Vol. 60
No. 1
January 1984
THE
Pan-Pacific Entomologist
EVANS, E. D.—A new genus and a new species of Dobsonfly from the far Western United
States (Megaloptera: Corydalidae) . 1
LATTIN, J. D. and G. M. STONEDAHL —Campyloneura virgula, a predacious Miridae not
previously recorded from the United States (Hemiptera). 4
THOMAS, D. B., JR.—A new species of Dendrocoris Bergroth from Mexico (Hemiptera:
Pentatomidae). 8
BOHART, R. M. — Gastrodynerus, a new genus of Eumenidae from western North America
(Hymenoptera: Eumenidae). 12
COYNE, J. A., I. A. BOUSSY, and S. BRYANT—Is Drosophila pseudoobscura a garbage
species?. 16
MUCHMORE, W. B.—Further data on Mucrochernes hirsutus (Banks) (Pseudoscorpionida,
Chemetidae)... 20
BROWN, H. P. and M. P. THOBIAS—World synopsis of the riffle beetle genus Leptelmis
Sharp, 1888, with a key to Asiatic species and description of a new species from India
(Coleoptera, Dryopoidea, Elmidae). 23
BAUMANN, R. W. and A. L. SHELDON— Capnia hornigi, a new winter stonefly from the
western Great Basin (Plecoptera: Capniidae). 30
POLHEMUS, D. A.—A new species of Dichaetocoris Knight from the western United States,
with notes on other species (Hemiptera: Miridae). 33
BARNBY, M. A. and V. H. RESH —Distribution and seasonal abundance of brine flies (Diptera:
Ephydridae) in a San Francisco Bay salt marsh. 37
STONEDAHL, G. M.—Two new conifer-inhabiting Phytocoris from western North America
(Hemiptera: Miridae). 47
BEZARK, L. G.—Description of the first Central American species of Slenopogon Loew (Dip¬
tera: Asilidae). 53
POLHEMUS, J. T. and D. A. POLHEMUS—Notes on Neotropical Veliidae (Hemiptera) VI.
Revision of the genus Euvelia Drake. 55
COPE, J. S.—A new species of Phymatodes from New Mexico (Coleoptera: Cerambycidae).... 63
SCIENTIFIC NOTES
TILLES, D. A.—Feeding behavior of Lacon profusa (Candeze) (Coleoptera: Elateridae) in
Carpenter Ant attended colonies of Cinara spp. (Homoptera: Aphididae). 65
TURNBOW, R. H., JR.—New records for two cerambycids from the Pacific Northwest. 67
SAN FRANCISCO, CALIFORNIA • 1984
Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
in cooperation with THE CALIFORNIA ACADEMY OF SCIENCES
The Pan-Pacific Entomologist
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OFFICERS FOR 1984
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This issue mailed January 27, 1984
The Pan-Pacific Entomologist (ISSN 0031-0603)
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PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 1-3
Published 27 January 1984
A New Genus and a New Species of Dobsonfly from the Far
Western United States (Megaloptera: Corydalidae ) 1
Elwin D. Evans
Michigan Department of Natural Resources, Stevens T. Mason Building, P.O.
Box 30028, Lansing, Michigan 48909.
During my study of the Megaloptera of the Pacific Coastal Region of the United
States, a new species of dobsonfly was discovered and a previously described
species was found to form a monotypic genus (Evans, 1972). I later noted that
this situation existed in this group of aquatic animals (Evans, 1978).
Orohermes, New Genus
Forewings with median vein 2-branched; posterior branch of radial sector forked
in both wings; hind wings with median vein 3-branched; wing expanse less than
110 mm. Larvae with ventral abdominal gill-tufts absent; last pair of abdominal
spiracles lateral and sessile, located above and slightly anterior to base of lateral
filaments.
Etymology. — From the Greek or os, mountain + Hermes, Greek God.
Type of genus.—Dysmicohermes crepusculus Chandler, 1954:107-110.
Chandler (1954) provisionally placed O. crepusculus in Dysmicohermes Munroe
and noted that it was not closely related to the other two species in this genus
which are larger in size and have the median vein of the hind wings four-branched.
These two genera differ from the other Chauliodinae which have the posterior
branch of the radial sector in both wings simple. Later, Chandler (1956) recognized
this larvae which is unique among the Chauliodinae with the last pair of abdominal
spiracles being lateral, instead of dorsal and raised or extended as in all other
known species of this subfamily.
