. One looks
amazingly like our Cysteodemus armatus LeConte (or like Gynaecomeloe females) ;
while another looks like many of our species of Eupompha [rr Calospasta accord¬
ing to MacSwain (1956)]. Yet the first mentioned has the short tarsal spine free
at the base and the other a long tarsal spine. These species represent additional
examples of extreme convergence in this family as previously shown for Hornia
in North America and Allendesalazaria from North Africa by Linsley (1942, Univ.
Calif. Publ. Entomol., 7: 169-188) and confirmed by further data revealed by
MacSwain (1956).
In all of the examples referred to above, the structures may eventually be cor¬
related with very similar habits and environments.—J. W. MacSwain, University
of California, Berkeley.
SOCIETY NOTICE
Special Sale of Volumes 1-16, Pan-Pacific Entomologist. —As a special offer
to members only of the Pacific Coast Entomological Society, volumes 1-16 of the
Pan-Pacific Entomologist are currently offered at the extremely reduced rate of
$1.50/volume or $24 for the entire first 16 volumes. These are on a first come,
first served basis as long as the supply lasts. Contact the Treasurer, P. H. Arnaud,
Jr., California Academy of Sciences, Golden Gate Park, San Francisco, California
94118.
152
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. DF, NO. 2
BOOK REVIEW
CURCULIONIDAE SUBFAMILY ApIONINAE OF NORTH AND CENTRAL AMERICA WITH
Reviews of the World Genera of Apioninae and World Subgenera of Apion
Herbst (Coleoptera) . By David G. Kissinger. Taxonomic Publications, South
Lancaster, Massachusetts. 559 pages, illus. 1968. $20.00.
In this revision of Apioninae, Kissinger has once again accomplished a
monumental task. The large number of species and small size of most of the in¬
dividuals make this a taxonomically difficult group. Included are keys to all 28
genera and the males of the 13 subgenera and 317 species. There are 221 pages
of figures and maps of species distribution. The many diagnostic photographs of
generic, subgeneric, and specific features, frequently including the male genitalia,
aid greatly in the identification of specimens. Some of the keys are difficult
to use and even weevil specialists will find many Apion are not easy to identify.
The subgeneric key and many of the specific keys are to males only, and it is
rather difficult to sex Apion. The list of male secondary sex characters is complex
and the small size of Apion complicates matters. A separate plate illustrating
these secondary sex characters would have aided the reader. Except for the
difficulties concerning the secondary sex characters, the sections on technique
and the discussion of taxonomically valuable structures of the adults are well
written.
There are lists of the genera and subgenera not available for study, the 12 species
known only from females, and the 6 species not recognized. There are indices to
all taxonomic categories covered in the paper; to the new taxa, new combinations,
and new synonymies; and to host and associated plants. One minor omission
noted is the lack of symbols for three species in the “Key to Symbols” for figure
188, and two species in figure 190, maps of species distributions.
This publication, though difficult to use, has drawn together in one place much
widely scattered data and has added a great deal of new information. It is
essential to anyone who wishes to identify Apioninae from North and Central
America.— Charles W. O’Brien, Purdue University, Lafayette, Indiana - 1
1 Present address: Texas Tech University, Lubbock.
NEW BULLETIN
The Insects of Virginia.— A new series of bulletins with this title has been ini¬
tiated by Michael Kosztarab and published by the Research Division, of the De¬
partment of Entomology, Virginia Polytechnic Institute. The objective is to
provide a systematic treatment including records on biology and ecology of the
different insect orders and families in Virginia. The general policy for the series
is described in the introductory article of the first bulletin which will be sent free
on request from the Publication Office, 405 Hutcheson Hall, Virginia Polytechnic
Institute, Blacksburg, Virginia 24061.
ADVEF
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Entomology Research Institute _ v
Chemagro Corporation_ vi
Entomological Exchanges and Sales_ vii
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Vol. 46 JULY 1970 No. 3
THE
Pan-Pacific Entomologist
STIBICK—A revision of the genus Leptoschema of California with a key to
related genera (Coleoptera: Elateridae) _ 153
LA RIVERS—A new Philippine Sagocoris (Hemiptera: Naucoridae) _ 167
WIRTH—A revision of the Neotropical genus Physemops Cresson (Diptera:
Ephydridae) _ 170
THORP—The type locality of Bombus franklini and notes on putative Arizona
records of other Bombini (Hymenoptera: Apidae) _ 177
WILLIAMS—A new species of scorpion belonging to the pumilis group of
genus Vejovis (Scorpionida: Vejovidae) _ 181
SCHUSTER AND GRIGARICK—Tardigrada of Santa Cruz Island, California 184
PAULSON—A list of the Odonata of Washington with additions to and
deletions from the state list ____ 194
BROTHERS—Notes on the saprophagous activity of Bufolucilia silvarum
(Meigen) (Diptera: Calliphoridae) _ 198
BOHART—New species and lectotype designations in North American Bem-
becini (Hymenoptera: Sphecidae) _ 201
GOODMAN—Observations on the behavior and biology of Microbembex cali-
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HURD—A new, narrowly polylectic, autumnal species of Dialictus from
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(Hymenoptera: Apoidea) _ 209
SMITH—Biology and structure of some California bristletails and silverfish
(Apterygota: Microcoryphia, Thysanura) _ 212
SCOTT—Resilin in the sound-organs of Pyralidae and Cicadidae (Lepidoptera;
Homoptera) _ 225
SCIENTIFIC NOTE_ 231
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The Pan-Pacific Entomologist
Vol. 46
July 1970
No. 3
A Revision of the Genus Leptoschema of California with
a Key to Related Genera
(Coleoptera: Elateridae)
Jeffrey N. L. Stibick 1
Arlington , Virginia
The comparatively rare elaterid genus Leptoschema possesses a strik¬
ing appearance which brought it to my attention several years ago. In
the course of this study it eventually became clear that this group is
properly placed in the Pomachiliini and not in the Denticollini where
it is currently listed. In addition, examination of available specimens
shows that a closely related complex of two species and four subspecies
exists throughout California. The results of the investigation are
presented in this paper.
I wish to express my appreciation to Miss C. M. F. von Hayek,
British Museum of Natural History (BM), who has assisted me through
her determinations of Chilean material in several genera of Pomachiliini
and discussed some diagnostic characters separating Leptoschema and
Deromecus. Mr. Hugh B. Leech, California Academy of Sciences (CAS),
kindly presented critical material, including the type of L. protractum
variety elegans Van Dyke, and provided indispensable information on
certain obscure localities. Thanks must also be expressed to Mr. T. J.
Spilman, U. S. Department of Agriculture (USNM), for the loan of
specimens; Prof. R. L. Fischer, Michigan State University (MSU), for
much of the Chilean material examined in this study; Dr. John Lawrence,
Museum of Comparative Zoology, Cambridge, Mass. (MCZ), for speci¬
mens; and Dr. N. D. Jago, Philadelphia Academy of Sciences (PAS),
for permission to see the lectotype of L. protractum (Horn). Finally,
I appreciate the efforts of Drs. C. W. and L. B. O’Brien, Purdue Uni¬
versity, for collecting several species of Deromecus and Medonia during
a recent visit to Chile.
1 Present address : D.A.S.F., Konedobu, Port Moresby, Papua—New Guinea.
The Pan-Pacific Entomologist 46: 153-166. July 1970
154
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Pomachiliini Candeze
Pomachiliites Candeze, 1859, p. 4; 1860; 1891.
Pomachiliini, Champion, 1895, p. 402; Schwarz, 1906; Heyne and Taschenberg,
1908.
Pomachiliinae, Schenkling, 1925, p. 189.
Pomachilinae, Laurent, 1965, p. 249. (Misspelling)
Head .—Mouthparts inferior; frons more or less vertical with distinct and
moderate to feebly arcuate carina between eyes, carina well separated from
labrum; antennae filiform to serrate; posterior part of eyes more or less covered
by pronotum.
Thorax .—Prothorax anteriorly somewhat broader than before base; prosternum
moderately lobed anteriorly, prosternal sutures double, often excavate anteriorly;
scutellum truncate anteriorly, noncordiform. Mesosternum separated from
metasternum by distinct suture; mesepisternum very narrowly open to mesocoxae,
touching mesotrochantin; mesocoxal cavities well separated from each other.
Metepimeron narrow. Legs with normally thickened femur and tibia (neither
compressed nor broadened towards the tip) ; tarsi simple or fourth broadened
and cordate or fourth very small and third broadened or lobed; first hind tarsal
segment from nearly as long as segments two and three to sometimes longer than
two, three and four; claws of all legs simple.
Abdomen .—Five visible segments.
Leptoschema, when first described by Horn (1884) was placed in
Candeze’s tribe Athoites. Agriotes protractus Horn was removed from
the Agriotini and designated as type species of Leptoschema. Horn,
however, made special note of the resemblance of A. protractus to a
figure of Psiloniscus borborurus Candeze, which occurs in Mexico
(Cordoba, Veracruz). Apparently there were no further efforts to
investigate any relationship to the Pomachiliini. Candeze removed
it to his Ludiites in 1891, associating it with Oxygonus and Agriotes ,
both now in the Agriotini. Schwarz (1906) moved it back to his
Athouini (later the Athouinae), and Arnett (1962) put Athous and
Denticollis in the same tribe (the Denticollini) with Leptoschema and
other genera. However, Van Dyke (1932), among other things, said
that Leptoschema was somewhere near Sericus (now in the Elaterini)
and not far from Agriotes. He pointed out that the short, free, elevated
clypeal margin and gradually dilated metacoxal plates separate it from
the Athoini (now Denticollini). The chief reason for confusion as
noted by him was Horn’s provisional inclusion of Athous bicolor
LeConte and Athous discalceatum (Say). These species do belong in the
Athouini, differing from Athous chiefly in the double prosternal sutures,
and he reestablished Reitter’s Elathous (placed in synonymy under
Leptoschema) which was based on congeneric European species. Most
European references to Leptoschema, then, actually refer to members
of this genus.
JULY 1970]
STIBICK-REVISION OF LEPTOSCHEMA
155
It is unfortunate that neither Drs. Horn or Van Dyke carried their
inquiries further. But during my investigation of Leptoschema I ran
across numerous examples of various genera of Pomachiliini from
Chile, all in the collection of Michigan State University. Struck by the
extremely close affinities between Leptoschema and especially Deromecus
species in the first section of that genus, I sent a number to Miss von
Hayek, who confirmed their generic placement and established some
specific identifications, based on material in the British Museum. Later,
fresh Chilean material was received from the O’Briens. The general
facies and other characters mentioned by Dr. Van Dyke, as well as
features given in the tribal description, leave no doubt on systematic
placement of Leptoschema in the Pomachiliini.
Since first proposed by Candeze, the Pomachiliini have been lately
elevated to subfamilial status. It seems to me that this is undesirable,
for there are strong relationships between this group and the Agriotini.
Consequently, I am retaining the Pomachiliini as a tribe of the Elaterinae
very near those Agriotini known as Dalopius, and the criddlei group
of Agriotes.
The name Pomachiliini is not new to North America (north of
Mexico). For a long time it was believed that one genus, Betarmon,
was represented in this area by Betarmon calijornicus Schaeffer and
B. higeminata (Randell). Brown (1933) put these species in a new
genus, Agriotella , which he placed near Agriotes and Dalopius
(Agriotini). Later Arnett (1968) put Agriotella in the Ampedini.
While there are strong reasons for keeping Agriotella in the Agriotini,
I do not wish to do more here than note that this genus need not be
considered where the Pomachiliini are concerned. Reestablishment of
this tribe in the United States is hence based on Leptoschema , which
may be likened to a northerly extension of the Pomachiliini into our
area from Mexico.
Leptoschema Horn
Leptoschema Horn, 1884, p. 50; Candeze, 1891, p. 200; Schwarz, 1906, p. 196;
Schenkling, 1925, p. 303; Van Dyke, 1932, p. 356; Arnett, 1962, p. 506.
Head .—Mouthparts inferior; head convex, deeply inserted; frontal ridge
moderately arcuate; eyes relatively small; antennal segments IV to X distinctly
longer than wide, II and III segments subequal in length, together about equal to
IV.
Thorax .—Prothorax anteriorly broader than before base; pronotum medially
convex, more or less broadly and shallowly concave laterally; hind angles pro¬
longed and acute; prostemal sutures double, distinctly curved inwards, feebly
excavated in front, inner carina narrow and thin; prosternal mucro slender and
straight. Mesosternum moderately declivous, margins of fossa feebly raised in
156
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO.
3
front. Elytra obtuse at tip. Metacoxae rather narrow, gradually dilated inwards,
inner angle very obtusely toothed at best. Tarsi simple, segments I to IV decreasing
in length, hind tarsal segment I subequal in length to segments II and III.
This genus is very closely related to Deromecus, especially to those
species forming the first group of the latter. Deromecus impressus
(Solier), in fact, is a close copy of Leptoschema protractum elegans
Stibick. A specimen of this species from Trovolhue, Chile (MSU)
was at first thought by me to be in or near Leptoschema. Miss von
Hayek, who identified it, pointed out that both this specimen and the
type species of Deromecus, D. angustatus (Solier), differ from
Leptoschema protractum as follows: Deromecus — prosternal suture
strongly grooved anteriorly and accompanied for % of its length by
smooth area on prosternum, prosternal mucro turned inward behind
mesocoxae, mesosternal groove with raised border, and metathorax with
lateral groove; Leptoschema —prosternal suture feebly grooved ante¬
riorly without smooth area on prosternum, prosternal mucro straight,
mesosternal groove with only feebly raised border at best, and
metathorax without lateral groove. In addition, the female and male
genitalia of Deromecus (Figs. 2—3) show some differences when com¬
pared with those of Leptoschema (Figs. 6-12). All these seem suf¬
ficient for separate genetic identity, the most important of which is
the prosternal mucro. Leptoschema , due to its curved and grooved
prosternal sutures, elongate antennae, and close affinity to Deromecus,
may be placed near the latter genus and Parapomachilius. It is not
particularly close to the aforementioned Psiloniscus which has (among
other things) straight and closed prosternal sutures. In geographical
terms it is of interest to note that its close relative Deromecus trivittatus
Champion (group one of Candeze) occurs in Oaxaca, Mexico, and that
Paranius mexicanus Champion is known from Ciudad, Durango, Mexico.
Leptoschema is presently known only from California (Map 1). The
specimens included in this study were collected between May and
October, with no indications that any of the species or subspecies appear
at any particular times. This paper greatly extends the known range,
including a previously unsuspected southward extension to Tulare
County. It is apparent that the various species and subspecies have
more or less well separated ranges arrayed east-west and north-south
with an important division of the subspecies of L. protractum in the
San Francisco Bay area. The reasons behind this pattern are not clear,
but it is remarkable that only one of the 35 specimens I have seen is a
female. This disparate sex ratio may be significant as a clue to specia-
tion and the present geographical arrangement of species and subspecies.
JULY 1970 ]
STIBICK—REVISION OF LEPTOSCHEMA
157
Map 1 . Distributions of Leptoschema species.
The role of San Francisco Bay, San Pablo Bay, and other physical
obstructions as barriers, assumes more importance in this light, but it
should be added that both male and female are fully alate.
Key to the Species of Leptoschema
1. Body solid brown; pronotal punctures shallow and close but not nearly
confluent (Fig. 4) ; middle California_ praelontactum Stibick, n. sp. 2
Body bicolored, pronotum darker than elytra or with median dark vitta;
pronotal punctures deep, nearly to sometimes confluent (Fig. 5) ; Coastal
and San Francisco Bay area _ protraction (Horn) 3
153
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
2. Antennae reaching two segments behind hind angles of pronotum _
_ p. praelontactum Stibick, n. subsp.
Antennae scarcely attaining apex of hind angles of pronotum_
___ p. gloris Stibick, n. subsp.
3. Pronotum uniformly dark or with very faint median vitta; east and west
of San Francisco Bay, south of Golden Gate _ 4
Pronotum with prominent median vitta, sides and elytra reddish-brown;
north of Golden Gate and south of San Pablo Bay__ 5
4. Along coast west and south of San Francisco Bay; aedeagus of moderate
length, without clear area at apex of paramere (Fig. 8) ; shorter (12-13
mm ) _ protractum (Horn)
Alameda County east of San Francisco Bay; aedeagus long and slender, basal
piece with notch in internal edge (Fig. 9) ; longer (14 mm) -
__ p. detractum Stibick, n. subsp.
5. North of San Francisco Bay along coast; aedeagus of moderate length, with
clear spot at apex of paramere (Fig. 10) _p. elegans Stibick, n. subsp.
Contra Costa County, south of San Pablo Bay; aedeagus long and slender,
basal piece V-shaped along internal edge (Fig. 11) _
_ p. analelegans Stibick, n. subsp.
Leptoschema protractum (Horn)
(Figs. 1, 5, 8)
Agriotes protractus Horn, 1871, p. 317; 1884, p. 50.
Leptoschema protractus (Horn); Schwarz, 1906, p. 197; Heyne and Taschenberg,
1908, p. 159; Schenkling, 1927, p. 303. (Error)
Leptoschema protractum (Horn) ; Horn, 1884, p. 51; Candeze, 1891, p. 200;
Van Dyke, 1932, p. 356.
Readily recognized by the deep and close, coarse punctation, bicolored appear¬
ance, and male aedeagus.
Male.— Length 12 mm, width 2.9 mm, shape elongate, scarcely convex; color
black to brownish-orange, generally bicolored with the pronotum pitchy black
and the elytra and sometimes the abdomen and legs brownish-orange; punctation
unisetigenous, coarse and very dense, finer on venter; surface generally rough
on pronotum, smoother and more lucidus elsewhere; vestiture moderate, lightly
flavous, wiry and decumbent.
Head. —About % as long as wide, nearly % median width of pronotum; front
arcuate at vertex, otherwise vertical; punctures large, vaguely hexagonal and
nearly confluent. Antennae reaching one segment beyond pronotal hind angles,
scape cylindrical, as long as pedicel and III segment combined; the latter short,
feebly cylindrical and compressed; IV to XI segments each nearly twice length
of preceding two, moderately serrate, save oblongly slender eleventh.
Thorax. —Median width/length as 6.25/9.25; sides straight, scarcely arcuate at
apex, feebly arcuate along hind angles; hind angles very prominent, prolonged and
acute, sides continued laterally by caudal extension of propleuron, bisecting dorsal
carina present, this prominent, thin, and sharp, continuing beyond base of angle;
disk moderately convex, sides abruptly declivous to low notopleural suture running
nearly ventral to eyes, caudal side very abruptly declivous in front of hind angles,
especially in median line where a short median canal is formed, this side abruptly
JULY 1970 ]
STIBICK—REVISION OF LEPTOSCHEMA
159
ending well before enlarged lateral plates of propleuron (consequently exposing
them to dorsal view) ; surface piceous, with vague longitudinal impressions near
sides, median area convex; punctures large, coarse, very closely spaced to con¬
fluent, vaguely hexagonal; setae directed caudad. Pleural region piceous with
band of siennous shading along pleurosternal suture, punctures moderately large,
moderately dense, rather circular in outline, surface lucidus; setae directed
anteriorly. Prosternal lobe rather abruptly turned ventrally and only moderately
arcuate; prosternum with moderately large punctures, these dense and rather
circular in outline, surface lucidus and convex, scarcely concave on sides, setae
directed anteriorly, prosternal suture feebly grooved anteriorly, prosternal mucro
slightly concave near base, straight. Sides of mesosternal fossa feebly raised
in front. Metasternum somewhat finely and closely punctate. Scutellum elongate,
width/length as 2.5/4, moderately inclined (30°), surface finely punctate, slightly
opaque and rather rugulose. Elytra striate, these more or less formed by moderate,
sometimes confluent punctures or grooves between punctures; interspaces with
three rows of very fine punctures; color lightly siennous.
Abdomen .—Moderately finely punctate, lighter in color towards middle.
Male genitalia .—Aedeagus (Fig. 8) slender, moderately elongate (1.9 mm).
Dorsal side flat, heavily sclerotized; ventral side nearly covered by lateroventral
sides of parameres, membranous under center piece except near apex. Apex of
center piece exceeding parameres; median strut slender, divided at apex and
fused to base of each paramere; lateral struts slender, elongate, nearly straight.
Parameres with strong lateral lobe; 12 to 13 loosely clustered setae on ventral
side of lobe, all confined to well-defined lightly sclerotized area at apex; lobe
acute, outer side straight (Fig. 8a). Pars basalis narrow and extremely convex,
V-shaped.
Female. —Unknown for the nominate subspecies. See under L. protractum
elegans.
Distibution. —California, found south of San Francisco Bay near
the coast. Thirty specimens were examined, all males.
No locality: 5, including types (CAS, MCZ, PAS). San Mateo Co.: 1,
Crystal Lake, 7 May 1916 (CAS); 2, Jasper Ridge (E. of Searsville Lake), 21
May 1950 (CAS, JNLS). Santa Clara Co.: 1, Stanford Univ., 21 May 1951 (CAS) ;
1, Stevens Creek (NW of Saratoga), 5 May 1940 (CAS). Santa Cruz Co.: 1, 9
June 1917, 600 ft. (CAS) ; 15, Ben Lomond, 1931, 16 and 17 May 1931, April 1932,
May, 25 May and 17 September 1932, 17 July 1933, 19 May and 3 July 1946 (BM,
CAS, JNLS, MCZ, MSU, USNM) ; 4, Soquel Creek, 30 May 1909 (CAS, JNLS).
The California Academy of Sciences, through Mr. Leech, kindly
presented me with seven specimens. I have in turn donated one to the
British Museum and one to Michigan State University. Five are retained
in my collection (JNLS).
This is the form first described by Horn. I have seen both male
cotypes in the Philadelphia Academy of Sciences. One has the following
labels: “Cal.,” “Lectotype, 3373,” “Leptoschema protractum Horn.”
The other is labeled “Cal.,” “Paratype 3373.” The first specimen may
be regarded as the lectotype of the species.
160
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 NO. 3
Fig. 1. Leptoschema protractum (Horn), length 12 mm. Fig. 2. Deromecus
impressus Solier, female genitalia. Fig. 3. Deromecus sp., group 1, male genitalia,
dorsal view. Fig. 4. Leptoschema praelontactum Stibick, section of pronotum,
showing punctation. Fig. 5. L. protractum, section of pronotum showing puncta-
tion. Fig. 6. L. praelontactum, male genitalia, dorsal view; 6a. Apex of left
paramere, ventral view. Fig. 7. L. praelontactum gloris Stibick, male genitalia,
dorsal view; 7a. Apex of left paramere, ventral view. Fig. 8. L. protractum,
male genitalia, dorsal view; 8a. Apex of left paramere, ventral view.
JULY 1970 ] STIBICK—REVISION OF LEPTOSCHEMA
161
When Dr. Horn described the genus Leptoschema in 1884, protractus
was corrected to protractum , since Leptoschema is a singular Greek
adjective, neuter gender. Other authors apparently mistook the generic
name for Latin, hence their use of the -us ending.
The relationship of a number of morphological features with locality
has indicated the necessity of recognizing a number of subspecies. Two
of these subspecies may yet prove to form a separate species.
Leptoschema protractum elegans Stibick, new subspecies
(Figs. 10, 12)
Leptoschema protractum var. elegans Van Dyke, 1932, p. 357.
Male.— Length 12-13 mm, width 2.8-3 mm. Color yellow (flavous) to orange
(aurantiacus), generally golden yellow (auratus) with median pitchy black
(piceous) to shining brown (fulvescent) vitta on pronotum, more fulvescent on
venter. Antennae extending one segment beyond pronotal hind angles, antennal
setae erect. Aedeagus (Fig. 10) slender, moderately elongate (1.9 mm) ; apex of
paramere with suboval clear spot, 14 to 15 setae, and feebly curved outer edge
(Fig. 10a).
Female. —Length 16.5 mm, width 3.6 mm. Pronotum (7/9) median width to
median length, sides straight, moderately arcuate near apex, scarcely arcuate along
hind angles. Antennal setae decumbent. Pleural region vaguely micro-reticulate.
Female genitalia (Fig. 12) with elongate, apically narrowing bursa copulatrix
adorned with numerous furca-like or simple spines in three major areas along back
and one area in U-turn. Colleterial glands elongate and narrow.
Distribution.— Found near the coast of California north of San
Francisco Bay.
Holotype male , Humboldt County, California, collected on 15 May
1911 by F. W. Nunenmacher (CAS, No. 3144, type specimen of Van
Dyke, 1932). Allotype, from Sylvania, Sonoma County, collected on
8 May 1898 by R. Haywood. Paratypes, four males as follows: 2,
California (PAS, MCZ) ; 1, Sonoma County (USNM) ; 1, Sylvania,
Sonoma County, May, collected by R. Ecker (CAS).
Originally described as a variety by Dr. Van Dyke, this form is
now raised to subspecies status on the basis of the male aedeagus and
distribution. It is apparently indistinguishable from L. protractum
analelegans Stibick; save for the aforementioned features. However,
the aedeagus shows it to be most closely related to the nominate sub¬
species (Figs. 8, 10).
I have not been able to determine the type locality, Humboldt County,
more precisely, since the late collector, Nunenmacher, did not leave
any pertinent notes. However, the similarity of the aedeagus in the
male examples from Sonoma and Marin counties to the type lead me to
place all these specimens in the same subspecies, including the one
162
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Fig. 9. L. protractum detraction Stibick, male genitalia, dorsal view; 9a.
Apex of left paramere, ventral view. Fig. 10. L. protractum elegans Stibick, male
genitalia, dorsal view; 10a. Apex of left paramere, ventral view. Fig. 11. L.
protractum analelegans Stibick, male genitalia, dorsal view; 11a. Apex of left
paramere, ventral view. Fig. 12. L. protractum elegans, female genitalia. Fig. 13.
Denticollis productus (Randell), ventral view of metasternum, mesosternum, and
mesocoxal cavities. Fig. 14. Limonius ectypus (Say), ventral view showing most
of mesosternum, tip of metasternum, pro- and mesocoxal cavities, and prosternal
mucro. Fig. 15. Athous nigropilis (Motschulsky), ventral view showing most of
mesosternum, tip of metasternum, pro- and mesocoxal cavities, and prosternal
mucro.
JULY 1970 ] STIBICK—REVISION OF LEPTOSCHEMA
163
female allotype. Sylvania, not on today’s maps, is near the present site
of Camp Meeker, Sonoma County (H. B. Leech).
From the only female Leptoschema known, we may possibly expect
other females of the genus to be somewhat larger and broader, and
without the stiffly erect antennal pubescence characteristic of the male.
Leptoschema protractum detractum Stibick, new subspecies
(Fig. 9)
Male.— Length 14 mm, width 3 mm. Pronotum piceous, elytra golden yellow,
antennae extending one segment beyond pronotal hind angles. Aedeagus (Fig. 9)
quite elongate (2.7 mm) ; apex of paramere with 17 to 19 setae, lobe acute, outer
side straight (Fig. 9a) ; basal piece with notch in internal edge.
Female.— Unknown.
Holotype male, San Leandro Hills, Oakland, Alameda County,
California, collected in June 1881 (CAS). “Hills back of Oakland,
Cal., VI-1881”; “4808”; “Van Dyke Collection.” One paratype male,
from Alameda County, collected on 21 May 1930 by F. E. Blaisdell
(CAS). “Hayward.” Type locality defined as San Leandro Hills,
Oakland, Alameda County, California.
Apparently indistinguishable from the nominate, save for a slightly
larger size, slightly lighter elytra, and range. The aedeagus is elongate,
so much so that this may form, with L. protractum analelegans, a distinct
species. However, since the shape of the aedeagus is otherwise some¬
what similar, and external features seem so feeble, for the present they
should be regarded as subspecies of the nominate.
Leptoschema protractum analelegans Stibick, new subspecies
(Fig. 11)
Male. —Length 13 mm, width 3 mm. Color generally golden yellow with median
pitchy black vitta on pronotum. Antennae extending one segment beyond pronotal
hind angles. Aedeagus (Fig. 11) quite elongate (2.7 mm) ; apex of parameres
with 14 to 15 setae, lobe acute, outer side straight (Fig. 11a) ; basal piece without
median notch.
Female. —Unknown.
Holotype male, Lafayette, Contra Costa County, California, col¬
lected on 24 May 1940 (CAS). The type locality defined as Lafayette,
Contra Costa Co., California.
Similar to L. protractum elegans, and best distinguished by the
aedeagus or by locality. But it is related to L. protractum detractum,
and may be only a feeble variety of the latter, although the prominent
median vitta, basal piece, and setae pattern of the parameres seem to
separate it.
164
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Leptoschema praelontactum Stibick, new species
(Figs. 4, 6 )
Male. —Length 15 mm, width 3 mm. Shape elongate, scarcely convex; color
uniformly shining brown, generally lighter beneath, somewhat darker on scutellum
and prosternum; punctation shallow, unisetigenous, moderately dense, finer on
venter; surface generally lucidus; vestiture moderate, lightly flavous, wiry and
decumbent.
Head —About % as long as wide, ^7 median width of pronotum; front arcuate
at vertex, otherwise vertical; punctures moderately large, circular in shape, close
but not confluent. Antennae reaching two segments beyond pronotal hind angles,
scape cylindrate, slightly longer than pedicel and III segment combined; the
latter short, feebly cylindrate and compressed; IV to XI segments each 2% times
length of preceding two, moderately serrate, save oblongly slender XI.
Thorax .—Median width/length as 5.6/7.5; sides straight, scarcely arcuate at
apex, very feebly arcuate along hind angles; hind angles very prominent, pro¬
longed, acute, sides continued laterally by caudal extension of propleuron, bisecting
dorsal carina present, this prominent, thin, and sharp, continuing beyond base
of angles; disk moderately convex, sides abruptly declivous to low notopleural
suture running nearly ventral to eyes, caudal side very abruptly declivous in front
of hind angles, especially in median line where short median canal occurs, this
side abruptly ending well before enlarged lateral plates of propleuron (consequently
exposing them to dorsal view) ; surface with vague longitudinal impressions, median
area convex, punctures moderately large, shallow, and numerous, generally well¬
spaced, circular in shape; setae directed caudad. Pleural region with moderate
punctation, surface lucidus; setae directed anteriorly. Prosternal lobe abruptly
turned ventrally and only moderately arcuate; prosternum with somewhat coarse
but well-separated punctures, surface highly lucidus, convex, scarcely concave on
sides, setae directed anteriorly; prosternal suture feebly grooved anteriorly;
prosternal mucro slightly concave near base, straight. Sides of mesosternal fossa
feebly raised in front. Metasternum somewhat finely and closely punctate.
Scutellum elongate ( 1 .3/2.4), moderately inclined (30°), surface tending to
piceous shade, finely punctate, slightly opaque and rugulose. Elytra striate, these
formed by moderate punctures or grooves between punctures; interspaces with
three rows of very fine punctures.
Abdomen .—Moderately finely punctate, color darker away from sides.
Male genitalia .—Aedeagus (Fig. 6 ) somewhat stout (1.9 mm) ; dorsal side flat,
heavily sclerotized; ventral side only partly covered by lateral-ventral sides of
parameres, membranous under center piece except near apex. Apex of center
piece exceeding parameres; median strut slender, divided at apex and fused
to base of each paramere; lateral struts moderately stout, nearly straight.
Parameres with strong lateral lobes; loose row of five setae on ventral side of
lobe, these confined to well-defined lightly sclerotized area at apex, lobe rather
obese in shape, outer side somewhat arcuate (Fig. 6 a). Basal piece convex,
U-shaped.
F emale. —Unknown.
Holotype male, Springfield, Tulare County, California, collected
on 24 May 1935 by F. T. Scott (CAS). “Springville, Cal., 5-24-35”;
JULY 1970 ] STIBICK-REVISION OF LEPTOSCHEMA
165
“Ft. Scott Coll.”; “R. Hopping Collection”; “Leptoschema sp. prob.
new, G. R. Hopping”; “n. sp. H.B.L.” One paratype male, creek near
Chico, Butte County, collected on 23 April 1922 by E. C. Van Dyke
(CAS). Type locality defined as Springville, Tulare Co., California.
A beautiful and unmistakable, but very rare species. Its distribution
(Map 1) greatly extends the known limits of Leptoschema throughout
California. There is one subspecies, described next.
Leptoschema praelontactum gloris Stibick, new subspecies
(Fig. 7)
Male.— Length 16 mm, width 3 mm. Uniformly shining brown, lighter beneath
save for darker prosternum and scutellum. Antennae scarcely attaining apex of
hind angles, IV to XI segments each nearly twice length of II and III combined.
Aedeagus (Fig. 7) somewhat stout (1.9 mm) ; apex of parameres with moderately
strong lateral lobes, row of five setae on ventral side below lightly sclerotized
apical area, lobe quite obese, outer side quite arcuate (Fig. 7a).
Female.— Unknown.
Holotype male , Putah Canyon, Yolo County, California, collected
on 14 May 1933 by E. C. Zimmerman (CAS). Type locality defined as
Putah Canyon, a section of Putah Creek in Yolo County, California
adjacent to the Napa County line. Mr. Leech informs me that the
Monticello Dam on the county line and the Putah Dam near Winters
have radically changed ecological conditions there.
The general shape of the aedeagus, shallow pronotal punctation, and
shiny brown appearance indicate its close relationship to the nominate.
But the very short antennae and specific features of the aedeagus,
especially the apex of the parameres, seem to warrant subspecific status.
The range (Map 1) is interesting in that it is midway between the
nominate and the various forms of L. protractum (Horn).
Generic Keys
The systematic reassignment of Leptoschema requires some changes
in Arnett’s 1968 key to the genera, couplets 34 to 38. I have taken
this opportunity to present some additional characters felt to be of value
in separating the various genera involved, and the following key may be
substituted for couplets 34 to 38 in Arnett (1968).
Tribe PoMACHlLUNI
1 . Head convex, front vertical, mouth inferior _ Leptoschema
Head and front inclined, mouth nearly prognathous (Denticollini) _ 2
166
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Tribe Denticollini
2(1). Metasternum pointed at apex (Fig. 13); mesocoxal cavities nearly con¬
tiguous; prosternal lobe not extending forward beyond hind margins
of eyes ___ Denticollis
Metasternum narrow, usually truncate (Figs. 14, 15) ; mesocoxal cavities
usually well separated; if metasternum apparently pointed and cavities
approximate, then prosternal lobe extending forward beyond hind
margins of eyes_ 3
3(2). Prosternal mucro moderately to strongly broadened behind procoxae
(Fig. 14), prosternal sutures double; antennal segments two and three
usually subequal_ 4
Prosternal mucro not widened behind procoxae (Fig. 15) ; prosternal
sutures single; antennal segment two usually smaller than three _ 5
4(3). Carina of frons thickened, frons deeply impressed behind margin_ Elathous
Carina of frons not thickened, often emarginate_ Limonius
5(3). Tarsal segments 1-4 with a membranous lobe beneath; carina of frons
weak_ Hemicrepidius
Tarsal segment one without a membranous lobe, others various; carina
of frons very well developed _ Athous
Literature Cited
Arnett, R. H., Jr. 1962. Beetles of the United States, Fascicle 46, Elateridae,
Catholic Univ. America, Press, Washington, D. C., pp. 497-512.
Brown, W. J. 1933. Studies in the Elateridae II. Can. Entomol., 65:173-182.
Candeze, E. 1859. Monographic des Elaterides II. Mem. Soc. Roy. Sci. Liege,
2:4-53.
1860. Monographic des Elaterides III. Mem. Soc. Roy. Sci. Liege, 15:1-49.
1891. Catalogue methodique des Elaterides connus en 1890. Liege, pp.
200 - 201 .
Champion, G. C. 1895. Insecta, Coleoptera (Elateridae). Biologia Centrali-
Americana, 3:402-407.
Heyne, A. and Taschenberg, O. 1908. Die exotischen Kafer in Wort und Bild.
Leipzig, 27:159.
Horn, G. H. 1871. Descriptions of new species of Elateridae, of the United
States. Trans. Amer. Entomol. Soc., 3:299-358.
1884. A study of some genera of Elateridae. Trans. Amer. Entomol. Soc.,
12:33-52.
Laurent, L. 1965. Les Elateridae de Belgique. Bull. Inst. Agron. Sta. Res.
Gembloux, 33:245-290.
Schenkling, S. 1925. Coleopterorum catalogus, pars 80, Elateridae I, 11:189.
1927. Coleopterorum catalogus, pars 888, Elateridae II, 11:303.
Schwarz, O. 1906. Coleoptera, fam. Elateridae. Genera Insectorum, 46a:
136-149, 196-197.
Van Dyke, E. C. 1932. Miscellaneous studies in the Elateridae and related
families of Coleoptera. Proc. Calif. Acad. Sci., sec. 4, 20:291-465.
JULY 1970]
LA RIVERS-SAGOCORIS USINGERI
167
A New Philippine Scigocoris
(Hemiptera: Naucoridae)
Ira La Rivers
Verdi, Nevada
During the course of monographic work with the naucorid fauna of
New Guinea, isolated specimens have come to hand from peripheral
areas. Since no unified treatment of these areas is contemplated in the
immediate future, I am submitting the following single description as
an interim contribution to what little we know of the naucorids of these
regions.
Sagocoris usingeri La Rivers, new species
General.— A moderately convex, contrastingly mottled species with some sheen,
micropunctation and roughness. Length 10.5 mm, width 6 mm. Head and pronotum
a bit lighter overall than hemelytra (i.e., with greater preponderance of background
light yellow). Venter a bit darker, with lighter plastron.
Male.—H ead—Rather square in interocular area, light yellow, darkening in pos¬
terior pattern; micropunctation cutting down noticeably on sheen; anteclypeal
area moderately and smoothly protuberant between eyes, latter rising noticeably
above general head surface when viewed obliquely from rear; inner eye margins
subparallel; outer margin curving smoothly into posterior edge, small hyperoche
(non-optical flap) evident at what would be the meeting point if that was angular.
Head set broadly and shallowly into leading edge of pronotum. Viewed anteriorly,
mouthparts (including labrum) set into distinct but shallow well formed by over¬
hanging edge of anteclypeus above, and by shallow extension of juga and maxillary
plates laterally. Labrum medium in size, yellowish, sharply rounded on end; ratio
of length:width 5:11. Gula keeled, but not sharply so, not pointed at either end.
Maxillary plates prominent, tips protruding free at edges of mouthparts-well.
Antennae 3-segmented, slender, basal segment bilobed. Head ratios (1) total
length:width (including eyes) 46:75, (excluding eyes) 23:21, (2) anterior dis¬
tance between eyes:posterior distance between eyes 41:50, (3) total length:length
of head posterior to line drawn between posterior eye edges 46:7.
Pronotum —Light yellowish, with usual double triangular pattern of dark brown
dots and suffusions occupying disc, producing, particularly, a prominent and light
posterior pronotal band. Side edges smoothly and slightly curved posteriorly to
sharply rounded posterolateral areas, lacking pilosity. Venter lighter in vicinity
of posterolateral areas, darkening inwardly. Prosternum prominently keeled in
anterior three-fifths, leading edge extending as rather blunt but definite point over
adjacent edge of gula. Propleural flaps just slightly free internally, but not ex¬
tending over any part of posterior prosternum. Pronotal ratios (1) width between
anterior angles:width between posterior angles 3:5, (2) median length:greatest
width 6:25.
Scutellum —Brownish-black with light yellow center. Ratio of three sides, ante¬
rior and two laterals, 15:11:11.
Hemelytra —Brownish-black over most of area, with lightening in emboliar,
The Pan-Pacific Entomologist 46: 167-169. July 1970
168
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 NO. 3
claval and corium-merabrane junction regions. Some glisten, but micropunctation
very evident. Extend full length of abdomen, with well-developed membrane sec¬
tions which overlap when wings closed. Claval, corial and emboliar areas well
marked. Rather narrowly exposing connexiva. Embolium typically lighter ante¬
riorly and laterally, crease anterointernally noticeable; ratio lengthrwidth 13:4,
making it rather narrow. Hind wings vestigial, reduced to one-third abdominal
length.
Venter —The pro thoracic venter has been considered. Abdomen a bit lighter than
sterna because of hydrofuge plastron. Mesosternal median tumidity bisinuous in
lateral view, (highly reminiscent of some oedipodine grasshoppers) with anterior
tumidity short and rounded, posterior one occupying remainder of mesosternum,
the two separated by a sulcus. Connexival margins thin, posterolateral angles
right-angulate but not spinose.
JULY 1970 ]
LA RIVERS-SAGOCORIS USINGERI
169
Proleg —Coxa and trochanter large, conspicuous, bearing very well-developed
incrassate femur, globular and swollen when viewed ventrad, rather flat in hori¬
zontal view. Femoral ratio of length :width 31:23. Tibia-tarsus a single narrow unit
closing against leading femoral edge, tarsus 1-segmented and grading into single
sharp claw or point.
Mesoleg —Coxa elongate-globular, connecting to long, narrow, flattened femur
by means of conspicuous trochanter. Legs of these sagocorines very reminiscent
of Ambrysus in the Ambrysinae. Femur somewhat rectangular in cross-section with
row of fine, short spines along each edge; ratio length:width 60:13, length 2.6 mm.
Tibia narrower, rounded in cross-section, bearing more prominent reddish spines
over its surface, these getting larger and more numerous distally; ratio length:
width 50:7, length 2.3 mm. Tarsus 3-segmented, first segment very short, remaining
two elongated and terminating in two large but weakly-curved claws.
Metaleg —Larger copy of mesoleg. Femur with ratio of length:width 82:15,
length 4 mm. Tibia more heavily spined than mesotibia, spines arranged more
systematically into longitudinal rows; ratio of length:width 95:8, length 4.1 mm;
with fringe of long swimming hairs on inner margin. Tarsi as in mesoleg, but
larger and darker.
Female. —Unknown.
Holotype male and 1 paratype Luzon, Mountain Province, Benguet,
Philippines, 15 July 1946, D. G. Frey (Amer. Mus. Nat. Hist.) ; 2
paratypes, Mountain Province, Mt. Data, April 1946, H. Hoogstraal &
D. Heyneman, 7,000 ft. (Chicago Nat. Hist. Mus.) ; 1 paratype, Moun¬
tain Province, Abatan, Buguias, 60 km S of Bontoc, 1 June 1964, H. M.
Torrevillas, c. 2,000 ft., light trap 1 (Bishop Mus.).
This distinctive species is named for the late Robert Leslie Usinger—
teacher, colleague and friend of 20 years, with whom I was working on
southeastern Asiatic naucorids at the time of his premature death in
San Francisco.
Comparative notes.— This is a well-defined species, in certain pro-
notal characteristics (such as the posterolateral angles) appearing more
like Sagocoris biroi Montandon 1911 than any other member of the
genus. However, it is rather sharply set off by the distinctive anterior
angulation of the tip of the prosternal median keel of xiphus. Very
likely the structure of the female subgenital plate will further distinguish
it when this is known.
1 The designation “light trap” on the label of this specimen can only be an error, since this is
a flightless species.
170 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
A Revision of the Neotropical Genus Physemops Cresson
(Diptera: Ephydridae)
Willis W. Wirth
Systematic Entomology Laboratory, Entomology Research Division,
Agr. Res. Serv., USDA 1
The genus Physemops Cresson has been known only from the type
species, P. nemorosus (Cresson), a small black fly with bright whitish
pubescent lower face, found around rocky streams in Neotropical rain
forests. The systematic position of this fly was dubious because of
several structural peculiarities which were thought to be of a generic
nature. The following quotation from the discussion accompanying
Cresson’s (1914) original description shows the quandary this species
has placed us in.
“This genus may be distinguished by the peculiar shield-like development of the
face which extends rather low, appearing somewhat subhemispherical, being evenly
and distinctly convex in all directions, including the cheeks. There are no
indications of the usual parafacial grooves or areas, and the face is destitute of
characteristic bristles or hairs. In general the face appears vaulted or arched,
as is typical with Ephydra and Parydra, but the mouth is not proportionately so
large. At first glance its head suggests affinity to Ilythea, but on further examina¬
tion it is seen to belong to an entirely different subfamily. The form of the head,
thorax and abdomen, and its chaetotaxy determines its relation to Hydrina and
Axysta. The abdomen suggests Gastrops, having apparently only four segments
in the male and five in the female with the second and third much longer than
the others. The apical margin of the third, in profile, is horizontal as in Lytogaster
and Axysta.”
Examination of much fresh material including five new species of this
genus results in an expansion of our concept of the generic characters
and indicates that some of the most striking characteristics of P.
nemorosus are of a specific nature. The presence in some of these species
of distinct series of dorsocentral and acrostichal bristles, and of a pair
of enlarged bristles on the medifacies suggests that Physemops belongs
in the subfamily Ephydrinae. Examination of the Oriental Psilephydra
cyanoprosopa Hendel reveals that it is related to Physemops on one hand
and to the Polynesian genus Apulvillus Malloch on the other. The
latter in turn is closely related to Scatella Robineau-Desvoidy and
Neoscatella Malloch.
Two terms coming into common use in Acalyptrate Diptera taxonomy
1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560.
The Pan-Pacific Entomologist 46: 170-177. July 1970
JULY 1970]
WIRTH-NEOTROPICAL PHYSEMOPS
171
should be mentioned here: The costal index is the length of the second
section of the costa (between apices of first and second veins) divided by
the length of the third section (between apices of second and third
veins). The fourth vein index is the length of the apical section of the
fourth vein (between posterior crossvein and wing margin) divided by
the length of the preapical section (between anterior and posterior cross¬
veins) .
I am especially indebted to J. Maldonado Capriles of the University of
Puerto Rico in Mayaguez, and to Marshall R. Wheeler of the University
of Texas in Austin, for the opportunity to study material from their
collections.
Genus Physemops Cresson
Physemops Cresson, 1914, Trans. Amer. Entomol. Soc., 60: 211. Type species,
Psilephydra nemorosa Cresson, by original designation.
Diagnosis. —Small, broad and stocky, blackish flies with long convex face
usually without bristles but with dense pubescence. Head broad, relatively high
and short, the frons broader than long; one pair of fronto-orbital bristles curving
laterad and caudad; inner verticals strong and also curving laterad and caudad;
outer verticals strong, erect, bent slightly forward and mesad; ocellars strong,
erect, bent slightly forward and laterad (absent in P. wheeled). (In P. nemorosus
and P. azul only the inner and outer verticals are strongly developed.) Face with
a pair of moderately long, divergent submedian bristles at apex of facial hump on
upper portion (these absent in P. nemorosus and P. azul) ; lower margin sharp at
the moderately wide mouth opening; no trace of clypeus present. Antenna
without spine on second segment; third segment usually broader than long, bearing
long arista with variable development of pubescence above and below, or of short
dorsal rays.
Thorax with one pair of strong dorsocentral bristles always present, some
species sometimes also with considerable development of up to three anterior
dorsocentrals and several pairs of acrostichals; two pairs of notopleurals and
variable development of other lateral bristles (presuturals, supraalars, interalars,
and postalars) ; two pairs of scutellars present; a strong mesopleural and a
strong sternopleural. (In P. nemorosus and some other species all but posterior
dorsocentral and marginal scutellars may be greatly or somewhat reduced.) Scutel-
lum broader than long and somewhat convex (except in P. panops).
Abdomen short and moderately convex, relative lengths of terga variable with the
species. Male abdomen usually with fifth tergum greatly convex and apex turned
under.
Legs without special modifications or strong bristles; femora moderately stout;
apical tarsomere may be expanded, especially in male; claws strong and curved,
empodium and pulvilli well developed.
Wing unmarked, apex of first vein sometimes strong and darkened; second
vein long, extending nearly to apex of wing; fifth vein straight in one group
of species, in the other markedly bowed or sinuate, discal cell thus convex behind.
172
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Key to the Species of Physemops
1. Antenna entirely black or dark brown___ 2
Antenna with third segment entirely or mostly yellowish _ 3
2 . Large species, wing 2.90 mm long; face entirely shining black, upper facials
well developed; arista with longer raylike branches above and below;
tarsi black _ wheeled Wirth, n. sp.
Small species, wing 2.00 mm long; face brown to black above, whitish to
golden below, upper facials absent; arista with short pubescence on both
sides; tarsi pale ___ nemorosus (Cresson)
3. Integument of face yellowish; dorsum of thorax coarsely granulose,
mesonotum greenish, scutellum bluish__ panops Wirth, n. sp.
Integument of face blackish; dorsum of thorax blackish, not coarsely
granulose _ 4
4. Small species, wing 1.75 mm long; lower face with bluish pubescence viewed
from above, upper facials absent; legs with brown markings _
_ azul Wirth, n. sp.
Large species, wing 2.10-2.34 mm long; lower face with whitish pubescence,
upper facials present; femora blackish _ 5
5. Tibiae yellow; only one pair of long dorsocentrals present; arista with
short dorsal rays ___ maldonadoi Wirth, n. sp.
Tibiae blackish; one pair of long dorsocentrals plus three shorter ones
present; arista virtually bare _ fairchildi Wirth, n. sp.
Physemops azul Wirth, new species
Female.— Length of wing 1.75 mm. Dark brown to black; mesonotum and
scutellum subshining, with sparse fine punctures; third antennal segment yellow;
legs yellowish, all of femora except apices, broad band on tibiae, and tarsomeres
3-5 brown; halter black, base of stem yellowish. Head bristles reduced except for
strong inner and outer verticals. Face relatively short and broad, shaped about as
in P. nemorosus, with scattered fine pale microscopic hairs, no enlarged submedian
bristles on upper face; with dense greenish pubescence, that on lower portion
bright aquamarine blue when viewed from above. Arista about as long as breadth
of vertex, with coarse pubescence above and below. Mesonotum with long bristles,
three pairs each in dorsocentral and acrostichal series. Scutellum broader than
long, moderately flattened; two pairs of strong marginal bristles plus two pairs
of finer marginal hairs. Relative lengths of abdominal terga 2-5 as 20:26:8:3.
Wing deeply brownish infuscated, the veins dark brown; fifth vein deeply bowed;
second vein long, costal index 5.0; fourth vein index 2.0.
Male.— Unknown.
Distribution. —Mexico.
Types. — Holotype female, 1 female paratype, Valle Nacional,
Oaxaca, Mexico, 15 May 1963, collector unknown (Type no. 70664,
USNM).
Discussion. —This small species is closely related to P. nemorosus
resembling it in general body shape and small size, wing venation, its
micropubescent arista, and lack of upper facials. It can readily be
JULY 1970]
WIRTH-NEOTROPICAL PHYSEMOPS
173
distinguished from that species by its yellow third antennal segment,
shining dorsum with numerous large mesonotal bristles, and by its
brilliant bluish face.
Physemops fairchildi Wirth, new species
Male. —Length of wing 2.10 mm. Large shining black species; third antennal
segment yellowish, becoming brownish on apex; tarsi brown distally, tarsomere
one yellowish on foreleg, one to three yellowish on mid and hind pairs; wing
yellowish brown hyaline with coarse microtrichia, veins yellowish brown; halter
whitish. Arista short and nearly bare. Face longer than in P. nemorosa, with
more pronounced facial hump, covered with blackish microscopic hairs, pair of
long submedian facials at apex of hump, face polished black above these and
pearly white pubescent below; cheek polished black, sharply contrasting with
whitish face. Frons, mesonotum and scutellum subshining granulose, scutellum
somewhat transversely microrugulose; abdominal terga smooth and shining, lengths
of terga 2-5 as 10:10:12:15. One pair of long dorsocentral bristles plus three
shorter ones anteriorly in series and some small scattered acrostichal hairs. Scutel¬
lum broader than long, moderately convex dorsally, the marginal bristles arising
from distinct tubercles; apical scutellars long, the anterior pair only half as long.
Wing long and narrow; second vein very long and paralleling costa, costal index
6.7, submarginal cell narrowed distally; fourth vein index 1.7; fifth vein straight
bordering discal cell.
Female. —Unknown.
Distribution.— Colombia, Panama.
Types.— Holotype male , 2 paratypes, Cerro Campana, Panama
Prov., Panama, July 1967, W. W. Wirth (Type no. 70665, USNM).
Colombia: Bogota vicinity, 8700 ft., February 1958, M. R. Wheeler, 1
male paratype; Medellin, 8000 ft., February 1958, M. R. Wheeler, 3
male paratypes.
Discussion.— This species resembles P. maldonadoi in wing venation,
pale halteres, long convex face with long facials, but differs in the nearly
bare arista, black tibiae, and the shining black upper facial area.
I take pleasure in naming this species in honor of Dr. G. B. Fairchild
of the Gorgas Memorial Laboratory in Panama City, Panama, in recogni¬
tion of his outstanding contributions to our knowledge of Neotropical
Diptera.
Physemops maldonadoi Wirth, new species
Male. —Length of wing 2.34 mm. Large shining black species; third antennal
segment, all of tibiae except apical spot, and first two tarsomeres yellow; halter
whitish; wing grayish hyaline, the veins brown. Face long, with pronounced
median hump, two moderately long submedian facials placed at the apex of the
hump; face clothed with scattered short, microscopic hairs; surface shining black,
with sparse pearly white pubescence becoming heavier towards lower margin.
Third antennal segment longer than broad; arista slightly shorter than width of
174
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
vertex, with short dorsal rays. Frons, mesonotum and scutellum with sparse but
coarse, scalelike tomentum, appearing moderately shining; only one pair of long
dorsocentrals, other hairs in dorsocentral and acrostichal series short and fine.
Scutellum broader than long, convex dorsally, basal scutellar a little more than
half as long as apical pair. Abdominal terga 2-5 with lengths as 8:13:10:3. Wing
with second vein long but running straight to costa distally, costal index 3.3;
fourth vein index 2.1; fifth vein in straight line behind distal cell.
Female.— Identical to male, except abdominal terga 2-5 with lengths as
10 : 8 : 8 : 8 .
Distribution. —Puerto Rico.
Types.— Holotype male , allotype female, 10 male and 6 female para-
types, Yauco-Lares Road km 29, Puerto Rico, 20 January 1954, J.
Maldonado and S. Medina (Type no. 70666, USNM).
Discussion. —The wing venation, pale halteres, long convex face with
long facials and yellow third antennal segment ally this species with P.
fairchildi , but that species differs in the nearly bare arista, black tibiae
and the shining black upper facial area.
This species is named for Dr. J. Maldonado Capriles of the University
of Puerto Rico in Mayaguez in recognition of his many contributions to
our knowledge of Puerto Rican Diptera.
Physemops nemorosus (Cresson)
Psilephydra nemorosa Cresson, 1914, Entomol. News, 25: 244 (Costa Rica; fig.
head) ; Cresson, 1918, Trans. Amer. Entomol. Soc., 44: 64 (redescribed; fig.
head).
Physemops nemorosus (Cresson) ; Cresson, 1934, Trans. Amer. Entomol. Soc., 60:
211 (combination; type species of new genus Physemops) ; Wirth, 1968, Cat.
Diptera Amer. S. of U. S. 77: 20 (distribution).
Male.— Length of wing 2.00 mm. Entirely shining black with faint submetallic
luster, or somewhat obscured by sparse brown pollen and granulose or scorbiculate
surface, especially of thorax and scutellum. Wing brownish hyaline; halter black;
legs black, trochanters, apices and bases of tibiae, and tarsomeres 1-4 yellowish.
Head bristles absent except for strong inner and outer verticals. Face densely
pubescent, appearing brownish above and whitish to golden on lower portion;
broadly convex without pronounced median hump, with scattered microscopic hairs,
no enlarged facials on upper portion. Antenna with third segment brown, broader
than long; arista very long, longer than breadth of vertex, with dorsal and ventral
feathering of minute hairs. Mesonotum and scutellum with dense, coarse granula¬
tion; one pair of long dorsocentral bristles, others in dorsocentral and acrostichal
series reduced to fine, short, seta-like hairs; scutellum broader than long, convex
dorsally, anterior scutellar half as long as apical pair. Abdomen dorsally with
coarse scalelike tomentum appearing subshining; terga 2-4 with lengths as
20:20:10, tergum 5 not visible from above, but quite prominent and convex from
caudal or ventral aspect. Fifth tarsomere of anterior leg dilated, with an apical
fan of eight or more hairs, the claws long and stout, so spread laterally as to
JULY 1970]
WIRTH-NEOTROPICAL PHYSEMOPS
175
oppose each other, their pulvilli also enlarged. Wing broad; second vein long and
paralleling costa; costa index 4.6; fourth vein index 1.7; fifth vein sinuate.
Female. —As for male, except abdominal terga 2-4 with lengths as 30:30:7,
and shape of fifth tarsomere of anterior leg.
Distribution. —Mexico to Ecuador and Brazil, Dominica.
Specimens Examined—Brazil : Juquia, Sao Paulo, December 1938, J. Lane, 1.
Costa Rica : Higuito, San Mateo, P. Schild, 1; La Suiza, 1923, P. Schild, 1.
Dominica: Fond Figues River, January-March 1965, W. W. Wirth, rain forest,
11; 17 March 1964, D. F. Bray, 1. Ecuador: Bugna, Chimborazo, July 1955, R.
Levi-Castillo, 1; Camarones, Manabi, 9 September 1955, R. Levi-Castillo, 3; Estero
Balsa, Manabi, 9 September 1955, R. Levi-Castillo, 3; Los Rios, Guare, August
1955, R. Levi-Castillo, 2; Santo Domingo de los Colorados, March 1958, M. R.
Wheeler, 1. El Salvador: Los Chorros, December 1953, W. B. Heed, 1; Santa
Tecla, 12 km nw, December 1953, W. B. Heed, 1. Honduras: Lancetilla, April
1954, W. B. Heed, 1. Mexico: Valle Nacional, Oaxaca, 16 May 1963, collector
unknown, 1. Nicaragua: El Recreo, June 1954, W. B. Heed, 1. Panama: Balboa,
Canal Zone, February 1958, M. R. Wheeler, 1; Barro Colorado Island, C. Z., July
1967, W. W. Wirth, 7; Cerro Campana, Panama Prov., July 1967, W. W. Wirth,
rocky stream, 11; Gamboa, C. Z., Rio Agua Salud, July 1967, W. W. Wirth, rocky
stream, 42; Hermita, Code Prov., 18 September 1952, F. S. Blanton, 1; Las Cruces
Trail, C. Z., February 1958, M. R. Wheeler, 5; Pedro Miguel, C. Z., 10 April 1923,
R. C. Shannon, 1.
Physemops panops Wirth, new species
Male. —Length of wing 2.40 mm. Large multicolored species; integument of
face yellowish; antenna, tibiae and first two tarsomeres yellow; wing deeply
yellowish including veins; halter creamy white. Body black; frons subshining
brown with fine granulosity; mesonotum and scutellum coarsely granulose;
scutellum and narrow median line on mesonotum and notopleura deep blue,
remainder of mesonotum greenish. Four pairs of dorsocentrals, last two becoming
well developed; acrostichal hairs short and fine. Scutellum as long as broad,
flattened above and pointed distally, apical bristles close together; basal scutellars
half as long as apical pair. Face long and markedly convex but hump low, sub¬
median pair of long facials on hump; face with scattered, short, fine, black hairs,
with white pubescence becoming thin on mid and upper portions. Antenna
with arista shorter than breadth of vertex, with moderately long dorsal rays.
Wing long and slender, with well developed anal angle and axillary lobe; second
vein long and paralleling costa, costal index 4.1; fourth vein index 2.0, fifth vein in
straight line behind discal cell. Abdomen with lengths of terga 2-5 as 16:20:15:20
in males; shining black, second tergum slightly granulose, the distal segments
with sparse, scalelike tomentum.
Female. —Identical to male, except abdominal terga 2-5 with lengths as
18:15:15:20.
Distribution.— Haiti.
Types. —Holotype male , allotype female, 1 male paratype, Kenscoff,
Haiti, 4000 ft., February 1956, W. B. Heed (Type no. 70667, USNM).
Discussion.— This species is closely related to P. maldonadoi from
176
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Puerto Rico with the same wing venation, pale halteres, long convex
face with long facials, and yellow antenna and tibiae, but the yellow
facial integument and the brilliant metallic blue and green granulosity
of the thorax are quite distinctive. The facial aspect suggests the name
panops, after Pan, the chimpanzee.
Physemops wheeleri Wirth, new species
Female.— Length of wing, 2.90 mm. Large uniformly dull black species, includ¬
ing tarsi and halteres; wing dark smoky brown with blackish veins, with very
coarse microtrichia. Third antennal segment much shorter than its breadth; arista
sparsely feathered with moderately long dorsal and ventral rays. Face long and
appearing more vertical than the other species of the genus, with submedian
upper facials well developed; integument transversely microrugulose, appearing
subshining, with scattered, fine short hairs, two widely spaced elongate hairs on
oral margin. Frons, mesonotum and scutellum finely granulose; ocellar bristles
absent. Only one pair of long dorsocentrals, the anterior dorsocentrals and the
acrostichals appearing as short fine hairs. Scutellum broader than long, posterior
margin rounded, basal scutellars very small, less than a third as long as apical
pair. Abdomen with sparse scalelike tomentum and scattered erect fine hairs;
lengths of terga 2-5 as 22:32:10:5. Wing broad, posterior margin rounded with
broadly rounded anal angle more or less paralleling the deeply bowed fifth vein;
marginal cell broad; costal index 10.0, this value so high due to the narrowing
of submarginal cell by the anteriorly bowed second vein; third vein nearly straight;
fourth vein index 1.7.
Male.—U nknown.
Distribution. —Ecuador, Panama.
Types. —Holotype female, Las Cruces Trail, Canal Zone, Panama,
February 1958, M. R. Wheeler (Type no. 70668, USNM). Paratypes, 25
females, as follows: Ecuador: Santo Domingo de los Colorados, March
1958, M. R. Wheeler, 1. Panama: Balboa, C. Z., February 1958, M.
R. Wheeler, 5; Barro Colorado Island, C. Z., July 1967, W. W. Wirth,
2; Gamboa, C. Z., Rio Agua Salud, July 1967, W. W. Wirth, 1; Las
Cruces Trail, C. Z., February 1958, M. R. Wheeler, 15; Las Cruces
Trail, ex bracket fungi, 21 July 1967, W. W. Wirth, 1.
Discussion. —This species is dedicated to Dr. Marshall R. Wheeler
of the University of Texas in recognition of his outstanding contributions
to our knowledge of Acalyptrate Diptera.
P. wheeleri belongs to the group with P. azul and P. nemorosus, all with
short stocky bodies, broad wing with posteriorly bowed fifth vein, dark
halteres and arista feathered or pubescent above and below. It differs
from the other two in its larger size, longer hairs on the arista, presence
of a well developed pair of facials and lack of pubescent vestiture on the
face.
JULY 1970]
THORP-PUTATIVE ARIZONA BOMBINI
177
Literature Cited
Cresson, E. T., Jr. 1914. Descriptions of new genera and species of the dipterous
family Ephydridae.—I. Entomol. News, 25: 241-250, 1 plate.
The Type Locality of Bombus franklini and Notes on Putative
Arizona Records of Other Bombini
(Hymenoptera: Apidae)
Robbin W. Thorp
University of California , Davis
The type locality and distribution of Bombus franklini (Frison) is an
enigma. Frison (1921) based his description on two queens sent to him
by Mr. E. J. Oslar and labelled as collected by Oslar at Nogales, Arizona
in July 1917. However, all subsequent specimens assignable to this
species (Frison, 1923, 1926; Stephen, 1957; and Thorp unpublished)
have been taken within a 60 mile radius of Grants Pass, Oregon.
Frison (1921, 1923) published additional records of Bombini
reputedly collected by Mr. Oslar in June and July 1917 at four localities
in southern Arizona (Nogales, Oracle, Catalina Mountains and
Patagonia Mountains).
It is my contention that these southern Arizona records, including
the type localities for B. franklini and the new varieties of B. sylvicola
and B. balteatus , published by Frison (1921, 1923) are not valid. Lines
of evidence used in reaching this conclusion include: the known dis¬
tributions of the species involved; the number of incongruous records
involved; and the lack of confirmation through subsequent collections
that any of these species occur in southern Arizona.
The Oslar material recorded by Frison (1921) contained the fol¬
lowing: Bombus (as Bremus ) o. occidentalis Greene, B. o. nigroscutatus
Franklin, B. appositus Cresson, B. edwardsii Cresson (as fernaldae
Franklin), B. vosnesenskii Radoszkowsky, B. flavifrons dimidiatus Ash-
mead (as ambiguus Franklin), B. californicus F. Smith, B. franklini
(Frison) and Psithyrus crawfordi Franklin. The Oslar material re¬
corded by Frison (1923) contained: B. balteatus (as kirbyellus ) vars.
alexanderi (Frison) and arizonensis (Frison), B. sylvicola var. lutzi
(Frison) and B. pleuralis Nylander.
Thanks to Dr. W. E. LaBerge of the Illinois Natural History Survey I
have had an opportunity to examine most of the types and specimens
The Pan-Pacific Entomologist 46: 177-180. July 1970
178
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
listed by Frison. Frison (1921) failed to recognize two additional
species included in the material he recorded: B. mixtus Cresson (mis-
identified as fernaldae in part) and B. caliginosus (Frison) (mis-
identified as vosnesenskii in part). I also found one queen each of two
other species with the same collection data on them. One was B.
nevadensis Cresson and the other B. morrisoni Cresson both correctly
identified by Frison.
Six of the species bearing the Oslar, June or July 1917, southern
Arizona labels are principally Boreal in distribution: B. balteatus
Dahlbom is circumpolar, in the Nearctic it extends southward only in
the higher mountains to California and New Mexico where it occurs
principally above 10,000 feet (Thorp, 1962) ; B. sylvicola Kirby occurs
from Alaska to Labrador with southward extensions similar to those of
B. balteatus (Thorp, 1962) ; B. pleuralis is known only from Alaska,
Northwest Territories, Yukon Territory, and northern British Columbia;
B. o. occidentalis occurs throughout the mountainous areas west of
103° W. longitude from Alaska south to the northern portions of
California, Arizona and New Mexico; B. appositus occurs throughout
the mountains west of 103° W. longitude from British Columbia and
Alberta south to the central portions of California, Nevada, Utah, and
New Mexico; B. mixtus occurs in the mountainous areas west of 105°
W. longitude from Alaska south to California, Nevada, Utah and
Colorado. The lower elevational limit for all these species increases as
their ranges extend southward so that at their southern limits they
occur only above 9,000 feet.
Bombus californicus is also principally Boreal, but extends into the
Transition zone in California and northwestern Baja California, Mexico.
It is found only sparingly in the Rocky Mountains of Idaho, Montana,
Wyoming and northern Colorado and appears to be replaced by B.
fervidus (Fabricius) in Nevada, Utah, southern Colorado, Arizona,
New Mexico and east of 100° W. longitude.
Most of the remaining species occur principally west of the crest of
the Sierra Nevada and Cascade Range in California, Oregon and Wash¬
ington, with two extending to northwestern Baja California and one into
Alaska: B. o. nigroscutatus is restricted to the San Francisco Bay region
of California; B. franklini is restricted to the Siskiyou Mountain region
of southern Oregon and northern California; B. caliginosus is restricted
to the coastal region of California, Oregon and Washington; B.
edwardsii occurs in northwestern Baja California, is widespread in
California and southwestern Oregon and is replaced by B. melanopygus
Nylander to the north; B. vosnesenskii ranges from northwestern Baja
JULY 1970]
THORP—PUTATIVE ARIZONA BOMBINI
179
California to southwestern British Columbia and extreme western
Nevada; B. flavifrons dimidiatus occurs from the Sierra Nevada and
North Coast Ranges of California through the Cascade Range into
southern British Columbia and is replaced to the north and in the
Rocky Mountain region by B. /. flavifrons Cresson; Psithyrus crawfordi
occurs in the North Coast Ranges, Sierra Nevada and Warner Mountains
of California, extreme western Nevada and southern and western Oregon,
and is replaced to the north and east by P. insularis (F. Smith).
Apparently Frison omitted the records of B. nevadensis and B.
morrisoni from his lists since both of these species had been recorded
in Arizona by Franklin (1913). Bombus nevadensis is a widespread
Boreal-Transition species with several color variants occurring from
British Columbia to New York and in the west south into the mountains
of California, Nevada, southern Utah and northern New Mexico.
Bombus morrisoni is principally a Great Basin-Sonoran species which is
abundant in Arizona and occurs in the area in question. Perhaps the
most significant omission is that of B. sonorus Say which is the most
abundant species in my experience in the area between Nogales and
Oracle. The lack of any specimens of this species bearing the same
collection data is another indication that the Oslar locality data reported
by Frison (1921, 1923) were based on specimens with erroneous labels.
Many of the species listed from the putative Oslar localities, coexist
in areas of comparable size (horizontal distance—85 miles and vertical
range—3,900 to 9,200 feet), but only west of the Sierran-Cascade crests.
Further, several of the species have mutually exclusive ranges separated
by 100 miles or more, e.g. B. pleuralis-franklini-o. nigroscutatus and
B. balteatus-franklini-o. nigroscutatus, and B. balteatus-caliginosus.
Frison (1923) evidently had more information than is available on
the specimen labels since he states “According to Mr. Oslar, the bumble¬
bees labelled ‘Patagonia Mountains, Arizona,’ were found feeding on
the blossoms of wild blackberry and mesquite at altitudes from 5,500
to 6,000 feet.” My own experience suggests that mesquite does not
reach this elevation in the Patagonias, but is replaced first by oaks
(4—5,000 feet) and then conifers (5-6,000 feet). In addition none of
the Bombini in question occur in or near areas containing mesquite.
According to Mr. R. Martin Brown ( pers. comm.) “He (Oslar)
exchanged widely with other professional and amateur collectors and
labelled his material to sell.” Neither Mr. Brown nor I have been able
to determine whether Oslar ever visited the area in question personally
or whether he received the material from another collector.
Among the 10 species of Bombus and one Psithyrus listed by Frison
180
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO.
3
(1921, 1923) as having been collected by E. J. Oslar in the Nogales to
Oracle area of southern Arizona, not one has been subsequently collected
in this area and only one species record has been definitely confirmed
for the state. The latter is based on specimens of B. o. occidentalis
taken above 9,000 feet in the San Francisco Mountains which is about
200 miles north of and 4,500 feet higher than Oracle. Of the additional
species I found in the Illinois Natural History Survey collection, only
B. morrisoni definitely occurs in the area between Nogales and Oracle.
Franklin’s (1913) putative record of B. nevadensis from Arizona based
on one male, and his supposition that B. californicus should occur in
Arizona have not yet been confirmed by subsequent collections.
In 1962 I discussed the distributions of B. balteatus and B. sylvicola
and suggested that the Patagonia Mountain records were incongruous
for varieties of these species. However, being unaware of any contrary
evidence, I accepted the records. Based on the evidence at hand I now
suggest that all the species recorded by Frison (1921, 1923) as being
collected by E. J. Oslar in 1917 in southern Arizona be disregarded
until confirmed by subsequent collections. Bombus o. occidentalis
should be retained in Arizona lists since it has been taken in the northern
part of the state. I also propose Gold Hill, Jackson County, Oregon
as the new type locality for B. franklini , since it is in the approximate
center of the known distribution of the species and specimens of all
castes have been taken there. Frison’s (1923) varieties of B. balteatus
and B. sylvicola most closely resemble color variants from the Rocky
Mountains of Colorado, however I do not consider these varieties as
valid subspecies and do not propose to select new type localities for
them.
Fiterature Cited
Franklin, H. J. 1913. The Bombidae of the new world. Trans. Amer. Entomol.
Soc., 38: 177-486.
Frison, T. H. 1921. New distribution records for North American Bremidae,
with the description of a new species (Hym.). Entomol. News, 32:
144-148.
1923. Systematic and biological notes on bumblebees (Bremidae:
Hymenoptera). Trans. Amer. Entomol. Soc., 48: 307-326.
1926. Descriptions and records of North American Bremidae, together
with notes on the synonymy of certain species (Hymenoptera). Ibid.,
52: 129-145.
Stephen, W. P. 1957. Bumble bees of western America (Hymenoptera:
Apoidea). Oregon State Coll. Agr. Exp. Sta. Tech. Bull. 40, 163 p.
Thorp, R. W. 1962. Notes on the distributions of some bumblebees of western
North America (Hymenoptera: Apidae). Pan-Pac. Entomol., 38:
21-28.
JULY 1970 ] WILLIAMS-A NEW MEXICAN VEJOVIS
181
A New Species of Scorpion Belonging to the Pumilis Group
of Genus Vejovis
(Scorpionida: Vejovidae)
Stanley C. Williams
San Francisco State College, California
During the summer of 1968 an entomological expedition through
Baja California, Mexico led by S. C. Williams and M. A. Cazier collected
a number of new species of scorpion. Of special interest was one sub¬
sequently described as Vejovis pumilis Williams. This was of special
interest because: it was one of the smallest species of the genus, ap¬
peared to be endemic to the Magdalena Plain (a terrestrial habitat of
relatively recent formation) and apparently was quite unrelated to the
other members of the genus. Also among the specimens collected on
this expedition, but not discovered until recently, was a much less
abundant and undescribed species of special interest because it appears
to be an ecological counterpart of Vejovis pumilis in the Viscaino Desert
of Baja California, and because it appears to be the only close known
relative of Vejovis pumilis. This new species is here described and
named Vejovis pseudopumilis.
Vejovis pseudopumilis Williams, new species
(Figures 1, 2)
Diagnosis.— Tiny pale yellow species of Vejovis. Only close relative known is
Vejovis pumilis Williams from which it differs in the following ways: pectines
distinctly larger and with more numerous teeth; telson shorter than metasoma
segment I; vesicle proportionally larger, twice aculeus length (three times aculeus
length in V. pumilis) ; telson light amber not contrasting orange; pedipalp fingers
not scalloped, but with tiny gap when chela closed (scalloped and with large gap
in V. pumilis).
Holotype male. Coloration: uniform pale yellow, pedipalp fingers light pink,
telson light amber, pectines whitish. Carapace with anterior border convex; each
group of lateral eyes with two large anterior eyes and third obsolescent one.
Mesosoma with stigma tiny, long oval; last sternite with one pair of irregularly
granular keels; last tergite with two pairs of irregularly granular lateral keels and
poorly developed median keel. Metasoma with dorsal and dorsolateral keels
not strongly raised, these irregularly crenulate to dentate; inferior lateral keels
smooth to faintly crenulate on segments I to III, irregularly crenulate on IV,
irregularly serrate on V; inferior median keels on segments I to III smooth, smooth
to faintly crenulate on IV, irregularly serrate on V; inferior median keels set
with 3/4/4/5 pairs of stout red hairs on segments I to IV respectively; segment I
slightly longer than wide, segment II definitely longer than wide. Telson elongate;
long slender vesicle almost twice aculeus length; entire telson shorter than
The Pan-Pacific Entomologist 46: 181-183. July 1970
182
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Figs. 1 and 2. Vejovis pseudopumilis Williams, new species, holotype male.
Fig. 1. Dorsal view. Fig. 2. Ventral view.
metasomal segment V; ventral surface of vesicle with several large rounded
tubercles including large subaculear tubercle; this surface with about 10 pairs
of long stout hairs. Chelicerae with inferior border of movable finger completely
lacking denticles. Pedipalps with small chelae, these slightly swollen and with
short fingers; movable finger % carapace length and shorter than carapace width
at median eyes; fingers essentially unscalloped inwardly but leave tiny proximal
space when chela closed; palm with smooth to crenulate keels.
Standard measurements .—Table 1.
The holotype male was collected at San Angel, 13 miles west of San
Ignacio, Baja California Sur, Mexico, 28 June 1968 by S. C. Williams,
JULY 1970 ]
WILLIAMS-A NEW MEXICAN VEJOVIS
183
Table 1 . Measurements (in millimeters) of Vejovis pseudopumilis
Williams, new species, holotype.
Holotype
(male)
Total length
27.0
Carapace, length
3.4
width (at median eyes)
2.3
Metasoma, length
12.9
segment I (length/width)
1.8/1.7
segment II (length/width)
2.1/1.6
segment III (length/width)
2.1/1.4
segment IV (length/width)
2.9/1.4
segment V (length/width)
4.0/1.4
Telson, length
3.6
Vesicle (length/width)
2.3/0.9
depth
0.8
Aculeus, length
1.3
Pedipalp
Humerus (length/width)
2.4/0.9
Brachium (length/width)
2.6/1.2
Chela (length/width)
4.2/1.4
depth
1.6
movable finger, length
2.0
fixed finger, length
1.4
Pectines
teeth (left/right)
18/18
M. A. Cazier and party. The holotype is permanently deposited in the
California Academy of Sciences.
This species is named “pseudopumilis” because it closely resembles its
own known close relative Vejovis pumilis Williams.
Acknowledgments. —The field and laboratory research involved in
the discovery, collection and study of this species was supported by the
systematics branch of the National Science Foundation by research
grant number GB 7679 and by a Faculty Research Leave from San
Francisco State College. Thanks are due Christie Steketee for clerical
assistance in the preparation of this manuscript.
134
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Tardigrada of Santa Cruz Island, California
Robert 0. Schuster and Albert A. Grigarick
University of California, Davis
Schuster and Grigarick, in a previous study of the Tardigrada fauna
of Western North America (1965), recorded 38 species from the Cali¬
fornia mainland. Since that report, extensive collections have been
accumulated from Santa Cruz Island, Santa Barbara County, and now
the major features of that insular fauna can be identified.
Santa Cruz Island is 25 miles south of Santa Barbara, and 18 miles
west of Port Hueneme. It is 25 miles long and over 2,400 feet in eleva¬
tion. A central valley and several permanent freshwater streams are
notable features of the island, and these are responsible for locally
modifying the climate and for increasing the variety of habitats.
Samples, mainly of algae, lichens, and mosses, were collected and
the tardigrades were extracted from them by using simple Baermann
funnels. Core samples of intertidal sand were washed and decanted
through sieves of 325 mesh per sq. inch. The specimens are mounted
on slides in a Hoyer’s type medium, and are in the museum of the
Department of Entomology at Davis.
Most of the collection sites may be found on United States Geological
Survey quadrangles Santa Cruz Island A, B, C, and D. These sites are
identified by number on the accompanying may (Map 1), and the
separate collections at each site bear letter designations. Some col¬
lections of plant material were too small or too intermixed to allow
easy identification, and the tardigrade species recovered were referred
simply to the site number.
Acknowledgments
We greatly appreciate the kindness of Dr. William A. Weber of the
University of Colorado Museum for providing determinations of the
lichens and mosses, and of Dr. Paul C. Silva, University of California,
Berkeley, for naming the marine algae. Michael R. Benedict of the
Channel Island Field Station, University of California, identified the
ecologically distinctive areas of the island and succeeded in transporting
us to most of them. Dr. Donald S. Horning, Jr. collected samples,
especially in the pines at Centinela, a habitat not sampled by the authors.
Joanne S. Wasbauer and Susan K. Senser also assisted in the survey.
The Pan-Pacific Entomologist 46: 184-193. July 1970
JULY 1970] SCHUSTER & GRIGARICK-SCI TARDIGRADA
185
Collection Data
Tardigrades were found at sixteen of the localities surveyed (Map 1)
and only a few unusual situations (for example, water troughs) did not
yield tardigrades. Specimens were recovered from approximately one-
half of the 45 plant species collected and these associations are sum¬
marized following the collection data.
Pelican Bay. —Collections labeled Pelican Bay were from the slopes east of the
Bay, from near sea level to about 600 feet, and were collected on 4 May 1969.
1 A Homalothecium pinnatifidum (Sull. et Lesq.) Lawton.
1 B Physcia sp.
1 C Ramalina ceruchis (Ach.) deNot.
1 D Ramalina homalea Ach.
1 E Ramalina reticulata (Noedh.) Krempelh.
Prisoners Harbor.- —These collections were made near the beach, at or to the
west of the pier. Sample (C) was collected on 9 May 1968, and the others
on 28 April 1969.
2 A Intertidal
2 B Physcia sp.
2 C Ramalina ceruchis (Ach.) deNot.
2 D Ramalina homalea Ach.
Coches Prietos.— The 1968 samples (A-D) were from moderately luxurious
growth on cliffs to the west of the beach. The 1969 collections (E-G) were
from the higher and somewhat drier hills to the east.
3 A Parmelia arnoldii Du Rietz.
3 B Ramalina ceruchis (Ach.) deNot.
3 C Ramalina homalea Ach.
3 D Selaginella bigelovii Underw.
3 E Caloplaca sp.
3 F Homalothecium pinnatifidum (Sull. et Lesq.) Lawton.
3 G Parmelia cristifera Say.
Johnston Canyon.—
4 Intertidal collections only were made at this locality.
Ciiristi Beach. —Numerous intertidal core samples did not reveal marine species.
Terrestrial species were recovered from lichens collected by D. S. Horning, Jr.,
14 May 1969, on the ridge above Christi airport.
5 A Parmelia caperata (L.) Ach.
5 B Ramalina reticulata (Noedh.) Krempelh.
Two miles east of Fraser Point. —This locality is a canyon protected from the
prevailing wind. A considerable quantity of material was processed from
both spring and fall collections with very few specimens being recovered.
Sample number 6 was collected 11 May 1968, and samples A and B were
collected 30 April 1969.
6 A Ramalina homalea Ach. on rocks.
6 B Selaginella bigelovii Underw.
Fraser Point. —A considerable amount of rocky shoreline is exposed at low
00
Ov
SANTA CRUZ ISL.
CALIF.
N
1 PELICAN BAY
2 PRISONERS HARBOR
3 COCHES PRIETOS
4 JOHNSTON CANYON
5 CHRISTI BEACH
6 2 MI. E. FRASER POINT
7 FRASER POINT
8 WEST POINT
9 LOS PINOS DEL SUR
10 LA ALAMEDA
11 CANADA DEL MEDIO
12 CASCADA
13 CENTINELA
14 LA CUESTA
15 CAMPO RATON
16 CANADA DE LOS SAUCES
0 / 2
mi/es
s
Ros 1969
Map 1. Santa Cruz Island tardigrade collection sites.
co
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO.
JULY 1970 ] SCHUSTER & GRIGAKICK-SCI TARDIGRADA
187
tide. Two genera of marine tardigrades were recovered from samples col¬
lected 30 May 1969, and 28 September 1969.
7 Intertidal algae (May).
7 A Mixed Rhodoglossum affine (Harv.) Kylin and Gigartina canaliculata
(Harv.) Harv., and Porphyra perforata J. Ag. (Sept.).
West Point. —Rocks on the cliffs at this locality had luxuriant lichen growth of
many species but few specimens of tardigrades were recovered. The samples
were collected on 21 September 1968.
8 Ramalina sp.
Los Pinos del Sur. —Collections at this locality were of lichens on branches of
Arctostaphylos insularis and A. tomentosa subcordata, in May, 1969.
9 Ramalina sp.
La Alameda. —Small rocks covered with lichen were collected at this locality on
11 May 1968.
10 Parmelia (Xanthoparmelia ) sp.
Canada del Medio.- —This locality includes samples from the floor of Canada del
Medio near the field station, but mostly from a south facing canyon opening
near the station. The canyon has a small stream and is shaded in the after¬
noon. Samples were taken from the lower elevations where Quercus agrifolia
was present, and were collected in May of 1968, and April and May of 1969.
11 A Algae in stream.
11 B Azola sp.
11 C Moss.
11 D Parmelia ( Xanthoparmelia ) sp.
11 E Physcia callosa Nyl.
11 F Moss near stream.
11 G Porella navicularis (Lehm. et Lindenb.) Lindb.
11 H Salix lasiolepis roots and algae.
11 I Selaginella bigelovii Underw.
Cascada. —Cascada is about a mile west of the field station. The situation differs
from that locality mainly in greater stream flow, extensive sandy stream
bottom, and a more open canyon with less shade.
12 A Cardionema ramosissima (Weinm.) A. Nels. & MacGr.
12 B Parmelia nr. mexicana Gyel.
12 C Selaginella bigelovii Underw.
Centinela.— This locality is in a coniferous forest and the samples were from
the bark and twigs of Pinus remorata and Heteromeles arbutifolia. Collections
were by D. S. Horning, Jr., 14 May 1969.
13 A Parmelia caperata (L.) Ach. on Pinus.
13 B Usnea sp. on Heteromeles.
La Cuesta.— La Cuesta is a grade between Campo Raton and Centinela. The col¬
lected area is east of, and at higher elevation than, Campo Raton. Col¬
lections were by D. S. Horning, Jr., 14 May 1969.
14 A Ramalina reticulata (Noedh.) Krempelh.
Campo Raton.-— This area is at the eastern end of Christi Canyon and consists
of a small stream shaded by oaks. Moss and lichens collected from oak in
1967 were not identified.
188
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
15 A Moss.
15 B Lichen.
Canada de los Sauces.— Sauces is a canyon south of, and parallel to, Christi
Canyon. Lichens were collected from Monterey Cypress on 14 May 1969 by
D. S. Horning, Jr.
16 Lichen.
Summary of Tardigrade and Plant Associations
Batillipes sp.: mixed Rhodoglossum affine (Harv.) Kylin and Gigartina
canaliculata (Harv.) Harv., Porphyra perforata J. Ag.
Echiniscoides sigismundi: marine algae, interstitial.
Echiniscus arctomys: Caloplaca sp., Parmelia arnoldii, P. caperata, P. cristifera,
Physcia callosa, Porella navicularis, Ramalina ceruchis, R. homalea, Selaginella
bigelovii, Usnea sp.
Hypsibius canadensis: Ramalina homalea, Physcia callosa.
Hypsibius dujardini: freshwater algae.
Hypsibius convergens: Porella navicularis, Selaginella bigelovii.
Hypsibius oberhaeuseri : Parmelia arnoldii, Porella navicularis, Ramalina ceruchis,
Selaginella bigelovii, Usnea sp.
Macrobiotus areolatus: Porella navicularis.
Macrobiotus dispar: freshwater algae, Azola sp.
Macrobiotus harmswortiti : Caloplaca sp., Parmelia caperata, Physcia sp.,
Ramalina ceruchis, Selaginella bigelovii, Usnea sp.
Macrobiotus hufelandii: Caloplaca sp., Cardionema ramosissima, Homalothecium
pinnatifidum, Parmelia cristifera, Physcia sp., Porella navicularis, Ramalina
ceruchis, R. homalea, R. reticulata, Selaginella bigelovii.
Macrobiotus intermedius: Physcia sp., Ramalina ceruchis.
Milnesium tardigradum: Cardionema ramosissima, Homalothecium pinnatifidum,
Parmelia caperata, P. cristifera, P. nr. mexicana, Physcia callosa, Ramalina
ceruchis, R. homalea, R. reticulata, Selaginella bigelovii.
The tardigrade species are neither equally abundant nor uniformly
distributed on the island. Marine, freshwater, and terrestrial environ¬
ments have characteristic tardigrade inhabitants. The majority of
species collected were terrestrial. Existing host-plant associations
appear inadequate to explain observed distributional patterns of these
terrestrial species. Habitat or host specificity may be expressed at a
particular site, but the same plant species may harbor different tardi¬
grade species at other localities.
Systematics
Batillipedidae Ramazzotti, 1962
Batillipes Richters, 1909
Specimens of this genus are either very uncommon or occupy a niche
consistently missed in random sampling of the intertidal habitat. More
JULY 1970] SCHUSTER & GRIGARICK-SCI TARDIGRADA
189
than 400 interstitial core samples, each of 100 cc minimum volume,
were collected from Prisoners Harbor, Coches Prietos, Christi Beach
and Fraser Point. These samples included sands of different texture,
were taken from high to minus tide levels, and were all tardigrade
negative. Three specimens were recovered from washings of three
algae (samples 7, 7A). This low density may be indicative of these
collections being from an atypical habitat for the genus.
Batillipes ? similis Schulz, 1955 (Figs. 1, 2). Three available speci¬
mens show instability in some taxonomic characters, particularly the
dorsal spines and the leg spines, and have not been identified with
certainty. The species is very similar to B. similis in respect to body
shape, and in size and form of the cephalic appendages.
Oreellidae Ramazzotti, 1962
Echiniscoides Plate, 1889
Echiniscoides sigismundi (M. Schultze, 1865). This marine tardi¬
grade is easily recognized by the number of claws, usually eight, on each
leg. It is most abundant in the green algae growing in and around
barnacles but infrequently may be found in washings from other
intertidal habitats. We have specimens from sites 2A, 4, and 7, and
presume the species may be found whenever the coast is rocky and
suitable algal growth is present.
Echiniscidae Thulin, 1928
Echiniscus C. A. S. Schultze, 1840
Echiniscus arctomys Ehrenberg, 1853. The genus Echiniscus is easily
recognized as it is the only one on Santa Cruz Island having the dorsum
covered by distinct plates. A single species, E. arctomys , is widely
distributed on the island. It has been recovered from samples IB,
3A, 3B, 3C, 3D, 3E, 3G, 5A, 11C, HE, 11G, and 13B. The species
appears to be equally prevalent in lichens, mosses, and hepatics.
Macrobiotidae Thulin, 1928
This family is represented on Santa Cruz Island by two genera,
Macrohiotus and Hypsibius. Macrobiotus is distinguished by the mouth
tube having a ventral supporting structure (fig. 7VS) and by the claws
of each leg being similar. Species of Hypsibius generally do not have the
buccal tube support and the two claws of each leg are dissimilar. The
species of both genera can be recognized by the diameter of the mouth
tube, by the placement of the stylet supports, by the number and relative
190
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
\ HI!
%\t
15
JULY 1970 ] SCHUSTER & GRIGARICK-SCI TARDIGRADA
191
size of the placoids (apophysis and placoids are inked solid in the figs.),
and by the shape of the claws. These structures are figured for the
relatively few Santa Cruz Island species.
Macrobiotus C. A. S. Schultze, 1834
Macrobiotus areolatus Murray, 1907 (Fig. 3). The specimens from
mainland California that were considered to be M. areolatus were
determined on the basis of eggs which were typical of that species. The
macroplacoids of these specimens were somewhat atypical, the second
being distinctly shorter than the first and third. The Santa Cruz Island
population has placoids of the same proportion, but differ slightly in
the location of the stylet supports which are often attached at a distance
from the base equal or greater to the mouth tube diameter. The eggs
of the Santa Cruz population are of the M. areolatus form although
the processes are somewhat shorter and more conical than those of
the mainland population. Most of the specimens have eyes and the
microplacoids are absent, typical of M. areolatus.
Specimens were found in samples 11F, 11G, 14, 15, and 16. The
only identified host was the hepatic, Porella navicularis , and the
remaining samples were unidentified moss.
Macrobiotus harmsworthi Murray, 1907 (Fig. 4). Macrobiotus
harmsworthi is easily distinguished from the other Santa Cruz species by
the wide mouth tube, the three subequal macroplacoids and the
presence of a microplacoid. Specimens segregated by this combination
of characters can be further separated into two distinct forms. One
form has no eyespots and the claws are typically Y-shaped with the
branches of each claw united in the basal one-half. This was the form
most frequently found, the one exception being at Coches Prietos.
The Coches Prietos specimens have eyespots and the macroplacoids
<-
Fig. 1 . Batillipes ? similis, cephalic appendages. Fig. 2. Batillipes ? similis,
leg IV. Fig. 3. Macrobiotus areolatus, mouthparts. Fig. 4. Macrobiotus
harmsworthi, mouthparts. Fig. 5. Macrobiotus hufelandii, mouthparts. Fig. 6.