The larvae of O. crepusculus are usually found in permanent, cold water streams
and rivers in the Sierra Nevada Mountains or the northern coast range of California
and in the Cascade Mountains of Oregon.
Protochauliodes cascadius , New Species
Male.— Wings and body similar to P. spenceri Munroe, alar expanse 67 mm;
genitalia with short, stubby claspers, less than twice as long as broad, convex
dorsally and ventrally, indented apically; aedeagus flattened in lateral view (Fig.
1); aedeagus in dorsal view narrowly notched apically, markedly expanded laterally
to about mid-point and sharply constricted basally (Fig. 2).
Female. — Similar to male, alar expanse 71 mm; claspers triangular shaped in
lateral view, slightly convex dorsally and ventrally (Fig. 3), trichobothrium slightly
raised, light in color and occupying most of clasper lateral surface. Claspers of P.
spenceri more elongate with basal width shorter than sides.
The claspers of the short, stubby male genitalia of P. cascadius resemble those
1 Collected with P. spenceri.
2
PAN-PACIFIC ENTOMOLOGIST
CLASPER
AEDEAGUS
LATERALIS
Figures 1-3. 1. Protochauliodes cascadius, lateral view of male genitalia. 2. P. cascadius, dorsal
view of aedeagus. 3. P. cascadius, lateral view of female genitalia.
of P. montivagus Chandler, another mountain species to the south in the Sierra
Nevada Mountains. Claspers of the males of other species of Protochauliodes
Weele in western North America have elongate genital claspers. The aedeagus of
P. cascadius with its apical notch and constricted base is distinct from that of P.
montivagus which is not constricted basally and is broadly indented apically.
Holotype. — Male, ca. 8 mi W Mill City, along Little Santiam R., Marion Co.,
Oregon, 26 July 1963 (S. Jewett). California Academy of Sciences.
Allotype. — Same as holotype.
VOLUME 60, NUMBER 1
3
Twenty paratypes .—All from Oregon as follows: Marion Co. 8, Silver Falls State
Park, 22 May 1957 (S. Jewett); Lane Co. 8, 40 mi E Eugene, 30 July 1965 (K.
Goeden); 8, 4 mi S Mackenzie Bridge, Horse Ck. (J. Bedea); Clackamus Co. 6 8,
vie. Firwood Rd., 4 mi W Oregon City, 5 August 1974 1 (S. Jewett); 2, 10 August
1974 (S. Jewett); 2, 30 July 1972 (S. Jewett); 2, 10 August 1972 1 (S. Jewett); 6, 5
August 1972 1 (S. Jewett); 3 8, 2, 4 August 1974 (S. Jewett); 8,9, 17 July 1972 (S.
Jewett); 8, 14 July 1972 1 (S. Jewett).
Paratypes will be distributed to the California Academy of Sciences, San Fran¬
cisco, California; Oregon State University Entomological Collection, Corvallis,
Oregon; United States National Museum, Washington, D.C.; Stanley G. Jewett
Collection, Rt. 1, Box 399, West Linn, Oregon and my personal collection.
This species is sympatric with P. spenceri, in the foothills of the Cascade Moun¬
tains of Oregon. The larval habitat of P. cascadius is unknown although adults
have been collected along permanent streams.
Acknowledgment
Financial support from a United States Department of Agriculture Research
Service grant, number 12-14.-9160(33).
Literature Cited
Chandler, H. P. 1954. Four new species of dobsonflies from California. Pan-Pac. Entomol., 30(2):
105-111.
-. 1956. Megaloptera. Pp. 229-233 in R. L. Usinger (ed.). Aquatic insects of California. Univ.
Cal. Press, Berkeley, 508 pp.
Evans, E. D. 1972. A study of the Megaloptera of the Pacific Coastal Region of the United States.
Ph.D. thesis, Oregon State Univ., Corvallis, 210 pp.
-. 1978. Megaloptera and aquatic Neuroptera. Pp. 133-145 in R. W. Merritt and K. W.
Cummins (eds.), Aquatic insects of North America. Kendall/Hunt Pub. Co., Dubuque, Iowa,
441 pp.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 4-7
Published 27 January 1984
Campyloneura virgula, a Predacious Miridae Not Previously
Recorded from the United States (Hemiptera ) 1
John D. Lattin and Gary M. Stonedahl
Systematic Entomology Laboratory, Department of Entomology, Oregon State
University, Corvallis, Oregon 97331.