Macrobiotus hufelandii, claws of leg IV. Fig. 7. Macrobiotus dispar, mouth¬
parts. Fig. 8. Macrobiotus dispar, claws of leg IV. Fig. 9. Macrobiotus
intermedius, mouthparts. Fig. 10. Hypsibius dujardini, mouthparts. Fig. 11.
Hypsibius dujardini, claws of leg IV. Fig. 12. Hypsibius convergens, mouth¬
parts. Fig. 13. Hypsibius convergens, claws of leg IV. Fig. 14. Hypsibius
canadensis, mouthparts. Fig. 15. Hypsibius canadensis, claws of leg IV. Fig.
16. Hypsibius oberhaeuseri, claws of leg IV. Fig. 17. Hypsibius oberhaeuseri,
mouthparts.
192
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
tend to be thicker. The claws are V-shaped, and the branches diverge
from less than 50% of the claw length. This form is present also on
San Nicholas Island and at Camarillo, Ventura Co. M. harmsworthi
was present in samples IB, 1C, 3D, 3E, 5, 6B, 8, 13A, and 13B. The
species was moderately common and no host preference was apparent
among terrestrial plants.
Macrobiotus hufelandii C. A. S., Schultze, 1834 (Figs. 5, 6). A wide
mouth tube, two macroplacoids of unequal length, and the presence of
a microplacoid characterize M. hufelandii. Our determination is based
on both adults and eggs. M. hufelandii was one of the most frequently
recovered species, and occurred in about the same number of samples
as Echiniscus arctomys and Milnesium tardigradum.
Specimens recovered from Selaginella at Cascada and at Coches
Prietos lack eyes, the first macroplacoid is constricted, and the secondary
branch of the claw is frequently reduced and more distal. The remaining
specimens have eyes, the first macroplacoid is somewhat shorter and
not constricted and the claws are of the typical M. hufelandii form.
Specimens were collected at 1A, 1C, ID, IE, 2B, 2D, 3D, 3E, 3F, 3G,
6A, 11G, 111, and 12A.
Macrobiotus intermedius Plate, 1888 (Fig. 9). This species is
represented in our collections by few specimens and no eggs. None
of the characters requisite for positive identification (the egg, or
granulation and punctation of the cuticle) are available in our material.
However, the general facies—small size, eyes, structure and size of
mouthparts—provides sufficient information for reasonably certain
identification. The specimens were recovered at Pelican Bay from
samples IB and 1C.
Macrobiotus dispar Murray, 1907 (Figs. 7, 8). This freshwater
species has been recovered from algae and Azola sp. in the canyon
across from the field station, and it could be expected to occur in the
other small streams. However, it was not found in numerous samples
from Cascada and Canada del Puerto. We considered this species to be
M. macronyx in our 1965 study, a species characterized by its eggs
being laid within the exuvia. Ramazzotti (1967) states that he has
never seen M. macronyx and doubts the existence of the species. We
have never seen eggs deposited within the exuvia, and now believe the
name dispar should be applied to the Californian populations. This
determination is based on the generally favorable agreement of our
specimens with the descriptive information on M. dispar , and partic¬
ularly on the presence of a chitinous band connecting the two claws
of each leg. Specimens were present in samples 11A, 11B and 11H.
JULY 1970 ] SCHUSTER & GRIGARICK-SCI TARDIGRADA
193
Hypsibius Ehrenberg, 1848
Four species of Hypsibius were identified from our collections. Also,
an apparently undescribed species similar to H. ( C .) castri was found
at Coches Prietos. Additional examples of this species will be necessary
to confirm the relationship.
Hypsibius ( Hypsibius ) oberhaeuseri (Doyere, 1840) (Figs. 16, 17).
The specimens of this species are less granular and less pigmented
than those from most mainland collections. However, the color and
sculpture of the cuticle are known to be quite variable and the mouth-
parts and claws are distinctly those of oberhaeuseri. Collections were
from 1C, 3A, 3D, 11G, and 13B.
Hypsibius ( Hypsibius ) convergens (Urbanowicz, 1925) (Figs. 12,
13). Hypsibius convergens is distinguished from H. oberhaeuseri by
the principal branch of the external double claw, which for H. convergens
gradually tapers from the base and appears thicker. Also, the placoids
are thinner. It was collected at 11G and 12C.
Hypsibius ( Hypsibius ) dujardini (Doyere, 1840) (Figs. 10, 11).
This is essentially an aquatic species. It is best recognized by the
elongate macroplacoids and the presence of a microplacoid. A single
specimen was recovered from collection 11 A.
Hypsibius ( Isohypsibius ) canadensis Murray, 1910 (Figs. 14, 15).
The three macroplacoids, which increase slightly in length from first
to third, and the absence of a microplacoid, distinguish this species.
It was present in samples 6A and HE.
Milnesiidae Ramazzotti, 1962
Milnesium Doyere, 1840
Milnesium tardigradum Doyere, 1840. The very wide mouth tube and
the absence of placoids allow easy recognition of this species. It was
recovered more frequently than any other species and was present in
samples 1A, IB, ID, 2C, 3G, 5A, 9, 10, 11E, 11F, 12A, 12B, 12C,
14A, and 15.
Literature Cited
Schuster, R. 0. and A. A. Grigarick. 1965. Tardigrada from Western North
America with Emphasis on the Fauna of California. Univ. Calif. Publ.
Zool., 76: 1-67.
Ramazzotti, G. 1967. Note per una revisione della sistematica dei Tardigradi.
Mem. 1st. Ital. Idrobiol., 21: 117-128.
194
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
A List of the Odonata of Washington with Additions to and
Deletions from the State List
Dennis R. Paulson
Department of Zoology, University of Washington, Seattle
The Odonata of Washington, like those of most of the western States,
have been little studied. Kennedy (1915) surveyed the odonates of the
Yakima River Valley, but no other section of the State has been sampled
adequately. The Seattle-Olympia region received much attention from
early entomologists, and probably most of the species occurring in that
region have been reported in scattered publications. My field work in
the State has extended thus far from September 1967 to June 1969,
mostly east of the Cascades. The present list was based on literature
records, specimens in the University of Washington Burke Memorial
Museum (UW), and specimens in my collection (DRP), many of
which have been deposited in the Florida State Collection of Arthropods
(FSC). I would like to thank my wife Mary Lynn, for her help in the
field and laboratory; Donald P. Frechin, who turned over to me
Odonata collected by him in western Washington; Melville H. Hatch,
who allowed me to examine specimens in the Burke Museum; and
Minter J. Westfall, Jr., who assisted me in searching the literature for
records of Washington Odonata. Much of my field work in Washing¬
ton was made possible by National Science Foundation Grant GB-7361
to Gordon H. Orians, to whom I am grateful.
Additions
Coenagrion resolutum Hagen. —Stevens Co., Coffin Lake, 3.8 mi. S Middle-
port, 3300 ft., 8 July 1968 (3$ 3$ DRP, 1$ FSC) ; Pend Oreille Co., Niles Lake,
5.8 mi. SW Tiger, 3400 ft., 8 July 1968 (5$ 1$ DRP, 4$ FSC) ; slough at Usk,
2200 ft., 9 July 1968 (2$ 12 DRP); Chelan Co., pond 1.2 mi. E Stevens Pass,
3700 ft., 7 August 1968 (3A 1 $ DRP, 2$ FSC).
Nehalennia Irene Hagen. —Okanogan Co., Black Pine Lake, 4200 ft., 18 June
1969 (10$ 5 2 DRP, 10$ 5 2 FSC); Chelan Co., Soda Springs, 25 June 1939
(12 UW) ; Pend Oreille Co., Niles Lake, 5.8 mi. SW Tiger, 3400 ft., 8 July
1968 (5$ 42 DRP, 4$ FSC).
Aeshna constricta Say. —Spokane Co., Turnbull National Wildlife Refuge,
19 September 1967 (12 DRP); 9 July-28 September 1968 (18$ 12 2 DRP,
11$ 5 2 FSC); also many exuviae. Previous records of this species (Calvert,
1901-08) from the State were referred to A. palmata and A. umbrosa by Walker
(1912).
Aeshna eremita Scudder.— Pend Oreille Co., Niles Lake, 5.8 mi. SW Tiger,
3400 ft., 8 July 1968 (3 exuviae DRP). Adults of a large Aeshna, presumably
this species, were seen at this locality.
The Pan-Pacific Entomologist 46: 194^198. July 1970
JULY 1970 ]
PAULSON—ODONATA OF WASHINGTON
195
Aeshna juncea Linnaeus.— King Co., pond 5.5 mi. W Stevens Pass, 2200
ft., 23 September 1967 (1$ DRP) ; Chelan Co., pond 1.2 mi. E Stevens Pass, 3700
ft., 7 August 1968 (2 $ DRP, 1$ FSC).
Aeshna tuberculifera Walker.— Mason Co., “oak patch,” 23-29 August 1959
(2$ 22 DRP, 1$ FSC).
Cordulegaster dorsalis Selys.—-“Western Washington” (l $ 1? UW).
Somatochlora minor Calverts— Stevens Co., Hande Creek, 1.0 mi. S and 0.6
mi. W Middleport, 3300 ft., 8 July 1968 (1$ DRP).
Somatochlora walshi Scudder.— Stevens Co., Hande Creek, 1.0 mi. S and 0.6
mi. W Middleport, 3300 ft., 8 July 1968 (1 2 DRP).
Leucorrhinia glacialis Hagen.— King Co., Redmond, 17 July 1966 (12 DRP).
Sympetrum danae Sulzer.— Spokane Co., Turnbull National Wildlife Refuge, 19
September 1967 (1 # DRP); 2-29 September 1968 (10$ 4 2 DRP, 8 $ FSC).
Trapezostigma laceratum Hagen.— Spokane Co., Turnbull National Wildlife
Refuge, 21 July 1968 (Id DRP) ; Grant Co., Columbia National Wildlife Refuge,
19 September 1967 (Id DRP); 5-7 July 1968 (Id 12 DRP). Emerged in
large numbers from Coot Lake on Columbia Refuge.
Of these species, Coenagrion resolution, Aeshna constricta, Cordule¬
gaster dorsalis, Somatochlora minor, Leucorrhinia glacialis, and
Sympetrum danae were to be expected in the State because of their
known distribution in the West. Aeshna eremita and A. juncea range
south in the Rockies to Utah and/or Colorado (Needham and Westfall,
1955; Kormondy, 1960) but were not known from the Pacific States.
Nehalennia irene and Somatochlora walshi occur entirely across south¬
ern Canada but have not been reported from the western United States.
Aeshna tuberculifera was known in the West from an isolated population
on Vancouver Island (Walker, 1958), and the present records indicate
it may be more widespread in this region. Trapezostigma laceratum
is the only southern species involved in the above list; it was known
previously as far north as California, Nevada and Utah (Needham and
Westfall, 1955).
Deletions
Archilestes grandis Rambur.—Calvert (1899) and Muttkowski (1910) listed
this species for Washington before Kennedy (1915) clarified the relationship
between it and A. californica McLachlan, to which all Washington records of
the genus should be referred.
Amphiagrion saucium Burmeister.-—All early records of A. abbreviation Selys
were listed under this name; A. saucium is eastern in distribution and does not
occur in the State.
Cordulegaster erronea Hagen.—This eastern species, recorded from Washing¬
ton by Fraser (1929), is very unlikely to occur in the State. Probably Fraser’s
specimens were C. dorsalis.
Lepthemis simplicicollis Say.—This species, recorded widely from the West
by Needham and Westfall (1955), does not occur west of the Great Plains, and
all records of it from this region should be referred to L. collocata Hagen. The
196
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
confusion between these two species was terminated by Gloyd (1958), and all
western specimens of this genus checked since that time by the author, L. K.
Gloyd, and M. J. Westfall, Jr., have proven to be L. collocata.
Sympetrum semicinctum Say.—Early records of this species from Washington
(Ris, 1909-1919; Kennedy, 1915) refer to S. occidentale Bartenef (Walker, 1951).
Washington List
The following list includes all species known to occur in Washington.
A “W” (west) or “E” (east) indicates species which occur on only one
side of the Cascades. Species known from only the boreal areas north
of Spokane or in the Cascades are indicated by an “S” or “C” respec¬
tively.
Calopterygidae : Calopteryx aequabilis califarnica (W) ; C. a. yakima (E).
Lestidae: Archilestes californica (E), Lestes congener, L. disjunctus disjunctus,
L. dry as, L. unguiculatus.
Coenagrionidae: Amphiagrion abbreviatum, Argia emma (E), A. vivida (E),
Coenagrion resolutum (S, C), Enallagma boreale, E. carunculatum, E. clausum
(E), E. cyathigerum, E. ebrium (E), Ischnura cervula, I. erratica (W), I.
perparva, Nehalennia Irene (S, C).
Petaluridae: Tanypteryx hageni (C).
Aeshnidae: Aeshna californica, A. canadensis (S), A. constricta (E), A. eremita
(S), A. interrupta interna, A. juncea (G), A. multicolor, A. palmata, A. tuber-
culifera (W), A. umbrosa occidentalis, Anax funius.
Gomphidae: Gomphus graslinellus (S), G. kurilis confraternus (W), Octogomphus
specularis (W), Ophiogomphus occidentis, O. severus (E), Stylurus sp. (prob¬
ably olivaceus) (E).
Cordulegastridae : Cordulegaster dorsalis (W).
Macromiidae: Macromia magnifica.
Corduliidae: Cordulia shurtleffi (S, C), Epitheca canis, E. spinigera, Somatochlora
albicincta (C), S. minor (S), S. semicircularis (S, C), S. walshi (S).
Libellulidae : Ladona julia, Lepthemis collocata, Leucorrhinia glacialis (W),
L. hudsonica, L. intacta, L. proxima (S, C), Libellula forensis, L. nodisticta
(known only from “Washington”), E. pulchella, L. quadrimaculata, Pachydiplax
longipennis, Plathemis lydia, Sympetrum corruptum, S. costiferum, S. danae,
S. illotum (W), S. internum (E), S. madidum, S. obtrusum, S. occidentale
occidentale, S. pallipes, S. vicinum, and Trapezostigma laceratum (E).
Discussion
Of the 69 species now known from the State, 13 occur only in montane
regions, of which three are restricted to the Cascades and five to the
mountains north of Spokane. Additional collecting may reveal that
these upland areas share more species than is presently known. Of
55 lowland species, 38 occur on both sides of the Cascades, seven
are known from only the west side, and ten are known from only the
east side. Thus 69% of the species are found on both sides of the major
JULY 1970 ] PAULSON-ODONATA OF WASHINGTON
197
biogeographic barrier in the State. With more field work this figure
may be raised, as only a few areas in the eastern part and none in the
western part of the State have been working intensively. This percentage
is higher than those of terrestrial vertebrates, both flying (59% of 138
land birds; 56% of 16 bats) and flightless (50% of 66 mammals, not
including bats; 33% of 21 reptiles; 22% of 18 amphibians).
Two factors are important in contributing to the widespread distribu¬
tion of odonate species in this region and in general. First, odonates,
especially anisopterans, have strong powers of flight, and many species
could fly across the Cascade mountain passes. Weaker-flying species
could be transported passively by the strong winds through these same
passes. Second, dragonflies are aquatic for much of their life history,
and when they attain sexual maturity the adults respond again to
aquatic habitats, which may be similar in regions of very different
terrestrial ecology. To an Aeshna, a shallow cattail-bordered pond in a
sagebrush desert may be equivalent to a similar pond in a humid
coniferous forest, yet few terrestrial organisms would inhabit both
of these environments. In this group, as in others, combinations of
historical and ecological factors have produced present-day bio-
geographies, and the differences in environments on either side of
physical barriers may be more important than the barriers themselves
in determining the respective biotas.
Literature Cited
Calvert, P. P. 1899. Odonata from Tepic, Mexico, with supplementary notes
on those of Baja California. Proc. Calif. Acad. Sci. (3) Zool. 1:
371-418.
1901-1908. Biologia Centrali-Americana, Vol. 50, Neuroptera. Porter &
Dulau, London, 420 p.
Fraser, F. C. 1929. A revision of the Fissilahioidae (Cordulegasteridae,
Petaliidae and Petaluridae) (Order Odonata). Part I.—Cordulegasteri¬
dae. Mem. Indian Mus., 9: 69-167.
Gloyd, L. K. 1958. The dragonfly fauna of the Big Bend region of Trans-Pecos
Texas. Occas. Pap. Mus. Zool. Univ. Mich., No. 593: 1-23.
Kennedy, C. H. 1915. Notes on the life history and ecology of the dragonflies
(Odonata) of Washington and Oregon. Proc. U. S. Nat. Mus., 49:
259-345.
Kormondy, E. J. 1960. New North American records of anisopterous Odonata.
Entomol. News, 71: 121-130.
Muttkowski, R. A. 1910. Catalogue of the Odonata of North America. Bull.
Public Mus. Milwaukee, 1: 1-207.
Needham, J. G., and M. J. Westfall, Jr. 1955. A Manual of the Dragonflies
of North America (Anisoptera). Univ. Calif. Press, Berkeley, 615 p.
Ris, F. 1909-1919. Libellulines. Cat. Systematique et Descriptif des Coll. Zool.
Edm. de Selys Longchamps, Fasc. 9-16, 1278 p.
198
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO.
3
Walker, E. M. 1912. The North American Dragonflies of the genus Aeshna.
Univ. Toronto Stud., Biol. Ser., No. 11: 1-213.
1951. SympetTum semicinctum (Say) and its nearest allies (Odonata).
Entomol. News, 62: 153-163.
1958. The Odonata of Canada and Alaska. Volume II. Part III. The
Anisoptera of Canada and Alaska. Univ. Toronto Press, Toronto, 318 p.
Notes on the Saprophagous Activity of Bufolucilia silvarum
(Meigen)
(Diptera: Calliphoridae)
Donald. R. Brothers
San Jose City Health Department, San Jose, California 1
Bufolucilia silvarum (Meigen) is known to be a parasite of toads and
frogs. Hall (1948) regarded the species as an obligatory parasite.
However, more recent literature suggests that B. silvarum may be a
saprophage and only a facultative parasite. Fly population data
obtained by Dicke and Eastwood (1952) and Schoof, Savage, and
Dodge (1956) show a substantially high population, much greater than
could be supported by parasitism alone. Dodge (1952) reported the
most significant evidence that B. silvarum may be a saprophage by
rearing this species from a rat carcass which had egg masses in its
fur at the time it was shot. To my knowledge, no other observations
supporting saprophagous behavior have been reported. The following
notes are based on observations I made during the spring of 1968
while studying fly populations in Santa Clara County, California.
Observations and Results.— Observations were made on a levee
separating a commercial salt flat and a tidewater slough at the south
end of the San Francisco Bay, one mile northeast of Alviso, Santa
Clara County. Commercial salt flats, tidal marsh, and a tidewater
slough occupy a one mile radius around the site.
On 24 May 1968, a fresh carcass of a young duck was found on
the slough side of the levee. The carcass was matted with numerous
egg masses around a mangled neck and other parts of the body. While
observing the flies attracted to the carcass, two B. silvarum were ob¬
served near it. On the premise that some of the eggs may be those of
B. silvarum, the duck carcass was taken for further study.
A few eggs were taken from each of five unhatched egg masses that
^Present address: 2610 Greenwood Circle, Boise, Idaho 83706.
The Pan-Pacific Entomologist 46: 198-200. July 1970
JULY 1970] BROTHERS-BUFOLUCILIA SILVARUM
199
Fig. 1. Position of Bufolucilia silvarum egg masses on the carcass of a young
duck found on a levee, one mile northeast of Alviso, Santa Clara County. Fig. 2.
Bufolucilia silvarum egg mass at A on Fig. 1 . Fig. 3. Bufolucilia silvarum egg
mass at B on Fig. 1.
were found on the duck carcass and placed separately on lean beef in
cotton-plugged test tubes. All eggs hatched, and the larvae developed
normally. As the larvae matured, crumpled paper toweling was placed
in the tubes to serve as pupation sites. After pupation the puparia were
placed in test tubes containing paper strips for use by the emerging
adults. Two of the five egg samples developed into B. silvarum adults.
The remainder were Phaenicia sericata (Meigen).
After the egg samples had been taken, the remainder of the eggs
were allowed to mature on the duck carcass in a gallon container. Seven
hundred and seven adults emerged. Of these, 103 were B. silvarum and
604 were P. sericata.
As a result of finding the duck carcass infested with B. silvarum
in that localized environment, I made additional observations in the
same area.
200
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO.
3
Two fly traps, after the design of Scott and Littig (1962), baited with
beef liver were employed from 30 May to 3 June 1968, to determine
the density of the B. silvarum population in the area. A total of 6,309
flies was taken. Bufolucilia silvarum was represented by 1,078 speci¬
mens, making up 17.1% of the total collection. Other flies taken were
as follows: P. sericata —4,798 or 76%; Calliphora spp.—295 or 4.7%;
and Phormia regina, Sarcophaga spp., and others—138 or 2.2%.
The saline surroundings support very few, if any, anuran populations
for parasitism by this species, however numerous carcasses of gulls,
ducks, shore birds, and fish litter the levees. Adults of B. silvarum
were found on all of these types of animal carcasses. While observing
a dead gull for a short time on 30 May, I collected ten adult female
B. silvarum that landed on it.
Conclusion.— The presence of a high population density of B. silva¬
rum in an environment with insufficient live anuran hosts to parasitize
led to further observations. The large number of animal carcasses avail¬
able, the attraction of these flies to them, and the rearing of this species
from a fly-blown duck carcass leads to the conclusion that B. silvarum
is primarily a saprophage in this area.
Bufolucilia silvarum adults collected during this study are deposited at
San Jose State College, Washington State University, and the California
Academy of Sciences in San Francisco.
Acknowledgments.— Special thanks are due to Mr. Tibor Banathy
of San Jose, who found the duck carcass and brought it to my attention.
I thank Dr. J. Gordon Edwards of San Jose State College and Dr.
Maurice T. James of Washington State University for critically reading
the manuscript.
Literature Cited
Dicke, R. J., and J. P. Eastwood. 1952. The seasonal incidence of blow flies
at Madison, Wisconsin (Diptera: Calliphoridae). Wis. Acad. Sci.,
Arts, Lett., 41: 207-217.
Dodge, H. R. 1952. A possible case of blowfly myiasis in a rat, with notes on
the bionomics of Bufolucilia silvarum (Mg.). Entomol News, 63:
212-214.
Hall, D. G. 1948. The blowflies of North America. Thomas Say Foundation,
Monograph IV. 477 pp.
Schoof, H. F., E. P. Savage, and H. R. Dodge. 1956. Comparative studies of
urban fly populations in Arizona, Kansas, Michigan, New York, and
West Virginia. Ann. Entomol. Soc. Amer., 49: 59-66.
Scott, H. G., and K. S. Littig. 1962. Flies of public health importance and
their control. U. S. P. H. S. Publication 772, Part 5. U. S. Govt. Pr.
Off. 40 pp.
JULY 1970]
BOHART—NEW BEMBICINI
201
New Species, Synonymy and Lectotype Designations in
North American Bembicini
(Hymenoptera: Sphecidae)
R. M. Bohart
University of California, Davis, California
During a survey of the bembicid fauna of California, several new
species have come to light and these are described below. Holotypes are
in the Museum at Davis and in the California Academy of Sciences. The
opportunity is taken also to establish lectotypes and point out synonymy
for some North American Bembicini. Remarks on the Handlirsch types
were made possible through the kindness of Dr. Claude Besuchet of the
Museum d’Histoire Naturelle in Geneva, Switzerland; and by Dr. Max
Fischer of the Natural History Museum in Vienna, Austria.
Bembix frommeri R. Bohart, new species
Male holotype.— Length 17 mm. Black, marked with whitish and pale yellow;
principal whitish areas on mandibles, clypeus, foretibia, foretarsus, scutellum,
metanotum and tergites. Pale markings: face except mandible tip, quadrate
frontal spot, and band across vertex connected with back of head; scape and
flagellum beneath; outer orbital stripe widened toward mandible; pronotum;
pleuron except for spots on mesopleuron; scutum laterally and 2 small discal
stripes anteriorly; posterior scutellar band and metanotum; posterior half of
propodeal enclosure followed by black band, then practically all yellow; legs
mostly, but with black stripes behind femora and small spots on tibiae; broad
apical tergal bands anteriorly tri-emarginate, 3 separate spots on V, double
median spot on VI and apical half of VII; sternite I except for 2 black spots, II
medially and basolaterally, III-V with lateral spots, VII-VIII apically. Wings
clear, veins mostly brown. Pubescence mostly pale, rather long and shaggy on
head.
Labrum a little shorter than eye height, slightly humped at basal third in
profile; mandible with strong denticle, scape two-thirds as long as clypeus,
midocellus a narrow but translucent lunule, flagellum with shiny tyloides beneath
articles II and following, last article with welt-like tylus on basal half; front
basitarsus with 8 rake setae; midtibia irregular beneath and with few minute
denticles toward apex; first intersubmarginal vein of forewing nearly straight;
tergite VII longitudinally wrinkled, distinctly angled out basolaterally, spiracular
lobe slender and reaching only as far as basolateral angle; sternites II and VI
without carinae or projections; gonostyle tapering gradually toward apex, covered
with rather fine hair; digitus slender, widening gradually to truncate and slightly
emarginate apex; cuspis slender, finger-like.
Holotype male (CAS), 5 mi. south of Deep Springs College,
Inyo County, California, 11 July 1967 (Saul and Suzy Frommer).
The Pan-Pacific Entomologist 46: 201-207. July 1970
202
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Paratype male, Wyman Canyon, Inyo Co., California, 25 June 1966
(Saul and Suzy Frommer, UCR).
A close relationship is indicated with B. dentilabris Handlirsch in
which the male also has an irregular clypeal profile, nearly straight
first intersubmarginal vein, wrinkled and basally angled tergite VII, and
simple sternites II and VI. The 2 species differ in markings, B.
dentilabris being yellow and having a u-shaped scutal mark. In B.
frommeri the markings are extensively whitish and the scutum is nearly
all dark discally. More significant are a few structural characters.
In B. frommeri the tylus beneath the last flagellomere is a shiny
welt but is confined to the basal half rather than tapering toward the
apex of the article. Also, the midfemur is sparingly spiculate beneath.
The genitalia show critical differences. In B. frommeri the gonostyle is
large, conical and finely haired, rather than ligulate and spinose apically.
Furthermore, the cuspis is narrowly finger-like rather than blade-like.
The species is named for Saul Frommer who has made many interest¬
ing collections of bembicids in California.
Microbembex californica R. Bohart, new species
Male holotype.— Length 11 mm. Black, extensively marked with yellow as
follows: lateral strip on labrum, pronotal band all across, lateral spot over wing
base, lateral third of scutellum, metanotum, broad band across propodeum,
lateral propodeal angles, irregular upper mesopleural spot, legs partly, foretibia
basally and posteriorly, midtibia except ventrally, hindtibia entirely, fore basitarsus
posteriorly, mid and hind tarsi mostly, tergites except narrow basal bands and 2
black dots at summit of I, arcuate band across sternite II, lateral spots on III-IV.
Wings lightly stained, veins light brown to black, basal half of forewing costa pale,
veins of first submarginal cell brown except narrowly at base. Pubescence short,
pale, mostly inconspicuous; moderately silvery on face, long and erect on frons
and propodeal angles. Punctation close, scutum and mesopleuron with punctures
separated by 1 diameter or less. Last 7 flagellar articles polished beneath; fore
basitarsus 3 times as long as broad; hindfemur regular in outline; median projec¬
tion of sternite II moderately large, keel-like, outline rounded at summit, then
nearly flat to an acute posterior point; tergite VII incised apically, incision about
half as broad as length of third hindtarsal article; sternite VIII curving gently,
bluntly spear-shaped. Digitus with broadly rounded apical part not projecting
backwards.
Female. —Length 8 to 13 mm. Black, marked with white or occasionally pale
yellow as follows: mandible and clypeus partly, labrum laterally or mostly, outer
orbits narrowly, pronotum laterally, rest of notum sparingly, spot on upper meso¬
pleuron usually, legs as in male, moderately narrow tergal bands on I-V, lateral
spots on sternites II—III. Wings as in male but usually darker. Pubescence as in
male except on mesopleuron where appressed silvery hair partially obscures puncta¬
tion. Tergite VI narrowly and sharply emarginate at apex.
JULY 1970]
BOHART-NEW BEMBICINI
203
Holotype male (UCD), Davis, Yolo County, California, 5 August
1964, in dry sand creekbed (R. M. Bohart). Paratypes, 29 males, 62
females (all from central California) : Davis, Sacramento, and Antioch.
Paratype dates are May to September. Additional specimens have been
examined from many other localities in California, and from other
western states east to Wyoming (Sweetwater Co.), Utah (Cornish), and
New Mexico (Embudo). It occurs also in Baja California Del Norte,
Mexico.
The species has previously been confused with M. monodonta (Say)
which occurs east of the Continental Divide. Females seem to be in¬
distinguishable but males of M. californica have the antenna shiny
beneath toward the apex rather than dull, the sternal keel nearly always
with a sloping flat top rather than evenly rounded, and the digitus with
the broadly rounded apical part not projecting backwards.
Microbembex argyropleura R. Bohart, new species
Male holotype.— Length 10 mm. Black, extensively marked with light yellow
as follows: pronotal band all across, broad lateral scutal stripe, pair of discal
scutal dots, scutellum and metanotum mostly, hand across summit and other small
dots on propodeum; legs partly and especially on hindleg, tergites mostly, and
lateral dots on sternites II—III. Abdominal venter mostly reddish brown; wing
membrane clear, costa mostly and most other veins basally white; first sub¬
marginal cell with radius white and other veins yellowish to light brown.
Pubescence silvery, moderately prominent on propodeum, mesopleuron, scutum
and frons. Punctation close, scutum and mesopleuron with punctures separated
by 1 diameter or less. Last 4 flagellar articles rather dull beneath, not polished;
fore basitarsus 4 times as long as broad; hindfemur regular in outline; median
projection of sternite II small, acute, keel-like, evenly curved; tergite VII incised
apically, incision about half as broad as length of third hindtarsal article; sternite
VIII curving gently, bluntly spear-shaped. Digitus with bluntly rounded apical
part not projecting backwards.
Female.— Length 9 to 12 mm. Black, marked with whitish yellow as follows:
mandible, labrum and clypeus mostly, thin lines along inner and outer orbits,
scape partly, thorax about as in male but scutum usually dark discally, tergites
with broad bands which are edged basally with dark brown or reddish brown,
tergite VI banded or with 2 spots, sternites with small lateral spots on II-TV or
II-V imposed on reddish brown background. Wings as in male. Silvery pubescence
forming dense coat on mesopleuron, scutum and frons. Tergite VI narrowly and
sharply emarginate at apex.
Holotype male (UCD), 18 mi. west of Blythe, Riverside County,
California, 16 October 1965 (R. M. Bohart). Paratypes, 43 males, 37
females (all from southern California) : near Blythe, Borrego Valley
(San Diego Co.) Cronise Valley (San Bernardino Co.), Palo Verde
(Imperial Co.) and 13 mi. east of Ocotillo Wells (Imperial Co.).
204
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Paratype dates are April to October. I have seen material also from
California (Heber, 1000 Palms Oasis, Palm Springs, Olancha, Antelope
Springs in Inyo Co.), Arizona (Yuma, Toltec, Eloy, Grand Canyon
floor, Willow Ranch in Mojave Co.), Nevada (Nixon, Pyramid Lake,
Wadsworth), Utah (12 mi. south of Eureka), and Baja California,
Mexico (28 mi. south of El Arco).
The species has previously been confused with M. hirsuta J. Parker
which occurs in New Mexico and Texas. I am unable to separate
females of the 2 species but males offer no problems. In M. argyropleura
males the sternites have short and rather inconspicuous pubescence,
whereas in M. hirsuta there is a dense erect pile which extends in
gradually increasing length from the sternal keel onto sternite VII. A
superficial resemblance exists between males of M. argyropleura and
M. californica. The white radius on the first submarginal cell and the
narrower fore basitarsus of M. argyropleura are distinguishing. The
silvery mesopleural mat of female M. argyropleura is distinctive among
Californian species.
Variation in color pattern occurs as would be expected. However,
the face of the male is always entirely black, contrary to the case in
M. californica. The scutum is usually black discally, the scutellum and
propodeum may each have 2 large spots rather than bands, and the
abdominal ground color varies from nearly black to brownish red,
especially in females.
Microbembex rufiventris R. Bohart, new species
Male holotype.— Length 13 mm. Black, extensively marked with yellow as
follows: pronotal lobes, spot over wing base, large scutellar spots, metanotal band
and one across summit of propodeum, legs partly but foreleg nearly all black,
tergites mostly, small lateral spots on sternites II-V; ground color of sternites
reddish brown. Wings nearly clear, forewing costa pale to stigma, veins of first
submarginal cell brown, a little yellowish at base of cell. Pubescence short, pale,
mostly inconspicuous; moderately silvery on face, long and erect on frons and
propodeal angles. Punctation close, scutum and mesopleuron with punctures
separated by 1 diameter or less. Last 4 flagellar articles not polished beneath; fore
basitarsus 3.5 times as long as broad; hindfemur regular in outline; median projec¬
tion of sternite II a raised carina, sharp posteriorly; tergite VII shallowly
emarginate apically, incision about as broad as length of third hindtarsal article;
sternite VIII curving gently, bluntly spear-shaped. Digitus stout, broadly rounded
apical part not projecting backwards.
Female.— Length 9 to 13 mm. Black, marked with greyish white as follows:
labrum and mandibles mostly, clypeus broadly, scape partly, narrow orbital
streaks, pronotal lobes, spot over wing base, metanotal spots, legs partly, narrow
apical tergal bands, broadened laterally, lateral dots on pygidium; sternites as in
male but redder; ground color of tergites mottled with reddish. Wings as in male
JULY 1970]
BOHART-NEW BEMBICINI
205
but a little darker. Punctation and pubescence as in male. Tergite VI narrowly
and sharply emarginate at apex.
Holotype male (UCD), Coalinga, Fresno County, California, 22
May 1936 (R. M. and G. E. Bohart). Paratypes, 1 male and 2 females,
same data as type; 1 male, Cawelo Junction, east of Shafter, Kern Co.,
California, 24 July 1952 (T. R. Haig, UCD).
There is considerable resemblance to darker forms of M. californica.
However, the male of M. rufiventris has the undersurfaces of the last
4 flagellomeres rather dull, the sternal keel cariniform, and tergite VII
much broader at the apex. The female has dorsal markings greatly
reduced. In the few females of M. californica which approach this con¬
dition the face is extensively dark in contrast to M. rufiventris. It
remains to be seen whether this difference in the females will hold up
when more specimens of M. rufiventris are collected.
Genus Bembix
Bembix amoena Handlirsch, 1893. The lectotype male here designated
is in the Vienna Museum, “Nevada.”
Genus Bicyrtes
Bicyrtes annulata J. Parker, 1917. The holotype female is in the
collection of the University of Kansas at Lawrence, “Oak Creek Canon,
Ariz. 6000 ft.” This is a synonym of B. capnoptera (Handlirsch), 1889.
It represents the more yellowish western variety.
Bembidula capnoptera Handlirsch, 1889. The lectotype female here
designated is in the Vienna Museum, “Kentucky.”
Bembidula capnoptera mesillensis Cockerell, 1898. The holotype male
is in the U. S. National Museum, “Las Cruces, N.M.” It is a synonym of
Bicrytes capnoptera (Handlirsch), 1889.
Bembidula diodonta Handlirsch, 1889. The holotype is a male in the
Geneva Museum, “Mexiq. Orizaba.” This is a distinctive species of
Bicyrtes related to B. discisa (Taschenberg) but with serrate midfemur,
broader foretarsus and dentate sixth sternite.
Bembidula fodiens Handlirsch, 1889. The lectotype male here
designated is in the Vienna Museum, “Dallas, Texas.”
Bicyrtes gracilis J. Parker, 1917. The holotype male is in the col¬
lection of the University of Kansas, Lawrence, “Santa Rita Mts., Ariz.,
5-8000 ft.” This is a synonym of B. viduata (Handlirsch), 1889.
Bembidula insidiatrix Handlirsch, 1889. The lectotype here desig¬
nated is a female in the Vienna Museum, “Kentucky.”
206
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Bembidula odontophora Handlirsch, 1889. The lectotype here desig¬
nated is a male in the Vienna Museum, Nauta, “E. Peru.”
Bicyrtes oribates Pate, 1936. The holotype is a male in the Cornell
University collection. “Compostela, Nayarit, Mexico.” This is a
synonym of B. diodonta (Handlirsch), 1889.
Bicyrtes parata Provancher, 1889. The holotype female is in the
collection of Laval University, St. Foy, Quebec, “Los Angeles (Coquil-
lett).” It is a synonym of B. ventralis (Say), 1824 and represents the
more yellow western variety.
Bicyrtes tristis C. Fox, 1923. The holotype is a male bearing no data
but presumably from La Paz, Baja California, in the California Academy
of Sciences, San Francisco. It seems to be a rather dark specimen of
B. capnoptera Handlirsch.
Bembidula viduata Handlirsch, 1889. The lectotype here designated
is a female in the Geneva Museum, “Huastec,” Mexico. This large
Bicyrtes is fairly abundant in southwestern United States and northern
Mexico.
Genus Microbembex
Microbembex monodonta deltaensis Johnson and Rohwer, 1908. The
lectotype here designated is a male in the U. S. National Museum
labeled “cotype” by Rohwer, “Delta, Col. 7 20 98.” This is a synonym
of M. nigrifrons (Provancher), 1889.
Microbembex monodonta neomexicana Johnson and Rohwer, 1908.
The lectotype here designated is a female in the U. S. National Museum
labeled “cotype” by Rohwer, “Las Cruces, N.M., flo. Solidago, Aug. 30
(Twns.).” This has rather shiny yellow spots on the scutum and is a
synonym of M. nigrifrons (Provancher).
Microbembex monodonta occidentalis Johnson and Rohwer, 1908.
The lectotype here designated is a female in the U. S. National Museum
labeled “cotype” by Rohwer, “Paris Tx XI-26 1904, C. R. Jones col¬
lector.” This is a synonym of M. monodonta (Say), 1824.
Bernbex nigrifrons Provancher, 1889. The lectotype here designated
is a female Microbembex in the collection of Laval University, St. Foy,
Quebec, “Los Angeles (Coquillett).” It has a pair of rather shiny discal
yellow spots on the scutum.
Genus Stictiella
Monedula mammillata Handlirsch, 1890. The lectotype here desig¬
nated is a male in the Geneva Museum, “Georgie.” It is a synonym of
Stictiella emarginata (Cresson), 1865, as previously supposed.
Monedula plana W. Fox, 1895. The holotype is a male in the U. S.
JULY 1970] GOODMAN—MICROBEMBEX CALIFORNICA BEHAVIOR 207
National Museum, “Custer, South Dakota (Aldrich)This is ap¬
parently a synonym of Stictiellci serrata (Handlirsch), 1890, differing
only in having the bands on tergites II—VI complete rather than nar¬
rowly broken medially.
Monedula serrata Handlirsch, 1890. The holotype is a male in the
Geneva Museum, “Georgie.” It is a rather abundantly yellow-marked
Stictiella agreeing with previous interpretation.
Observations on the Behavior and Biology of Microbembex
calif or nica Bohart
(Hymenoptera: Sphecidae)
W. G. Goodman
University of California, Davis
During July 1969, observations were made on a colony of Micro¬
bembex californica R. Bohart in the dunes 5 miles south of Samoa, Hum¬
boldt County, California. M. californica , limited mostly to the Pacific
coastal states, is a general scavenger, taking dead arthropods as “prey”
for nest provisioning. Although no previous work dealing with M.
californica has been recorded, the behavior and biology of the wasp
is very similar to that of an eastern species, Microbembex monodonta
(Say) described by Evans (1966) and Parker (1917).
The nesting site, approximately 6 X 15 meters, was located in a
large blowout free of vegetation and surrounded by plants typical of
the dune ecology: Artemisia pycnocephala, Eriogonum latifolium , and
Haplopappus racemosus. The area was shared with three other species
of wasps: Bernbix americana comata Parker, Oxybelus uniglumis
quadrifasciatus Say, and Philanthus pacificus Cresson.
The female, usually in the early afternoon, begins to dig at a series of
sites, particularly in crusted sand, before choosing the true nest site.
A few females may begin digging in uncrusted sand and may spend
more than an hour digging. The digging action of the female resembles
that of a teeter-totter. Balanced on her middle legs, the female bobs her
head down, scrapes sand loose and kicks it out behind her. After
several scuffs, she bobs back to a normal position and continues to bob
up and down until she has started the burrow. Every few minutes she
The Pan-Pacific Entomologist 46: 207-209. July 1970
208
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
returns to the entrance to scrape away any sand that may have built
up during her excavation of the burrow and to make flights about the
nesting site. The expelled sand is kicked outward approximately 10 cm,
which disperses it and leaves no trace of a mound. It appears that the
nests of M. calif or nica are one-celled, as in the other known North
American species. The burrow, descending at a 30 degree angle from
the surface, is approximately 25 cm in length. The cell is approximately
2.5 cm long and 1.0 cm high and lies 5 to 17 cm below the surface,
depending upon the moisture level. Construction of the nest and the
initial closure takes about 2 hours.
The female lays the egg (in 2 cases examined) before any attempts
at provisioning. Temporary outer closures of the nest during provi¬
sioning are merely a few scuffs of sand thrown behind her into the
entrance, leaving the entrance still visible. An inner closure near the
cell was found in a single burrow. It is highly unlikely that the closure
was an artifact due to my excavation, but possibly the beginning of the
final closure of the burrow. Even this explanation seems questionable
since the female was still attempting to provision the nest after my
excavation had begun. All other burrows were free of an inner closure,
which indicates that normally no inner closures are maintained.
Evidence that M. californica is a general scavenger, provisioning the
nest with dead arthropods, is exhibited by the prey recorded from the
cells. In one nest, representatives of the following families were found:
Cercopidae, Carabidae, Curculionidae, Asilidae, Therevidae, Formicidae,
and a spider. Earwigs were quite commonly used as larval food, and
in every case the appendages had been removed before the female flew
into the nesting area. The body of the earwig seemed flexible as if it
were recently killed. The females appear to hunt most often within 10
meters of the colony in the peripheral areas of the blowout where
vegetation is most abundant. When hunting, the female skims several
cm above the sand, rapidly zig-zagging back and forth across a small
area. When “prey” is discovered the female hovers over it, picks it
up with her forelegs, then rolls it end over end several times as if to
inspect it. She then tucks the carcass under her and flies off. The
simulation of the stinging of prey as observed by Evans (1966) and
Parker (1917) in other Microbembex species was not observed in M.
californica . The female upon entering the nest continues inward with¬
out releasing the carcass, if it is small enough. However, if the nest
entrance is too large to accommodate both, she drops the carcass
at the entrance, enters the burrow and pulls it in with her mandibles.
Females return to the nest with provisions about every 40 minutes.
JULY 1970 ]
HURD-A NEW DIALICTUS
209
Male M. californica , as in other species in the genus, appear to con¬
struct some type of burrow for sleeping. Male activity, other than
burrow construction, is limited to feeding and mating.
Microbembex californica at the Samoa site appeared to be free of
parasites. However, several cocoons contained unidentified nematodes.
Literature Cited
Evans, H. E. 1966. The comparative ethology and evolution of the sand wasps.
Harvard Univ. Press, 526 p.
Parker, J. B. 1917. A revision of the bembicine wasps of America north of
Mexico. Proc. U. S. Nat. Mus., 52: 1-155.
A New, Narrowly Polylectic, Autumnal Species of Dialictus
from the Flowers of Jepsonia heterandra , an endemic
California saxifrage
(Hymenoptera: Apoidea)
Paul D. Hurd, Jr.
University of California , Berkeley
During the autumns of 1967, 1968 and 1969, Professor Robert
Ornduff has intensively investigated the reproductive biology of the
fall-blooming Jepsonia heterandra Eastw. (Saxifragaceae) at two locali¬
ties within its restricted range along the western foothills of the Sierra
Nevada in central California. The results of these studies, which are
being reported elsewhere (Ornduff, 1970), record the discovery and
role of an undescribed species of the genus Dialictus which is involved
in the pollination of this locally endemic and late flowering perennial
(Ornduff, 1969). The bee has not been observed to visit the flowers of
/. heterandra before the anthers dehisce in mid or late morning. At the
two study sites there is a paucity of other plant species in flower when
/. heterandra flowers (late September and October) and repeated sur¬
veillances of these potential pollen and nectar sources (Compositae) have
revealed the presence of only an occasional male or female bee. It is
thus unclear whether these visits represent a temporary and opportunistic
departure from the usually observed pollen and nectar collections source
(Jepsonia ) or if in fact such sources are less attractive than is Jepsonia.
The Pan-Pacific Entomologist 46: 209-212. July 1970
210
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
At the latter source both the males and females are conspicuously abun¬
dant suggesting that the flowers of that plant provide the preferred
nectar and pollen source of the bee. Although repeated attempts to
locate the nests of this bee have been disappointingly unsuccessful, it is
anticipated that their discovery will make it possible to determine the
sources of pollen and their relative amounts by analysis of cell pro¬
visions. Analysis of pollen from the scopae and metasoma of females
collected from the flowers of /. heterandra revealed pollen of Compositae
in only one of the five individuals examined. This individual was taken
at 12:30 p.m. by which time most of the available pollen of Jepsonia
had already been collected (Ornduff, 1970). These observations sug¬
gest that Dialictus ornduffi is a narrowly polylectic species. Its apparent
restriction to Jepsonia heterandra as the primary source of pollen and
nectar and its apparently secondary dependence upon the pollen and
nectar of certain Compositae may simply reflect the relative competitive
attractiveness of the late flowering plants present in the flora super¬
imposed upon the lack in the flora of other potentially competitive
sources of pollen and nectar.
It is with much pleasure that I dedicate this new species to Professor
Robert Ornduff, University of California, Berkeley, who introduced me
to this interesting bee-plant relationship and with whom I have spent
many enjoyable and profitable hours in the field on this and other pol¬
lination projects.
Dialictus ornduffi Hurd, new species
Female.— Length 4-4.5 mm. Head, and thorax, dark bluish-green; metasoma
brownish-piceous without metallic reflections; wings with three submarginal cells,
subhyaline, veins including stigma mainly testaceous though somewhat more brown¬
ish along anterior wing margin; tegulae brownish, not enlarged, rounded behind,
minutely punctured except on posterior third where glabrous, impunctate and
shining; legs piceous except brownish tarsal segments; pubescence short, thin,
chiefly white, somewhat more dense along sides of metasoma and on discs of terga
III-V; head slightly longer than broad, densely and very nearly confluently punc¬
tured except vertexal areas adjacent to ocelli and on lower parocular areas where
more sparse and shallowly punctate; eyes moderately convergent below (33:25) ;
cheeks about as broad as eyes in width as seen in lateral view, finely and reticu-
lately punctate; maximum interorbital width somewhat less than distance from
anterior ocellus to apical margin of clypeus (38:42) ; lateral ocelli much nearer to
eyes than to each other and removed from posterior margin of vertex by more
than their diameters; clypeus piceous, without metallic reflections, broader than
long, slightly convex, projecting about two-thirds below suborbital line, punctures
large, coarse, irregular in outline, separated by less than maximum diameter;
supraclypeal area piceous, faintly metallic, somewhat swollen, much wider than
long, punctures rather coarse and sparse, interspaces finely tessellated; hypostomal
JULY 1970 ]
HURD—-A NEW DIALICTUS
211
carinae parallel, adjacent ventral surfaces of head shining and sparsely punctate;
mesoscutum and scutellum shining, interpunctational spaces tessellate, punctures
fine, deep, well separated medially, especially on scutellum, but becoming quite
close and crowded at extreme lateral sides; pleura coarsely reticulo-punctate an¬
teriorly, more finely rugoso-punctate medially and becoming striato-punctate along
posterior margin; posterior face of propodeum rugosely striato-punctate, very
nearly though incompletely, delimited by lateral and dorsal carinae, carinate
dorsal rim low, weakly defined medially, lateral carinae obsolete dorsally; dorsal
area of propodeum with striations incomplete, not attaining posterior rim, rather
incomplete and irregularly striate medially, but becoming more distinct laterally;
lateral sides of propodeum dull, finely and irregularly roughened; metasomal
terga rather shining, almost imperceptibly though finely striato-reticulate, punc¬
tures minute, scarcely evident, sparse except near base of tergum II, apical rims
neither impressed nor glabrous.
Male. —Length 3.5-4 mm. Head and thorax greenish, propodeum darker above
rather bluish, metasoma piceous somewhat paler ventrally and along apices of
metasomal terga; clypeus entirely piceous; antennae yellowish-brown below,
fuscous above; tarsi yellowish-brown; pubescence similar to that of female, though
considerably denser on face, but sparser generally on metasomal terga medially;
punctation of head, thorax including propodeum and metasoma essentially like that
of female, though punctation on metasomal terga less obscure and not as sparse;
pleura with punctures more distinct and less rugose, medial area shining between
sharply defined punctures; sculpturing of propodeum, especially that at dorsal
and lateral sides of posterior face, less well defined, but that on dorsal area and
sides essentially as in female; antennal segments short, less than twice as long
as broad.
Holotype female , allotype and 17 paratypes (14 $ $ , 3 2 2), along
Rock Creek, 4 miles east of Milton, Calaveras County, Califor¬
nia, 1 October 1969, at flowers of Jepsonia heterandra Eastw. (R.
Ornduff). The primary types are deposited in the collections of the
California Academy of Sciences, San Francisco. Additional paratypes
(17) were collected at the type locality on 8 October 1967 by R. W.
Thorp (11$ 5 2 2 ) and by R. Ornduff {!$) on 10 October 1968,
from the same population and 2 other specimens (lc?, 12 ) also desig¬
nated as paratypes were collected on 10 October 1968, by R. Ornduff
and the author at Sullivan Creek, 3 miles south of Stent, Tuolumne
County, California.
Dialictus ornduffi closely resembles D. brurmeiventris (Crawford,
1907:194) and is apparently its nearest described relative. It differs
most conspicuously from that species in having the disc of the third
metasomal tergum of the female entirely pubescent and in having the
clypeus of the male entirely dark in coloration. Further D. brwnneiven-
tris, which was originally described from Ormsby County, Nevada, is a
summer species to judge from the available records.
212 THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Literature Cited
Crawford, J. C. 1907. New Halictinae from the western United States. Inverte-
brata Pacifica, 1: 190-197.
Ornduff, R. 1969. Ecology, morphology, and systematics of Jepsonia (Saxi-
fragaceae). Brittonia, 21: 286-298.
1970. The reproductive system of Jepsonia heterandra. Evolution, (in press).
Biology and Structure of Some California
Bristletails and Silverfish
(Apterygota: Microcoryphia, Thysanura)
Edward Laidlaw Smith
Chico State College, Chico, California
A variety of observations were made recently on Microcoryphia and
Thysanura, both in the field and among laboratory rearings. These
ectognathous Apterygota are relict descendents of the Pterygote Pre¬
cursor, and closely resemble immature Hemimetabola. The Micro¬
coryphia (bristletails) are the more generalized, and include 2 extant
families: the Machilidae and Meinertellidae. As a whole, machilids are
morphologically the more primitive. Modern Thysanura, the silverfish
or firebrats, include the Lepidotrichidae, Lepismatidae and the Nico-
letiidae. The taxonomy of both orders is unsettled, and the unidentified
species considered here have been referred to the genera “ Machilis 5,1
and Mesomachilis (Machilidae) ; Machilinus (2 species: Meinertel¬
lidae) ; and two species of Clenolepisma (Lepismatidae).
Machilids (Fig. 1) bear “styli” on either or both the meso- and meta-
thoracic coxae, and abdominal coxites II-IX (the thoracic and abdomi¬
nal styli are not homologous—the former being coxal endites or exites,
the latter telopodite rudiments). “ Machilis” strenua has thoracic styli
on both the meso- and meta-thoracic coxae. There are 2 eversible vesicles
on each abdominal coxite II through V, and 1 vesicle on I, VI and VII.
The males lack gonapophyses VIII. The species of Mesomachilis had
thoracic styli only on the metathoracic coxae and but 1 vesicle on each
1 The “ Machilis” here referred to apparently was described as Machilis strenua Silverstri. How¬
ever, since the genus Machilis now has been restricted to the European fauna, this California species
presumably belongs to an undescribed genus (Wygodzinsky, pers. comm. 1969). Taxonomic confusion
led me originally to put this species in Pedetontus by mistake, and designate it as such in F'ig. 2
of Smith 1969.
The Pan-Pacific Entomologist 46: 212-225. July 1970
Fig. 1. Male Mesomachilis (Microcoryphia: Machilidae). The posture is natural but for the following. The maxillary endites have
been pulled back slightly to show the mandible, and 1 vesicle has been illustrated everted. Gonopophyses VIII and IX and the penis are
outlined and shaded. The entire abdominal coxite VI is stippled. The mesothoracic stylus found in some other machilids but lacking
in this genus is outlined. Abbreviations: AN, anus; CE, cercus; CE-CX, cereal coxite; CX, coxite; EV, everted vesicle (endite?) ; GC, gono-
coxite; GP, gonapophysis (endite?); GS, gonostylus (telopodite) ; GL, glossa (outer endite) of 1st maxilla; LM, labrum; LP, palp
(telopodite) of 2nd maxilla (labium) ; MD, mandible (coxite-f- endite) ; MF, median filament (tergite XI) ; MP, palp (telopodite) of 1st
maxilla; MS, mesothorax; MSS, mesothoracic coxal stylus (endite?); MT, metathorax; MTS, metathoracic coxal stylus (endite?); PG,
paraglossa (endite) of labium; PN, penis (fused papillae of IX) ; PR, prothorax; RV, retracted vesicle; ST, abdominal stylus (telopo¬
dite). Abdominal segments are identified by numbered circles.
JULY 1970] SMITH-BRISTLETAILS AND SILVERFISH 213
214
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
abdominal coxite I—VII. Males possessed grossly reduced gonapophyses
VIII. The antennal flagella of these 2 genera are scaly, and this char¬
acteristic separates them from perhaps the other most common genus
( Pedetontus ) in the Sacramento area, the flagella of which lack scales.
Members of Machilidae have 1 or 2 eversible vesicles on each of the
abdominal coxites. Females (and some males) bear 2 sets of multi-
segmented (or annulated) gonapophyses on the coxites of the genital
segments (abdominal somites VIII, IX). These structures are homolo¬
gous with the eversible vesicles of pregenital somites. The 2 pairs of
gonapophyses of the female remain interlocked as is usual among the
more primitive insect groups. Those of males are disengaged, and
gonapophyses VIII greatly reduced or entirely absent (Smith, 1969 and
unpublished). The gonostyli (telopodites of coxites VIII and IX) are
prominent and heavily musculated in both sexes. The abdominal ster-
nites are distinct and may wholly separate the coxites on either side
from each other. The coxites, however, are flexibly attached more or
less completely to the sternites along at least their proximomesal margins.
This overall pattern of abdominal appendages is the most generalized
known among both Apterygota and Pterygota.
California meinertellids (Fig. 2) have no abdominal vesicles, but
some found in other areas have 1—never 2—on some coxites. All
Meinertellidae lack thoracic styli. They have the usual abdominal styli
on segments II through IX. Male Meinertellidae lack either pair of
gonapophyses, and only a bilobed median penis remains. A penis is
present in Machilidae as well, but it lies between either gonapophysis IX.
The abdominal sternites of Meinertellidae are reduced to narrow straps
at the bases of the coxites, an ordinal character of the family.
Thysanura have no thoracic styli. According to Wygodzinsky (1961),
the relict Tricholepidion gertschi Wygodzinsky (Lepidotrichidae) re¬
tains abdominal styli on segments II—IX, and eversible vesicles on II-
VII. This is the microcoryphian pattern. Other Thysanura have ab¬
dominal styli only on segments VII, VIII and IX, or just VIII and IX.
Some Nicoletiidae have eversible vesicles as well. When present, gona¬
pophyses IX of male Thysanura apparently are reduced to a single seg¬
ment (radix), the pterygote condition. A median phallus similar to that
in Microcoryphia Meinertellidae usually replaces gonapophyses IX. Ex¬
cept for Lepidotrichidae, male Thysanura lack gonapophyses VIII.
Microcoryphia have prominent ocelli and large, often holoptic com¬
pound eyes which give them a remarkably intent expression. Their eyes
glow mysteriously in the dark when illuminated with a flashlight.
Thysanura have reduced or no compound eyes and, except for Lepido-
JULY 1970 ]
SMITH-BRISTLETAILS AND SILVERFISH
215
Fig. 2. Laboratory colony of Machilinus Sp. (Microcoryphia: Meinertellidae)
clustering on pine bark under a bright light. Apart from the dark scales on the
mesothorax, this species is relatively unpatterned. There are 4 individuals in the
top picture and 5 in the lower. The average size is 8 mm, excluding appendages.
trichidae, no ocelli. Microcoryphia have maintained the primitive hy-
pognathous head posture, and retain the prominent telopodite of the
(1st) maxillae as a locomotory and grasping appendage. The maxillary
palpi are used most often to manipulate food and to examine the sub¬
strate, but sometimes they may be used in climbing. These “maxil-
216
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
lipeds,” taken with the long, mobile abdominal styli (especially the 9th),
give the creatures a most “un-insectan” aspect as they scuttle about.
Except for Lepidotrichidae, Thysanura are hypognathous in early instars,
but become prognathous after a few instars. Both groups continue to
moult periodically after reaching sexual maturity.
Eversible Vesicles: Water Absorbing Organs
Many myriapodan and insectan arthropods can protrude 1 or 2 mem¬
branous sacs from some of the coxites. Among these structures are the
individual eversible vesicles of Protura, Microcoryphia, Symphyla, and
Diplopoda, and the combined pair forming the collophore of Collembola.
All are everted by blood pressure and withdrawn by 2 to 4 muscles. In
machilids, 1 muscle set originates within the subcoxa and the other in
the coxa. The corium of machilid vesicles is opaque in preserved
material, but the everted sacs in life are so transparent that blood cells
can be seen within them. The protrusible vesicles are apparently homolo¬
gous with the coxal vesicles of Megaloptera: Corydalidae, and the fleshy
prolegs of larval Hymenoptera, Mecoptera and Lepidoptera.
The exact homology of the vesicles themselves, and their counterparts
on the genital segments (the gonopophyses) is problematical. Gustafson
(1950) suggested they were modified coelomoducal papillae which
originally bore the gonopores. Sharov (1966) suggested that both
vesicles and gonapophyses were coxal exites (epipodites). I suspect that
both the vesicles and the gonapophyses represent reduced coxal endites.
All of these conclusions assume the vesicles and their presumed counter¬
parts are homologous in all groups where they appear, and the evidence
overwhelmingly favors this (Smith, 1969). Regardless of structural
origin, however, the purpose of these eversible sacs is clear: water ab¬
sorption. Heeg (1967a) reviewed some of the commentary supporting
this. I should like to describe the actual use of the protrusible vesicles
to sponge up water.
Around midnight during a dry August weekend in the California
Coast Ranges west of Sacramento, I collected many large (13 mm body
length, 40mm antennae + body + terminal filament) machilids of at
least 2 genera. They were put in several jars with only paper toweling.
The following afternoon I began to place each lot in a common terrar¬
ium. This container had lichens and dry forest litter in the bottom, and
several pieces of wet maple and cascara bark. As the insects scattered,
the majority sooner or later came in contact with the damp bark, and
halted abruptly. After a moment’s examination of the moist surface,
they would lower their abdominal coxites against the surface, and pro-
JULY 1970 ]
SMITH-BRISTLETAILS AND SILVERFISH
217
trude the vesicles with a pronounced front to rear contraction of the
body. After a minute or two of inactivity, all vesicles would be retracted
simultaneously, and the eversion process repeated, often with a change
of body position. They sponged only on the heavily fissured, rough
maple bark, or on damp, compact clay. They consistently rejected
moistened smooth bark, moss, lichens, leaves, or sand. Actual drops
of water were avoided, and simulated rain caused an immediate rush
for cover.
The following evening I returned to the collection site, and poured
water down the bark of a maple stump on which I had seen the insects
running up and down. In less than 5 minutes, 2 machilids were busy
sponging up the water. Such was their preoccupation that I was able to
pick them up with my fingers. The same preference for substrate texture
seen in the laboratory was demonstrated in the field.
It is significant that most nocturnal, forest-floor dwellers in the insect
and myriapod groups possess water absorbing vesicles. Moreover, many
diurnal members of these taxa such as the North American Machilinus
lack them. The machilids I reared did not appear to drink water, and
they avoided drops of it. However, Heeg (1967a) reported Machiloides
(a vesicle-bearing Meinertellid) normally drank water, and seemed to
use vesicles only when no other source was available. My Meinertellid
Machilinus did drink, and commonly died within 2 or 3 days if deprived
of water in droplet form, whether or not moist earth or bark were pres¬
ent. While most Thysanura lack vesicles, those I was rearing showed
greater tolerance to dessication than Machilinus. Heeg (1967b) found
Ctenolepisma capable of cuticular absorption of water vapor at levels
as low at 60% relative humidity, but no such ability in Machiloides.
The Uses of Styli
While the abdominal styli of both apterygote adults and pterygote
immatures are undoubtedly telopodite rudiments, the identity of the
unmusculated styli borne more or less laterally on the thoracic coxae
is uncertain. They superficially resemble both the musculated lateral
abdominal styli as well as the mesal unmusculated styli of Symphyla.
Sharov (1966) felt they were exites and thus not homologous with the
abdominal styli. However, some past workers have also suggested that
all styli are exites. There is still another possibility: the thoracic styli
are not truly lateral, but angle about 45° away from the body axis.
They describe a considerable arc as the insects walk. Machilids use
the thoracic styli (which, while unmusculated, are well-supplied with
sensilla) to orient themselves in crevices: they try to keep at least 1 on
218
the pan-pacific entomologist
[vol. 46 NO. 3
0.5 mm
JULY 1970 ]
SMITH-BRISTLETAILS AND SILVERFISH
219
either side in contact with their surroundings. A slight rotation of the
coxae (or minor shift in the position of the styli) would place them in
the same position as the Symphylan stylus. One could make at least a
consistent argument that the mesal stylus and adjacent eversible vesicle
of symphylans, and the thoracic styli and paired eversible vesicles of
machilids are endite derivations and therefore homologous with the
gnathal endites. Such an arrangement would make the palpi, legs and
abdominal styli all telopodites or their rudiments. This proposal has as
much morphological support as the all-exite or exite-telopodite theories,
which is to say all are inconclusive at present.
While they are movable, the abdominal styli of Microcoryphia and
Thysanura act as rigid skis upon which the abdomen is borne. The
actual gliders are the scale-like hairs covering the anterior faces of the
styli (Fig. 3, ST). However, the gonostyli on VIII and IX can be truly
locomotory. When these animals climb objects, the 9th styli actually
push, and bristletails trying to climb a smooth surface such as glass
prop themselves up on the rear styli and grapple overhead with the
thoracic legs and maxillary palps. Some males of Nicoletiidae (Thysa¬
nura) may use the 9th styli as clasping organs during reproduction, the
characteristic role of the appendages among male Pterygota.
Female Pterygota usually employ the 9th gonostyli as sensory palps
or as a scabbard for the ovipositor, which apparatus is derived from the
8th and 9th gonapophyses (8th gonostyli are absent in male and female
pterygotes). However, Grylloblattodea, Orthoptera and Raphidida use
the gonostyli along with the gonapophyses to penetrate the substrate
during oviposition. Machilids may use the gonostyli while egg-laying
in a manner reminiscent of these groups. The female will back up in
loose duff, and vigorously excavate along the surface with the 9th
gonostyli by flipping the appendages laterally in unison. The styli move
synchronously much as one uses the arms to swim the breaststroke. Dirt
and litter may be tossed over an inch away. The flexible, interlocked
<-
Fig. 3. Tergal lobe (TL) on ventrolateral margin of abdominal tergite III of
male “Machilis” Strenua Silvestri (Machilidae). Fig. 4. Idealized machilid abdomi¬
nal coxite, ventral aspect. Abbreviations : Fig. 3, 4: CL, claw; CX, coxa; EV,
eversible vesicle; EV-L, lateral vesicle; EV-M, mesal vesicle; RV, retracted vesicle;
S, sternite; ST, stylus; SX, subcoxa; T, tergite; TL, tergal lobe. Fig. 5. Collection
jar (Machilitrap) for Microcoryphia and Thysanura. Abbreviations-, a , plastic vial
approx. 8 cm tall, 5 cm dia. (e.g., Thornton Plastic Co., Salt Lake City, Utah,
No. 55-40) ; b, polyeurethane skirt approx. 1 cm thick glued to free edge and pro¬
jecting about 2 cm beyond lip.
220
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
gonapophyses extend between the longer gonostyli, and twist and thrust
side to side between the earth particles as the styli rake along before
and to either side of them.
Aggregation and Other Behavior
Delany (1959) found a weakly developed asexual sociality in Ptero-
bius (Machilidae), and apparent sleep aggregations. The microcory-
phians I observed (but not the thysanurans) behaved similarly. While
there was ample room in containers and terraria for relatively uniform
dispersion under individual shelters, invariably the entire population
of machilids (some 25 males and females) gathered under 2 or 3 shelters.
Their antennal tips might be crossed, and body contact was common.
All of the machilids would aggregate together, regardless of genus, but
the Machilinus in the same terrarium would cluster elsewhere. There
was a definite avoidance reaction between members of the 2 families.
Aggregation in favored localities was seen both in the laboratory and
field. If the insects were removed forcibly from these refuges, they
eventually would return. There appears to be some trail marking sub¬
stance involved: in nature, the machilids (but not the meinertellids or
thysanurans) followed definite paths on both tree trunks and on the
forest floor. There were some areas where they would run back and
forth in a seemingly aimless fashion. On occasion they would chase
each other. If one scuffed away the litter in the middle of one of these
trails, the machilids using them would halt and cast about. Some turned
around and ran back the way they came; others crossed the disrupted
zone and continued on. Such habits can be a boon to the collector: after
such a machilid run is located, one can sit down and catch them as they
scamper by.
A peculiar exhibition may be related to emission of sex pheremones.
Several times I found solitary male machilids (genus undetermined)
sitting on exposed rocks near the trails they frequented. They had their
abdomens raised in an arc and deflexed over their thoraces, exposing the
abdominal sternites. The posture occasionally was accompanied by a
grotesque rowing motion of the styli and a flipping of the cerci. Con¬
ceivably, some secretion was being emitted. This exhibition was ob¬
served during the course of the oviposition period. Unfortunately, I
did not observe actual courtship or mating.
A bilobed tergal process (Fig. 3, TL) which may have something to
do with reproduction can be found on the proximoventral edge of either
tergite III of “Machilis” strenua. The upper % of the bifurcate projec¬
tion is covered by a dense mat of black, scale-like hairs. It is filled with
JULY 1970 ]
SMITH-BRISTLETAILS AND SILVERFISH
221
fat body and apparently is not movable. Since Microcoryphia are gen¬
erally believed to mate by spermatophore transfer without direct body
contact, a sexual role for these pairs of recurving lobes is difficult to
envision. I suggest it mainly because I can think of no other, and since
they are found only in adult males.
The arched position of the abdomen, but without the stylar move¬
ment, is the usual offensive posture in both sexes of Microcoryphia and
some Thysanura. The cerci and longer caudal filament (tergite XI) are
thrust far beyond the head in a rather scorpion-like pose, and then
rapidly vibrated. The antennae may be pulled back along the sides of
the body into what is normally a resting position for the insect. Bristle-
tails I have seen in motion hold their antennae at an acute angle ahead
of themselves and scan at intervals by swinging the long flagellum a
half a dozen times in a tight circle. At rest, the antennae are held next
to the body but with the projecting tips often crossing the antennae
of an adjacent machilid.
ECDYSIS
One of the most bizarre habits of Microcoryphia is their moulting
behavior: they glue themselves down before shedding. The cement ap¬
pears to be fecal material, and it attaches the underside of the end of the
abdomen to the substrate. Then the head and thorax emerge through the
usual median slit in the vertex and thorax as the legs and abdominal
styli thrust the body forward. Next, the legs and maxillary palps are
used literally to walk the abdomen and its appendages out of the old
integument. I have seen several instances with both machilids and
meinertellids where either the cement failed or the wrong substrate was
chosen (e.g., sand). None of these unfortunates completed ecdysis
(normally undertaken at dawn), and all died by mid-morning regardless
of temperature and humidity. Entangled specimens I attempted to free
also died: the appendages did not expand when the stuck cuticle was re¬
moved, apparently from rapid irreversable damage due to the tourniquet
effect of the uncast integument.
Microcoryphians normally are covered with scales and scale-like
hairs, often in elaborate patterns. For instance, there is a very common
Machilinus throughout the drier areas of northern California with a buff
background, brown reticulations, and paired black bands running down
its back. The effect is ludicrously like a miniature Volkswagen with
racing stripes. The scales of bristletails and silverfish generally provide
excellent crypic coloration amid forest litter. That Machilinus mentioned
is nearly invisible among pine needles. If one descales a microcoryphian,
222
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
and it has not recently moulted, it will shed in a few days. The detach¬
able scales also may foil a predator since the aggressor most likely would
get only scales unless a sclerite or appendage were gripped. Apart from
the gliding surfaces provided by the scaly hairs on the styli, the pre¬
ceding appear to be the only major roles played by the pelage on the
integument of bristletails. Heeg (1967b), for instance, found they had
nothing to do with water conservation.
Histological Differences between Apterygota and Pterygota
The tissues of Microcoryphia and Thysanura have fixation and stain¬
ing properties very different from Pterygota. Living or fixed unabraded
pterygote cuticle (sclerotized or otherwise) which has not been treated
with a lipid solvent cannot be stained easily in most water-based histo¬
logical dyes. This is apparently the effect of hydrophobic wax layers
in the pterygote epicuticle. However, methylene blue applied to these
apterygotes (living or dead) rapidly penetrates scales, sclerotized cuticle,
corium and, given sufficient time, the internal organs. Heeg (1967b)
found that whatever cuticular wax layer (if any) is on these insects, it
is of little significance in preventing water loss. In a similar vein, the
poor preservation of most apterygotes in alcohol seems related to basic
biochemical differences in their tissues. Indeed, fixation of living speci¬
mens in alcoholic Bouin’s solution 2 or F.A.A. 3 is the only practical way
to obtain satisfaction preserved material. In alcohol alone, the cuticle
softens and shreds, and the hypodermis and muscles fragment and pull
away from the integument.
Collection, Rearing and Preservation
Thysanura, and Microcoryphia except Meinertellidae, are crepuscular
or nocturnal, and generally photonegative. Machilinus spp. in northern
California are not only diurnal and phototropic, but can be captured
most easily at night by aiming a bright flashlight beam on the forest
floor where they have been seen during the day. After about 20 minutes,
they will appear in numbers and crawl toward the light. They can be
collected during the day, but it means much searching under rocks and
in leaf litter. They are very partial to warmth, and will cluster against
2 Alcoholic Bouin’s solution: picric acid, 5.5 gm (saturated) ; absolute 2-propanol, 105 ml; formalde¬
hyde, 50 ml; glacial acetic acid, 15 ml; water, 45 ml. Transfer specimens after 6-12 hrs. at room
temperature into 70% 2-propanol. Fixation can be enhanced by heating, but specimens harden rapidly.
For sectioning or dissections, the picric acid must be removed before staining is possible. Soak speci¬
mens or sections in 10% ammonium acetate in 70% 2-propanol until the yellow disappears (5 min.
or longer).
3 F.A.A.: 85% 2-propanol, 80 ml; glacial acetic acid, 10 ml; formaldehyde, 10 ml. Specimens may
be kept in solution indefinitely without undue hardening if kept from heat. Material can be im¬
bedded without additional fixation after 24 hrs.
JULY 1970 ]
SMITH—BRISTLETAILS AND SILVERFISH
223
the walls of a container warmed by a lamp. A similar response is ob¬
tained with a heating element. They sometimes can be found in con¬
siderable numbers on warm surfaces away from direct sunlight. The
largest daytime aggregation of Machilinus I have ever seen was inside
a well-lit men’s privy in the Lava Beds National Monument (Modoc
County, California) one hot July afternoon!
The machilids studied were not unduly disturbed by strong light at
night, and were easily caught by stationing oneself by a rock, stump or
run they frequented. Most Thysanura take alarm at light and, while a
bristletail will ignore a beam and allow leisurely collection, one must
move very rapidly to capture a silverfish. Ctenolepisma lineata pilifera
(Lucas) often enters homes in rural areas of northern California, and can
be spotted when the light is turned on. They prefer warmer areas, and the
most fruitful collections I have made of the species were in my garage dur¬
ing the summer. This place is warm during the day since it lacks windows,
has west-facing doors often hot to the touch from the inside, and is a
veritable silverfish rookery. The insects can be found at all hours, but
especially in the late afternoon, scurrying to and fro over the warm
doors.
All of these apterygotans are caught easily by use of the vial illus¬
trated in Fig. 5, or an aspirator. The latter has obvious disadvantages
with the agile creatures except when they are cornered. The % to % in.
thick polyeurethane foam wrapped around the vial lip and extending
about % in. from it permits complete edge contact of the container on
uneven surfaces. If the inner surface of the plastic vial is sanded about
% in. down from the outer lip, microcoryphians will climb up the side as
far as they can get a grip when the bottle is inverted over them. Thysa-
nurans usually will not climb, and a 3 X 5 card or similarly sized piece
of stiff cellulose acetate must be worked between the vial and insect, and
the substrate. Once contained, when the vial is quickly turned right side
up, the captive will drop to the bottom. If there is a groove around the
edge in the bottom of the vial, most individuals will freeze in the crevice
and not make an immediate attempt to escape. The insects then can be
dropped into a jar by inverting the vial again and tapping it. The lid
should be replaced quickly if Microcoryphia are involved and the con¬
tainer is for live storage: the creatures may leap when they land, and
can jump a foot.
As noted, Apterygota preserve poorly in alcohol. F.A.A. is the pre¬
ferred solution, with alcoholic Bouin’s solution 2nd choice. Specimens
must be dropped in alive. Vesicles can be made to evert by pinioning
the insect upside down, pushing down on the abdominal coxites, then
224
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
flooding with F.A.A. as the vesicles inflate. Sometimes the sacs will
evert of their own accord when killed in solution. Scale patterns are
obliterated in preservative if not during capture, and photography pro¬
vides the only practicable record. Specimens descaled when collected
can be held until they shed and appear in new vestiture.
The diet of Microcoryphia and Thysanura in nature includes lichens,
terrestrial algae, molds, and decaying fruits and insects. They can be
reared easily in a jar or terrarium on dried or moist live yeast, and
rolled oats. Lichens and wilted lettuce leaves should be added occasion¬
ally. A few milliliters of water periodically squirted on rough bark suffice
for machilids, but meinertellids require drops of water daily. The en¬
vironment, however, should be essentially dry. Excess water or food
results in mite and Collembola infestations. Loose duff with bark and a
few mossy stones provide a satisfactory ground cover.
Species of Machilinus like to bask in the “sun,” and should be pro¬
vided with a few hours of it with a lamp, preferably beamed over a rock.
They will cluster on warm glass, stones, etc. regardless of the intensity
of the visible light, suggesting a definite infra red tropism. This aggre¬
gating tendency provides the most convenient time for feeding, observa¬
tion and photography (e.g., Fig. 2). No Thysanura or Microcoryphia
I know of can climb unetched glass, but some screen (not a tight fitting
glass cover) should be provided.
Acknowledgments
Without the advice of Robert 0. Schuster, Department of Entomology,
University of California, Davis, I would never have learned where or
how to collect these fascinating beasts. Determinations, references, and
needed encouragement and comment came from Pedro Wygodzinsky,
American Museum of Natural History, New York. To both, my deepest
thanks.
Literature Cited
Delany, M. J. 1959. Group formation in the Thysanura. Anim. Behav., 7: 70-5.
Gustafson, J. F. 1950. The origin and evolution of the genitalia of the Insecta.
Micro entomology, 15(2) : 35-67.
Heeg, J. 1967a. Studies on Thysanura. I. The water economy of Machiloides
delanyi Wygodzinsky and Ctenolepisma longicaudatcc Escherich. Zool.
Afr., 3(1) : 21-41.
1967b. Studies on Thysanura. II. Orientation reactions of Machiloides delanyi
Wygodzinsky and Ctenolepisma longicaudata Escherich to temperature,
light and atmospheric humidity. Zool. Afr., 3(1) : 43-57.
Sharov, A. G. 1966. Basic arthropodan stock. Pergamon Press, Oxford. 271 p.
JULY 1970]
SCOTT—TYMPANAL RESILIN
225
Smith, E. L. 1969. Evolutionary morphology of the insect genitalia. 1. Origin
and relationships to other appendages. Ann. Entomol. Soc. Amer., 62:
1051-79.
Wygodzinsky, P. 1961. On a surviving representative of the Lepidotrichidae
(Thysanura). Ann. Entomol. Soc. Amer., 54: 621-7.
Resilin in the Sound-Organs of
Pyralidae and Cicadidae
(Lepidoptera; Homoptera)
James A. Scott
University of California, Berkeley
Introduction
The purpose of this paper is to chart the distribution of resilin in the
tympanal structures of selected insects, and to explain its function where
it occurs in these structures.
Resilin is a rubberlike protein, found in the cuticle of insects and
crustaceans, which functions as a natural spring. Anderson and Weis-
Fogh (1964) give a recent review. Resilin is the most “perfect” rubber
known; only 3% of the energy of deformation is lost as heat, and resilin
returns to its normal state faster than any other rubber. Dry resilin is
unstretchable, but it swells and becomes rubberlike in many solvents
including water. Resilin can be dried and hydrated repeatedly without
losing its rubberlike properties, and can be heated to 140° C without
degrading. Ethanol and histological fixatives harden it, but this effect
is reversible in water. It is easily digested by proteases. Resilin absorbs
ultraviolet light maximally between 280 and 320 mg depending upon
pH, and fluoresces between 350 and 600 mg, with a maximum fluores¬
cence at 420 mg.
Procedure
The sound-organs of Galleria mellonella L. (Pyralidae) and Cacama
valvata (Uhler) (Cicadidae) were tested for the presence of resilin. The
tympana of undetermined specimens of Acrididae, Arctiidae, Noctuidae,
and Tettigoniidae were also examined, but no large resilin deposits were
found, and these insects are not considered further. The structures of
Galleria and Cacama and a known sample of resilin, the prealar arm of
The Pan-Pacific Entomologist 46: 225-231. July 1970
226
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Locusta migratoria L. (Acrididae) were sectioned after fixing in Lillie’s
AAF (Lillie, 1954). Before staining, the sections were examined for
fluorescence using a Zeiss microscope with exciter filter IIII-UG-1 and
barrier filter no. 41. This system illuminates the section with ultra¬
violet light between 3000 and 4000 A, and permits all light above
4200 A to pass through to the observer. Resilin fluoresced blue. Sec¬
tions were examined on slides for fluorescence before staining since the
process of wax removal and staining destroyed fluorescence. The slides
were then stained with Mallory’s Triple Stain and mounted with Per-
mount. Resilin appears yellow, endocuticle and muscle appear blue,
and exocuticle appears red. The presence of resilin was further con¬
firmed by staining dissected specimens with methylene blue or toluidine
blue-light green, and by digesting with trypsin for 60 minutes at 37° C
and observing the change in fluorescence and elasticity. Staining is less
useful than fluorescence. The toluidine blue-light green stain and the
methylene blue stain caused endocuticle to appear blue-green and resilin
to appear greenish blue. Although resilin stained yellow after Mallory’s
Triple Stain, other tissues which did not fluoresce were also stained
yellow. Fluorescence was more selective, but many small areas within
the endocuticle were found to fluoresce. This may only indicate that
endocuticle contains resilin, however. Examining sections or whole
dissections alone was insufficient to determine the presence of resilin;
a combination of the two produced reliable results.
Pyralidae
Several small, fluorescent, membranous areas were found upon sec¬
tioning and staining fresh Galleria, but these areas have not been located
on dissected specimens. In the sections a fluorescent, small, rodlike
structure was found between the anterolateral corner of each tympanum
(which is on the anteroventral part of the first abdominal segment) and
the posterior part of the metathorax. This structure ceased to fluoresce
after trypsin digestion. The presence of resilin in this structure was
also indicated after staining with toluidine blue-light green and with
methylene blue. It would seem that this rodlike structure functions as
a spring in the hinge formed by the junction of thorax and abdomen;
it does not appear to have a role in sound reception by the tympanum.
ClCADIDAE
Sections of the anterior part of the abdomen of a male Cacama were
stained with Mallory’s Triple Stain. Fluorescence microscopy revealed
the presence of a large membranous area and a large solid pad in addi-
JULY 1970]
SCOTT—TYMPANAL RESILIN
227
tion to many smaller areas of fluorescence for which location remains
unknown. These two large areas are located around the tymbal and
beneath the toe of the “foot” of the chitinous V (Figs. 1 & 2). The
presence of resilin in these areas was confirmed by staining with tolui-
dine blue-light green and methylene blue, and by digesting a dissection
with trypsin and comparing it to a non-digested control in buffer. The
resilin stained blue, and digestion caused much more freedom of move¬
ment, especially in the resilin pad. Pringle (1954) found an elastic
tendon in the cicada which he said was stretched when the tensor muscle
contracted. He later reinterpreted this as chordotonal organ 1 (Fig. 1).
It contains no resilin and is not truly elastic. Furthermore, it is relaxed
when the tensor muscle contracts.
The following summary of the structure and function of the sound-
organs in the cicada will help the reader to understand the function of
resilin in the cicada. The summary is based on Pringle (1954, 1957),
Vogel (1923), Wakabayashi and Hagiwara (1953), Hagiwara (1956),
and Hagiwara et al. (1954). Pringle speculates that these organs arose
in copulatory behavior from transmission of weak sound from one insect
to the other or by gradual modification of stridulatory organs.
Sound reception is due to the tympanum (Fig. 1). The ventral open¬
ing to the tympanum is covered by a backward extension of the meta¬
thorax, the operculum. The tympanum receives the sound and transmits
it to the auditory “spatel,” which enters the auditory capsule. In the
auditory capsule is an invaginated cuticular auditory horn, and between
the auditory “spatel” and horn are stretched up to 1500 scolopidia which
detect the sound but transmit only the pulse pattern (the “modulation
envelope”) of the sound, not the primary frequency. The detensor
tympani muscle (side view Fig. 2) pulls the chitinous V rearward,
buckling the tympanum. Pringle states that this muscle contracts prior
to sound production and speculates that buckling may prevent the
tympanum from being ruptured by the sound. The ventral muscle (an
intersegmental muscle from thorax to abdomen) pulls the chitinous V
forward and would tend to oppose the detensor muscle and stretch the
tympanum. Vogel found a tensor tympani muscle which originated
from a dorso-ventral muscle of the nymph which would stretch the
tympanum; however, Pringle finds that the muscle degenerates after
emergence in the adult male, and is of importance perhaps only in
stretching the tracheal air sac after emergence. Expansion of the air
sacs is also aided by many small muscles which degenerate after emer¬
gence and by the excretion of large amounts of fluid, which creates a
space into which the air sacs expand. Tracheal air sacs cover the tymbal
228
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
Tracheal air sac
Resilin
Chordotonal
organ 1
Spiracle
Spiracular
muscle
Cho rdotonal
organ 2
■Dorsal muscle
/Tymbal apodeme
/Tymbal ring
/Tymbal cover
-Tymbal
Invagination of
auditory horn
'Auditory
capsule
'Process of auditory spatel
v 'Chitinous V
^erculum \ \ Resilin pad
y Foot of chitinous V
dll juC \
Sternum
/Tympanum
•Folded membrane
Tensor muscle
Chordotonal organ 1
Spiracle 2
-Spiracular muscle
Chordotonal organ 2
s Operculum
Foot of chitinous V
x Resilin pad
Detensor tympani muscle
Ventral muscle
■Intersegmental membrane
Figs. 1-2. Sound-organs of the cicada, Cacama valvata (Uhler). Fig. 1 . Anterior
view of cross-section between thorax and abdomen, X 10. Fig. 2. Right-lateral view
of chitinous V, abdominal pleuron and tergum removed, showing points of attach¬
ment of tympanum on chitinous V and pleuron, Xl5.
muscle, the inner surface of the tympanum and tymbal, and the postero-
ventral surface of the “folded membrane.” The folded membrane (inter¬
segmental membrane between thorax and abdomen) stretches from the
anterior border of the chitinous V to the metathorax and serves as a
barrier to hemolymph penetration of the tympanal air sac. The chitinous
V is a very large brace consisting of sternum 1 and the anterior part of
sternum 2. It attaches solidly to the cuticle wall dorsolaterally and rests
ventrally on two small “feet.” The chitinous V serves as the dorsal
JULY 1970]
SCOTT-TYMPANAL RESILIN
229
attachment of the tympanum and, especially, as the origin of the tymbal
muscle.
Sound production is due to the inward and outward buckling of the
tymbal, which is covered outwardly by the tymbal cover, an anteriorly
extending flap of the abdomen. The tymbal is surrounded by a ring and
consists of a thin cuticular area reinforced by struts, and is buckled
along a line through the center. It is moved by the huge tymbal muscle,
which wraps around the chitinous Y dorsally. The tymbal muscle is
thought to be a metathoracic flight muscle which has migrated backward,
or a first abdominal dorsoventral muscle. The backward migration of
the tymbal muscle (if it occurred) has left the metathorax with inade¬
quate flight muscles; Pringle cut the ventral nerve cord in front of the
metathorax and got normal flight due to the wing-coupling apparatus
of fore and hindwings. One sound is emitted during the in-buckling,
and another during the out-buckling of the tymbal. There is alternate
innervation of the two tymbal muscles, so there are twice as many sounds
as would be expected if the two muscles contracted simultaneously.
Some cicada tymbal muscles are not myogenic; one stimulation of such
a muscle results in a damped bimodal pulse of sound. Other cicadas
have myogenic tymbal muscles; one stimulation results in a series of
bimodal pulses to produce a train of pulses. The frequency and damping
of each train of sound is due to the resonating frequency of the tymbal
(which is usually the same as the resonating frequency of the air sacs),
which depends on the elasticity of the tymbal and the tensor and dorsal
muscles. The tensor muscle (originally in the first abdominal segment)
arises from the metathorax and inserts on the upper edge of the tymbal.
Tetanic contraction of this muscle increases the curvature of the tymbal
and therefore requires a greater force to be exerted to cause its buckling.
The dorsal muscle appears to act in a similar manner by altering the
angle of attack of the tymbal muscle. Thus the tensor and dorsal muscles
appear to amplify the sound, especially the in-click, and to decrease the
primary frequency. In cicadas which have myogenic tymbal muscles,
these muscles prevent tetanic contraction of the tymbal muscle by in¬
creasing the force required to buckle the tymbal. The tensor muscle
along with many other muscles helps raise the abdomen during sound
production. Associated with the tensor muscle are two chordotonal
organs, which presumably register the change in tymbal concavity pro¬
duced by the tensor muscle. There are many variations in sounds due
to the tensor and dorsal muscles and due to changes in resonant fre¬
quency of the air sacs produced by abdominal muscles and by opening
the opercula to different degrees (opercular opening results from lifting
230
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 NO. 3
the abdomen, since the opercula are rigidly fixed to the thorax). A
single species of cicada may have three types of song: 1) calling or con¬
gregating song; 2) courtship song; 3) disturbance squawk.
The function of resilin in the cicada can now be understood. Resilin
is obviously useful in the tymbal because of the buckling and distortion
which this structure undergoes; it prevents cuticle “fatigue” and enables
more of the energy to be channeled into sound production and less into
heat. It is remarkable that such a massive structure as the chitinous V
has only a very small ventral attachment. The resilin pad beneath the
“toe” of the “foot” of the chitinous V permits anteroposterior movement
of the chitinous V, which increases or relaxes the tension on the tym¬
panum. The tympanum is attached medially to the chitinous V, laterally
to the sternum and to a brace supporting the tymbal ring. Around its
circumference the foot of the chitinous V is the only point at which
movement can occur. Slackening the tympanum by detensor muscle
contraction during sound production would compress and rotate the
resilin pad; elastic forces would cause restretching of the tympanum
again when the detensor muscle relaxes. Another hypothesis that has
not been previously proposed is that the ventral muscle contracts in
opposition to the detensor muscle, thus stretching the tympanum. The
location of the ventral muscle is ideally suited for this purpose. Another
speculation is that the ventral muscle alters the tension of the tympanum
during the reception of a cicada song. The tympanum receives sound
waves which produce standing waves over its surface. Vibration is
maximal at the resonant frequency so that reception of sound is probably
strongest at the resonant frequency of the tympanum. Therefore, if the
tympanum could be stretched and tensed to various degrees by the ven¬
tral muscle, a wider range of frequencies could be monitored. The ar¬
rangement of scolopidia within the auditory capsule is such that some
scolopidia are moved a much greater proportion of their length than
are others, so that the scolopidia might be differentially sensitive to
various frequencies. The fact that recordings of the auditory nerve have
shown that only the “modulation envelope” or pulse pattern is trans¬
mitted does not support this hypothesis however, and mere closing or
opening of the operculum could accomplish the same effect since the
resonant frequency of the air sacs and presumably the tympana also are
altered. Clearly these hypotheses need to be tested.
Acknowledgments
The author would like to thank Dr. R. Pipa for providing most of the
equipment and for reading the manuscript.
JULY 1970 ]
SCIENTIFIC NOTE
231
Literature Cited
Anderson, S. 0., and T. Weis-Fogh. 1964. Resilin, a rubberlike protein in
arthropod cuticle. Advan. Insect. Physiol., 2: 1-65.
Hagiwara, S. 1956. Neuro-muscular mechanism of sound production in the
cicada. Physiol. Comp. Oecol., 4: 142-153.
Hagiwara, S., H. Uchiyama, and A. Watanabe. 1954. The mechanism of sound
production in certain cicadas with special reference to the myogenic
rhythm in insect muscles. Bull. Tokyo Med. Dent. Univ., 1: 113-124.
Lillie, R. D. 1954. Histopathologic Technique and Practical Histochemistry.
McGraw-Hill, N.Y.
Pringle, J. W. S. 1954. A physiological analysis of cicada song. J. Exp. Biol.,
31: 525-560.
1957. The structure and evolution of the organs of sound production in
cicadas. Proc. Linnean Soc. London, Session 167, 1954-55, pt. 2, March
1957, pp. 144-159.
Vogel, R. 1923. Ueber ein tympanales sinnesorgan, das mutmassliche hororgan
der singzikaden. Z. Ges. Anat., Abt. 1, Z. Anat. Entwicklungsgesch.,
67: 190-231.
Wakabayashi, T., and S. Hagiwara. 1953. Mechanical and electrical events in
the main sound muscle of cicada. Jap. J. Physiol., 3: 249-253.