Abstract. —Campyloneura virgula (Herrick-Schaeffer), a European species, is
recorded from the United States, based on specimens from western Washington
and Oregon and northwestern California. C. virgula is predacious on small ar¬
thropods and is found on a variety of trees and shrubs including alder, filbert and
rhododendron.
Campyloneura virgula (Herrick-Schaeffer) is a predacious species of Miridae
whose natural range includes Europe, North Africa, Asia Minor and Turkestan
(Wagner and Weber, 1964). It has been reported from a variety of trees and shrubs
including Alnus, Corylus, Crataegus, Fagus, Fraxinus, Quercus and Tilia (Wagner
and Weber, 1964; Southwood and Leston, 1959). The latter authors report that
C. virgula overwinters as an egg on the British Isles, whereas the adult overwinters
on the continent of Europe. Males are very rare and it is assumed that partheno¬
genesis is the usual mode of reproduction. The food of this species includes spider
mites, Psocoptera, greenflies and occasionally honeydew (Southwood and Leston,
1959).
Downes (1957) reported C. virgula from Vancouver Island and the lower main¬
land of British Columbia, Canada. His earliest collections were made in 1949.
Specimens were collected on alder, poplar, nettles and other plants. He mentioned
the potential value of this species as a predator, especially of mites. Scudder (1960)
again referred to this species while discussing various introduced Hemiptera.
The present paper was prompted by the submission of a specimen for identi¬
fication by Duane Hatch, County Agent for Lane County, Oregon. The bug had
bitten a person on the ankle, raising a hard welt. Rykman (1979) reviewed the
literature on Hemiptera and Homoptera bites. He cited a number of references
for Miridae but C. virgula was not included.
All of the specimens we have examined from the United States were females,
further supporting the likelihood of parthenogenetic reproduction being the nor¬
mal mode. Specimens examined include: CALIFORNIA: Humboldt County:
Shiveley, 4 August 1964, pear and apple (T. K. Haig, F. Spallini) (USNM).
OREGON: Benton County: Corvallis, 11 August 1962; Corvallis, 21 July 1968
(Evans); Corvallis, 30 July 1968, at light (Lattin); Corvallis, 24 June 1979, Quercus
garryana Dougl., Corylus cornuta Marsh., Ulmus sp., apple (Stonedahl) many
specimens (all OSU); Lobster Valley, 15 mi SW Alsea, 24 July 1979, Alnus rubra
1 Oreg. Exp. Sta. Paper No. 5837.
VOLUME 60, NUMBER 1
5
Bong., Corylus cornuta (Lattin) (OSU); 4 mi W Philomath, Hwy 20, 26 July 1979,
Quercusgarryana (Stonedahl) (OSU); Rock Creek, nr. Philomath, 23 August 1961,
Ribes sp. (Lattin) (OSU); Curry County: Alfred A. Loeb State Park, 10 August
1979, Lithocarpus densiflora (H. & A.) Rehd., Umbellularia californica (H. &
A.) Nutt. (Stonedahl) (OSU); Little Redwood campground, T39S-R12W-S29-
NW‘/ 4 , 11 August 1979, Lithocarpus densiflora, Umbellularia californica (Stone¬
dahl) (OSU); Linn County: 2 mi NE Green Peter Dam, 15 July 1979, Castanopsis
chrysophylla (Dougl.) A. DC. (Stonedahl) (OSU); Polk County: 6 mi E Indepen¬
dence, 10 July 1957, apple, (Lattin) (OSU). WASHINGTON: King County: Se¬
attle, Shipcanal, 4 August 1977 (Leffler) (Leffler Collection); Seattle, U. Wash.
Campus, 14 July 1977, leaves of Corylus sp. (Leffler) (Leffler Collection); Lewis
County: Chehalis, Urquhart Rd., 22 July 1979, Corylus cornuta (Stonedahl) (OSU);
Clark Co.: Vancouver, 20 July 1975 (Oman) (OSU); Whatcom County: Belling¬
ham, 19 July 1979, Alnus rubra, Rhododendron sp. (Stonedahl) (OSU); 4 mi NE
Welcome, N. Fork Rd., 22 July 1979, Corylus cornuta, Alnus rubra, Acer macro-
phyllum Pursh, Acer circinatum Pursh, Rubus spectabilis Pursh, Rubus parviflorus
Nutt. (Stonedahl) (OSU).