SCIENTIFIC NOTE
Unique structures in members of Tachys sensu lat. (Coleoptera: Carabi-
dae). —The species of the tachyine groups Tachylopha Motschulsky, Tachyphanes
Jeannel, and Sphaerotachys Miiller are distributed from Africa to East Asia,
while those of B ary tachys Chaudoir are confined to the New World and distributed
from Canada to Brazil. Bruneau de Mire has recently revised the African species
of Tachylopha (1966, Rev. Zool. Bot. Afr., 73(1-2): 59-100), and I am presently
studying Barytachys and other New World groups. Bruneau de Mire illustrated
and noted an interesting structure in the species of Tachylopha in his study. I
have found this same structure, though less well developed in members of
Tachyphanes and Sphaerotachys and made a further discovery, hitherto unre¬
ported, of an interesting structure in some members of Barytachys. The purpose of
this note is to describe these structures and speculate on their function.
If a specimen of any species of Tachylopha , Tachyphanes, or Sphaerotachys is
examined from either side, one will immediately note a small to large hole (Fig.
1) in the body wall at the juncture of the mesepisternum and mesepimeron. This
hole is an infolding of the entire body wall forming a tube which passes through
the beetle, over the gut, and meets the infolding from the other side. In the
specimens I examined, no septum was apparent (but see Bruneau de Mire, p. 63),
thus the beetle has a transverse “tunnel” completely through the body. In some
specimens one can actually see through the beetle. Inside the tube the thickly
sclerotized wall is ridged, the ridges being continuous from one side of the beetle
to the other. The tube does not have any macro-openings into the coelum of the
beetle and no glands seem to be associated with the structure.
The Pan-Pacific Entomologist 46: 231-232. July 1970
232
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 NO. 3
Fig. 1. Tachylopha ovatus Motschulsky, male, left lateral aspect. Fig. 2.
Tachyura tripunctatus Say, male, ventral aspect, left side.
Since I have neither collected these beetles nor seem them alive, I can only
speculate on the function of this unique tachyine structure. My first thought
was that this may be an exudation source for ants. Many tachyines are
myrmecophilous. The lack of glands and internal openings might exclude this
explanation. The tube certainly strengthens the structure of the exoskeleton, but
this seems an insufficient explanation because no other tachyine has any structure
remotely like this one. My last hypothesis must be tested in the field. These
holes may be handles whereby ants could carry the beetles about. The ant
mandibles would be inserted into these holes and the beetle lifted in this fashion.
This is supported by the fact that the elytra of Tachylopha members have another
unique structure. Above the hole, the humerous of the elytron is deeply notched,
forming a tooth. The ant mandibles, when inserted in the hole, could fit over this
notch and reinforce the hold.
J. F. Lawrence informed me (per. comm.) that Lea (1917, Trans. Proc. Roy.
Soc. South Australia, 41: 121-322, fig. 8a) reported a similar structure in the
ptinid beetle Polyplocotes perforatus Lea. In these beetles the “tunnel” occurs
through the head beneath the clypeus, and its entrances are covered by the
antennae when in repose. It is interesting to note that these beetles are
myrmecophiles, but again no function has been assigned to the structure.
Some members of Barytachys have perforations (Fig. 2) of the prosternum,
anterior to the coxae, one on each side between the prosternum and proepisternum.
This hole is smaller than those in Tachylopha and are directed longitudinally
rather than transversely. The “pouch” ends abruptly at the dorsal wall of the
coxal cavity and has no openings into the coelum. It is much harder to visualize
ants using these longitudinal pouches for handles, but it is interesting that many
of the Barytachys members with pouches are associated with ants. Field work is
being planned to study this problem.— Terry L. Erwin, Museum, of Comparative
Zoology , Cambridge , Massachusetts.
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Vol. 46 OCTOBER 1970 No. 4
THE
Pan-Pacific Entomologist
EVANS —Thalccssotrechus barbarae (Horn) and the Santa Barbara oil spill
(Coleoptera: Carabidae) - 233
WILLIAMS—A redescription of the scorpion Vejovis bilineatus Pocock (Scor-
pionidae: Vejovidae) - 238
PARKER—New North American Microdynerus with notes on the nests of two
species (Hymenoptera: Eumenidae) _ 241
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reticulata McDunnough in Oregon (Ephemeroptera: Heptageniidae) _ 268
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INDEX TO VOLUME 46 _ 309
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Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
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The Pan-Pacific Entomologist
Vol. 46
October 1970
No. 4
Thalassotrechus barbarae (Horn) and the Santa Barbara
Oil Spill
(Coleoptera: Carabidae)
William G. Evans
Department of Entomology, University of Alberta, Edmonton, Canada
When oil spills from tanker wrecks or off-shore wells occur near the
coast, the subsequent effect on marine life may be very conspicuous,
such as the high mortality of oil soaked birds (at least 10,000 dead
birds were collected after the Torrey Canyon wreck according to Bourne,
1968) or the decimation of extensive littoral communities of rocky
shores due to oil smothering (Ranwell, 1968). Because of the stranding
of oil after high tides an off-shore spill can seriously affect plants and
animals in the higher intertidal zone (the high littoral) of the rocky
shore habitat, where there are often periods of days between the times
of the modifying effects of water. The midlittoral and the infralittoral
zones are more subject to wave action and are only temporarily affected
by oil (Riitzler and Sterrer, 1970). The dominant inhabitants of the
high littoral in California consist of those forms found in high tide
pools and those on rock surfaces, e.g. littorine molluscs, limpets, barna¬
cles, isopods, and amphipods along with such algal forms as Enteromor-
pha, Porphyra, Pelvetia , Fucus, Gigartina, and Ulva. All of these orga¬
nisms are susceptible to smothering doses of oil, but they could probably
survive light doses mainly because of their hydrophilic surfaces. After
the Torrey Canyon wreck lightly polluted areas of Cornwall, where
emulsifiers had not been used, showed only low mortality and limpets
and periwinkles apparently grazed away thin layers of oil (Nelson-
Smith, 1968).
Another distinct animal association of the high littoral zone, the
crevice fauna (Lewis, 1964; Glynne-Williams and Hobart, 1952; Evans,
1968) , is probably more susceptible to oil than any other animal
association of the marine littoral. The crevices, which can extend into
the rock from an inch to perhaps more than a foot, harbor a wide variety
of animals that are not normally seen due to their cryptozoic habits.
The Pan-Pacific Entomologist 46: 233-237. October 1970
234
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Many of these animals, particularly the young ones, use crevices as a
protection against wave action, predation, or desiccation while others
live on detritus and plankton stranded in the cracks when the tide re¬
cedes. Others emerge at night at low tide and scavenge or prey on
crustaceans and insects associated with the algae found in this zone.
Though the kinds as well as the numbers of species may vary geographi¬
cally from one place to another, certain groups of animals are char¬
acteristically found in high intertidal crevices throughout the world.
Pseudoscorpions, mites, and centipedes for instance, are typical crevice
dwellers in many areas, including California and, among the insects,
carabid, melyrid and staphylinid beetles and chironomid flies are domi¬
nant forms. In addition, the usual marine animals such as the ubiquitous
isopod Ligia, various amphipods and the young of littorine molluscs
are common members of the high crevice fauna.
The preponderance among this fauna of terrestrial, air-breathing-
arthropods which have become adapted to the high intertidal zone is
typical for most areas and it is this group that can be extremely sensitive
to oil contamination. Some of the reasons for this are: a. oil accumu¬
lates in crevices rather than being washed away, exposing the inhabi¬
tants to a continual dose; b. heavy fractions of oil can block the crevice
entrances completely, preventing food from being washed in and pre¬
venting passage of the inhabitants; c. the cuticle of many of the terres¬
trial forms, particularly of the adults, repels water due to wax layers
or dense hair piles but these hydrofuge properties are also oleophilic
properties making the cuticle readily wetted by oil. (Petroleum oils are
potent insecticides and historically have been used for this purpose very
extensively.)
The crevice fauna of the high tide zone could therefore be sensitive
indicators of the extent of oil pollution. (There is circumstantial evi¬
dence to support this view. In the heavily polluted oil port of Talara
in northern Peru I found the high littoral crevice fauna of nearby rocky
shores to be completely absent during a survey made in April 1968,
though it seemed to be unaffected at the unpolluted region of Paita
which is about 40 miles further south.) An opportunity arose to examine
this idea after the 28 January 1969 Santa Barbara oil spill. Though the
area of Santa Barbara had not been surveyed for the high crevice fauna
just previous to the oil spill (an unavoidable shortcoming of many
studies of this sort as is illustrated by the extensive surveys taken after
the Torrey Canyon wreck) there was no reason to believe that the com¬
position of the crevice fauna of this region would be much different than
that of central California which I had previously surveyed (Evans,
OCTOBER 1970 ] EVANS-THALASSOTRECHUS AND OIL SPILL
235
1970). However, because of this uncertainity, I selected Thalassotrechus
barbarae (Horn) as a representative indicator species of this crevice
community. This carabid beetle is distributed on the Pacific coast from
northern California to central Baja California and since the adults
emerge from the crevices in the evening and disperse over the rocks
where they prey on algal-feeding dipterous larvae or small crustaceans,
they can be found without destroying the habitat by searching with a
flashlight at this time. This method is preferable to splitting open
crevices in the daytime, a procedure also followed to obtain this beetle
as well as other members of the community. Finally, because these
beetles forage widely at night, they are more likely to come into contact
with oil, particularly if it is patchily distributed. Thus, they could re¬
ceive a dose of the oil by walking through it or by contamination of
their food, but the oil would probably be fatal in either case.
A survey of the distribution of T. barbarae after the oil spill was made
from June 1969 through the middle of August 1969 and was confined
to rocky shores between Shell Beach, San Luis Obispo County, in the
north to Leo Carillo State Beach (about 20 miles north of Malibu), Los
Angeles County, in the south, resulting in a discontinuous sampling-
pattern along the coast because of the prevalence of sandy beaches along
this region of coastline. Public beach areas were also sampled more
frequently than other areas because of their accessibility. Each locality
sampled was visited several times and observations were made at night
at low tide for foraging beetles or series of crevices were split open
with a geological hammer during low tides in the daytime over what
was thought to be representative areas of rocks.
According to Allen (1969a), irregular stretches of coastline amount¬
ing to more than 100 miles in length were affected by the oil in the two
month period following the oil spill (Fig. 1). “Heavy” deposits (several
inches of dark black crude oil over the entire tidal zone) occurred near
the Santa Barbara-Ventura area while “moderate” to “light” and “very
light” deposits occurred in other areas. The heavy deposits of oil on
the intertidal rocks were very evident in the Santa Barbara area even
in August, six months after the spill and it was also quite evident that
the crevice fauna of the high, oil-covered rocks had been completely
killed. The flora and fauna of the mid- and infralittoral zone did not
appear to be much affected at this time. Moderate deposits consisted of
isolated patches of tar while the light and very light deposits which
appeared as iridescent films when the area was surveyed in February
and March (Allen, 1969a) were no longer visible.
Thalassotrechus barbarae was not found in the following localities
236
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Fig. 1. Extent of oil pollution during the first two months after the 28 January
1969, Santa Barbara oil spill (after Allen, 1969a) and sampling localities for T.
barbarae for the June-August 1969 survey.
(from west to east) in the region of Santa Barbara: Refugio Beach,
El Capitan Beach, Ellwood Beach, Goleta point, Arroyo Burro Beach,
Carpinteria and Ventura (Fig. 1). Since all of these localities were sub¬
ject to heavy or moderate deposits of oil it is very probable that popu¬
lations of T. barbarae, which quite likely existed in these areas before,
were killed by the oil along with other members of the high littoral
crevice community. And judging by tar residues still remaining on the
rocks in most of these areas it may take many years before this com¬
munity can re-establish itself.
Some natural oil seepage occurs off Coal Oil Point, approximately
10 miles west of Santa Barbara, and other areas along the California
OCTOBER 1970 ]
EVANS—THALASSOTRECHUS AND OIL SPILL
237
coast and this could also contribute to oil deposits on nearby rocks and
beaches. But the flow rate from these sources differs by a factor of
about 100 when the conservative estimate of five thousand barrels per
day for the first 11 days of the oil spill is considered (Allen, 1969a,
1969b). So it is doubtful if these natural seeps could have affected the
crevice fauna so seriously in the Santa Barbara region.
Thalassotrechus barbarae was named by Horn (1892) after the type
locality, Santa Barbara, California. It seems ironic that this beetle can¬
not now be found anywhere near Santa Barbara and I think it is about
time for biologists and conservationists to become just as concerned
about the effects of pollution on insects and other invertebrates as about
those on the more familiar larger animals.
Literature Cited
Allen, A. A. 1969a. Santa Barbara Oil Spill. Statement prepared for U. S.
Senate Interior Committee, Subcommittee on minerals, materials, and
fuels. 20 May 1969.
1969b. Estimates of surface pollution resulting from submarine oil seeps at
Platform A and Coal Oil Point. Tech. Memo. 1230, November 1969,
General Research Corporation, Santa Barbara, California.
Bourne, W. R. P. 1968. Oil pollution and bird populations, pp. 99-121. In J. D.
Carthy and D. R. Arthur, Eds. The Biological Effects of Oil Pollution
on Littoral Communities. Field Studies 2, Suppl. Field Studies Council,
London.
Evans, W. G. 1968. Some intertidal insects from western Mexico. Pan-Pac.
Entomol., 44: 236-241.
1970. Insects and relatives. In Morris, R. H., and D. P. Abbott. Marine
Invertebrates of the California Shores. Stanford Uiiiv. Press (in press).
Glynne-Williams, J., and J. Hobart. 1952. Studies on the crevice fauna of a
selected shore in Anglesey. Proc. Zool. Soc. Lond., 122: 797-824.
Horn, G. H. 1892. Random studies in North American Coleoptera. Trans. Amer.
Entomol. Soc., 19: 40^8.
Lewis, J. R. 1964. The Ecology of Rocky Shores. English Universities Press,
Ltd., London.
Nelson-Smith, A. 1968. Biological consequences of oil pollution and shore
cleansing, pp. 73-80. In J. D. Carthy and D. R. Arthur, Eds. The Bio¬
logical Effects of Oil Pollution on Littoral Communities. Field Studies
2, Suppl. Field Studies Council, London.
Ranwell, D. S. 1968. Extent of damage to coastal habitats due to the Torrey
Canyon incident, pp. 39-47. In J. D. Carthy and D. R. Arthur, Eds.
The Biological Effects of Oil Pollution on Littoral Communities. Field
Studies 2, Suppl. Field Studies Council, London.
Rutzler, K., and W. Sterrer. 1970. Oil Pollution. Damage observed in tropical
communities along the Atlantic seaboard of Panama. BioScience, 20:
222-224.
238
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46, NO. 4
A Redescription of the Scorpion Vejovis bilineatus Pocock
(Scorpionida: Vejovidae)
Stanley C. Williams
Department of Biology, San Francisco State College, California 94132
R. I. Pocock described eight new species of North American Vejovis
in a brief but very important paper (Pocock, 1898). Among these was
Vejovis bilineatus Pocock, from “San Diego, Texas.” The identity of
this species has caused some confusion among scorpion workers ever
since the original description. Even Hoffmann, in his systematic mono¬
graph of the scorpion fauna of Mexico (Hoffmann, 1931) was not sure he
knew this species, and indicated the species he called Vejovis bilineatus
possibly might not be the same species as that described by Pocock,
although he believed them to be the same.
To clarify the status of this species, Willis J. Gertsch, of the Ameri¬
can Museum of Natural History, borrowed the types from the British
Museum of Natural History. Knowing of my interest in this group also,
Dr. Gertsch permitted me to examine the specimens while they were in
his custody. Since knowledge of this species is important in under¬
standing the composition of the scorpion fauna of the United States
and Northern Mexico, a redescription is here made.
Much appreciation is due W. J. Gertsch for providing me an oppor¬
tunity to study the type, and to the British Museum of Natural History
for the courtesy of making it possible to borrow the type. Thanks to
Christie A. Steketee and Charlene F. Williams for clerical assistance.
This work was partially supported by research grant GB 7679 from the
systematics section of the National Science Foundation.
Vejovis bilineatus Pocock, 1898
(Figs. 1, 2)
Diagnosis. —Small species of Vejovis belonging to the “spinigerus” sub-group.
Probably seldom attains body size longer than 35 mm. Coloration: Mesosomal
dorsum with one pair of distinct dark, longitudinal stripes; carapace with dark
underlying marbling; metasoma with dark stripes in position of inferior median
and inferior lateral keels; ventral surface of metasomal segment V somewhat dark.
Metasoma with obsolete inferior median keels; inferior lateral keels irregularly
smooth and fairly developed; dorsal and dorsolateral keels on segments II to
III more crenulate than serrate; ventral surface of vesicle not granular, but may
have punctiform depressions; pedipalp hand small, palm not very swollen, movable
finger about % carapace length; pectines with 13 to 16 teeth in females and 15
to 17 teeth in males.
The Pan-Pacific Entomologist 46: 238-241. October 1970
OCTOBER 1970] WILLIAMS—SCORPION REDESCRIPTION
239
Figs. 1 and 2. Vejovis bilineatus Pocock. Holotype female. Fig. 1. Dorsal view.
Fig. 2. Ventral view.
Holotype female. — Coloration. —Color pattern somewhat obscured by poor
preservation. Base color of cuticle deep yellow; some dark underlying markings
on carapace; one pair submedian dark longitudinal stripes on dorsum of mesosoma.
Carapace .—Anterior margin straight; lateral eyes 3 per group; median eyes
on very subtly elevated ocular tubercle; diad % carapace width at that point.
Carapace surface granular, slightly lustrous.
Mesosoma .—Tergites granular, somewhat lustrous; tergite 7 with 2 pair of
crenulate to serrate lateral keels. Sternites smooth; last sternite lacking lateral
keels.
240
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Table 1 . Measurements (in millimeters) of Vejovis bilineatus Pocock,
holotype.
Holotype
(female)
Total length
35.0
Carapace, length
3.5
width (at median eye)
2.9
Metasoma, length
11.5
segment I (length/width)
1.6/2.2
segment II (length/width)
1.8/2.0
segment III (length/width)
1.9/2.0
segment IV (length/width)
2.4/2.0
segment V (length/width)
3.8/2.0
Telson, length
3.5
Vesicle (length/width)
2.3/1.6
depth
1.2
Aculeus, length
1.2
Pedipalp
Humerus (length/width)
2.5/1.0
Brachium (length/width)
2.8/1.2
Chela (length/width)
4.1/1.1
depth
1.2
movable finger, length
2.4
fixed finger, length
1.7
Pectines
teeth (left/right)
14/14
Metasoma .—Dorsal and dorsolateral keels of segments I to III irregularly crenu-
late; dorsal keel of segment IV crenulate, dorsolateral keel of IV smooth to crenu-
late; dorsolateral border of segment V essentially smooth; dorsal and dorsolateral
keels of segments II and III terminate posteriorly in an enlarged spine. Inferior
lateral keels irregularly smooth and faint on segments I to IV, inferior median
keels obsolescent on segments I to IV. Inferior intercarinal spaces of segment V
with many granules.
Telson .—Vesicle smooth and lustrous with several punctate depressions. Aculeus
very short, broadly attached to vesicle. Very subtle suhaculear tubercle.
Chelicerae .—Inferior border of movable finger completely lacking denticles.
Pedipalps .—Palm slightly swollen inwardly. Palm surface smooth and lustrous,
lacking keels. Fingers internally unscalloped, opposing teeth of fingers meet
closely along entire length when chela closed.
Standard measurements .—Table 1.
Type DATA. —Holotype female, “Texas, North America (San Diego),
W. Taylor 1889.” Type depository, British Museum. Type specimen
poorly preserved, telson essentially separated from metasoma, one pedi-
OCTOBER 1970 ] PARKER-MICRODYNERUS REVIEW
241
palp and one walking leg completely separated from prosoma. Specimen
darkened from poor preservation.
The original description was very brief, composed of two short para¬
graphs and not including many important characters. The details of this
redescription essentially agree with those of the original description in
all but the following ways: pectinal teeth 14 per comb, not 15; total
length 35 mm, not 25 mm (however, some of the membranes appear to
be stretched now). Pocock did not define the sex which is clearly a
female and probably mature.
Hoffmann (1931) discussed a population which he called Vejovis
bilineatus Pocock, with some reservation, from the state of Agua-
scalientes, Mexico. Hoffmann had neither opportunity to study the type
nor to see material from near the type locality in Texas. Comparison
of the holotype with Hoffmann’s careful diagnosis indicates his speci¬
mens from Aguascalientes, Mexico were Vejovis bilineatus.
Literature Cited
Hoffmann, C. C. 1931. Los scorpiones de Mexico. Primera parte, Diplocentridae,
Chactidae, Vejovidae. (In) Monografias para la entomologia medica de
Mexico. An. Inst. Biol. Univ. Nac. Auton. Mex., 2(4) : 291-408.
Pocock, R. I. 1898. The scorpions of the genus Vaejovis contained in the collec¬
tion of the British Museum. Ann. Mag. Natur. Hist., Ser. 7, 1: 394-400.
New North American Microdynerus with Notes
on the Nests of Two Species
(Hymenoptera: Eumenidae)
Frank D. Parker
Entomology Research Service, Agr. Res. Serv., USDA,
Columbia, Missouri 65201
The North American species of Microdynerus Thomson have been
treated as species groups in the genus Leptochilus Saussure (Bohart,
1955). Parker (1966) removed the trinodus and lissosomus groups
from Leptochilus and placed these species in the European genus Micro¬
dynerus. Additional specimens have accumulated since Bohart’s paper
was presented and among this material were the 4 new species described
below.
The nesting habits of the Nearctic species are undescribed, except
The Pan-Pacific Entomologist 46: 241-253. October 1970
242
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Parker and Bohart (1968) recorded 2 species in this genus that nested
in trap stems.
BIOLOGY
M. BAKERIANUS (Cameron)
(Figs. 51, 52)
Nesting site. —All nests of M. bakerianus were recovered from trap
stems with a bore diameter of Vk—ViQ in. Holes drilled in the end and side
of the trap stems were utilized by these wasps.
Nest construction. —The females did not modify the nest, except for
removing pith. The prey was placed rather loosely at the end of the
burrow. A partition was made above the prey by cementing pieces of
gravel and sand with what appeared to be a salivary secretion. Addi¬
tional prey was stored above the partition and another made so that a
series resulted. The length of the cells varied from 3 to 8 mm; generally
shorter cells were found in wider burrows.
The number of cells found in each nest varied from one to seven.
There were three nests with one cell, 22 with two, 27 with three, four
with four and one with seven.
The last partition in most nests served as the entrance plug; which
was usually located slightly below the entrance. A typical M. bakerianus
nest is shown in Fig. 51.
Prey. —Lepidoptera larvae.
Larval habits. —The cocoon was formed by lining the cell with opaque
silk that adhered to the cell walls.
Seasonal occurrence. —Collecting records indicate that there are at
least 2 generations a year.
Sex ratio. —From the nests observed, 13 males and 22 females emerged.
Parasites. —Fourteen (9.2%) of the 75 M. bakerianus cells found con¬
tained parasites. In 22 (14.5%) additional cells the wasp failed to
emerged; some of these may also have been parasitized. The following
six species of parasites and one predator were reared from M. bakerianus
nests.
Chrysididae. —A nest of M. bakerianus , collected from 9 mi. S. of
Reno, Nevada, during the winter of 1964, contained three cells from
which emerged: cell A, female M. bakerianus ; cell B, male Chrysis
parkeri Moore; cell C, male Chrysis stenodyneri Krombein. At the same
locality the next winter, three cells from three separate nests were
parasitized by Chrysis pattoni Aaron.
All these chrysidids probably ate the cell provisions and/or early host
instars, because no host cocoons were observed in parasitized cells.
OCTOBER 1970 ] PARKER-MICRODYNERUS REVIEW
243
Bombyliidae .—Two species of bombyliids were reared from M.
bakerianus nests: Toxophora virgata Osten Sacken and Anthrax irroratus
Say. The beeflies killed the prepupal stage of the host. In one nest
collected during the winter of 1965, one cell contained a tiny larva and
another a mature larva of Toxophora virgata. This suggests that T.
virgata overwintered in either a late or early instar.
Mutillidae .—One cell in a M. bakerianus nest from Banning, Cali¬
fornia, contained an unidentified species of mutillid that had killed the
prepupa.
Predator .—A larva of Trichodes ornatus Say was found in one of 3
cells in a nest from Banning, California.
Supersedure .—One bakerianus nest from Patrick, Nevada was super¬
seded by a nest of Solierella sp. which had emerged before the nest
was collected. Three M. bakerianus nests were constructed above bee
nests, two above Proteraides bullifacies (Michener) and one above
Ashmeadiella sp.
M. singulus (Bohart)
Nesting site .—Two nests of M. singulus were recovered from trap
stems with a bore diameter of Vig in.; both nests were in holes drilled in
the end of the stem.
Nest construction -—In one nest the cells were initiated 11 mm above
the bottom of the burrow and in the other the first cell was started at
the end of the burrow. Cell partitions and entrance plugs were made
by cementing pieces of gravel and sand with what appeared to be a
salivary secretion. Additional prey were stored above the partition and
another partition was made. The length of the cells varied from 7 to
10 mm. The number of cells found in each nest was two.
Prey .—Lepidoptera larvae.
Larval habits .—-The cocoon was formed by lining the cell with opaque
silk that adhered to the cell walls.
Seasonal occurrence .—Collecting records indicate that there are at
least 2 generations a year.
Parasites .—None of the M. singulus nests contained parasitized cells.
SYSTEMATICS
Key to Species of North American Microdynerus
Males
1. White or yellow markings extending at least % length of scape_ 6
Scape not pigmented or with small basal spot _ 2
2. Antennal hook thick, blunt (Figs. 2, 11) _ 3
Antennal hook slender, pointed (Figs. 1, 4) _ 5
3. Upper propodeal lamellae blunt apically, in profile, lower lamellae without
244
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
apical notch (Fig. 18) ; frons swollen between midocellus and compound
eye (Fig. 2) _ monolobus (Bohart)
Upper propodeal lamella pointed apically, lower lamella with apical notch
(Fig. 14) ; frons not swollen between midocellus and compound eye
(Figs. 5, 8) - 4
4. Vertex with 3 shiny, knob-like swellings, one between lateral ocelli, one at
upper posterior margins of each compound eye; vertex round in profile,
(Fig. 5) _ trinodus (Bohart)
Vertex with single shiny swelling between lateral ocelli; vertex flat in pro¬
file, (Fig. 8) ___ singulus (Bohart)
5. Head 1% times longer than broad (Fig. 4) ; upper propodeal lamella blunt,
short; tergite II without lateral spot connected to apical band -
_ hurdi Parker, n. sp.
Head as wide as long (Fig. 1) ; upper propodeal lamella slender, pointed;
tergite II with lateral spot connected to apical band_ bolingeri Parker, n. sp.
6. Antennal hook thick, round apically, U/4 times as long as flagellomere IX
(Fig. 12) ___ bakerianus (Cameron)
Antennal hook slender, pointed apically, not longer than flagellomere IX
(Figs. 13, 16) _ 7
7. Antennal hook minute, only % length of flagellomere IX (Fig. 16) ; flagel-
lomeres VII-IX thick, wider than long; vertex without tubercles, abdomen
shiny, without deep macropunctures _ 8
Antennal hook as long as flagellomere IX (Fig. 13), apical flagellomeres as
long or longer than broad; vertex with tubercles; abdomen with deep large
macropunctures _ 9
8. Apical clypeal margin with small, deep U-shaped notch (Fig. 6) _
_ lissosumus (Bohart)
Apical clypeal margin broadly excised (Fig. 17 _ inusitatus Parker, n. sp.
9. Body marked with red _12
Body without red markings_10
10. Lateral spot on tergite II isolated or attached to apical band; diameter of
midocellus 4 times width of macropunctures on frons; propodeum armed
with small tubercle _ schlingeri (Bohart)
Tergite II without lateral spot; diameter of midocellus twice width of macro¬
punctures on frons; propodeum armed with prominent tooth or carina_11
11. Frons, vertex densely pitted, margin of macropunctures connected (Fig. 49) ;
in profile, tergite I without subapical indentation; pronotal lamella raised
medially_ arenicolus (Bohart)
Frons, vertex with scattered pits, punctures not uniformly connected; in pro¬
file, tergite I indented subapically; pronotal lamella not raised medially
_-_ pcctagoniae Parker, n. sp.
12. Sternite II with median basal depression (Fig. 27) _13
Sternite II without depression (Fig. 25) _14
13. Pronotal macropunctures as large as lateral ocellus; flagellomeres dark
apically, ventrally; dorsal surface of tergite II shiny_ bechteli (Bohart)
Pronotal macropunctures not more than *4 diameter of lateral ocellus; flagel¬
lomeres yellow ventrally; dorsal surface of tergite II finely shagreen _
_ cavatus (Bohart)
14. Lateral swellings on vertex prominent; frons between compound eye and
OCTOBER 1970 ]
PARKER—MICRODYNERUS REVIEW
245
midocellus swollen (Fig. 7) ; propodeum mostly red; clypeus about as
long as wide (Fig. 19) _ gibboceps (Bohart)
Frons, vertex without raised swellings (Fig. 3) ; propodeum mostly black;
clypeus wider than long (Fig. 20) _15
15. Apical clypeal margin broadly, shallowly excised (Fig. 10) ; _
_ sayi (Cameron)
Apical clypeal margin angularly excised (Figs. 20, 36) _ umbifer (Bohart)
Females
1. Sternite II with median basal depression (Fig. 28) _ 2
Sternite II smoothly curved basally, without depression _ 4
2. Thorax black with yellowish-white markings; head as long as broad (Fig. 35) 3
Thorax red with white markings; head longer than broad (Fig. 34) _
_ hurdi Parker, n. sp.
3. Lower propodeal lamella with subapical notch (Fig. 47) ; sternite II shiny;
pronotal macropunctures as large as lateral ocelli _ bechteli (Bohart)
Lower propodeal lamella continuous with upper lamella (Fig. 18) ; sternite
II densely pitted, dull; pronotal macropunctures % as large as lateral
ocelli _ monolobus (Bohart)
4. Median clypeal lobe with shallow, semicircular emargination (Figs. 38, 46) ;
head longer than broad (Fig. 34) _ 5
Median clypeal lobe with deep, narrow V- or U-shaped emargination (Figs.
39, 40) _________ 9
5. Vertex with 2 or more shiny knob-like swellings (Figs. 29, 33) _ 6
Vertex without swellings, or if present, then low and punctured _ 7
6. Lower propodeal lamella subapically notched (Fig. 48) ; vertex with 3
swellings (Fig. 33) ; pronotum black with creamy-white markings _
_ cavatus (Bohart)
Lower propodeal lamella continuous with upper lamella (Fig. 37) ; vertex
with 2 swellings (Fig. 30) ; pronotum red with light markings_
--- sayi (Cameron)
7. Median clypeal lobe minute, not longer than diameter of midocellus (Fig. 46)
- lissosomus (Bohart)
Median clypeal lobe wide, at least 3 times diameter of midocellus_ 8
8. Body sculpturing smooth, shiny, macropunctures shallow; head as long as
broad; vertex not enlarged (Fig. 31) _ inusitatus Parker, n. sp.
Body sculpturing coarse, macropunctures deep; head longer than wide;
vertex enlarged (Fig. 29) - gibboceps (Bohart)
9. Vertex with swellings only between lateral ocelli _10
Vertex with swellings between lateral ocelli and compound eye; interocellar
area with or without swellings _11
10. Vertex swollen between lateral ocelli, depressed medially (Fig. 49) ; median
clypeal lobe with V-shaped notch; upper propodeal lamella broad; body
black with yellow to white markings_ arenicolus (Bohart)
Vertex with single shiny welt between lateral ocelli (Fig. 50) median clypeal
lobe with U-shaped emargination; upper propodeal lamella narrow, blunt;
body marked with red_ singuhis (Bohart)
11. Vertex with 3 shiny knob-like swellings _12
Vertex without distinct raised swellings_13
246
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
12. Median clypeal lobe as in Fig. 42; lower propodeal lamella continuous with
upper lamella; sternite II with lateral spot attached to apical band; thorax
black_ bolingeri Parker, n. sp.
Median clypeal lobe as in Fig. 39; lower propodeal lamella with subapical
notch; sternite II without lateral spot; thorax with red markings__
_ trinodus (Bohart)
13. Head as long as broad; lower propodeal lamella with subapical notch _
_14
Head longer than broad; lower propodeal lamella continuous with upper
lamella _ umbifer (Bohart)
14. Median clypeal lobe with U-shaped emargination (Fig. 43) marginal cell
stained_ schlingeri (Bohart)
Median clypeal lobe with V-shaped emargination (Fig. 44) marginal cell
with apical % stained_ bakerianus (Cameron)
Microclynerus bolingeri Parker, new species
(Figs. 1, 22, 42)
Male.— Black; the following with yellowish-white markings: clypeus, spot medi¬
ally on frons, spot in eye emargination, spot behind compound eye, mandible, leg
partly, divided spot basally on scutellum, anterior margin of pronotum, spot on
mesopleuron, tegula, apical band on tergites I, II, IV, V, tergite II with lateral
spot attached to apical band, sternite II with apical band; the following bright
red: ventrally on flagellomeres, legs partly, summit of tergite I, wing veins basally.
Punctation close, coarse; pits on frons, vertex close, deep, vertex with 3 shiny,
knob-like areas above and behind compound eye and between lateral ocelli; rest
of body rather uniformly pitted, pits close, deep, surface between pits with fine
microsculpture; tergite II finely micropunctate basally, coarsely macropunctate
apically. Pubescence white, short, erect on head, notum; setae mat-like on rest
of thorax; abdomen with fine, short, appressed setae. Median clypeal lobe with
deep V-shaped excision; flagellomeres about as broad as long apically; antennal
hook thin, pointed, reaching base of flagellomere IX; vertex with 3 raised knob¬
like swellings; head as long as broad; pronotal lamella raised along entire margin;
hind face of propodeum with stout shiny lateral swellings; upper propodeal lamella
projecting, blunt apically, lower lamella rounded, slightly indented apically; in
profile, abdominal segment II broader than long; genitalia (Fig. 22) ; length 6 mm.
Female.— Essentially as for male except: mandibles, apical clypeal margin black.
Figs. 1-5. Front or side view of heads of Microdynerus males: 1, M. bolingeri ;
2, M. monolobus; 3, M. sayi ; 4, M. hurdi; 5, M. trinodus. Fig. 6. Clypeus male
M. lissosomus. Figs. 7-8. Side view of heads of Microdynerus males: 7, M. gib-
boceps ; 8, M. singulus. Fig. 9. Apical flagellomeres of male M. bechteli. Fig. 10.
Clypeus of male M. sayi. Figs. 11-13. Apical flagellomeres of Microdynerus males:
11, M. monolobus ; 12, M. bakerianus ; 13, M. sayi. Fig. 14. Propodeal outline of
male M. inusitatus. Figs. 15-16. Apical flagellomeres of Microdynerus males:
15, M. trinodus-, 16, M. lissosomus. Fig. 17. Lower part of head of male M. inusita¬
tus. Fig. 18. Propodeal outline of male M. monolobus. Figs. 19-20. Lower part
of heads of Microdynerus males: 19, M. gibboceps ; 20, M. umbifer.
OCTOBER 1970 ]
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20
243
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
Holotype male and 3 paratypes (2$, 12) 20 mi. W. Durango,
Durango, Mexico, June 1964 (D. Bolinger), 7,100 ft. Holotype de¬
posited in the type collection of the California Academy of Sciences.
Superficially M. bolingeri and M. schlingeri are similar in size and
markings. The red markings and narrow clypeal excision will separate
the former species.
Microdynerus inusitatus Parker, new species
(Figs. 14, 17, 24, 31, 45)
Male.— Black; the following with creamy white markings: mandibles basally,
clypeus, scape in front, inner orbital spot, postocular spot, front margin of pro-
notum, mespoleural spot, tegula except central spot, 2 spots on scutellum, legs
partly, apical band on tergites I-IV, lateral spots on sternite II; light red ventrally
on flagellomeres, legs partly, abdominal segment I, most of II, basally on III-IV;
wings stained light brown. Punctation not uniform; micropunctures rather shallow;
pits on head, thorax shallow, not well formed, widely spaced; pits on abdomen ob¬
scure. Clypeal lobe truncate apically (Fig. 17); antennal hook thin, needle-like reach¬
ing base of flagellomere IX; head broad, ocelli large; pronotal lamella not raised
laterally, medially; summit of metanotum round in profile; upper, lower propodeal
lamellae continuous, not indented; in profile, tergite I weakly depressed sub-
apically; tergite II not indented subapically in profile; genitalia (Fig. 24) ;
length 5 mm.
Female.— Essentially as for male except: less white on clypeus, abdomen.
Holotype male, “Gavid” = the Gavilan?, California, 2 April 1956
(P. H. Timberlake). Paratype female, Perris, California, 14 May 1948
(P. H. Timberlake). Holotype in the type collection of the California
Academy of Sciences.
The smooth body sculpturing and the round metanotum separate M.
inusitatus and M. lissosomus from other Nearctic Microdynerus. Dif¬
ferences in clypeal and mandibular configuration will separate M.
inusitatus (Figs. 14, 31) from M. lissosomus (Figs. 6, 32).
Microdynerus hurdi Parker, new species
(Figs. 4, 21, 27, 34)
Male.— Black; the following with white markings: mandibles, clypeus, medio-
lateral spot on frons, postocular spot, anterior margin of pronotum, mesopleural
spot, tegula, divided hand on scutellum, legs partly, apical hand on tergites I—II,
Figs. 21-24. Genitalia of Microdynerus males: 21, M. hurdi ; 22, M. bolingeri ;
23, M. patagoniae ; 24, M. inusitatus. Figs. 25-28. Abdominal segments I—II of
Microdynerus spp.: 25, male M. sayi; 26, female M. sayi; 27, female M. hurdi-,
28, male M. cavatus.
OCTOBER 1970 ]
PARKER-MICRODYNERUS REVIEW
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250
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
medially on III, spot laterally on sternite II; the following bright red: flagello-
meres, pronotum, parts of mesopleuron, metanotum, legs partly, propodeum, sternite
II, abdomen except sternite II laterally, most of tergite II. Punctation moderate;
pits on head, thorax deep, separated by about their diameter; sternite II coarsely
pitted medioapically. Pattern and color of pubescence same as M. bolingeri.
Clypeus with wide U-shaped median excision, antennal hook slender, pointed,
reaching base of flagellomere IX; head longer than broad; interocellar area swollen
with flat summit, lateral ocelli sloping toward compound eye; single knob-like
swelling between lateral ocelli, compound eye; pronotal lamella raised laterally,
fading medially; propodeal armature weak, punctured; upper, lower propodeal
lamellae merging; abdominal segment II as long as wide; sternite II with medio¬
basal depression; genitalia (Fig. 21) ; length 4.5 mm.
Female.-— Essentially as for male except: mandibles, clypeus, thorax bright red.
Holotype male, 18 mi. N. Rodeo, Hildalgo County, New Mexico,
25 August 1958 (C. G. Moore). Paratypes, 1$, 2 2 from 18 mi. E.
Douglas, Cochise Co., Arizona, 5 September 1958 (P. D. Hurd),
Eriogonum abertianum neomexicanum. Holotype deposited in the type
collection of the University of California, Davis campus.
Metatypes, 9 from the following North American localities: Arizona:
12 mi. N. Yuma, 2 October 1958 (T. R. Haig) ; 5 mi. N. Wickenburg,
11 September 1957 (T. R. Haig) ; Madera Canyon, 13 July 1958 (R. M.
Bohart) ; 17 mi. E. Douglas, 4 August 1958 (R. M. Bohart) ; Tucson,
24 October 1939 (R. H. Crandall) ; 29 mi. E. Ajo, 20 September 1966
(R. Rust). New Mexico: 5 mi. N. Rodeo, 21 August 1962 (Rozen,
Statham, Hessel) ; 11 mi. N. Rodeo, 2 August 1961 (J. G. Rozen),
Euphorbia ; 4.8 mi. N. Rodeo, 4 September 1961 (P. D. Hurd), Sphaer-
alcea angustifolia var. cuspidata. Baja California: La Paz, 7-8 Octo¬
ber 1955 (F. X. Williams). Sonora: Empalme, 26 October 1965 (G. E.
and A. S. Bohart) ; 10 mi. E. Navajoa, 13 August 1959 (Nutting &
Werner).
The long head, distorted interocellar area and the shiny knobs on the
vertex distinguish M. hurdi from other Nearctic Microdynerus. The
bright red females could be mistaken for the red color forms of both
Figs. 29-35. Front or side view of heads of Microdynerus females: 29, M.
gibboceps; 30, M. sayi; 31, M. inusitatus; 32, M. lissosomus ; 33, M. cavatus; 34,
M. hurdi ; 35, M. bechteli. Fig. 36. Clypeus of female M. umbifer. Fig. 37. Pro¬
podeal outline of female M. sayi. Figs. 38-46. Clypeus of Microdynerus females:
38, M. cavatus ; 39, M. trinodus; 40, M. arenicolus ; 41, M. singulus; 42, M. bo¬
lingeri; 43, M. schlingeri; 44, M. bakerianus; 45, M. inusitatus; 46, M. lissosomus.
Figs. 47-48. Propodeal outlines of Microdynerus females: 49, M. arenicolus; 50,
M. singulus.
OCTOBER 1970 ] PARKER-MICRODYNERUS REVIEW
251
50
252
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
M. smgulus and M. bakerianus. The longer head of M. hurdi will sepa¬
rate it from the latter species.
Microdynerus patagoniae Parker, new species
(Fig. 23)
Male. —Black; the following yellowish-white markings: mandibles, clypeus,
scape in front, medial bar on lower frons, spot in eye emargination, postocular spot,
anterior margin of pronotum, mesopleural spot, tegula partly, band on scutellum,
apical band on tergite I, II, IV, V, sternite II; flagellomeres yellowish beneath;
wings lightly stained, marginal cell dark apically. Punctation not uniform; macro¬
punctures deep, scattered except tergite II with shallow, obscure, close pits.
Pubescence as for M. bolingeri. Median clypeal lobe with deep U-shaped excision;
antennal hook thin, pointed, reaching base of flagellomere IX; head longer than
broad; interocellar area raised, distorted, ocelli slanted; lateral tubercle on vertex
indistinct; pronotal lamella not raised medially; semicircular carina on upper
lateral surface of propodeum; propodeal lamella fused, not indented; in profile
tergite II with subapical indentation; tergite II smooth in outline; genitalia (Fig.
23) ; length 4.3 mm.
Female. —Unknown.
Holotype male , Patagonia, Santa Cruz County, Arizona, 10 August
1958 (F. G. Werner). Holotype deposited in the type collection of the
California Academy of Sciences.
This species is closely related to M. arenicolus, but the two can be
separated by differences in sculpture on the frons and vertex. The con¬
figuration of tergite I and the upper propodeal lamella are additional
characters that differentiate these species.
Microdynerus arenicolus (Bohart)
(Figs. 40, 49)
Leptochilus arenicolus Bohart, 1955: 299.
This species was described from a series of specimens collected at
Antioch, California. Additional material has extended the range of M.
arenicolus to include the following localities: California: Dutch Flat,
San Jacinto Mts., 14 August 1934 (C. D. Michener). Arizona: 13 mi.
N. W. Nogales, 8 September 1967 (T. R. Haig) ; 2.5 mi. W. Pena
Blanca, 26 August 1961 (Noller, Bequaert, Santana) ; 5 mi. W. Bisbee,
15 August 1958 (P. M. Marsh) ; 5 mi. E. Fort Apache, 28 August 1964
(M. E. Irwin) ; S. W. Research Station, 6 September 1958 (P. D. Hurd) ;
Rucker Cyn., Chiricahua Mts., 25 September 1961 (J. C. Bequaert).
New Mexico: Rodeo, 6 August 1958 (P. M. Marsh).
The markings on the Arizona and New Mexico specimens are ivory-
white whereas California specimens tend to be a creamy-yellow.
OCTOBER 1970 ] PARKER—MICRODYNERUS REVIEW
253
Figs. 51-52. Nests of M. bakerianus in trap stems: 51, 4-celled nest made in
side of stem. Note gravel entrance plug. 52, prey in 2-celled nest.
MlCRODYNERUS BECHTELI (Bohart)
(Figs. 9, 35, 47)
Leptochilus bechteli Bohart, 1955: 294--5.
Bohart (1955) recorded a single specimen from Red Rock, Arizona;
additional distribution records from this state are: 2 mi. N. E. Portal,
30 August 1959 (M. Statham) ; 7 mi. N. W. Gila Bend, 10 April 1961
(R. H. & E. M. Painter).
Literature Cited
Boiiart, R. M. 1955. North American Leptochilus of the trinodus group. Proc.
Entomol. Soc. Wash., 57: 287-299.
Parker, F. D. 1966. A revision of the North American species in the genus
Leptochilus. Misc. Publ. Entomol. Soc. Amer., 5: 151-229.
Parker, F. D., and R. M. Bohart. 1968. Host-parasite association in some twig¬
nesting Hymenoptera from western North America. Pan-Pac. Entomol.,
44: 1-6.
254
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Coexistence of Desert Scorpions by Differential
Habitat Preference
Stanley C. Williams
Department of Biology, San Francisco State College, California 94132
The scorpion fauna of Arizona is rich both in abundance and in
number of species. About 19 species have been reported from Arizona
in the literature, and these represent four of the six living families in
the order Scorpionida.
Most of the published research on scorpions has been concerned with
systematics, behavior and venoms. Ecological research has been con¬
spicuously lacking in the literature, with population and community
studies being especially uncommon. Baerg (1954) recognized popula¬
tion periodicity in relation to reproduction and seasonal activity while
observing Jamaican scorpions. Alexander and Ewer (1958) discussed
the use of temperature-adaptive behavior. Sreenivasa-Reddy (1959)
discussed the function of the pectines, and analyzed pectine morphology
in relation to burrowing and humidity. Cloudsley-Thompson (1956,
1961, 1962, 1964) discussed biological rhythms, heat death and adapta¬
tion to desert environments. Gertsch and Allred (1965) made an eco¬
logical contribution by showing habitat affinities, relative abundances
and sex ratios of nine species occurring on the Nevada Test Site.
McAlister (1966) discussed the aggregating tendency of Centruroides
vittatus (Say) and Smith (1966) discussed the life history of Urodacus
abruptus Pocock with emphasis on its home sites. Williams (1966) dis¬
cussed the obligate burrowing activities of Anuroctonus pliaeodctctylus,
and related this to its ecology. Williams and Hadley (1967) and Hadley
and Williams (1968) discussed the habitat preferences, community
compositions and nocturnal activities of some populations occurring in
Sonora, Mexico.
The purposes of this project were two-fold. The first was to study
the influence of several different habitat types on the abundance, dis¬
tribution and community structure of scorpions. The second was to
analyze the changes in scorpion distribution and abundance which occur
from year to year.
The field work was carried out at Phoenix South Mountain, Maricopa
County, Arizona from September 1964, to January 1967.
Special thanks are due to Mont A. Cazier, David I. Rasmussen,
Wendell L. Minckley, and Frank F. Hasbrouck of the Department of
Zoology, Arizona State University, for their advice and suggestions.
The Pan-Pacific Entomologist 46: 254-267. October 1970
OCTOBER 1970] WILLIAMS—COEXISTENCE OF DESERT SCORPIONS 255
Thanks also are due William S. Parker lor assistance in the field, to
Charlene F. Williams, Christie Steketee, and William Azevedo for cleri¬
cal help. This project was partially supported by the National Science
Foundation through research grant GB 7679, and by the National
Institutes of Health through predoctoral fellowship number 5-F1-GM-23,
794-02.
Materials and Methods
An area approximately three square kilometers in size, located at the
eastern end of Phoenix South Mountain (30° 26' N. Lat., 112° 01' W.
Long.), Maricopa County, Arizona, was selected for the field study.
This area begins about one kilometer west of the town of Guadalupe,
along Guadalupe Road. Selection of this study area involved the follow¬
ing considerations: the ecology was relatively undisturbed; access was
convenient; several distinct desert habitats of interest were represented
in close proximity; and, the scorpion fauna was abundant and diverse.
The general study area was composed of four subareas, each repre¬
senting a distinct ecological situation. These subareas differed from one
another in topography, soil composition, slope, plant community and
sun exposure. Essentially, they represented the following ecological
situations: an open desert basin near the base of a mountain; a moun¬
tain canyon slope with a north-facing exposure; a mountain canyon
floor; and a mountain canyon slope with a south-facing exposure.
Sampling was carried out exclusively by means of unbaited pitfall
traps, which were in place for the duration of the study. Number 10
food cans served as traps. These were deep enough to prevent the escape
of even the largest scorpion species, and to retain the captured scorpions
well below the environmental extremes occurring on the soil surface.
Each can was buried so that its open end was flush with the ground
surface, and was covered by a suitably sized rock. The trapping method
employed has been described in detail by Williams (1968a).
A sampling scheme was designed so the four subareas studied could
be compared by equivalent trap catch units. For this purpose, the pooled
catch of twenty traps was used as a sample unit. Twenty traps were,
therefore, established in each subarea. In addition, the data from all
subareas could be pooled together to give one sample unit of 80 traps
which would reflect the total scorpion activity of the whole area during
any given time interval. Because of the seasonal periodicity of the
populations in surface activity, the data for trapping samples were
pooled for analysis, by 52-weeks. All data were gathered during 104
consecutive weeks from 29 December 1964 through 25 December 1966.
256
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
The assumption was made that relative catch numbers were an index
of relative abundance for Hadrurus arizonensis (Ewing), Vejovis con-
fusus Stahnke and Vejovis spinigerus (Wood).
Traps were checked at seven day intervals throughout the year to
preserve the equivalence of catch units. The pattern of trap distribution
employed was a line-transect along which the traps were placed at vary¬
ing intervals but never closer than 10 meters apart. Scorpions were
removed and preserved (Williams, 1968b) for future reference.
Throughout the study, trap maintenance was carried out as needed.
This maintenance included removal of soil and animals from traps, re¬
moval of water after rain, keeping the lip of the trap flush with the soil
surface, replacement of traps as they became rusty, and keeping trap
sides clean and smooth in order to prevent escape.
In addition to the 80 traps sampling the subareas, 100 additional traps
were established in other similar areas to test the effect of scorpion
removal upon subsequent catches and to test the influence of trap covers
on catches (Williams, 1968a).
Results
Description of study area. Phoenix Mountain is a long, northeast
to southwest oriented mountain range, isolated from neighboring moun¬
tains by expanses of flat desert basin. At the eastern foot of the moun¬
tain the elevation is 382 meters above sea level while the highest peak
reaches an elevation of 758 meters above sea level. The mountain is
rugged, rocky and dissected by many irregular valleys and canyons.
The mountain belongs to the Basin and Range Physiographic Province
of Arizona and is of heterogeneous geologic structure and origin. Most
of the eastern end of the mountain is composed of metasedimentary rock
of Precambrian origin represented by gneiss and schist associated with
dikes of metamorphosed basalt. The remainder of the surface is domi¬
nated by broad felsic dikes of Tertiary origin and subangular, poorly
indurated gravel Quaternary origin (Avedisian, 1966). The soils of
the mountain slopes are generally shallow while those of the open desert
surrounding the mountain are deeper, often stony and with a relatively
water-impervious, subsurface mineral layer.
Climatic studies (Kangieser, 1966) provide the following data for
the past 28 years. Air temperature extremes during this period ranged
from —13° to 46° C with the lowest temperatures in December and
January and highest temperatures during June, July and August. Rain¬
fall was very irregular from year to year, but generally was greatest in
August, and varied during this month from 0.17 to 14.12 cm with an
OCTOBER 1970] WILLIAMS—COEXISTENCE OF DESERT SCORPIONS 257
average of 2.84 cm. May and June were the months of lowest rainfall,
with monthly rainfall varying from 0.00 to 2.41 cm.
The daily high temperature coincided with the occurrence of the daily
low relative humidity, this point occurring at about 4:00 p.m. (MST)
during the summers. The daily low temperature coincided with the
occurrence of the daily high relative humidity, this point being reached
at about 6:00 a.m. (MST) during the summers.
A number of distinct environmental situations occur because of the
physical influence of the mountain. As the mountain is ascended, a
heterogeneous array of habitats differentiated by various sun exposures,
degrees of slope, water drainages, soil types and plant associations occur.
Ecotone effects occur as judged by plant heterogeneity, where two habi¬
tat types intergrade. Such ecotones are apparent in the canyon bottoms
and where the mountain grades into open desert.
To study how the various habitats existing at South Mountain affect
the distribution and abundance of scorpion populations, four distinct
habitats were sampled for comparison. These are described below.
Open desert basin. —This area was characterized by a Larrea-Franseria
community on open, relatively flat desert (Fig. 1). Since the plant com¬
munity was more diverse and complex than would have been expected,
this area perhaps showed some ecotone effect.
The site of this study area was one kilometer west of the town of
Guadalupe, along Guadalupe Road. The terrain was basically flat but
was occasionally divided by shallow washes. The soils had a texture
similar to sandy loam and were well compacted. In most places a hard,
subsurface, mineralized layer was present.
The vegetation was dominated by creosote bush ( Larrea tridentata
(D.C.) Coville) and bur-sage ( Franseria deltoidea Torr.), but, paloverde
(Cercidium microphyllum (Torr.) Rose and Johnston), ironwood
('Olneya tesota Gray), white bur-sage ( Franseria dumosa Gray), Mor¬
mon tea ( Ephedra sp.) and saguaro ( Carnegiea gigantea (Engelm.)
Britt, and Rose) also occurred, although in less abundance.
North-facing slope. —This study area was located 1.5 kilometers west
of the open desert basin study area, and was a canyon side with a due
north exposure (Fig. 2). The terrain had a slope approximating 25°.
Plants were more abundant and larger in this habitat than in other
nearby habitats. The vegetation was bushy, but grasses, mosses, liver¬
worts, and a number of annuals were also common. The most abundant
and characteristic plants were: bur-sage, brittle bush ( Encelia farinosa
Gray) and paloverde. The following plants also occurred, but were
less abundant: creosote bush, white bur-sage, apricot mallow ( Sphaeral -
258
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
F
Fig. 1. Open desert basin habitat. In the background, the end of Phoenix South
Mountain may be seen. Fig. 2. North-facing slope habitat. This habitat is charac¬
terized by a high degree of shielding from sun exposure. Ground temperatures are
generally lower and soil moisture higher in this habitat throughout the year in
comparison to the other study areas.
cea ambigua Gray), ironwood, Mormon tea, wolf berry ( Lycium sp.),
ocotillo ( Fouquieria splendens Engelm.), saguaro, staghorn cactus
(Opuntia versicolor Engelm.), barrel cactus ( Ferocactus wislizeni
OCTOBER 1970 ] WILLIAMS—COEXISTENCE OF DESERT SCORPIONS 259
Fig. 3. South-facing slope habitat. This habitat is characterized by an extreme
exposure to the sun. Ground temperatures are higher and soil moisture is lower in
this habitat than in the others studied throughout the year.
(Engelm.) Britt, and Rose), Christmas cactus (Opuntia leptocaulis
D.C.), and teddy bear cholla (Opuntia bigelovii Engelm.).
The soil, although somewhat sandy, was well packed, considerably
finer in texture, and deeper than on the opposing slopes. Large rocky
outcrops occurred over the slopes, and between outcrops, the soil was
abundantly studded with smaller surface rocks.
South-facing slope .—The south-facing slope studied was the canyon
side directly opposite the north-facing slope study area (Fig. 3). This
hillside had a slope approximating 25° and was of the same height as
the opposite slope. Presumably, this area should have differed from
the north-facing slope only in sun exposure and in the environmental
effects caused by this intensive exposure.
The whole hillside was very rocky and large outcrops were abundant.
Soils were very shallow and coarse, consisting mostly of gravel. This
area was sparsely vegetated throughout most of the year, and the
weathering effects of the harsh sun were apparent.
The plant community of this habitat was distinctive because of the
reduction of both numbers and species of plants. Teddy bear cholla
was the dominant species and occurred in stands of varying density.
Creosote bush, brittle bush, barrel cactus and ocotillo were moderately
260
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
t i . v J. i
Fig. 4. Canyon bottom habitat. This photograph was taken from the north-facing
slope, looking down the canyon bottom. This was the most heterogeneous habitat
both in terms of physical and biotic diversity.
abundant, while paloverde, ironwood, wolf berry and saguaro were less
abundant.
Canyon bottom .—This was the intermediate area between the north-
and south-facing slope study areas, and was the most irregular and
physically diverse of all the study areas (Fig. 4). Here were found
rocky outcrops, alluvial soils, a deep sandy wash and a seasonal spring.
The physical heterogeneity was matched by the heterogeneity of the
flora. No one plant species appeared dominant; instead, the flora was
a mixture of those plant species found on the adjacent hillsides and
open desert floor.
Scorpion populations present at South Mountain. Eight scorpion
species from three families occurred at South Mountain. The Buthidae
OCTOBER 1970 ] WILLIAMS—-COEXISTENCE OF DESERT SCORPIONS 261
Table 1 . Relative numbers of scorpions captured in four habitats at
Phoenix South Mountain. Data were based on captures by 80 traps
during a two year interval (104 weeks). Each habitat was sampled by
20 traps.
Catch Numbers
H. arizonensis V. confusus V. spinigerus Habitat Total
Habitat 1965 1966 1965 1966 1965 1966 1965 1966 Total
desert basin
51
22
50
53
10
11
111
86
197
northern exposure
8
3
0
0
52
61
60
64
124
canyon bottom
23
12
2
1
53
75
78
88
166
southern exposure
25
26
6
0
40
61
71
87
158
species totals
107
63
58
54
155
208
320
325
645
was represented by one species, Centruroides sculpturatus Ewing.
The Vejovidae was represented by six species: Hadrurus arizonensis
(Ewing), Vejovis confusus Stahnke, Vejovis stahnkei Gertsch and Sole-
glad, Vejovis spinigerus (Wood), Vejovis wupatkiensis Stahnke, and
Vejovis lapidicola Stahnke. One small scorpion was found in the
stomach of a whip-tail lizard ( Cnemidophorus tigris Baird and Girard)
and was identified as a member of the Chactidae because of the occur¬
rence of six eyes. Due to poor physical conditions, this specimen was
identified only to family .
Only three species were captured in sufficient numbers to yield sig¬
nificant ecological information: V. spinigerus, V. confusus and H. ari¬
zonensis. Only the data on these three species were considered in sub¬
sequent ecological analyses unless otherwise mentioned. Centruroides
sculpturatus was found under rock chips and dead vegetation in low
numbers, especially on the northern and eastern exposures. Only two
specimens were collected by traps during the study. Five V. stahnkei
were collected by traps in open desert basin areas. One V. wupatkiensis
was collected in a trap on the southern exposure, and one V. lapidicola
was collected under a rock chip at the base of an eastern exposure.
During the two-year period, 645 scorpions were captured in the 80
traps (Table 1). No significant difference occurred between the total
number of scorpions captured in 1965 (320 scorpions) and those cap¬
tured in 1966 (325 scorpions) when the yearly totals were compared by
Chi Square Analysis (x 2 (i) = 0.024; P > 0.80).
Relative numbers of scorpions in relation to habitat type.
Examination of the total catch data indicated that scorpions were more
262
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Table 2. Comparisons of the relative numbers of scorpions captured
in each of four different habitats. The captures for all areas were placed
in an abundance array and adjacent numbers were compared by Chi
Square Analysis using Yates’ Correction for Continuity. These data
were obtained from 80 traps during 1965 and 1966 (104 weeks). Only
H. arizonensis, V. confusus and V. spinigerus were considered.
Habitat
Observed
Numbers
Expected
Numbers
Sample
Size
X 2 (d.f.)
P
Desert basin
197
181.5
363
2.48(d
0.20 > P > 0.10
Canyon bottom
166
181.5
Canyon bottom
166
162
324
0.152 ( d
0.70 > P > 0.50
Southern exposure
158
162
Southern exposure
158
141
282
3.86(d
P < 0.05
Northern exposure
124
141
abundant in some habitats than in others. The total scorpion catches
from each habitat, placed in an abundance array, gave the following rela¬
tionship: 197 (desert basin) ; 166 (canyon bottom) ; 158 (southern
exposure) ; and 124 (northern exposure). Analyses of these data gave
highly significant Chi Squares (x 2 (3) = 16.74; P<0.01), indicating
that scorpions were significantly more abundant in some habitats.
To ascertain which areas supported a more abundant scorpion fauna,
habitats which occupied adjacent positions in the abundance array were
compared by a series of Chi Square Analyses (Table 2). It then became
evident that three habitats, the desert basin, canyon bottom and southern
exposure, each supported not significantly different numbers of scor¬
pions and each supported a significantly more abundant scorpion fauna
than did the northern exposure habitat.
Differential relative abundances of the species in relation to
habitat. Examination of the catches (Table 1) suggested that the com¬
munity structure in regard to numerical dominance and relative abun¬
dances may have been unique to each habitat. Therefore, the relative
abundances of the different scorpion species were analyzed in each hab¬
itat separately.
Examination of the pooled, 104 week, scorpion catches clearly indi¬
cated that H. arizonensis, V. spinigerus and V. confusus were not equally
abundant in any habitat (Table 3). A very obvious, non-random, rela¬
tive abundance array occurred among the three species in each habitat.
Vejovis confusus was the numerically dominant species in the habitat
characterized by non-rocky, fine textured soils, while V. spinigerus was
OCTOBER 1970] WILLIAMS-COEXISTENCE OF DESERT SCORPIONS 263
Table 3. Comparison of the relative numbers of H. arizonensis,
V. confusus and V. spinigerus captured in each of four desert habitats.
Data pooled for 104 consecutive weeks, and analyzed by Chi Square
Analysis.
Observed
Numbers
Expected
Numbers
Sample
Size
Analysis
Open desert
V. confusus
103
65.7
197
x 2 (3) = 52.40; P<0.01
H. arizonensis
73
65.7
V. spinigerus
21
65.7
Northern exposure
V. spinigerus
113
62
124
x 2 (d — 82.26; P<0.01
H. arizonensis
11
62
Canyon bottom
V. spinigerus
128
55.3
166
x 2 (2) = 154.39; P<0.01
H. arizonensis
35
55.3
V. confusus
3
55.3
Southern exposure
V. spinigerus
101
52.7
158
x 2 (2) = 85.70; P<0.01
H. arizonensis
51
52.7
V. confusus
6
52.7
the dominant species in the rocky habitats (northern exposure, canyon
bottom, and southern exposure). The large species, H. arizonensis,
occurred in each habitat, but was never a numerically dominant species.
From the standpoint of relative abundance and numerical dominance
of the species, three types of community structures occurred in the four
habitats. The open desert was characterized by the numerical dominance
of V. confusus , the relative lack of the congeneric V. spinigerus, and
the intermediate abundance of H. arizonensis. The northern exposure
was unique in that V. spinigerus was the numerical dominant, H. ari¬
zonensis was relatively uncommon and V. confusus was absent. The
canyon bottom and southern exposure had essentially the same scorpion
community structures. Here V. spinigerus was always the numerical
dominant, H. arizonensis was common, and V. confusus was relatively
rare.
Changes in the population size of the species between 1965 and
1966. The population sizes of V. spinigerus, V. confusus, and H. arizo¬
nensis remained remarkably stable during the two years of the study. In
only two situations did the trap catches indicate a significant increase or
264
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Table 4. Comparison of the scorpion catches of 1965 with those of
1966 in each of four habitats at South Mountain. Each habitat was
sampled by 20 traps for 52 weeks each year. Yearly catches were com¬
pared by Chi Square Analysis using Yates’ Correction for continuity.
Catch Numbers
Habitat
Year
V.
confusus
V.
spinigerus
V.
arizonensis
Open desert:
1965
50
10
51
1966
53
11
22
X 2 d)
0.03
0.00
10.74
0.90 > P > 0.80 P = 1
P < 0.01
Northern exposure:
1965
0
52
8
1966
0
61
3
v 2
* (1)
—
0.57
1.45
—
0.50 > P > 0.30
0.30 > P > 0.20
Canyon bottom:
1965
2
53
23
1966
1
75
12
X 2 <i)
—
3.44
2.86
— •
0.10 > P > 0.05
0.10 > P > 0.05
Southern exposure:
1965
6
40
25
1966
0
61
26
-y2
(1)
-—
3.96
0.00
—
0.05 > P > 0.025
P = 1
decrease in species numbers (Table 4). In these cases, the H. arizonensis
catch decreased from 51 in 1965 to 22 in 1966 (P < 0.01) in the open
desert habitat, while V. spinigerus increased from 40 in 1965 to 61 in
1966 (P < 0.05) in the southern exposure habitat. The catches also
indicated that V. confusus may have been declining in the canyon bot¬
tom and southern exposure habitats but catch numbers were not high
enough to permit statistical analysis.
Discussion and Conclusions
Species diversity. —Eight scorpion species were found at South Moun¬
tain. Three species ( Hadrurus arizonensis , Vejovis confusus and Vejovis
spinigerus ) were very abundant, while the other five ( Vejovis lapidicola,
Vejovis stahnkei, Vejovis wupatkiensis, Centruroides sculpturatus and
the unidentified chactid species) were rare. This same relationship
between the abundant and rare species was reported in the scorpion
fauna of the Nevada Test Site (Gertsch and Allred, 1965).
OCTOBER 1970] WILLIAMS-COEXISTENCE OF DESERT SCORPIONS 265
The South Mountain scorpion fauna (eight species) compared favor¬
ably in diversity with the regional scorpion faunas reported in other
North American areas. Nine species were reported from the Nevada
Test Site (Gertsch and Allred, 1965), five species were reported from
Puerto Penasco, Sonora, Mexico (Williams and Hadley, 1967), and five
species were reported from Durango, Durango, Mexico (Baerg, 1929).
Based on these regional reports, it appears that even diverse scorpion
faunas are composed of a relatively few species in North America. The
record for scorpion diversity is the nine species reported from the
Nevada Test Site.
The regional species diversity of scorpions is considerably less than
that reported for other related arachnid groups. For example, 14 species
of Araneida were reported from Portal, Arizona (Chew, 1961) while 28
species of Solpugida were reported from the Nevada Test Site (Gertsch
and Allred, 1965, citing Muma, 1963). The most likely explanation for
the occurrence of scorpion faunas composed of so few species as com¬
pared with other related groups was the apparent similarity, within the
group, of food sources, habitat requirements, morphology and general¬
ized behavior. No spectacular ecological radiation has occurred in the
Scorpionida as compared with the Araneida, Acari and Insecta. The
similarities among species of the Scorpionida should tend to cause
competitive interactions among these species. Assuming that this were
true, the limited number of scorpion species within a regional fauna
could be explained by competitive exclusion. The coexistence of scorpion
species does indicate, however, that at least a few mechanisms exist
which permit escape from competitive exclusion.
The two main mechanisms permitting coexistence were habitat
specialization and choice of different sized prey. Hadrurus arizonensis
avoided competition with the other species by capturing prey too large
to be useful to other species. The other species that feed on the same
kinds and sizes of prey avoided competition by occupation of different
types of habitat. Vejovis conjusus and V. spinigerus clearly utilized the
available space at South Mountain in such a way that V. conjusus domi¬
nated the open desert habitats surrounding the mountain while V.
spinigerus was dominant on the rocky slope and in the canyons of the
mountain. The spatial distribution of these two species did overlap to
some extent, but they never shared the dominant position in any given
habitat. Such spatial overlap of these two congeneric species was per¬
haps due to some ecotone effect which occurs at the base of desert moun¬
tains and their peripheral canyons.
Relative abundance and habitat preference. —Each of the four habitats
266
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
had a unique and well structured scorpion community, characterized by
species composition and relative abundances. Scorpions were not equally
abundant in each of these habitats. They were significantly more abun¬
dant in the open desert, canyon bottom and southern exposure habitats
than in the northern exposure habitat.
Hadrurus arizonensis was moderately abundant in all habitats except
on the northern exposure. The abundance was greatest in open desert
habitats characterized by the Larrea-Franseria plant association. Gertsch
and Allred (1965) also found H. arizonensis in greatest abundance in
this type of habitat at the Nevada Test Site.
Vejovis confusus was most frequently captured in the open desert
habitat with H. arizonensis, but unlike H. arizonensis, it was infrequently
found in the rocky habitats more intimately associated with the moun¬
tain. The complete absence of V. confusus in the northern exposure
habitat was striking.
Vejovis spinigerus was equally abundant in all the rocky habitats of
the mountain, but was relatively uncommon in the open desert habitat.
The few individuals collected in the open desert suggested that these
were either migrants or were peripheral representatives of their popu¬
lation.
In 1965, V. confusus and H. arizonensis shared equally the numeri¬
cally dominant position in the open desert habitat. This codominance
could exist because the two species avoided competitive exclusion by
feeding on prey of different kinds and sizes, and thereby avoided direct
competition for food. However, the abundance of H. arizonensis in the
open desert habitat in 1965 may have been unstable since it was sig¬
nificantly decreased in 1966.
Two significant fluctuations in population density occurred during
the two years of the study. Hadrurus arizonensis significantly decreased
in abundance in the open desert habitat and V. spinigerus significantly
increased in abundance in the southern exposure habitat.
Literature Cited
Alexander, A. J., and D. W. Ewer. 1958. Temperature adaptive behavior in the
scorpion, Opisthophthalmus latimanus Koch. J. Exp. Biol., 35(2) :
349-359.
Avedision, G. E. 1966. Geology of the western half of Phoenix South Mountain
Park, Arizona. Unpublished Master’s Thesis, Arizona State Univsity,
Tempe, 54 pp.
Baerg, W. J. 1929. Some poisonous arthropods of North and Central America.
IV Int. Congr. Entomol., 2: 418-438.
1954. Regarding the biology of the common Jamaican scorpion. Ann. Entomol.
Soc. Amer., 47(2) :272-276.
OCTOBER 1970 ] WILLIAMS-COEXISTENCE OF DESERT SCORPIONS 267
Chew, R. M. 1961. Ecology of the spiders of a desert community. J. N. Y.
Entomol. Soc., 69(1) : 5^11.
Cloudsley-Thompson, J. L. 1956. Studies in diurnal rhythms, VI. Bioclimatic
observations in Tunisia and their significance in relation to the physi¬
ology of the fauna, especially woodlice, centipedes, scorpions and beetles.
Ann. Mag. Nat. Hist., Ser. 12, 9: 305-329.
1961. Rhythmic activity in animal physiology and behavior. Acad. Press,
N. Y. 236 pp.
1962. Lethal temperatures of some desert arthropods and mechanism of heat
death. Entomol. Exp. Appl., 5: 270-280.
1964. Terrestrial animals in dry heat: Introduction. (In) Handbook of Physi¬
ology, Sect. 4. Amer. Physiol. Soc., Washington, D. C., p. 447-449.
Gertscii, W. J., and D. M. Allred. 1965. Scorpions of the Nevada Test Site.
Brigham Young Univ. Sci. Bull., Biol. Ser., 6(4) : 1-15.
Hadley, N. F., and S. C. Williams. 1968. Surface activities of some North
American scorpions in relation to feeding. Ecology, 49(4) : 726-734.
Kangieser, P. C. 1966. Climates of the states, Arizona. (In) Climatography of
the United States, Nos. 60-62. U. S. Dept, of Commerce, Environmental
Services Administration, Environmental Data Service. 14 pp.
McAlister, W. H. 1966. The aggregating tendency of Centruroides vittatus Say
(Arachnida: Scorpionidae). Tex. J. Sci., 18(1) : 80-84.
Muma, M. H. 1963. Solpugida of the Nevada Test Site. Brigham Young Univ.
Sci. Bull., Biol. Ser., 3(2) : 1—13.
Smith, G. T. 1966. Observations of the life history of the scorpion Urodacus
abruptus Pocock (Scorpionidae), and an analysis of its home sites.
Aust. J. Zool., 14(3) : 383-398.
Sreenivasa-Reddy, R. P. 1959. A contribution towards the understanding of the
functions of the pectines of scorpions. J. Anim. Morphol. Physiol.,
6(2) : 75-80.
Williams, S. C. 1966. Burrowing activities of the scorpion Anuroctonus phaeo-
dactylus (Wood), (Scorpionida: Vejovidae). Proc. Calif. Acad. Sci.,
34(8) : 419-428.
1968a. Methods of sampling scorpion populations. Proc. Calif. Acad. Sci.,
Ser. 4, 36(8) : 221-230.
1968b. Scorpion preservation for taxonomic and morphological studies. Was-
mann J. Biol., 26(1) : 133-136.
Williams, S. C., and N. F. Hadley. 1967. Scorpions of the Puerto Penasco area
of Sonora, Mexico, with description of Vejovis baergi, new species.
Proc. Calif. Acad. Sci., 35(5) : 103-115.
268
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Observations on the Biology of Cinygmula
reticulata McDunnough in Oregon 1
(Ephemeroptera: Heptageniidae)
D. M. Lehmkuhl 2 and N. H. Anderson
Department of Entomology, Oregon State University,
Corvallis, 97331
Mayflies of the genus Cinygmula are widespread and frequently
abundant in the lotic waters of the western United States. The immature
stages are undescribed at the species level and aside from scattered
notes, no details have been published on the biology of any species
(Berner, 1959). Information reported here was gathered as a back¬
ground for studies of benthic communities in lotic waters.
This genus has been included in the works of Traver (1935), Burks
(1953), Day (1956), and Jensen (1966). A drawing of Cinygmula sp.
nymph is provided by Edmunds (1959, p. 909). Three of the 11 North
American species were recorded from Oregon by Allen and Edmunds
(1956). Cinygmula par (Eaton) and C. tioga (Mayo) have been col¬
lected in the vicinity of Mt. Hood, whereas C. reticulata McDunnough
is widespread in western Oregon. Allen (1955) collected the latter only
west of the Cascade Mountains. Jensen (1966) reported it from east-
central Idaho.
Taxonomic Considerations
Jensen (1966) used the following combination of characters to distin¬
guish Cinygmula nymphs from those of other genera of Heptageniidae:
(1) three well developed caudal filaments present; (2) gills on abdominal
segments one and seven not enlarged nor extending beneath the abdomen,
and similar to those on intermediate segments, only smaller; (3) fibrilli-
form portion of gills absent or reduced to a few tiny filaments; (4) front
of head distinctly emarginate medially.
According to Jensen (1966) C. reticulata is included in the minimus
group which is the most difficult section of the genus, with much varia¬
tion in size and color.
The mature nymph is 6 to 8 mm in length with caudal filaments 5 to
7 mm long. The fibrilliform portion of the gill of C. reticulata is absent.
The color of the dorsum varies from light brown to dark brown to red,
1 Technical Paper 2817 Oregon Agricultural Experiment Station. From a thesis by D. M. Lehmkuhl
submitted in partial fulfillment of the requirements of the Ph.D. degree. This research was supported
by National Science Foundation grants GB-3643 and GB-7958 to N. H. Anderson.
2 Present address: Department of Biology, University of Saskatchewan, Saskatoon, Canada.
The Pan-Pacific Entomologist 46: 268-274. October 1970
OCTOBER 1970 ] LEHMKUHL & ANDERSON—CINYGMULA BIOLOGY 269
Fig. 1. Femur of Cinygmula reticulata illustrating color pattern.
and hence is unreliable as a taxonomic character (see below). Day
(1956) used the color pattern on the femur to separate species of
Cinygmula nymphs. We found that the femur of C. reticulata has a
basal, central, and sometimes apical white area, with the basal and
central areas usually connected by a longitudinal white patch; there
are 10 to 15 brown spots in the white area of the basal half of the
femur (Fig. 1). Based on the examination of a number of specimens
(some of which were not C. reticulata) it appears that the brown spots
on the femur may be useful for separating the species of nymphs. An
unidentified species collected in Montana had nearly 40 of these spots.
Nymphs and adults from the same collection may vary in color (repre¬
sentatives are deposited in the Oregon State University insect collection).
The basic color of nymphs ranges from ochreous to grayish-brown and
the nymphs occasionally have a distinct dorsal stripe, or they may be
marked strongly with red. In reared material, red nymphs retain the
red markings from one moult to the next. Within the family Hepta-
geniidae, this occurrence of red nymphs is not restricted to Cinygmula
as Day (1957) commented on the occurrence of 15 to 18% of the
nymphs of Rhithrogena with varying degrees of reddish color. Based on
laboratory rearing of C. reticulata, adults from red nymphs do not differ
morphologically from other adults.
In a single series collected in Oregon, adults varied considerably in
size, and the wings varied from clear to strongly tinged with yellow.
The dark markings forming the wing reticulation may be strong or en¬
tirely absent. Since some species in this genus are distinguished only
by differences in size or color (Traver, 1935), a revision of the genus
would likely yield many synonyms.
270
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Seasonal Size Class Distributions
The life cycle was studied in two streams. Samples were taken from
July 1967 to June 1968; monthly in Oak Creek, Benton County, in the
Willamette Valley and in alternate months in the Metolius River, Jeffer¬
son County, on the eastern side of the Cascade Range. On each sampling
date, three samples were taken from each biotope described below.
Benthos samples were taken by embedding the end of a length of stove¬
pipe (6 in. dia.) into the substrate of the stream. The larger stones were
removed by hand, and then the water was stirred vigorously and the
contents were transferred to a pail with a fine mesh (0.2 mm openings)
aquarium net.
Oak Creek is a small, densely-shaded woodland stream. The sampling-
station was five miles northwest of Corvallis in the foothills of the Coast
Range at an elevation of 500 feet. Conditions of flow and temperature
varied considerably as a result of normal seasonal changes. Stream
width in the sample area varied from 3.5 to 16 feet, depending on rain¬
fall, and water temperature varied from 1° C in December and January
to 21° C in August. Benthos samples were taken from the following bio¬
topes in Oak Creek: still backwater (protected from main current by
stumps and other obstructions) ; glide (water up to 2 feet deep with a
smooth flow, impacted silt and rubble substrate) ; and riffle (turbulent
flow with much splashing, clean rubble substrate).
Samples from the Metolius River were taken at a site between Camp
Sherman and the headwater springs at an elevation of 3,000 feet. In the
sample area, volume of flow and water temperatures were fairly constant
year round (water temperatures at mid-day were: March, 8° C; May, 10°
C; July, 12° C; August, 10° C; September, 10° C; November, 9.5° C; and
January, 9° C). The bed of the river is broad (about 100 feet), flat, and
shallow, the water seldom being over 2 feet deep. Most of the gravel
substrate is less than 2 inches in diameter. Dense beds of Ranunculus
aquatilus L., islands of sedge, and growths of Lemna near the shore, are
conspicuous features. Samples were taken from both the gravel and
the plant beds.
C. reticulata , in Oak Creek, hatches in the fall, grows slowly during
the winter and adults emerge from April to June (Fig. 2). Eggs ap¬
parently require several months to hatch. Some small nymphs occurred
in the June 1968, sample but from the present data we cannot determine
their significance. It is possible that these emerged from eggs deposited
in early spring. Sampling was not continued after June so data are not
available on the fate of this group. In the previous year, no active stages
OCTOBER 1970] LEHMKUHL & ANDERSON-CINYGMULA BIOLOGY 271
NO.
COLLECTED MONTH
Fig. 2. Size class distribution of Cinygmula reticulata in monthly benthos sam¬
ples, Oak Creek, Benton Co. Ore. A = adults collected. Nymphs absent in July.
of C. reticulata were collected during July, and the major hatching period
did not start until October.
Figure 3 shows the life cycle of C. reticulata in the Metolius River.
All size classes were present on nearly all sampling dates. Adults, or
nymphs with dark wing pads, were present from April to November
indicating that adults emerge during eight or more months of the year.
In spite of the long hatching and emergence periods, the overlapping
generations can be distinguished by noting the increase in nymphal
density in the autumn, and the change in percentage of newly-hatched
nymphs (from 4% to 65%) between July and September. The < 2 mm
size group in September and the < 3 mm group in November represents
the bulk of a new generation hatching in the fall (Fig. 3). The over-
272
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
NO.
COLLECTED MONTH
872 M
J
BWP 434 M
Fig. 3. Size class distribution of Cinygmula reticulata nymphs in benthos sam¬
ples, Metolius River, Jefferson Co. Ore. BWP mature nymphs with black wing
pads. A = adults collected.
lapping generations are separated by a diagonal line, indicating a clear
univoltine cycle (by adding those to the left of the diagonal line to the
top of the graph).
While the streams differ in many respects, the life cycle of the species
is similar in both. The duration and temporal spacing of events differ,
probably as a result of differences in air and water temperature. Speci¬
mens from the Metolius River tend to be larger and darker in color than
those from Oak Creek.
Habitat Preferences
According to Day (1956), the nymphs of Cinygmula spp. are often
found in water only 1 to 2 inches deep at the foot of riffles, in crevices,
and on the lower surfaces of small stones in small streams in California.
Jensen (1966) stated that the nymphs of C. mimus (Eaton) are found
in small to moderate streams, usually above 5,000 feet, on rocks and
among gravel; nymphs of C. ramaleyi (Dodds) are found in small to
moderate streams above elevations of 4,000 feet; and those of C. par
occur in small, cold, swiftly flowing streams above elevations of 5,000
feet where they cling to the undersides of rocks in the swiftest portions
of the current.
OCTOBER 1970] LEHMKUHL & ANDERSON-CINYGMULA BIOLOGY 273
As is apparent from our study, C. reticulata occurs at considerably
lower altitudes and in larger streams than Jensen reports for the other
species. Nymphs of C. reticulata live in gravel or rubble substrates, and
while they tolerate some silt and organic matter, they are usually absent
from muddy areas and plant beds. Nymphs are found in moderate cur¬
rents, i.e., they are absent from still water, and uncommon in extremely
fast riffles. Small nymphs (presumably newly hatched) and mature
nymphs (those with dark wing pads) were almost entirely restricted to
riffle areas. Of 140 from Oak Creek and 1,269 from the Metolius River,
81% and 99%, respectively, of the small nymphs were from the gravel
riffle area. Of the ultimate instar nymphs, 15 of 24 from Oak Creek and
37 of 40 from the Metolius River were from gravel, as opposed to plant
bed areas.
We have no detailed data on food habits of the species but nymphs
apparently feed both on diatoms and on detritus. Chapman and Demory
(1963) carried out gut analysis of a species of Cinygmula (as Cynigmula)
in western Oregon; they found that the nymphs fed heavily on diatoms
in spring and early summer but turned to allochthonous detritus from
late summer to mid-winter.
Mating and Oviposition
Adults of this species were observed in the field many times. At the
Metolius River, swarms comprised of hundreds of adults were observed
both in direct sunlight at midday and in the evenings. At Oak Creek,
most mating flights were seen on cloudy days or in the shade. The
mating flight usually occurred 5 to 8 feet above the surface of the water,
and the rising and falling of individuals covered a vertical interval of
less than 2 feet.
Thirteen males were collected from a swarm and kept in a cage at
outside temperatures. Eight were alive after 24 hours, five after 48
hours, three after 52 hours, and all were dead at 64 hours. Thus, under
artificial conditions, males live long enough to swarm more than one day.
A female was observed laying eggs at Oak Creek. She maintained a
stationary position by fluttering at the height of one foot over the sur¬
face of a riffle. The female extruded a cluster of yellow eggs, dropped
to the water and touched the mass to the surface. The eggs scattered in
the water and the female flew back to the former position above the riffle.
She fluttered there 1 to 2 minutes, more eggs appeared on the underside
of the abdomen, and she again dropped to the water. This was repeated
four times, after which the insect flew off to the vegetation along the
stream bank.
274
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Summary
Cinygmula reticulata McDunnough (Ephemeroptera: Heptageniidae)
was studied in Oak Creek (yearly water temperature range 1° to 21° C)
and in the spring-fed Metolius River (yearly water temperature range 8°
to 13° C) in Western Oregon. In Oak Creek, eggs hatched mainly in
October and adults emerged from April to June. In the Metolius River
most eggs hatched from September to November and adults emerged
from April to November. The species was univoltine in both, and the
abbreviated period of adult emergence in Oak Creek was presumably
due to high summer temperatures. Nymphs were most numerous in
moderate current or silt-free substrates. The most stable character for
identification of the nymphs is the color pattern on the femur.
Literature Cited
Allen, R. K. 1955. Mayflies of Oregon. Unpubl. M.S. Thesis, Univ. Utah, 162 pp.
Allen, R. K., and G. F. Edmunds, Jr. 1956. A list of the mayflies of Oregon.
Proc. Utah Acad. Sci. Arts and Lett., 33: 85-87.
Berner, L. 1959. A tabular summary of the biology of North American mayfly
nymphs (Ephemeroptera). Bull. Fla. State Mus., 4: 1-58.
Burks, B. D. 1953. The mayflies, or Ephemeroptera, of Illinois. Bull. Ill. Nat.
Hist. Surv., 26: 1-216 .
Chapman, D. W., and R. L. Demory. 1963. Seasonal changes in the food ingested
by aquatic insect larvae and nymphs in two Oregon streams. Ecology,
44: 140-146.
Day, W. C. 1956. Ephemeroptera. In: Aquatic Insects of California, ed. by R. L.
Usinger. Univ. Calif. Press, Berkeley, pp. 79-105.
1957. The California mayflies of the genus Rhithrogena (Ephemeroptera).
Pan-Pac. Entomol., 33:1-7.
Edmunds, G. F., Jr. 1959. Ephemeroptera. In: Freshwater Biology, ed. by W. T.
Edmundson. Wiley, New York, 2nd ed., pp. 908-916.
Jensen, S. L. 1966. The mayflies of Idaho (Ephemeroptera). Unpubl. M.S.
Thesis. Univ. Utah, 367 pp.
Traver, J. R. 1935. Systematics. Part II. In: The Biology of Mayflies, by J. G.
Needham, J. R. Traver, and Yin-Chi Hsu. Comstock, Ithaca, pp. 239-751.
OCTOBER 1970 ]
MARSH—ODONTOBRACON REVIEW
275
The Nearctic Doryctinae, IX.
The Genus Odontobracon and Notes on Related Genera
(Hymenoptera: Braconidae)
Paul M. Marsh
Systematic Entomology Laboratory, Agriculture Research Service,
U.S. Department of Agriculture 1
Within the Doryctinae is a group of genera all of which possess one
constant character shared by no other genera, namely the extension of
the postnervellus of the hind wing distally toward the wing apex (Fig.
12). They also have a characteristic oval-shaped raised area on abdomi¬
nal tergum 2 + 3 (Fig. 11), but this is somewhat more variable than the
wing venation. The teeth on the dorsum of the hind coxae were previ¬
ously considered diagnostic, but they do not occur in all of these genera,
and are present in other doryctine genera which do not possess the ex¬
tension of the postnervellus or the characteristic sculpturing of the
abdominal terga.
The genera involved are Odontobracon Cameron, Zombrus Marshall,
Liodoryctes Szepligeti, Nervellinus Roman, Holcobracon Cameron, and
Lophogaster Granger. Various authors in the past have expressed dif¬
ferent views concerning these genera, particularly Zombrus and Odonto¬
bracon ; some have kept them as separate genera, others have synony-
mized one with the other. However, it appears that each of the above
genera is not only distinct morphologically, but geographically—e.g.,
Zombrus is exclusively Oriental and Ethiopian, Odontobracon is ex¬
clusively Neotropical and Nearctic, etc.—and I believe it is advisable to
retain each as a distinct genus. They can be distinguished by the follow¬
ing key:
1. Hind coxae with two teeth dorsally, one long and one short _ 2
Hind coxae with one long tooth or no teeth dorsally _ 3
2. Notauli present; occipital carina present_ Zombrus Marshall
Notauli absent; occipital carina absent _ Liocloryctes Szepligeti
3. Hind coxae with one long tooth dorsally _ Odontobracon Cameron
Hind coxae without teeth dorsally _ 4
4. Radiellen cell of hind wing divided by a spurious cross vein _
_ Nervellinus Roman
Radiellen cell without a cross vein _ 5
5. Cubitella of hind wing arising from junction of postnervellus and basella_
_-_-_ Holcobracon Cameron
Cubitella arising from middle of basella_ Lophogaster Granger
1 Mail address: c/o U. S. National Museum, Washington, D. C. 20560.
The Pan-Pacific Entomologist 46: 275-283. October 1970
276
THE PAN-PACIFIC ENTOMOLOGIST f VOL. 46, NO. 4
Zombrus Marshall
Zombrus Marshall, 1897, p, 10. Type species: Zombrus antsopus Marshall, in Hun¬
garian Natural History Museum, Budapest. Monotypic.
Trimorus Kriechbaumer, 1894, p. 60. Preoccupied by Foerster, 1856. Type species:
7 rimorus nigripennis Kriechbaumer, in Transvaal Museum, Pretoria, South
Africa. Monotypic.
Neotrimorus Dalla Iorre, 1898, p. 100. New name for Trimorus Kriechbaumer.
Acanthobracon Szepligeti, 1902, p, 47 (not Acanthobracon Kriechbaumer, 1900).
Type species: Acanthobracon fuscipennis Szepligeti, in Hungarian Natural
History Museum, Budapest. Designated by Viereck, 1914, p. 2.
Trichtobracon Cameron, 1905b, p. 104. Type species: Trichiobracon pilosus
Cameron, in British Museum. Monotypic.
Trichodoryctes Szepligeti, 1906, p. 599. Type species: Acanthobracon striolatus
Szepligeti, in Hungarian Natural History Museum, Budapest. Monotypic and
original designation.
Oriental, Ralearctic, Ethiopian, and Australian Regions. About 40
species have been described in this genus.
Liodoryctes Szepligeti
Liodoryctes Szepligeti, 1906, p. 599. Type species: Acanthobracon australiensis
Szepligeti, in Hungarian Natural History Museum, Budapest, Monotypic and
original designation.
Ncotrimoroides Strand, 1911, p. 104. Type species: Neotrimor aides dentifer
Strand, in Zoological Museum. Humboldt University, Berlin. Monotypic NEW
SYNONYMY. '
Australian Region. Five species have been described.
Holcobracon Cameron
Holcobracon Cameron, 1905a, p. 89. Type species: Holcobracon fulvus Cameron,
in British Museum (Natural History). Monotypic.
Neotropical and Oriental Regions. Three species have been described.
N ervellinus Roman
Smelhnus Roman , 1924, p. 5. Type species: Nervellinus subdivUus Roman, in
Swedish Natural History Museum, Stockholm. Monotypic.
Neotropical Region. Only one species has been described. The place-
ment of this genus here is based solely on the extension of the post-
nervellus. rhe sculpturing of the abdominal tergum 2 + 3 is much dif-
ferent from that in the other genera.
Lophogaster Granger
Lophogaster Granger, 1949, p. 93. Type species: Lophogaster
Museum National d'HUtoire Naturelle, Paris. Monotypic.
Madagascar. Only one species has been described.
seyrigi Granger, in
OCTOBER 1970 ] MARSH—ODONTOBRACON REVIEW
277
ODONTOBRACON Cameron
Odontobracon Cameron, 1887, p. 384. Type species. Odontobracon nigricep
Cameron, in British Museum (Natural History). Designated by Viereck,
p. 103.