Campyloneura belongs to the tribe Dicyphini of the Bryocorinae (Schuh, 1976).
The genus may be recognized as a member of the subtribe Dicyphina by the
following characters: (1) setiform parempodia, (2) anterior pronotal margin in the
form of a well developed, rounded collar, (3) pseudopulvilli present, (4) parallel¬
sided tarsi and (5) two cells in the membrane of the forewing.
This genus runs to couplet 160 ( Cyrtopeltis ) in Slater and Baranowski (1978).
The eyes are contiguous with the anterior margin of the pronotum in Campylo¬
neura, while the eyes are removed from the anterior margin by a distance equal
to or greater than the thickness of the second antennal segment in Cyrtopeltis
Reuter, Dicyphus Fieber and Macrolophus Fieber. Other distinctive features of
C. virgula include a bright yellow cuneus with a red apex and a large brown spot
on either side of the ninth abdominal segment. The accompanying figure shows
the habitus of the species. The nymphs are yellow with the wing pads and sides
of the pronotum red, while the antennae are banded with red.
In June 1980, nymphs, and later adults of C. virgula were abundant on a
deciduous species of rhododendron heavily infested with the aphid Illinoia ( Ma -
sonaphis) lambersi (MacGillivary) (determined by D. Hille Ris Lambers, June
1980). As indicated in the information on specimens examined, this mirid occurs
on a variety of trees and shrubs. Where observed, it appears to be a common
associate of a variety of species of Aphidae. On filbert this species is part of a
rather substantial complex of predatory Miridae (5 spp.) and Anthocoridae (3
spp.).
Campyloneura virgula is a useful addition to our western mirid fauna. Its pre¬
dacious habits, its occurrence on a variety of economically important plants, and
its reproduction via parthenogenesis all are beneficial attributes. Since so many
of our introduced species of Hemiptera occur on both coasts, it seems likely that
C. virgula will be collected on the east coast as well. Other introduced species of
predacious Miridae known to occur in the Pacific Northwest include: Blephari-
dopterus angulatus (Fallen), Campylomma verbasci (Meyer), Heterotoma meriop-
terum (Scopoli), and Phytocoris tiliae (Linn.). A species of Hyaliodes occurs in
western Oregon and may represent an introduction from eastern North America.
6
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Adult female Campyloneura virgula (Herrick-Schaeffer).
VOLUME 60, NUMBER 1
7
Based upon collection records most of the introductions into the Pacific North¬
west appear to be relatively recent. As the landscape becomes more disturbed,
the likelihood of successful colonization by selected species from other regions
seems to increase. Thus, we may look forward to additional adventitious species
being found here.
Acknowledgments
Our thanks to Bonnie B. Hall for the fine illustration of the adult, to the General
Research Fund, Oregon State University, for the funds for the illustration, to Dr.
Richard C. Froeschner of the Smithsonian Institution for assistance in examining
specimens under his care, to Dr. S. LefRer, Seattle, Washington, for the opportunity
of examining specimens in his collection, and to Dr. D. Hille Ris Lambers,
Bennekom, The Netherlands, for the identification of Aphidae.
Literature Cited
Downes, W. 1957. Notes on some Hemiptera which have been introduced into British Columbia.
Proc. Entomol. Soc. Brit. Columbia, 54:11-13.
Henry, Thomas J., and A. G. Wheeler, Jr. 1979. Palearctic Miridae in North America: Records of
newly discovered and little-known species (Hemiptera:Heteroptera). Proc. Entomol. Soc. Wash.,
81(2):257-268.
Ryckman, Raymond E. 1979. Host reactions to bug bites (Hemiptera, Homoptera): A literature
review and annotated bibliography. Parts I and II. California Vector Views, 26:1-49.
Schuh, R. T. 1976. Pretarsal structure in the Miridae (Hemiptera) with a cladistic analysis of rela¬
tionships within the family. Amer. Mus. Nat. Hist., Novitates, No. 2601:1-39.
Scudder, G. G. E. 1960. Dictyonota fuliginosa Costa (Hemiptera:Tingidae) in the Nearctic. Proc.
Entomol. Soc. Brit. Columbia, 57:22.