Head cubical; face very coarsely rugose; frons, vertex and temples smooth; from,
excavated, divided by longitudinal ridge from median ocellus to between antennae;
occipital carina distinct, not meeting hypostomal carina on ri es, notau
sternaulus crenulate, shallow, nearly as long as mesopleuron; second cubital ce o
fore wing quadrate, nervulus postfurcal; first segment of me ie a o
longer than second, postnervellus angled distally toward wing apex '
abdominal tergum 2+3 with large raised basal oval median area t e inn *«
crenulate grooves (Fig. 11).
Nearctic and Neotropical Regions. Nine species have been described,
six of which occur in the Nearctic Region.
Kf.y to the Nearctic Species of Odontobracon
1. Radius of fore wing reaching wing margin well before wing apex, radial «dl
along wing margin at most as long as stigma I Fig. 1» «
Radius reaching wing margin nearer apex of wing, ra ia ce r, ^ 2
margin longer than stigma. < Fig. 12) ~ , /
2. Malar space with distinct groove extending from d ^ ° oemeovorut R u hwer
mandible -—- 3
Malar space without such groove .“ : " fivers ei V striated or areolated
3. Notauli meeting posteriorly in wide, rugo.t, ^
area, usually without two longitudinal
Notauli meeting posteriorly in narrow longitu i y 5
usually with two strong posteriorly converging cannue ,g.
4. Ovipositor longer than abdomen; firs, ahdommal tergum ^lyaslong
apical width; subalar groove nearly smooth; large
15 mm or more —---, horter - first abdominal
Ovipositor a. most as long as abdomen, usually Jt^. dis , inclly
tergum considerably wider at apex than . *=’ niericcDS Cameron
-nulate; smaller species, body length less Hum 15
5. Propodeum aerolate posteriorly, basa carin mont anus Cameron
usually black----, , F“*7L ■nosteriorlv; mesonotum
Propodeum rugose posteriorly, basa carina ca lijornicus Rohwer
red-- ---—..
Odontobracon californicus Rohwer
(Figs. 4, 9)
Odontobracon caUfornicus Rohwer, 1917, p. 169. Holotype female, in U. S. National
Museum.
n c jyivv. Color orange or red ex-
Female—L ength of body, 6 mm; oviposi , _ and , egs Qcellocular dis-
cept black head, prothorax, upper part o ™ es0p spacc % eye height, without
lance about four times as long as latera oce us, n d anter ior tentorial
distinct groove from eye to mandible; frontoclypeal suture
278
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46, NO. 4
X /!
I-res. 1 5, & 9th tergum, side view: I, Odontobracon montanus Cam.; 2, O. cel-
ulus, n. sp.; 3, 0. nigrkeps Cam.; 4, 0. californicus Roh.; 5, 0. oemeavorns Roh.
''’V 1 genitalia, ventral view, somewhat diagramatie; 7, 0. oemeovorus ;
, . cellulus. Figs. 6, 8 & 9, £ gonobase, ventral view: 6, O. nigriceps; 8, 0,
montanus; 9, 0 . californicus, Fic. 11, abdominal tergum 2 + 3, dorsal view, 0 .
f!® 8, 12 & 13 ’ fore and hiad wings: 12, O. nigrkeps; 13, O. cd»u/w.
FlGS - U & 15 ’ meS ° notum ^ dor&a l view: 14, O. nigrkeps; 15, O. montanus.
OCTOBER 1970 ]
MARSH-ODONTOBRACON REVIEW
279
pits not deep; two posteriorly converging longitudinal carinae at junction of notauli
before scutellar furrow; subalar groove crenulate; propodeum rugose on posterior
surface, basal carina indistinctly forked posteriorly, sides of propodeum weakly
punctate on lower half; second segment of radius of fore wing less than twice
length of first segment; recurrent vein entering first cubital cell; cubitus between
recurrent and first intercubitus about as long as first segment of radius; fore
tarsus 1.4 times as long as fore tibia; first abdominal tergum wider at apex than
long, tooth at basolateral corner not prominent, basal longitudinal carinae distinct;
tergum 2 3 weakly striate beyond raised basal oval area; ovipositor slightly
shorter than abdomen.
Male.- —Essentially as female; ninth tergum and gonobase (Figs. 4, 9), ninth
tergum finely longitudinally striate, setae on distal edge scattered; ninth sternum
with few scattered setae medially.
Type locality. —Santa Cruz Mountains, California.
Distribution. —Known only from the type locality in California.
This species is easily distinguished by its size, color and thoracic
sculpturing.
Oclontobracon cellulus Marsh, new species
(Figs. 2, 10, 13)
Female. —Length of body, 10-12 mm; ovipositor, 4-6 mm. Color of head, pro¬
thorax, mesonotum and sides of propodeum varying from red to black; mesopleural
disc, propodeum dorsally and abdomen always red; legs usually black, rarely
femora and coxae red. Ocellocular distance about twice as long as lateral ocellus;
malar space % eye height, without groove from eye to mandible; anterior tentorial
pits deep; triangular area where notauli meet before scutellar furrow transversely
striate or rugose, rarely with two indistinct converging longitudinal rugae; subalar
groove crenulate; propodeum areolated, basal carina present only dorsally; fore
tarsus about 1.5 times as long as fore tibia; dorsal spine on hind coxa usually
abruptly angled at apex; first segment of radius of fore wing less than twice as
long as second segment; radial cell along wing margin at most as long as stigma,
usually shorter, radius reaching wing margin well before wing apex (Fig. 13) ;
recurrent vein entering first cubital cell; cubitus between recurrent and first inter¬
cubitus about as long as first segment of radius; first abdominal tergum wider at
apex than long, tooth at basolateral corner very weak; tergum 2 -f- 3 usually striate
beyond raised oval area, rarely smooth; fourth abdominal tergum sometimes striate
at base ; ovipositor nearly as long as abdomen.
Male. —Essentially as female; ninth tergum and genitalia (Figs. 2, 10) ; sculptur¬
ing and setae on apical edge of ninth tergum as in 0. nigriceps ; ninth sternum
with few scattered setae medially.
Holotype female , Alamo Crossing, Yuma County Arizona, 7 Sep¬
tember 1959, Carl E. Benson, collector. U. S. National Museum type
number 70853.
Paratypes.—ARIZONA: 12, Alamo Crossing, Yuma Co., 22 Septem¬
ber 1959, Carl E. Benson; 12, Dobbs Spring, reared 21 April 1919,
280
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Hopk. no. 10087k, G. Hofer, coir.; 1 2, mouth Bear Cyn., Sta. Catalina
Mtns., 3 July 1961, Werner-Nutting; 1$, Douglas, Cochise Co., 15
August 1958, P. M. Marsh; 1$, Arizona-Son. Desert Mus., Pima Co.,
5-8 August 1962, Nutting-Oman; 12, 2 $, Picture Rock Pass, Tucson
Mtns., 25 July 1961, Werner-Nutting; 12, Portal, Cochise Co., 27 June
1958, W. F. Barr; 22, Sabino Canyon, 10 August 1953, 6 August 1959,
Butler, Krombein; 22, Santa Rita Mtns., 4 September 1914; 12, near
Sabino Canyon, Pima Co., R. H. Arnett, Jr.; 1<$, 8 mi. N. Vail, Pima
Co., 30 August 1962, Werner-Nutting. CALIFORNIA: 12, Vallecitos,
San Diego Co., 24 September 1936. NEW MEXICO: 12, Las Cruces,
26 July 1961. TEXAS: 12,3 mi. E. Presidio, 1 May 1963, H. E. Evans.
MEXICO: 1 $, MacDougal Crater, Penacate Mtns., 28 November 1959,
G. D. Butler. Paratypes are deposited in the U. S. National Museum,
University of Arizona, Canadian National Collection, and the Museum
of Comparative Zoology.
This species is very similar to O. nigriceps, but the short radial cell of
the forewing, the abruptly curved spine on the hind coxae, and the
structure of the male genitalia distinguish O. cellulus.
Odontobracon grandis Ashmead
Odontobracon grandis Ashmead, 1894, p. 122. Holotype female, in U. S. National
Museum.
Female. —Length of body, 16-20 mm; ovipositor, 11-14 mm. Color of head,
prothorax, lateral mesonotal lobes, upper part of mesopleuron and legs black;
abdomen, rest of thorax and area around mouth opening red. Ocellocular distance
about 1.5 times as long as lateral ocellus; malar space about % eye height, without
distinct groove from eye to mandible; frontoclypeal suture and anterior tentorial
pits very deep, face very coarsely rugose; notauli obscured posteriorly by strong
transverse striations in V-shaped area before scutellar furrow; subalar groove
smooth or very weakly crenulate; propodeum areolated dorsally and posteriorly,
rugose laterally, basal carina indicated dorsally only; fore tarsus 1.5 times as long
as fore tibia; second segment of radius of fore wing twice as long as first segment;
recurrent vein interstitial, or nearly so, with first intercubitus; cubitus between
recurrent and first intercubitus, if present, much shorter than first segment of
radius; first abdominal tergum longer than wide at apex, tooth at basolateral corner
very prominent; tergum 2 + 3 strongly striate beyond raised basal oval area; fourth
tergum striate at base; ovipositor longer than abdomen.
Male. —Unknown.
Type locality.— San Jose del Cabo, Baja California.
Distribution. —Arizona; Baja California.
This species, one of the largest in the genus, can be separated from
O. nigriceps by its size, longer ovipositor and first abdominal tergum,
and its thoracic sculpturing.
OCTOBER 1970 ]
MARSH-ODONTOBRACON REVIEW
281
Odontobracon montanus Cameron
(Figs. 1, 8, 15)
Odontobracon montanus Cameron, 1887, p. 384. Holotype female, in British Mu¬
seum (Natural History).
Odontobracon crassiventris Cameron, 1887, p. 385. Holotype female, in British
Museum (Natural History). NEW SYNONYMY.
Female.- —Length of body, 7-14 mm; ovipositor, 3-7 mm. Color of head, thorax
and legs black; abdomen red; mesonotum occasionally and dorsum of propodeum
frequently red; base of middle tibia white. Ocellocular distance 2+2.5 times as long
as lateral ocellus; malar space % eye height, without groove from eye to mandible;
anterior tentorial pits deep; two parallel longitudinal carinae at junction of notauli
before scutellar furrow (Fig. 15) ; subalar groove weakly crenulate, sometimes
smooth; propodeum areolated dorsally and posteriorly, laterally punctate or oc¬
casionally smooth, basal carina obscured on posterior face of propodeum; fore
tarsus 1.6 times as long as fore tibia; second segment of radius of fore wing about
2.5 times as long as first segment; recurrent vein entering first cubital cell; cubitus
between recurrent and first intercubitus about as long as first segment of radius;
first abdominal tergum wider at apex than long, tooth at basolateral corner weak;
tergum 2 + 3 striate beyond basal raised oval area; fourth tergum striate at base;
ovipositor about as long as abdomen.
Male.- —Essentially as female; mesopleuron, propodeum and hind coxae red;
third and fourth abdominal terga rugose at base; ninth tergum and gonobase (Figs.
1, 8) ; sculpturing and setae on distal edge of ninth tergum similar to 0. californicus ;
ninth sternum with scattered setae medially.
Type locality. —Irazu, Costa Rica.
Distribution. —Alabama, Florida, Georgia, North Carolina, South
Carolina, Texas; Costa Rica; British Honduras; Guatemala; Panama.
Specimens of this species have been previously confused with 0.
elaphidiovorus (now a synonym of 0. nigriceps ) but can be distinguished
by the different sculpturing where the notauli meet on the mesonotum.
Odontobracon nigriceps Cameron
(Figs. 3, 6,11,12,14)
Odontobracon nigriceps Cameron, 1887, p. 385. Holotype female, in British Mu¬
seum (Natural History).
Odontobracon elaphidiovorus Rohwer, 1917, p. 168. Holotype female, in U. S.
National Museum. NEW SYNONYMY.
Female. —Length of body, 8-13 mm; ovipositor, 3-6 mm. Head black, rarely
marked with red; pro- and mesothorax varying from entirely red to black; pro¬
podeum and abdomen always red; legs black. Ocellocular distance twice as long
as lateral ocellus; malar space about % eye height, without groove from eye to
mandible; anterior tentorial pits not deep; notauli obscured posteriorly by V-shaped
transversely rugose area before scutellar furrow (Fig. 14) ; subalar groove strongly
crenulate; propodeum areolated, basal carina present only dorsally; fore tarsus
1.25 times as long as fore tibia; dorsal tooth on hind coxa broadly curved, not
232
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
abruptly angled at apex; second segment of radius of fore wing about twice as
long as first segment (Fig. 12) ; radial cell along wing margin longer than stigma,
radius reaching wing margin near wing apex; recurrent vein entering first cubital
cell; cubitus between recurrent and first intercubitus about as long as first seg¬
ment of radius; first abdominal tergum wider at apex than long, tooth at basolateral
corner very weak; tergum 2 3 usually striate beyond raised oval area, occasionally
smooth (Fig. 11) ; fourth tergum sometimes striate at base; ovipositor at most as
long as abdomen, usually shorter.
Male. —Essentially as female; ninth tergum and gonobase (Figs. 3, 6) ;
sculpturing on dorsum of ninth tergum transverse, fingerprint-like, distal edge
with dense fringe of setae; ninth sternum with few scattered setae medially.
Type locality. —St. Geronimo, Guatemala.
Distribution. —Arizona, Arkansas, Connecticut, Florida, Georgia,
Illinois, Massachusetts, Mississippi, Nevada, New Jersey, New Mexico,
North Carolina, Pennsylvania, Texas, Utah, West Virginia, Wisconsin;
Mexico; Guatemala.
Host.— Elaphidion villosum F.
Odontobracon oemeovorus Rohwer
(Figs. 5, 7)
Odontobracon oemeovorus Rohwer, 1917, p. 167. Holotype female, in U. S. National
Museum.
Female. —Length of body, 7-9 mm; ovipositor, 2-3 mm. Color of head, thorax
and legs black; abdomen red; propodeum occasionally red. Ocellocular distance
twice as long as lateral ocellus; malar space about % eye height, with distinct
groove extending from base of eye to base of mandible; anterior tentorial pits and
frontoclypeal suture not deeply excavated; two longitudinal carinae on mesonotum
converging before scutellar furrow at junction of notauli with rugae or carinae
between; subalar groove strongly crenulate; propodeum entirely areolate, some¬
what rugose on sides, basal carina present only dorsally; fore tarsus 1.3 times as
long as fore tibia; second segment of radius of fore wing slightly less than twice
as long as first segment; recurrent vein entering first cubital cell; cubitus between
recurrent and first intercubitus about as long as first segment of radius; first
abdominal tergum wider at apex than long, tooth at basolateral corner not distinct;
abdomen usually smooth beyond raised oval area of tergum 2 + 3; ovipositor
shorter than abdomen.
Male. —Essentially as female; abdominal tergum 2 + 3 striate beyond raised
oval area; ninth tergum and gonobase (Figs. 5, 7) ; sculpturing and distal fringe
of setae on ninth tergum as in nigriceps ; ninth sternum entirely covered with setae.
Type locality. —Appalachicola, Florida.
Distribution. —Alabama, Florida, Illinois, Maryland, Michigan,
Pennsylvania, Texas.
Host.— Oeme rigida (Say).
The groove on the malar space will immediately distinguish this
species.
OCTOBER 1970]
MARSH-ODONTOBRACON REVIEW
>OQ
!oo
Acknowledgments
I wish to thank the following museums and persons for allowing me
to study certain types associated with this study: Museum of Compara¬
tive Zoology, Harvard University (H. E. Evans); Swedish Natural
History Museum, Stockholm (K.-J. Hedqvist); British Museum (Natural
History), London (G. E. J. Nixon); Zoological Museum of the Humboldt
University, Berlin (E. Konigsmann); Hungarian Natural History Mu¬
seum, Budapest (L. Moczar); Institut Royal des Sciences Naturelles de
Belgique, Brussells (P. Dessart); Polish Academy of Sciences, Warsaw
(B. Pisarski and E. Kierych); Museum National d’Histoire Naturelle,
Paris (S. K. Pillault and J. R. Steffan); Musee Royal de 1’Afrique Cen-
trale, Tervuren, Belgium (J. Decelle); South African Museum (Natural
History), Cape Town (A. J. Hesse).
Literature Cited
Asiimead, W. H. 1894. Some parasitic Hymenoptera from Lower California.
Proc. Calif. Acad. Sci. (Ser. 2) 4: 122-129.
Cameron, P. 1887. Family Braconidae. Biologia Centrali-Americana, Hymenop¬
tera. 1: 312-419.
1905a. On the phytophagous and parasitic Hymenoptera collected by Mr. E.
Ernest Green in Ceylon. Spolia Zelan., 3: 67-97.
1905b. A third contribution to the knowledge of the Hymenoptera of Sarawak.
J. Straits Brit. Asiat. Soc., 44: 93-168.
Dalla Torre, C. G. 1898. Nomenclatorisches fiber Braconiden-Gattungen. Wien.
Ent. Zeit., 17: 99-100.
Granger, C. 1949. Braconides de Madagascar. Mem. Inst. Sci. Madagascar, (A)2:
1-428.
Kriechbaumer, J. 1894. Hymenoptera ichneumonidea a medico nautico Dr. Joh.
Brauns in itinere ad oras Africae occidentalis lecta. Berk Ent. Z.,
39: 43-68.
Rohwer, S. A. 1917. Descriptions of thirty-one new species of Hymenoptera.
Proc. U. S. Nat. Mus., 53: 151-179.
Roman, A. 1924. Wissenschaftliche Ergebnisse der Schwedischen entomologischen
Reise des Herrn Dr. A. Roman in Amazonas 1914M5. 10. Hymenoptera:
Braconidae, Cyclostomi pro p. Ark. Zook, 16: 1-40.
Strand, E. 1911. Zur Kenntnis papuanischer und australischer Hymenopteran,
insbesondere Schlupwespen. Int. Ent. Z., 5: 103-105.
Szepligeti, G. V. 1902. Tropische Cenocoelioniden und Braconiden aus der Samm-
lung des ungarischen National-Museums. Term. Fiiz., 25: 39-84.
1906. Braconiden aus der Sammlung des ungarischen National-Museums, I.
Ann. Hist. Natur. Mus. Nat. Hung., 4: 547-618.
Viereck, H. L. 1914. Type species of the genera of ichneumon flies. Bulk U. S.
Nat. Mus. no. 83, 186 pp.
284
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Biological Notes on some Nearctic Lepturinae
(Coleoptera: Cerambycidae)
L. M. Gardiner
Forest Research Laboratory, Canadian Forestry Service, Department of
Fisheries and Forestry, Sault Ste. Marie, Ontario
Collecting cerambycids for larval taxonomic study has produced much
new biological information on these insects. Ordinarily, this information
would be published with larval descriptions for logical groupings such
as genera. However, as the North American Lepturinae are presently
under revision by J. A. Chemsak and E. G. Linsley, University of Cali¬
fornia, Berkeley, it is thought worthwhile to make the following notes
available for reference now.
Most of the species noted were collected in Ontario and western
Quebec in recent years, but some were collected in the Kananaskis Valley
near Seebe, Alberta, in 1968.
W. R. Mason and G. S. Walley, Entomology Research Institute,
Ottawa, identified the braconid and ichneumonid parasites, respectively.
Pidonia ruficollis (Say).-—In Ontario and Quebec, adults are found in June
and early July on flowers of Rosa, Cornus, Heracleum and other plants, particu¬
larly in partial shade. The host is unknown, but the insect almost always is found
in or near aspen stands. Caged females oviposited readily in decaying Populus
wood.
Grammoptera exigua (Newman).—This species was reared from decaying
limbs of Tilia americana L. from Rondeau Park, Lake Erie, Ontario. Boring and
pupation occurred in the bark. Some parasitism by an unidentified braconid of
the genus Eubadizon was noted.
Grammoptera subargentata (Kirby).—Adults have been collected from flowers
of Rosa, Cornus and other shrubs in late June and early July in Quebec, Ontario
and western Alberta. Caged females laid eggs on decaying Populus, cementing
them to bark or wood, or in cracks. Hatching occurred in about 10 days. Some¬
times the chorion was partly ingested.
Leptura plebeja (Randall).—This species appears to be rather rare in Ontario
and Quebec, but may be found on flowers of Heracleum and Spiraea, from June to
August.
Leptura subhamata (Randall).-—Adults are fairly common on flowers of
Spiraea, Achillea and other plants in Ontario and Quebec during July and August.
They also are attracted to freshly split pine and birch firewood. Females were ob¬
served ovipositing in cracks in 4-year-old, peeled pine logs.
Typocerus sparsus (LeConte).—Adults have been taken on flowers of Chrysan¬
themum, Achillea and Spiraea in July in western Quebec and eastern Ontario.
Caged females oviposited in decaying pine, forcing the ovipositor into the rotten
wood.
The Pan-Pacific Entomologist 46: 284-288. October 1970
OCTOBER 1970 ] GARDINER—NEARCTIC LEPTURINAE
285
Typocerus vehitinus (Olivier).-—In southern and central Ontario, adults are
common on flowers of Spiraea and Asclepias from July to September. This species
infests decaying wood of various conifers and hardwoods, laying eggs in small
checks in the relatively sound wood.
Anoplodera aspera (LeConte).—Adults are fairly common on flowers, especially
of Heracleum, in western Alberta during late July and early August.
Anoplodera canadensis (Olivier).—Adults are very common on flowers, espe¬
cially of Spiraea, in Ontario and Quebec in July and August. Groups of up to 75
eggs are laid in cracks in logs and stumps that have begun to decay. Hatching
occurs in about 20 days, the larvae ingesting the chorion. Larval galleries often are
excavated between sapwood and heartwood. Coniferous wood seems to be preferred
but ovipositing females also have been found on decaying Acer and Ostrya.
Anoplodera chrysocoma (Kirby).—This species is very common during June
and July in Ontario, Quebec and western Alberta, on flowers of many plants in¬
cluding Cornus, Viburnum, Sorbus, Heracleum, Rosa, Ledum, Chrysanthemum,
Achillea and Epilobium. Near Laniel, Quebec, it was found breeding in decaying
stems of fallen black spruce, Picea mariana (Mill.) BSP. The larvae mined ex¬
tensively in the decaying heartwood, packing the gallery with granular and fibrous
frass. Pupation occurred at the end of the gallery and the adult bored its way
to the outside. Near Sault Ste. Marie, Ontario, larvae were found working in the
base of a dead Populus. In black spruce, this species was parasitized by the
ichneumonid Coleocentrus quebecensis Provancher.
Anoplodera instabilis (Haldeman).—Adults were plentiful on flowers of Achil¬
lea in western Alberta. They also have been taken in window traps at Black Stur¬
geon Lake in northwestern Ontario.
Anoplodera minnesotana (Casey).—In Ontario and Quebec, adults have been
collected from flowers of Spiraea, Achillea, Viburnum, Sorbus, Heracleum, Rosa,
Rubus and Cornus. They also have been reared from decaying Betula alleghaniensis
Britt, and Fagus grandifolia Ehrh. The larvae mine extensively in relatively sound
wood at the edge of decay. Pupation occurs near the wood surface, the larva first
preparing an exit hole plugged with frass. At least three years seem to be required
for development. The same wood is attacked repeatedly.
Anoplodera mutabilis (Newman).—Adults are very common on flowers of
Cornus, Viburnum, Crataegus, Sorbus, Achillea, Spiraea, etc., during June and
July in Ontario and Quebec. They also occur in western Alberta. The pink, scaly
eggs are laid under bark and in cracks in decaying hardwoods. The larva mines in
the wood, and the adult chews a circular exit hole. This species has been reared
from decaying Alnus, Populus, Betula, Fagus, Quercus, Prunus, Ostrya, Acer and
Amelanchier. At Laniel, Quebec, larvae in Alnus were parasitized by the ichneu¬
monid Coleocentrus rujus Provancher and a braconid, Eubadizon sp.
Anoplodera nigrella (Say).—Adults have been reared from standing and fallen
dead Pinus banksiana Lamb, and P. strobus L. Larvae excavated galleries in the
decaying heartwood.
Anoplodera proxima (Say).—Adults have been found, often in company with
A. minnesotana, in Ontario and Quebec on flowers of Cornus, Achillea, Chrysanthe¬
mum and Spiraea.
Anoplodera pubera (Say).—Adults are fairly common in Ontario and Quebec
on flowers of Cornus, Chrysanthemum, Achillea and Spiraea during June and July.
One adult emerged from a dead limb of black walnut, Juglans nigra L.
286
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
Anoplodera sangninea (LeConte).—This species seems rare in Ontario and
Quebec, but a few adults have been captured on flowers of Viburnum and Chrysan¬
themum leucanthemum L. In western Alberta, adults are more plentiful on flowers
of Achillea and Galium boreale L. Caged females oviposited readily in the softened
sapwood of decaying Pinus.
Anoplodera sexmaculata (Linnaeus).—This species is fairly common on
flowers of Rosa, Achillea, Heracleum, Viburnum, Sorbus, Amelanchier and other
species in June and July in western Quebec, Ontario and western Alberta. Caged
females laid eggs in profusion on decaying Pinus, but also on the cage floor.
Anoplodera tibialis (LeConte).—In Ontario adults are rather rare on Spiraea
flowers in July. In western Alberta, they were found on flowers of Heracleum up
to an altitude of 6,500 feet. Caged females laid small groups of eggs in cracks in
decaying Pinus.
Anoplodera vagens (Olivier).—In central Ontario, adults have been taken
occasionally on flowers of Heracleum and Spiraea in July.
Anoplodera vittata (Swederus).—In Ontario and Quebec, adults have been
collected in July from flowers of Viburnum, Rosa, Ranunculus, Chrysanthemum and
Spiraea. One adult was found in a pupal cell in a decaying stump of Betula
alleghaniensis.
Toxotus obtusus (LeConte).—Adults were fairly common during July and early
August on Achillea, Heracleum, Galium and other flowers in subalpine meadows
near Seebe, Alberta.
Rhagium inquisitor (Linnaeus).—This species flies in May and June in Ontario
and Quebec but has not been found on flowers. Adults are readily attracted to
turpentine bait (Gardiner, 1957). The females lay eggs in groups of four to 10
under bark scales on stumps and logs of Pinus and Picea, Hatching takes place
in 17 to 20 days. In northern Ontario, development requires two years, the larva
feeding entirely in the phloem. The larvae construct a broad oval or circular pupal
cell with fibres cut from the wood surface. The second winter is passed in the
adult stage in the cell.
Centrodera decolorata (Harris).—The adults are nocturnal and are attracted
to light. Larvae have been found in a decaying branch stub on a living Quercus
rubra L. near Sault Ste. Marie, Ontario.
Anthophilax attenuatus (Haldeman).—In Ontario and Quebec this species
has been found breeding in the relatively solid parts of decaying Betula, Acer,
Ostrya, Fagus and Populus. At least two years are spent in the host. The last-instar
larvae leave the wood in June and burrow into the grey, leached layer below the
humus of the soil, where they pupate in cells in August. A month later they com¬
plete development and overwinter as adults, which emerge in late May and June,
and feed on the male flowers of Pinus. Eggs are laid in fan-shaped clusters of
from 20 to over 100 in cracks in solid wood or among fibres of decayed wood.
Some mortality in the soil is caused by the entomophagous fungus, Isaria farinosa
(Dicks.) Fr.
Anthophilax malachiticus (Haldeman).—This species has been found breed¬
ing in well decayed, fallen trunks of Acer saccharum Marsh, and Betula papyrifera
Marsh, near Bruce Mines, Ontario. It also has been found in the decaying base of
an Amelanchier stem 3 cm in diameter. Eggs are placed singly and in groups in
cracks of decaying wood.
OCTOBER 1970 ] GARDINER—NEARCTIC LEPTURINAE
287
Sachalinobia rugipennis rugipennis (Newman).—Little is known of this
rather rare species. In Ontario it appears to be associated with dead spruce. One
adult was found on a dead black spruce and another emerged from a caged black
spruce “club top.” Others have been captured at turpentine bait.
Pachyta lamed (Linnaeus).—In early August, adults were captured in flight
near Seebe, Alberta, in a spruce-pine forest. One caged female laid more than 275
eggs in and under a piece of moist absorbent cotton.
Evodinus monticola (Randall).—Adults fly in late May and throughout June
in Ontario and Quebec, visiting flowers of Trilium, Cornas, Viburnum, Ledum, etc.
Eggs are laid under bark scales of dead Pinus, Picea and Tsuga. The larvae feed
in the phloem for 12 to 14 months, then drop to the soil in which they pupate at
the onset of cold weather. It is not known if they pass the second winter as adults
or pupae. Occasionally this species is infected with Isaria farinosa in the soil. It
is also parasitized by the ichneumonid Rhyssa persuasoria (L.).
Pseudogaurotina abdominalis (Bland).—This species is very common in west¬
ern Quebec in June on flowers of Rosa, Viburnum, Sorbus, Cornus, Heracleum,
Ledum and Diervilla. It does not appear to occur in the region around Sault Ste.
Marie, Ontario. Attempts to rear it have been unsuccessful, although it has been
offered wood of many species and conditions.
Pseudogaurotina cressoni (Bland).—Near Seebe, Alberta, adults were rather
rare on Rosa flowers in July, at 4,500 feet above sea level. In August, they were
more plentiful at 6,500 feet on flowers of Heracleum.
Acmaeops bivittatus (Say).'—In 1968, a few adults were found on flowers of
Galium near Seebe, Alberta.
Acmaeops pratensis Laicharting.—Adults are common in June and July in
Ontario, Quebec and western Alberta on flowers of Achillea, Sorbus, Viburnum,
Ledum, Erigeron, Heracleum and Galium. In Sweden, it breeds under bark of
spruce and pine, pupating in the soil (Palm, 1956).
Acmaeops proteus (Kirby).—This is one of the most common cerambycids in
eastern Canada and it was also collected in western Alberta. It breeds in all conif¬
erous trees except cedars, attacking recently killed material. Adults fly from early
May through July, feeding on pollen of various flowers, especially Achillea. They
are attracted readily to turpentine. Eggs are laid under bark scales and lichen;
hatching occurs in about 16 days; and the young larvae ingest the chorion before
boring into, and feeding on the phloem. At the end of the second summer, some
larvae leave the host and construct pupal cells in the soil, pupating the following
spring. The remainder spend a third summer in the host, so that part of each
population has a 2-year and part a 3-year life cycle. Larvae in the soil are some¬
times infected with Isaria farinosa (Gardiner & MacLeod, 1959).
Bellamira scalaris (Say).—In Ontario and Quebec adults are fairly common
on flowers, especially Spiraea and Rhus typhina L. in July and August. This species
has been found in decaying Populus and Betula alleghaniensis. Eggs are laid in
groups of up to 40 in cracks in the wood. The larvae mine extensively in the de¬
caying material, pupating close to the wood surface. Two ichneumonid parasites
attack this species in western Quebec: Arotes amoenus Cr. and Spilopteron jormo-
sum formosum (Cr.).
Desmocerus palliatus (Forster).—Larvae develop in the lower stems and roots
of living Sambucus, and adults feed on the flowers and foliage of the same shrub
throughout the summer. The larvae bore upwards in the stem and pupate up to 30
288
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
cm from the ground. The adults emerge hy chewing a circular hole, sometimes
through an old branch scar. The brown, scaly, longitudinally striated eggs are
distinct from all other known lepturine eggs.
Cortodera subpilosa (LeConte).—Adults are plentiful during July in western
Alberta on flowers of Achillea.
(Acmaeops) rufula (Haldeman).—This species was assigned by Swaine and
Hopping (1928) to Acmaeops, but it obviously does not belong in that genus. It
is rare in collections. Adults appear in June on flowers of Ledum groenlandicum
Oeder, and in smaller numbers on Cornus stolonifera Michx. and Antennaria
canadensis Greene. Distribution appears to he boreal; it has been taken at Laniel,
Quebec, and Sault Ste. Marie and Black Sturgeon Lake, Ontario. Attempts to rear
this species have been unsuccessful and the host is unknown. It is invariably asso¬
ciated with black spruce.
Literature Cited
Gardiner, L. M. 1957. Collecting wood-boring beetles by turpentine and smoke.
Can. Dep. Agr., Bi-mon. Progr. Rep., 13(1) : 2.
Gardiner, L. M., and D. M. MacLeod. 1959. An entomogenous fungus on
Acmaeops proteus (Kby.) (Coleoptera: Cerambycidae). Can. Entomol.,
91: 62-63.
Palm, T. 1956. Zur Kenntnis der Biologie von Acmaeops pratensis Laich. und
verwandten Arten (Col., Cerambycidae). Opuscula Entomol., 21: 233-245.
Swaine, J. M., and R. Hopping. 1928. The Lepturini of America north of Mexico,
Part I. Nat. Mus. Can. Bull. No. 52, Biol. Ser., No. 14.
Synopsis of Anamphidora Casey With a New Species
From Baja California
(Coleoptera: Alleculidae)
James D. Marshall
Biology Department, The College of Idaho, Caldwell, 83605
Casey (1924) originally described Anamphidora in the Tenebrionidae
and included a single species, A. parvula, from Mexico. Spilman (1958)
discovered the misplacement and transferred the genus to the Alleculidae.
Marshall (1967) recorded the genus from Texas and described a new
species which he named A. campbelli. The present paper summarizes
the available information on Anamphidora, places it systematically, and
describes a third species from Baja California.
The terminology in general accordance with that of Campbell (1966,
1968) is used in the following account. The size of the eyes is expressed
as the ocular index, a quantitative character discussed fully by Campbell
and Marshall (1964).
The Pan-Pacific Entomologist 46: 288-295. October 1970
OCTOBER 1970 ] MARSHALL—ANAMPHIDORA SYNOPSIS
289
Figs. 1-14. Key characters for distinguishing Central and North American
genera of Lystronichini. Figs. 1-2. Dorsal view of heads. (1) Erxias, (2) typical
Lystronichini. Figs. 3-4. Forelegs, tb = tibia, fm = femur. (3) Prostenus, (4)
typical Lystronichini. Figs. 5-7. Antennae. (5) Prostenus, (6) Xystropus, (7)
Lystronichus. Figs. 8-10. Metacoxae and abdominal processes, abd. pr. = abdomi¬
nal intercoxal processes. (8) typical Alleculini, (9) Lystronichus, (10) Anamphi-
dora. Figs. 11-14. Dorsal views of left halves of pronota and elytra, prn — pro-
notum, ely = elytra. (11) Cteisa, (12) Xystropus, (13) Lystronichus piliferus,
(14) Lystronichus scapularis.
290
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Systematic Placement
The family Alleculidae is currently represented in the New World by
four tribes of the subfamily Alleculinae (Campbell, 1966, pp. 4-5).
One of these tribes, Lystronichini Lacordaire, includes seven New World
genera characterized by a combination of unlobed tarsi and somewhat
wide, apically acute intercoxal processes between the metacoxae. Camp¬
bell (1966), never having seen Anamphidora, placed this genus in his
classification, along with a few others, into a category of genera of un¬
known tribal affinity. On the basis of the rather wide abdominal process
(compare Fig. 10, Anamphidora, and Fig. 9, Lystronichus, with Fig. 8,
typical alleculini), the present author places Anamphidora in the Ly¬
stronichini. As Campbell (1966) has stated, the morphological studies
reported hy Crowson (1955) indicate that the Lystronichini may eventu¬
ally have to be given subfamily status.
Anamphidora somewhat resembles Phedius in the tribe Alleculini
Seidlitz and would probably be included with it were it not for the
complete absence of tarsal lobes in the former. In addition to the super¬
ficial similarities both genera are flightless, with the species being either
apterous or subapterous.
The following key separates the known Central and North American
alleculid genera assignable to the Lystronichini. In couplet 4 a decision
will have to be made regarding the presence or absence of functional
wings. The apterous or subapterous versus alate condition of most Alle¬
culidae can usually be determined without lifting an elytron, because
normally the elytra are sufficiently dehiscent apically so that the wings
are visible underneath. If, as in certain rare instances, the elytra of
the specimen in question are not dehiscent, then it becomes necessary to
lift an elytron to determine the presence or absence of wings.
Key to Genera of Central and North America Lystronichini
1. Head prolonged in front of eyes (Fig. 1) _ Erxias Champion
Head not as above (Fig. 2) ___ 2
2. Femora distally clavate (Fig. 3) ; antennae with distal joints expanded and
greatly flattened (Fig. 5) _ Prostenus Sober
Femora not clavate (Fig. 4) ; antennae not as above_ 3
3. Pronotum in dorsal view with caudal margin as wide as cephalic margin of
elytra (Fig. 11) __ Cteisa Sober
Pronotum with caudal margin distinctly narrower than cephalic margin of
elytra (Figs. 12, 13, 14) _ 4
4. Wings rudimentary ____ Anamphidora Casey
Wings developed _ 5
OCTOBER 1970 ] MARSHALL—ANAMPHIDORA SYNOPSIS
291
Map 1 . Distribution of known species of Anamphidora. Goode Base Map No.
202 reproduced by permission of the Department of Geography, the University of
Chicago. Copyright by the University of Chicago.
5. Antennae with distal joints transverse, about as broad as long (Fig. 6) _
___ Xystropus Sober
Antennae with distal joints longer than broad (Fig. 7) _ Lystronichus Latrielle
Anamphidora Casey
Anamphidora Casey, 1924: 330; Spilman, 1958: 288; Campbell, 1966: 5.
Type: Anamphidora parvula Casey, monobasic.
Diagnosis.— The combination of the slender, filiform antennae and
rudimentary wings serve to separate Anamphidora from all other ly-
stronichine genera of Central and North America.
Definition of Genus.— Small to moderate in size, subovate, convex in cross-
section; integument shining, punctate, glabrous or pubescent. Head exserted, visible
from above; distinctly, transversely impressed line on front between bases of
antennae; terminal segment of maxillary palpi robust and triangular; antennae
slender, filiform, third segment as long as or only slightly longer than fourth, re¬
maining segments slender, elongate, each segment gradually expanded distally;
eyes small; prothorax subquadrate; cephalic margin subtruncate; sides feebly
rounded throughout, subparallel, only slightly narrower than caudal margin. Sub-
apterous, wings rudimentary. Elytra wider at base than caudal margin of pronotum;
elongate, ovate; striae obsolete or impressed; intervals flat or convex. Abdomen
with intercoxal process of hind coxae wide, apex somewhat acute, angulate. Legs
with tarsal segments not lobed.
Distribution. —United States and Mexico (Map 1).
Discussion. —At present no males of Anamphidora are known. The
known species can be distinguished by the following key.
292
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Fig. 15. Anamphidora campbelli Marshall.
Campbell, (line r= 5.0 mm.)
Photograph courtesy of J. Milton
Key to Species of Anamphidora
1. Elytra with striae pronounced (Fig. 15) ; glabrous above; (Texas) _
____ campbelli Marshall
Elytra with striae not at all, or only slightly impressed (Figs. 16, 17) ; long,
erect setae above _ 2
2. Highly polished throughout; setae sparse; (Mexico) _ parvula Casey
Only slightly shining throughout; setae moderately dense; (Baja California)
-__ kimberleei Marshall, n. sp.
Anamphidora campbelli Marshall
(Fig. 15)
Anamphidora campbelli Marshall, 1967:209.
Diagnosis.- —This species is known only from western Texas, and is
easily distinguished from other species of the genus by the glabrous
nature of the dorsal surface and the deeply impressed elytral striae.
Location of type: United States National Museum.
Type locality: Alpine, Texas.
Discussion. —In addition to the type, only one other specimen of A.
campbelli is known, and it was collected near Ft. Davis, Texas (Map 1).
This latter specimen resides in the collection of J. Milton Campbell.
OCTOBER 1970 ] MARSHALL—ANAMPHIDORA SYNOPSIS
293
Fig. 16. Anamphidora parvula Casey, (line = 5.0 mm.)
Anamphidora parvula Casey
(Fig. 16)
Anamphidora parvula Casey, 1924: 330; Spilman, 1958: 288.
Diagnosis. —This species is known only from Mexico, and can be dis¬
tinguished from the other members of the genus by the highly polished
integument, the sparsely distributed, long erect setae, and the lack of
impressed elytral striae.
Description.— General : Convex in cross-section; integument piceous, highly
polished throughout; body with sparsely distributed, long, erect setae above,
appendages sparsely pubescent. Head : Elongate, narrowing gradually behind eyes,
punctures moderately impressed, loosely distributed; terminal segment of maxillary
palpi with angle formed at junction of inner and basal sides approximating 90°,
inner side about as long as outer side; antennae almost one-half as long as body,
segment three subequal to fourth, four through eleven about two times as long as
greatest distal width; eyes small, ocular index = 58.0. Thorax: Subquadrate in
shape, one-fourth wider than long, sides arcuate; caudal margin broadly and
feebly sinuate; basal angles rounded. Pronotum with punctation moderately im¬
pressed; punctures sparse, majority separated by much more than their diameters.
Elytra : Sparsely and confusely punctate; one and one-half times wider at base
than caudal margin of pronotum; sides rounded, becoming wider near middle;
about twice as long as greatest width; striae on disc unimpressed, sutural striae
one and two feebly impressed, intervals flattened, those near suture becoming
feebly convex apically. Abdomen : polished, piceous throughout. Length: 5.3 mm.
Location of type: United States National Museum.
Type locality: Tepehuanes, Durango, Mexico (Map 1).
294
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46, NO. 4
Fig. 17. Anamphidora kimberleei Marshall, (a) female holotype; (b) close-
up of apical portion of right elytron showing setae, (line = 5.0 mm.)
Discussion.— This species is known only from the female holotype
in the Casey collection at the United States National Museum. It was
described originally in the Tenebrionidae, and Spilman (1958) properly
placed it in the Alleculidae.
Anamphidora kimberleei Marshall, new species
(Fig. 17 a, b)
Diagnosis.— This species is known only from Baja California. The
very conspicuous, moderately dense, erect black setae on the dorsal
surface distinguish A. kimberleei from the other members of the genus.
Description of female holotype. — General: Moderately convex in cross-
section; integument brown, only slightly shining throughout; body conspicuously,
somewhat coarsely setate above; setae black, erect, each seta about equal to length
of third joint of antenna; appendages moderately, finely pubescent. Head: Nar¬
rowing gradually behind eyes, not constricted; punctures irregularly impressed,
rather shallow and sparse laterally and posteriorly, becoming more densely, strongly,
and slightly more irregularly punctate on front anteriorly and between eyes; terminal
segment of maxillary palpi with angle formed at junction of inner and basal sides
about 90°, inner side feebly sinuate, approximately same length as outer side;
antennae one-third as long as body, third joint 1.2 times as long as fourth, joints
four through eleven about two times as long as greatest distal width; eyes small,
ocular index = 54.5. Thorax: subquadrate, cephalic margin three-fourths as wide
as caudal margin; caudal margin rather narrowly and very feebly sinuate; basal
foveae lacking; basal angles almost obtuse, only faintly rounded. Pronotal punc-
tation moderate in density; punctures rather deep and unevenly distributed, be¬
coming more shallowly so toward cephalic margin. Elytra: One and two-thirds
times wider at base than caudal margin of pronotum; sides rounded, becoming
OCTOBER 1970 ] MARSHALL-ANAMPHIDORA SYNOPSIS
295
wider postei'iorly; about twice as long as greatest width; striae not deeply im¬
pressed but plainly marked with serial punctures; punctures bearing setae, sepa¬
rated by distance averaging not quite the diameter of punctures; intervals feebly
and irregularly punctate, feebly convex on disc, becoming more so apically, espe¬
cially toward suture. Abdomen : Shining throughout; setae rather sparse, equal in
length to those on dorsal surface, appressed, pointing toward apex. Length : 8.3 mm.
Holotype female, San Quintin, Baja California, Mexico, 12 July
1922 (Map 1). To be deposited in the collection of the California
Academy of Sciences.
Discussion. —This species is known only from the female specimen
described above, and is named for the author’s daughter, Kimberlee
Marshall.
Acknowledgments
Several people have assisted the author in this study. T. J. Spilman,
United States National Museum, kindly permitted access to the Casey
collection and obtained the ocular index of A. parvula. Hugh B. Leech,
California Academy of Sciences, loaned the specimen described as a
new species in the preceding account. J. M. Campbell, Canadian Na¬
tional Collection, furnished the photograph of A. campbelli. Jan Boles,
The College of Idaho, assisted with the photograph of the new species.
P. M. McKnight, Department of Geography, University of Chicago,
granted permission to reproduce the North American map used in show¬
ing distribution records. Miss Alexis Eichman, The College of Idaho,
typed the manuscript.
Literature Cited
Campbell, J. M. 1966. A revision of the genus Lobopoda (Coleoptera, Allecu-
lidae) in North America and the West Indies. Ill. Biol. Monogr.,
37: 1-203.
1968. A revision of the Mexican and Central American species of Isomira
(Coleoptera, Alleculidae). Can. Entomol., 100 : 449-469.
Campbell, J. M., and J. D. Marshall. 1964. The ocular index and its applica¬
tion to the taxonomy of the Alleculidae (Coleoptera). Coleopt. Bull.,
18: 42.
Casey, T. L. 1924. Additions to the known Coleoptera of North America. Mem.
Coleopt., 11: 1-347.
Crowson, R. A. 1955. The natural classification of the families of Coleoptera.
Nathaniel Lloyd & Co., Ltd., London, 187 pp.
Marshall, J. D. 1967. A new species and new United States record of Anamphi-
dora Casey (Coleoptera, Alleculidae). Entomol. News, 78: 209-210.
Spilman, T. J. 1958. The transfer of Ancimphidora from the Tenebrionidae to
the Alleculidae. Proc. Entomol. Soc. Wash., 60: 288.
296
THE PAN-PACIFIC ENTOMOLOGIST
[VOL. 46 , NO. 4
The Cerambycidae of the Panamint Mountains, California
(Coleoptera)
W. H. Tyson 1
823 Cashew Way, Fremont, California
The Panamint Mountains form the western edge of Death Valley
National Monument, California. They are an isolated range of desert
mountains that vary in elevation from below sea level at their base, in
Death Valley proper, to over 11,000 feet on Telescope Peak.
The flora of the upper mountains has been divided into three distinct
groups (Ferris, 1962). The Upper Desert Slope Flora consists of desert-
type shrubs common to higher elevations throughout the Sonoran Desert.
In the Panamint Mountains, this floral group is found primarily be¬
tween 3,500 and 6,000 feet elevation. Above 6,000 feet the influence
of the Piny on-Juniper Woodland begins. Single-leaf piny on pine and
Utah juniper are the dominant plants in this region. Near the lower
range of the woodland the plants are restricted to the washes and ravines
with the juniper being dominant. Above 7,000 feet the pine becomes
dominant and both plants are found on all slope aspects. Common
tree-like shrubs in this region are the curlleaf mountain mahogany and
antelope brush. Above 8,000 feet the Limber-Bristlecone Pine Woodland
is found. This is perhaps the least collected of the three floral groups.
The mountains above 5,000 feet are for the most part inaccessible
except for a few gravel roads. Because of their inaccessibility most
collecting has been restricted to the regions bordering these roads.
Mahogany Flats (8,000 feet) is the highest point one can reach by road.
From this point only a trail traverses the slopes to Telescope Peak. Even
with this road and trail, little or no concentrated cerambycid collecting
has been attempted and the only record from these mountains was by
Dillon and Dillon (1941), when Monochamus linsleyi was described
from a single specimen.
During the springs of 1965—1968, trips were undertaken to this region
to collect wood infested with cerambycid larvae. The wood was returned
to San Jose State College and placed in rearing containers (for tech¬
niques see Tyson, 1966). Plant species were identified via Munz and
Keck (1963) and Ferris (1962). Beetles were identified via Dillon
& Dillon (1941), Dillon (1956), and Linsley (1935, 1962a, 1962b, 1963,
and 1964).
1 Presently with the Southeast Asia Mosquito Project, Smithsonian Institution, Washington, D. C. 20012.
The Pan-Pacific Entomologist 46: 296-299. October 1970
OCTOBER 1970] TYSON—PANAMINT MOUNTAIN CERAMBYCIDAE 297
The following is an alphabetical list of the cerambycid species reared
during the study, and those seen in other collections. With those reared,
biological data is given on host, larval habits, and collection localities.
Emergence dates are not given since the change in both elevation and
climate may have altered the normal life cycles.
The results of this study are not intended to be complete, but to arouse
entomological interest in this unique region. It is probable that future
collections will produce Cerambycidae not here recorded.
Aneflomorpha lineare (LeConte).—This species was reared from girdled
branches of antelope brush ( Purshia tridentata ) collected at Mahogany Flats.
The majority of plants in the region were in very poor condition due to the
severe girdling by this species. It appears that the larvae work down the
branches girdling them several times before pupating. The larvae mine down
the center of the branch, but before pupating form a pupal chamber just be¬
neath the thin bark. Although only one adult specimen was reared, many larvae
were encountered. Limbs up to 4 cm in diameter were found girdled, but
smaller limbs were most commonly utilized.
Arhopalous rusticus hesperus Chemsak and Linsley.—This recently described
subspecies was reared from the root-crown of a dead single-leaf pinyon pine
(Pinus monophylla) . The wood was collected near the Charcoal Kilns (7,200
feet) and the adults emerged over a three year period. The larvae work in the
sound heartwood and pupate in same.
Callidium antennatum hesperum Casey.—This common and wide spread species
was reared in large numbers from single-leaf pinyon pine. Larvae were collected
between 7,000 and 8,500 feet, being most common at 8,000 feet. The larvae
work beneath the bark of small to medium sized dead limbs, and produce
copious amounts of frass, some of which is exited through small holes cut in
the bark by the larvae. The mature larvae enter the heartwood and before
pupating cut an exit hole through the bark. This form is an early emerger,
being caught as adults as early as April at 8,000 feet.
Callidium texanum Schaeffer.-—This species was reared from Utah juniper
(Juniperus osteosperma ) wood collected below Mahogany Flats. The larval
habits are as those described for C. antennatum, but smaller limbs were at¬
tacked. Limbs varying from 5 cm to less than 1 cm were found infested. It
appears that all western Callidium attack only recently killed wood. This species
is extremely variable throughout its range and the examples reared were gen¬
erally smaller in size than the normal range.
Eucrossus villicornis LeConte.—Specimens of this species were reared from
single-leaf pinyon pine. The larval habits conform to those described by Tyson
(1966) and more recently by Wickman and Seminoff (1968). The immatures
are very susceptible to moisture changes, and if the bark is removed from wood
with the larvae in their pupal chambers, they usually die from dehydration.
However, if the adult stage is reached, many will die within the pupal chamber
being unable to exit through the dried frass plug.
Hesperanoplium antennatum (Linsley).—Curlleaf mountain-mahogany ( Cerco-
carpus ledifolius) is the primary host of this species. The larvae deeply score
the heartwood beneath the bark. The work usually begins at the apical end
298
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
of a branch and the larvae work toward the base with frass being exited from
the gallery along its length. Before pupation each larvae enters the heartwood
and constructs a long, frass-free, pupation chamber. This chamber is sealed
with a thin calcarious plug, and just behind this plug is another which is
thicker and composed of thin strips of wood. Larval work was also observed in
the small branches of a low-growing willow ( Salix ) in the same region the
mahogany was collected (Mahogany Flats).
Haplidus testaceus LeConte.—Small to medium sized branches of single-leaf
pinyon pine were most frequently found infested by this species. The infested
wood was collected at the Charcoal Kilns and Mahogany Flats. The larvae work
beneath the bark of dry limbs (usually dead for two seasons) and enter the
heartwood to pupate. The outer surface of the wood, just beneath the bark, is
sealed with a calcarious plug. Limbs infested are usually those that die and
remain on the tree. No wood in contact with the soil was found infested.
Neacanthocinus obliquus pacificus Casey.—The host of this species is single¬
leaf pinyon pine. The larger dead branches or the main trunk of dead trees
are most commonly found infested. The wood was collected at the Charcoal
Kilns. For larval habits see Tyson (1966).
Monochamus linsleyi Dillon & Dillon.—Until recently the unique type specimen
was the only known representative of this species. To date over 40 have been
reared from single-leaf pinyon pine. The larval work is characteristic of other
members of this genus with the larger limbs and the main trunk being attacked.
Infested wood was collected at Mahogany Flats and the Charcoal Kilns. A
large number of the immature forms are destroyed by predaceous beetle larvae
(Ostomatidae) and woodpeckers.
Oeme costata costata LeConte.—The larval work in single-leaf pinyon pine is very
characteristic. The galleries are beneath the bark and are very wide, much wider
than the larvae. The sides of the galleries are heavily reticulated and resemble
work of the Buprestidae. The larvae enter the heartwood to pupate, and may
or may not cut exit holes through the bark.
Osmidus guttatus LeConte.—Several specimens in the Park Service collection at
Furnace Creek, California, are labeled Furnace Creek. These specimens were
reared from mesquite (Prosopis juliflora ) by the Park Naturalist. Since mes-
quite is found in at least one valley of the Panamint Mountains (Wild Rose
Canyon) it is possible that Osmidus does occur there.
Poliaenus schaefferi Linsley.—The larvae mine the small limbs (1 to 2 cm in
diameter) of dead branches of single-leaf pinyon pine. They etch both the
bark and the sapwood and enter the heartwood to pupate. The pupal chamber,
as with the majority of the members of this genus, is sealed with a tight plug
of coarse frass. This record extends the range of this species which was thought
to be only found west of the Sierra Nevada Mountains.
Prionus californicus Motschulsky.—One specimen in the Death Valley Park
Service collection labeled Wild Rose Canyon.
Semanotus ligneus amplus (Casey).—This species was reared from dead Utah
juniper wood collected in the region of the Charcoal Kilns. The larvae etch
both the hark and the sapwood and their galleries are tightly filled with coarse
frass. The larvae enter the heartwood to pupate and seal the opening with
a tight plug of frass.
OCTOBER 1970 ] CHEMSAK & LINSLEY-MEXICAN CERAMBYCIDAE 299
My thanks to James Cope, Richard Main, and G. G. Bianconi of San
Jose State College for valuable assistance on collecting trips, and to
the National Park Service for making these collections possible.
Literature Cited
Dillon, L. S. 1956. The Nearctic Components of the Tribe Acanthocinini. Part
II. Ann. Entomol. Soc. Amer., 49: 207-235.
Dillon, L. S., and E. S. Dillon. 1941. The Tribe Monochamini in the Western
Hemisphere. Reading Public Museum and Art Gallery. Reading, Pa.,
135 pp., 5 pis.
Ferris, R. S. 1962. Death Valley Wildflowers. Death Valley Natural History
Association, Death Valley, Calif., 141 pp. illus.
Linsley, E. G. 1935. A Revision of the Pogonocherini of North America. Ann.
Entomol. Soc. Amer., 28: 73-103, 1 pi.
1962a. The Cerambycidae of North America. Part II. Taxonomy and Classi¬
fication of the Parandrinae, Prioninae, Spondylinae, and Aseminae.
Univ. Calif. Publ. Entomol., 19: 1-102, 1 pi.
1962b. The Cerambycidae of North America. Part III. Taxonomy and classi¬
fication of the subfamily Cerambycinae, tribes Opsimini through Mega-
derini. Univ. Calif. Publ. Entomol., 20: 1-188, figs. 1-56.
1963. The Cerambycidae of North America. Part IV. Taxonomy and classi¬
fication of the subfamily Cerambycinae, tribes Elaphidionini through
Rhinotragini. Univ. Calif. Publ. Entomol., 21: 1-165, figs. 1-52.
1964. The Cerambycidae of North America. Part V. Taxonomy and classifi¬
cation of the subfamily Cerambycinae, tribes Callichromini through
Ancylocerini. Univ. Calif. Publ. Entomol., 22: 1-197, figs. 1-60.
Munz, P. A., and D. D. Keck. 1959. A California Flora. Univ. Calif. Press,
Berkeley and Los Angeles, 1-1681 pp., figs. 1-134.
Tyson, W. H. 1966. Notes on Reared Cerambycidae. Pan-Pac. Entomol., 42:
201-207.
Wickman, B. E., and S. G. Seminoff. 1968. Notes on the Biology of Eucrossus
villicornis LeConte. Pan-Pac. Entomol., 44: 321-324.
New Mexican Species of Eburia Serville and
Eburodacrys Thomson
(Coleoptera: Cerambycidae)
John A. Chemsak and E. G. Linsley
Division of Entomology, University of California, Berkeley, 94720
The hesperophanine genera Eburia Serville and Eburodacrys Thom¬
son are represented by numerous species in the Neotropical regions. Re¬
cent field expeditions to Mexico have resulted in the collection of a num¬
ber of undescribed species, primarily with the use of ultraviolet lights.
The Pan-Pacific Entomologist 46: 299-305. October 1970
300
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
The west coast area, especially around Mazatlan, Sinaloa, is quite rich
in this group both in species and in numbers of specimens. For the
purpose of making their names available for use in other projects, the
following three species of Eburia and one of Eburodacrys are described
at this time.
This and related studies have been supported in large part by the
National Science Foundation through Grant GB-4944X.
Eburia juanitae Chemsak and Linsley, new species
Male.— Form elongate, parallel-sided; integument dark reddish-brown, elytra
slightly paler; eburneous elytral fasciae usually absent; pubescence fine, grayish,
appressed, obscuring surface. Head with front small, with arrow-shaped glabrous
spot at middle; median line deep, extending onto neck; vertex very small with
median line becoming a glabrous carina; antennal tubercles not prominent; obtuse
above; pubescence dense, appressed; antennae extending over two segments be¬
yond elytra, segments three to ten externally produced at apices, scape to basal
half of third segment sparsely pubescent, shining, remaining segments densely
clothed with very short appressed pubescence, third segment longer than first,
fourth subequal to third, fifth longer than fourth, remaining segments gradually
increasing in length, eleventh longest, appendiculate. Pronotum about as broad
as long, slides obtusely tuberculate; disk with two prominent antemedian glabrous
calluses, smaller median callus and two basal ones toward sides; surface irregular,
densely clothed with appressed pubescence, longer erect hairs present at sides;
prosternum deeply impressed, coarsely deeply punctate over apical half, pubescence
dense; procoxal process abruptly declivous toward apex, coxal cavities open be¬
hind; meso- and metasternum minutely punctate, densely pubescent, scent glands
prominent at apex of metepisternum. Scutellum densely pubescent. Elytra about
three times as long as broad; eburneous fasciae usually absent, occasionally with
small basal pair and/or small median pair on each side, the outer longer; punc¬
tures irregular, becoming obsolete toward apex; pubescence short, dense, appressed,
obscuring surface, few longer erect hairs sparsely interspersed; apices bispinose,
spines short, subequal in length. Legs slender; posterior femora attaining elytral
apices; middle and hind femora acutely spined internally, spines longer than
those of elytra. Abdomen minutely densely punctate, densely clothed with short
appressed pubescence with a number of long suberect hairs interspersed; apex
of last sternite subtruncate. Length, 20-26 mm.
Holotype male (California Academy of Sciences) from 5 miles N.
Mazatlan, Sinaloa, Mexico, 1 July 1965, at black and white lights
(J. A. and M. A. Chemsak, E. G. and J. M. Linsley). Eleven male para-
types same data; one male paratype, 27 June 1965; two male paratypes,
27 July 1964 (J. A. Chemsak and J. Powell) ; one male paratype from 8
miles S. Elota, Sinaloa, 2 July 1963 (F. D. Parker, L. A. Stange) ; and
one male from 7 miles N. Mazatlan, 7 July 1962 (Sleeper, Anderson,
Hardy, Somerby).
Paratypes will be deposited in the California Insect Survey, Berkeley;
OCTOBER 1970 ] CHEMSAK & LINSLEY-MEXICAN CERAMBYCIDAE 301
California Academy of Sciences; University of California, Davis;
Canadian National Collection, Ottawa; United States National Museum;
British Museum (Natural History) ; and the collection of E. Sleeper.
This is one of the most distinctive species of Eburia. The usual lack
of eburneous fasciae, the short, dense pubescence, and subserrate ap¬
pearing antennae will readily separate it. One of the specimens in the
type series has small basal and median fasciae and several have vague
ones at the base only. The dorsal callosities of the pronotum are usually
clothed with pubescence.
We take pleasure in dedicating this species to Juanita M. Linsley
for her field assistance in various cerambycid projects.
Eburia powelli Chemsak and Linsley, new species
Male.— Form slender, slightly tapering; integument pale reddish-brown, mandi¬
bles, genae, antennal scape partially, front margin of pronotum, pronotal calluses,
stripes extending from outside of basal eburneous fasciae, apical halves of femora,
apices of tarsal segments, and parts of sternum black; elytra with a usually single
pair of slightly curved eburneous fasciae and a pair of contiguous fasciae on each
side at middle, the outer slightly longer; pubescence very dense, short, appressed.
Head with front deeply impressed, densely pubescent; median line moderately
deep, not extending onto neck; vertex small, plane; antennal tubercles prominent,
obtuse above; pubescence whitish, dense, appressed; antennae extending over
two segments beyond elytral apices, basal segments shining, outer segments mod¬
erately densely clothed with very short, appressed pubescence, third segment
longer than first, fourth shorter than third, fifth subequal to fourth, remaining
segments gradually increasing in length, eleventh narrow, appendiculate. Pro¬
notum slightly broader than long, sides with a short, rather blunt spine; disk
densely, coarsely, irregularly punctate, black antemedian tubercles prominent;
each side with glabrous callus slightly behind anterior margin; pubescence short,
appressed, obscuring surface, sides with several long erect hairs; prosternum
deeply impressed, rectangular areas at apical two-thirds coarsely deeply punctate,
pubescence dense, appressed; procoxal process abruptly declivous, coxal cavities
open behind; meso- and metasternum very finely punctate, densely pubescent,
scent glands prominent at apex of metepisternum. Scutellum densely pubescent.
Elytra about three times as long as broad; inner basal eburneous fasciae slightly
curved, shorter than median ones, outside basal fasciae, if present, small and sepa¬
rated from inner pair; median fasciae contiguous, outer pair slightly longer; basal
punctures moderately coarse, black areas behind median fasciae coarsely punc¬
tured; each side bicostate from base of median fasciae but not to apex; pubescence
fine, appressed, obscuring surface, black spots before and behind median fasciae
usually glabrous; apices spined at outer margin, spines short. Legs slender; hind
femora extending to apex of elytra; middle and hind femora internally spinose,
spines slightly longer than those of elytra. Abdomen densely clothed with re¬
cumbent pubescence, surface obscured; apex of last sternite broad, slightly
emarginate. Length, 11-18 mm.
Female.— Form similar. Antennae slightly longer than body. Abdomen with
apex of last sternite narrower, emarginate. Length, 14*—19 mm.
302
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Holotype male, allotype (California Academy of Sciences) from 5
MILES N. Mazatlan, Sinaloa, Mexico, 27 July 1964, at black and white
lights (J. A. Chemsak, J. A. Powell). Paratypes as follows: 9 males, 12
females, same data, 28 July 1964 (Chemsak and Powell) ; 24 July 1964
(Chemsak, Powell, H. F. Howden) ; 30 July 1964 (W. C. McGuffin) ;
5 August 1964 (Chemsak, Powell, Howden) ; 11 August 1965 (G. H.
Nelson). Additional paratypes: 2 males, 7 miles N. Mazatlan, 7 July
1962 (E. Sleeper, R. Anderson, A. Hardy, R. Somerby) ; 3 males, 3
females, Yenedio (= Venedillo), Sinaloa, 27 June 1918, 5 July 1918,
10 July 1918, 27 July 1918 (Van Dyke Collection) ; 1 male, 40 miles N.
Mazatlan, 27 July 1952 (J. D. Lattin) ; 1 female, 26 miles N. Pericos,
Sinaloa, 13 August 1960; 1 female, 34 miles N. Los Mochis, Sinaloa,
27 August 1963 (F. D. Parker, L. A. Stange) ; 1 female, 6 miles S.
Culiacan, Sinaloa, 6 August 1964 (Chemsak and Powell) ; 1 female,
Minas Neuvas, Sonora, 7 August 1952 (C. & P. Vaurie) ; 1 female, Rio
Cuchuhaqui, 7 miles S. Alamos, Sonora, 25 July-7 August 1953 (F. S.
Truxal).
Paratypes are deposited in the collections of the California Insect
Survey, Berkeley; Canadian National Collection, Ottawa; California
Academy of Sciences; American Museum of Natural History; United
States National Museum; British Museum (Natural History) ; Los
Angeles County Museum; G. H. Nelson; and E. Sleeper.
This species resembles E. aegrota Bates but the two may be separated
by the darker color, shorter femoral spines, longer and coarser elytral
pubescence and by differences of the eburneous fasciae of E. powelli.
The inner basal pair of fasciae of E. aegrota are reduced or lacking
while E. powelli has the outer pair usually missing.
This species is named for J. A. Powell in appreciation for his ceramby-
cid collecting efforts.
Eburia sinaloae Chemsak and Linsley, new species
Male. —Form moderate-sized, subparallel; integument pale reddish-brown,
mouthparts, pronotal calluses, apices of antennal segments, areas surrounding
eburneous fasciae, parts of coxae, and tips of femora darker; each elytron with
a pair of short, separated, eburneous fasciae at base and a median, narrowly sepa¬
rated pair at middle, outer significantly longer than inner; pubescence pale, short,
appressed, obscuring surface. Head with front small, median triangular area
deeply impressed, median line fairly shallow, extending from vertex to impressed
glabrous spot; vertex small, with glabrous shallow carina extending from median
spot onto neck; antennal tubercles small, obtuse above; pubescence short, dense,
recumbent, mandibles and labrum with long stout setae; antennae extending about
five segments beyond body, scape short, stout, segments rather densely clothed
with very short recumbent pubescence, segments three to five with dense fringe
OCTOBER 1970 ] CHEMSAK & LINSLEY-MEXICAN CERAMBYCIDAE 303
of suberect hairs along inner margin, third segment about twice as long as first,
segments four to seven subequal to third, eight to ten slightly shorter, eleventh
very long, appendiculate. Pronotum inflated, much broader than long, sides with
small obtuse tubercle; disk with two prominent, glabrous antemedian calluses,
two flattened glabrous calluses also present at sides, one behind apical margin
and another at lateral tubercle, irregular large callus present at center behind
middle and each side with longitudinal swellings extending from base to ante-
median calluses; surface coarsely, irregularly punctate; pubescence dense, re¬
cumbent, sides with some long erect hairs interspersed; prosternum shallowly
impressed, densely, deeply punctate except for basal one-fourth, pubescence dense;
procoxal process abruptly declivous, coxal cavities open behind; meso- and meta¬
sternum densely, minutely punctate, densely pubescent, metepisternal scent gland
prominent. Scutellum moderately densely pubescent. Elytra over twice as long
as broad; basal ehurneous fasciae short, separated, subequal in length, median
pair separated, outer pair much longer; punctures fine, irregular, asperulate with
larger asperites bearing erect setae interspersed throughout; pubescence fine, re¬
cumbent, partially obscuring surface; apices bispinose, spines short, subequal in
length. Legs slender; posterior femora extending beyond apices of elytra; middle
femora with a short broad tooth internally, hind femora with subequal short broad
teeth at apices. Abdomen densely pubescent; apex of last sternite emarginate.
Length, 13-21 mm.
Holotype male, allotype (California Academy of Sciences) from 5
miles N. Mazatlan, Sinaloa, Mexico, 5 August 1964, at black and
white lights (J. A. Chemsak, J. Powell). Paratypes as follows: 18 males,
24 females, same data, 5 August 1965 (Chemsak and Powell) ; 27 July
1964 (Chemsak, Powell, H. F. Howden) ; 28 July 1964 (Chemsak and
Powell) ; 24 July 1964 (Chemsak, Powell, Howden) ; 27 June 1965
(J. A. and M. A. Chemsak, E. G. and J. M. Linsley) ; 1 July 1965
(Chemsaks and Linsleys). Additional paratypes: 1 male, 10 miles S.
Mazatlan, 3 July 1965 (Chemsaks and Linsleys) ; 2 males, 1 female, 7
miles N. Mazatlan, 7 July 1962 (E. Sleeper, R. Anderson, A. Hardy,
R. Somerby) ; 2 males, Mazatlan, 27 June 1962 (A. E. Michelbacher) ;
1 male, Mazatlan, 20 July 1960 (R. B. Loomis, J. Maris).
Paratypes are deposited in the collections of: California Insect Sur¬
vey, Berkeley; Canadian National Collection, Ottawa; California Acad¬
emy of Sciences; United States National Museum; British Museum
(Natural History) ; and E. Sleeper.
This species is closely related to E. brevispinus Bates but differs by
the lack of prominent spines on the middle and hind femora and the
much finer pubescence of the elytra. Additionally E. sinaloae has very
distinct seta-bearing asperate punctures on the elytra.
The type series is rather uniform except for the obvious size differ¬
ences. The integumental color is paler in some individuals and there is
some variation in the size of the eburneous fasciae.
304
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Eburodacrys hesperidis Chemsak and Linsley, new species
Male. —Form moderate sized, elongate, parallel-sided; integument shining,
testaceous, head, pronotum and antennae basally more orangish, tips of mandibles
and genae, eyes, three dorsal and two lateral pronotal spots, areas around eburne-
ous fasciae, tips of femora, and some sternites black; each elytron with single
short, basal eburneous fascia and narrowly separated median pair, the outer below
the inner. Head with front deeply impressed, median line deep, extending only
onto vertex; vertex small, coarsely, irregularly, rugosely punctate; antennal tuber¬
cles moderate, obtuse above; pubescence sparse, short, depressed with few erect
setae on mandibles, labrum and frons; antennae extending about four segments
beyond body, basal segments shining, segments from apex of fourth moderately
densely clothed with very short pubescence, long erect hairs dense along inner
margin of third segment, hairs gradually diminishing in number to seventh seg¬
ment, third segment with few long hairs at apices, these gradually decreasing in
length, third segment much longer than first, fourth shorter than third, fifth
slightly longer than fourth, segments six to eight subequal to fifth, nine and ten
shorter, eleventh longest, appendiculate. Pronotum as broad as long, sides obtusely
tuberculate; disk with two prominent antemedian glabrous calluses and an elongate
glabrous median callus extending beyond middle; surface around tubercles coarsely
transversely wrinkled; pubescence sparse, mostly long and erect; prosternum mod¬
erately impressed, transversely rugose, pubescence sparse, procoxal process abruptly
declivous, coxal cavities wide open behind; meso- and metasternum shining, mod-
episternal scent gland distinct. Scutellum glabrous. Elytra about 2% times as
erately coarsely punctate at middle, sides minutely punctate, subopaque; met-
long as broad; eburneous fasciae usually surrounded by black markings, outer
median pair longer than inner, both longer than basal pair; each elytron with
four costae, middle pair extending along eburneous fasciae; basal punctures coarse,
subconfluent, punctures becoming obsolete toward apex; pubescence sparse, short,
with numerous long erect setae interspersed along sutural one-half; apices spinose
at outer margins, inner margins dentate, outer spines short. Legs slender; hind
femora extending beyond apices of elytra; middle and hind femora internally
spinose, spines long and subequal in length. Abdomen sparsely pubescent; apex
of last sternite emarginate. Length, 15-20 mm.
Female.— Form similar. Antennae about as long as body. Abdomen with apex
of last sternite subtruncate. Length, 17-20 mm.
Holotype male, allotype (California Academy of Sciences) from 5
miles N. Mazatlan, Sinaloa, Mexico, 1 July 1965 (J. A. and M. A.
Chemsak, E. G. and J. M. Linsley). Five paratypes (3 males, 2 females)
same data (one female on 27 June 1965).
Paratypes are deposited in the California Insect Survey, Berkeley
and the British Museum (Natural History).
Eburodacrys sticticollis Bates appears to be closely related to this
species. Eburodacrys hesperidis differs by the smooth median callus of
the pronotum, smaller and thinner eburneous fasciae, and by the more
densely punctate basal area of the elytra.
OCTOBER 1970]
CHEMSAK & LINSLEY-DEATH-FEIGNING
305
The type series is fairly uniform in color and structure except for
the small lateral pronotal tubercles of the type. The other six specimens
have longer, more acute tubercles.
Death-Feigning among North American Cerambyciclae
(Coleoptera)
John A. Chemsak and E. G. Linsley
Division of Entomology, University of California, Berkeley, 94720
In reviewing the literature on North American Cerambycidae we
have encountered very few accounts of “death-feigning.” Townsend
(1884) reported that the adult of the diurnal Mecas inornata (Say)
takes wing and flies away when it observes someone approaching, but
drops to the ground and feigns death when unexpectedly disturbed, and
Frost (1916) states that Physocnemum brevilineum (Say) also drops
when disturbed.
The most informative account of “death-feigning”’ among North
American Cerambycidae of which we are aware is that of Vogt (1949),
with reference to the flightless cactus-feeding lamiine Moneilema ulkei
LeConte. He reported that on numerous occasions this species was
found to feign death with the appendages arranged in a curious and
characteristic manner as follows: Antennae extended forward, hind
legs outstretched and directed partially backwards, middle legs directed
straight backward between the hind legs, and the front legs outstretched
anteriorly. He further commented that two similarly flightless cactus-
feeding lamiines which occur in the same general area of the lower Rio
Grande Valley of Texas, Moneilema armatum LeConte and Parmenosoma
griseum Schaeffer “were observed never to latisimulate in such a man-
55
ner.
In view of Vogt’s account, it may be of interest to report the attitude
assumed by two species of nocturnal cerambycines, Eburia ulkei Horn
and Eburia juanitae Chemsak & Linsley when disturbed after being
attracted to light near Mazatlan, Sinaloa, Mexico. Although both species
assume rigid positions when disturbed and will remain rigid when laid
on their backs, sides, or ventral surface, in light or in darkness, even
when placed in the intense heat range of a gasoline lantern, the two
sexes of Eburia ulkei were by far the most persistent in maintaining
their rigid positions.
The Pan-Pacific Entomologist 46: 305-307. October 1970
306
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Fig. 1. Ventral and lateral views of female Eburia ulkei Horn showing positions
of appendages during “death-feigning.”
OCTOBER 1970 ]
CHEMSAK & LINSLEY—DEATH-FEIGNING
307
The typical position assumed by Eburia ulkei (Fig. 1) involved ex¬
tending the antennae at an angle so that each was separate, pulling the
front legs in close to the body with the femoral spine exposed and lower¬
ing the posterior femora exposing the paired apical spines. With the
posterior part of the body protected by the terminal spines of the elytra,
the anterior portion by lateral thoracic spines, and the antennae sepa¬
rated in a manner that would leave one expendable, if necessary, we
conclude that death-feigning by this dark colored cerambycid would not
only render it inconspicuous among the debris of the forest floor but
would find it utilizing to the fullest its defensive structures. Further,
when picked up before feigning death, both males and females emit a
loud “squeaking” sound.
In a series of tests, involving numerous males and females, “death-
feigning” was maintained for periods twenty minutes to more than two
hours in darkness, for shorter periods in the light on the white collect¬
ing sheet. While rigid, individuals of both sexes could be picked up by
the fourth or fifth segment of one antenna and held horizontally.
Although more than fifty species of cerambycids were attracted to
the light, including the closely related Eburia powelli Chemsak and
Linsley, E. sinaloae Chemsak and Linsley, E. aegrota Bates, E. nigrovit-
tata Bates, Eburodacrys callixantha Bates, and E. hesperidis Chemsak
and Linsley, death-feigning could be induced in none but these two
species. Additionally, this behavior has not been observed in other
cerambycines, such as Aneflus, Aneflomorpha, Cacophrissus, Gnapha-
lodes, Elaphidion, Achryson, Eustromula, and several Ibidionini.
Literature Cited
Frost, C. A. 1916. Collecting Coleoptera in a Maine sawmill yard. Can. Entomol.,
48: 381-390.
Townsend, C. H. T. 1884. Notes on some Coleoptera taken in South Louisiana.
Psyche, 4: 219-222.
Vogt, E. B. 1949. Notes on Cerambycidae from the Lower Rio Grande Valley,
Texas. Pan-Pac. Entomol., 25: 137-144, 175-184.
308
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
GDbtttrary Notire
John Winslow MacSwain, a past President of the Society, died in
Berkeley on 9 September 1970. Dr. MacSwain, teacher and biologist,
was well known for his systematic studies on bees and first instar larvae
of meloid beetles and his studies on the ecology and pollination relation¬
ships of bees. He was Professor of Entomology at the University of Cali¬
fornia, Berkeley, until his retirement in June 1970.
Notice of Dues Change
Due to increased costs of publication of the Pan-Pacific Entomologist,
the Executive Board of the Pacific Coast Entomological Society has
voted to increase the dues and subscription rates for the journal. An¬
nual dues for regular members will be $7.50, those for student members
will be $5.00. Members in each category will receive the journal. Sub¬
scriptions to the journal for non-members will be $10.00 per year. These
new rates will go into effect beginning with volume 47, number 1,
January 1971.
MAILING DATES FOR VOLUME 46
No. 1 19 March 1970
No. 2 29 May 1970
No. 3 10 August 1970
No. 4 12 November 1970
OCTOBER 1970 ]
INDEX TO VOLUME 46
309
INDEX TO VOLUME 46
Anamphidora kimberleei, 294
Anderson, Cinygmula biology, 268
Arachnida
Acari, Mesostigmata, 91
Araneida, Antrodiaetidae, 74
Sicariidae, 74
Scorpionida, Vejovidae, 1, 77, 181,
238, 254
Arnaud, Dr. C. P. Alexander—Honored
Member, 65
note on Cerogenes identification, 68
note on Noctuid prey of Carabid, 69
note on Hilara flight, 72
note on “biting” thrips, 76
Baker, Malacosoma ecology, 27
Bembix frommeri, 201
Bentzien, Atypoides habitat, 74
Blanc, note on Oriental fruit fly in Cali¬
fornia, 72
Bohart, new Bembicini, 201
Book review, 152
Brothers, Bufolucilia silvarum, 198
Buckingham, note on Rhagoletis para¬
sites, 78
Bulletin, new, 152
Chemsak, New Mexican Cerambycids,
299
death-feigning, 305
Choriolaus aurifer, 139
similis, 140
sulcipennis, 139
Chromaphis hirsutustibis, 123
Clifford, mayfly variability, 98
Coleoptera
Alleculidae, 288
Carabidae, 69, 111, 231, 233
Cerambycidae, 33, 128, 284, 296, 299,
305
Curculionidae, 42, 152
Elateridae, 153
Meloidae, 151
Pselaphidae, 36
Scarabaeidae, 106
Staphylinidae, 12
Dialictus ornduffi, 210
Diptera
Calliphoridae, 198
Empididae, 72
Ephydridae, 170
Scenopinidae, 96
Tephritidae, 78
Trypetidae, 72
Eburia juanitae, 300
powelli, 301
sinaloae, 302
Eburodacrys hesperidis, 304
Edwards, note on Yucatan Margarodid,
68
Ephemeroptera
Heptageniidae, 124, 268
Leptophlebiidae, 98
Erwin, Nearctic Leistus, 111
note on unique tacliyine structures,
231
Evans, Thalassotrechus and oil spill, 233
Gardiner, immatures of Spondylis, 33
Nearctic Lepturinae, 284
Goodman, Microbembex californica be¬
havior, 207
Grigarick, new Pselaphid genus, 36
Santa Cruz Island Tardigrada, 185
Grotea athenae, 55
lokii, 60
perplexa, 61
Haddock, note on Leptocella biology, 79
Halstead, a new Largus, 45
Hemiptera
Leptopodidae, 74
Naucoridae, 167
Pyrrhocoridae, 45
Reduviidae, 74
Homoptera
Aphididae, 120
Cicadidae, 225
Fulgoridae, 68
Margarodidae, 68, 79
Huffaker, new biological control orga¬
nization, 150
Hurd, a new Dialictus, 209
Hymenoptera
Apidae, 177
Apoidea, 209
Braconidae, 275
Eumenidae, 241
Icheumonidae, 50, 85
Sphecidae, 201, 207
Isoptera
Kalotermitidae, 74
Nasutitermitinae, 12
Termitidae, 39
Jenkins, note on fat-yielding Coccid, 79
Kelsey, a new Scenopinus, 96
Kinn, a new genus of Celaenopsidae, 91
Kistner, new termitophiles, 12
Kumar, new Callipterine aphids, 120
Langston, new Philotes localities, 74
Largus semipunctatus, 45
La Rivers, Sagocoris usingeri, 167
310
THE PAN-PACIFIC ENTOMOLOGIST [VOL. 46 , NO. 4
Lavigne, new Callipterine aphids, 120
Leech, note on Gnophaela trapped by
milkweed flower, 71
note on grooved entomology, 76
Lehmkuhl, Rhithrogena life cycle, 124
Cinygmula biology, 268
Leistus madmeridianus, 117
(Neoleistus) , 112
Lepidoptera, 81, 82
Lasiocampidae, 27
Lycaenidae, 74
Noctuidae, 69
Pericopidae, 71
Pyralidae, 225
Leptoschema praelontactum gloris,
165
Leptoschema protractum analelegans,
163
detractum, 163
elegans, 161
Linsley, new lepturine Cerambycids, 128
new Mexican Cerambycids, 299
death-feigning, 305
Longipedisymbia bouceki, 24
bradburni, 24
carlislei, 26
Longipedoxenus emersoni, 17
shinneri, 19
Lycomorphoides simulans, 133
Macrochoriolaus elegans, 135
MacSwain, note on Tesla Canyon in¬
sects, 74
note on Meloid convergence, 151
Marsh, Odontobracon review, 275
Marshall, Anamphidora synopsis, 288
Megachoriolaus chemsaki, 129
imitatrix, 131
Megaloptera
Corydalidae, 142
Microbembix argyropleura, 203
californica, 202
rufiventris, 204
Microcoryphia, 212
Microdynerus bolingeri, 246
hurdi, 248
inusitatus, 248
patagoniae, 252
Notices, Society, 151
dues change, 308
Nutting, Tenuirostritermes foraging, 39
Obituary notice, 308
O’Brien, Pselactus in California, 42
book review, 152
Odonata, 194
Odontobracon cellulus, 279
Parker, Microdynerus review, 241
Paulson, Odonata of Washington, 194
Physemops azul, 172
fairchildi, 173
maldonadoi, 173
panops, 175
wheeleri, 176
Pilactium summersi, 37
Platerosida howdeni, 137
Pleocoma octopagina, 107
Powell, note on Humboldt Bay moths,
81
Proceedings, Pacific Coast Entomologi¬
cal Society, 65
Publications, Pacific Coast Entomologi¬
cal Society, 64
Robertson, a new Pleocoma, 106
Sagocoris usingeri, 167
Scenopinus galapagosensis, 96
Schizoeyrtillus lathrius, 92
Schuster, new Pselaphid genus, 36
Santa Cruz Island Tardigrada, 184
Scott, tympanal resilin, 225
Sears, Macrolepidoptera emergence
peaks, 82
Slobodchikoff, Grotea revision, 50
Grotea ratios, 85
Smith, Neohermes biology and struc¬
ture, 142
bristletails and silverfish, 212
Stibick, revision of Leptoschema, 153
Tardigrada, 184
Thorp, putative Arizona Bombini, 177
Thysanoptera
Thripidae, 76
Thysanura, 212
Trichoptera
Leptoceridae, 79
Tuberdefectus eastopi, 120
Tyson, Panamint Mountain Cerambyci-
dae, 296
Vejovis deserticola, 4
gramenestris, 1
pseudopumilis, 181
shulovi, 7
Wasbauer, Calif. Dep. Agr. types, 47
Williams, new California scorpions, 1
note on Vejovis sting, 77
a new Mexican Vejovis, 181
scorpion redescription, 238
coexistence of desert scorpions, 254
Wirth, Neotropical Physemops, 170
Zoological Nomenclature, 63
Published by the
Pacific Coast Entomological Society
in cooperation with
The California Academy of Sciences
VOLUME FORTY-SIX
1970
EDITORIAL BOARD
R. W. THORP, Editor
R. 0. SCHUSTER, Assistant Editor
E. G. LINSLEY
HUGH B. LEECH
E. S. ROSS
P. D. HURD, JR.
P. H. ARNAUD, JR., Treasurer
W. H. LANGE, Advertising
PUBLICATION COMMITTEE
1970
1971
1972
D. D. Linsdale, Chairman P. D. Hurd, Jr. H. V. Daly
D. P. Furman R. F. Wilkey A. A. Grigarick
San Francisco, California
1970
11
CONTENTS FOR VOLUME 46
Arnaud, P. H., Jr.
Dr. Charles P. Alexander—Honored Member _ 65
“Flying mouse” identified as Cerogenes auricoma (Burmeister) 68
Chlaenius tomentosus zunicmus preying on an adult Lacinipolia
spiculosa _ 69
Flight of Hilara sp. at Webber Lake, California_ 72
Thrips “biting” man _ 76
Baker, B. H.
Occurrence of Malacosoma incurvum discoloratum in Zion Can¬
yon, Utah _ 27
Bentzien, M. M.
Habitat of Atypoides river si ___ 74
Blanc, F. L.
Status of Oriental fruit fly in California __ 72
Bohart, R. M.
New species, synonymy and lectotype designations in North
American Bembicini _ 201
Brothers, D. R.
Notes on the saprophagous activity of Bufolucilia silvarum
(Meigen) - 198
Buckingham, G. R.
Searching for a parasite of the walnut husk fly Rhagoletis com-
pleta Cresson _ 78
Chemsak, J. A. and E. G. Linsley
New Mexican species of Eburia Serville and Eburodacrys Thom¬
son _ 299
Death-feigning among North American Cerambycidae _ 305
Clifford, H. F.
Variability of linear measurements throughout the life cycle of
the mayfly Leptophlebia cupida (Say) _ 98
Edwards, J. G.
Giant Margarodid scales from Yucatan_ 68
Erwin, T. L.
The Nearctic species of the genus Leistus Frolich_ 111
Unique structures in members of Tachys sensu lat. _ 231
Evans, W. G.
Thalassotrechus bar bar ae (Horn) and the Santa Barbara oil spill 233
Gardiner, L. M.
Immature stages and habits of Spondylis upiformis Mannerheim 33
Biological notes on some Nearctic Lepturinae_ 284
Goodman, W. G.
Observations on the behavior and biology of Microbembex cali-
fornica Bohart _ 207
Grigarick, A. A. and R. 0. Schuster
A new genus in the tribe Euplectini_ 36
Haddock, J. D.
Biological traits of two species of Leptocella _ 79
Halstead, T. F.
A new species of the genus Largus Hahn with a key to the species
of the genus in the southwestern United States_ 45
Huffaker, C. B.
New biological control organization_ 150
Hurd, P. D., Jr.
A new narrowly polylectic autumnal species of Dialictus from the
flowers of Jepsonia heterandra, an endemic California saxi¬
frage _ 209
Jenkins, K. D.
The fat-yielding coccid Llaveia, a monophlebine of the Margaro-
didae _ 79
Kelsey, L. P.
A new Scenopinus from the 1964 Galapagos International Scien¬
tific Project _ 96
Kinn, D. N.
A new genus of Celaenopsidae from California with a key to the
genera _ 91
Kistner, D. H.
Two new genera of termitophiles associated with Longipedilermes
longipes (Haviland) _ 12
Kumar, R. and R. Lavigne
A new genus and new species of Callipterinae from India
120
IV
Langston, R. L.
Philotes enoples bayensis and tildeni Langston—four new locali¬
ties _ 74
La Rivers, I.
A new Philippine Sagocoris _ 167
Leech, H. B.
Gnophaela latipenrris trapped by flowers of milkweed_ 71
Grooved entomology _ 76
Lehmkuhl, D. M.
The life cycle of Rhithrogena morrisoni (Banks) in western
Oregon _ 124
Lehmkuhl, D. M. and N. H. Anderson
Observations on the biology of Cinygmula reticulata McDun-
nough in Oregon _ 268
Linsley, E. G.
New genera and species in the lepturine complex related to
Euryptera and Choriolaus _ 128
MacSwain, J. W.
A naturalist returns to Tesla Canyon___:_ 74
Examples of convergence between blister beetles of California
and others from Chile _ 151
Marsh, P. M.
The Nearctic Doryctinae, IX. The genus Odontobracon and notes
on related genera _ 275
Marshall, J. D.
Synopsis of Anamphidora Casey with a new species from Baja
California _ 288
New Bulletin _ 152
Notice of Dues Change _ 308
Nutting, W. L.
Free diurnal foraging by the North American nasutiform termite,
Tenuirostritermes tenuirostris _ 39
Obituary Notice_ 308
O’Brien, C. W.
Pselactus spadix (Herbst), a European cossonine weevil new to
California _ 42
V
Book Review: Curculionidae subfamily Apioninae of North and
Central America with reviews of the world genera of Apio¬
ninae and world subgenera of Apion Herbst_ 152
Parker, F. D.
New North American Microdynerus with notes on the nests of
two species _ 241
Paulson, D. R.
A list of the Odonata of Washington with additions to and dele¬
tions from the state list_ 194
Powell, J. A.
Moths of the Humboldt Bay area_ 81
Proceedings, Pacific Coast Entomological Society__ 65
Publications of the Society_ 64
Robertson, J.
A new species of Pleocoma from southern California_ 106
Schuster, R. 0. and A. A. Grigarick
Tardigrada of Santa Cruz Island, California _ 184
Scott, J. A.
Resilin in the sound-organs of Pyralidae and Cicadidae_ 225
Sears, T. A.
A possible explanation for seasonal emergence peaks of Macro-
lepidoptera in southwestern Chihuahua_ 82
Slobodchikoff, C. N.
A revision of the genus Grotea_ 50
Ratios: an intuitive vs. a quantitative approach in Grotea _ 85
Smith, E. L.
Biology and structure of the dobsonfly Neohermes californicus
(Walker) _ 142
Biology and structure of some California bristletails and silver-
fish _ 212
Society Notice _ 151
Stibick, J. N. L.
A revision of the genus Leptoschema of California with a key to
related genera _ 153
VI
Thorp, R. W.
The type locality of Bombus franklini and notes on putative Ari¬
zona records of other Bombini_ 177
Tyson, W. H.
The Cerambycidae of the Panamint Mountains, California_ 296
Wasbauer, M. S.
The type material in the collection of the California Department
of Agriculture _ 47
Williams, S. C.
Three new species of Vejovis from Death Valley, California_ 1
The effects on man of a natural sting by the scorpion Vejovis
confusus Stahnke _ 77
A new species of scorpion belonging to the pumilis group of
genus Vejovis _ 181
A. redescription of the scorpion Vejovis bilineatus Pocock_ 238
Coexistence of desert scorpions by differential habitat preference 254
Wirth, W. W.
A revision of the Neotropical genus Physemops Cresson_ 170
Zoological Nomenclature _ 63
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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” 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
C. Rentz and James D. Birchim. This 173-page revision of Nearctic
shield-back katydids includes descriptions of one new genus and
13 new species and subspecies. Published July 1968. Price $4.00.
(Plus 250 postage.)*
* (Add 5% sales tax on all California orders. Members of the Society will
receive a 20% discount.)
Send orders to:
Pacific Coast Entomological Society
c/o California Academy of Sciences
Golden Gate Park
San Francisco, California 94118