Slater, J. A., and R. M. Baranowski. 1978. How to know the true bugs. W. C. Brown Co., Dubuque,
Iowa, 256 pp.
Southwood, T. R. E., and D. Leston. 1959. Land and water bugs of the British Isles. Frederick Warne
and Co., Ltd., London, 436 pp.
Wagner, E., and H. H. Weber. 1964. Faune de France. No. 67. Heteropteres Miridae. Paris, 589 pp.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 8-11
Published 27 January 1984
A New Species of Dendrocoris Bergroth from Mexico
(Hemiptera: Pentatomidae)
Donald B. Thomas, Jr.
Department of Entomology, University of Arizona, Tucson, Arizona 85721.
The genus Dendrocoris Bergroth was most recently revised by Nelson (1955).
That revision provided keys, descriptions, and figures of the male genitalia for
the eight known species. A review of the literature and a discussion of the biology
of the species was also included. In a subsequent article (Nelson, 1957), a ninth
species was added.
Rolston (1978) discussed the position of Dendrocoris in relation to some other
pentatomine genera and provided a key for its separation. Dendrocoris is related
to Odmalea Bergroth, Brepholoxa Van Duzee, and Thoreyella Spinola. From
these, Dendrocoris is distinguished by the combination of an elongated ostiolar
canal and obtuse femoral apices.
A new species is described here based on two specimens from Jalisco, Mexico.
It is readily distinguished from all other species of Dendrocoris by the presence
of spots on the legs. The description below follows Nelson’s format except that
the terminology for the genitalic structures follows McDonald (1966). All mea¬
surements are from the holotype male.
Dendrocoris maculosus. New Species
(Figs. 1-5)
Appearing dull gray in dorsal view, with alternate connexivum. Distinctive
features: legs maculate; spiracles ringed with black. Related to D. variegatus Nel¬
son.
Color. — Pale ochraceous dorsally; yellowish tan ventrally. Posterolateral angles
of abdominal sternites with broad, black blotch. Spiracles ringed with black. Legs
pale yellow except tarsi and apical sixth of tibiae orange; femora and tibiae densely
black maculate. Antennae orange. Eyes reddish-brown; ocelli clear, transparent;
an irregular, short, black, supra-antenniferal vitta present beginning at anterior
margin of each eye. Each connexival segment with broad, posterior, marginal,
black stripe and an anterior, submarginal stripe; stripes connected along mesial
border by narrow, irregular band of black punctures. Hemelytral membrane in-
fuscated, translucent, veins reticulate.
Punctations dense dorsally; black, except those on discs of coria and pronotum
which are dark brown. Ventral punctures concolorous with surface except for row
of 6-7 black punctures near lateral margin of each propleura and short coalescent
series near lateral margin of each metapleura.
Structure.— Length 7.4, width 4.5 mm; widest point of abdomen and humeral
width subequal. Head flat dorsally; vertex and posterior half of tylus weakly tumid.
Sides of jugae edentate, parallel for short distance before apex; apices overlapping
tylus and contiguous anteriorly. Length of head 1.2, width 1.2 mm. Antennifers
prominent and distinctly visible from above. Antennal segments III, IV, and V
VOLUME 60, NUMBER 1
9
with short, dense pilosity; I and II glabrous; segmental ratios 10:11:24:22:24.
Rostrum in repose just attaining metacoxae; bucculae longer than first rostral
segment, lobed posteriorly.
Pronotum broad, flat, somewhat explanate laterally; humeri prominent but not
as produced as in D. humeralis (Uhler); anterolateral margins irregularly subrec-
tilinear in dorsal view. Thoracic width 4.5, mesial length 1.6 mm. Hemelytral
coria reach to middle of connexival segment VI.
Abdominal tubercle vestigial in male.
Male genitalia .—Pygophore similar in structure to D. variegatus. Posterolateral
Figures 2-5. D. maculosus genitalia. 2. Left paramere, ental aspect. 3. Aedeagus, ventral view. 4.
Pygophore, caudal view. 5. Female terminalia, ventral view.
“hypopygial” plates fused with sides of pygophore; ventral margin deeply, rect¬
angularly emarginate mesially; nadir of emargination with pair of rounded cusps
either side of median; pygophoral rim either side of mesial emargination with
black, spinose, laterally directed tooth. Posteroventral face with pair of black,
triangular teeth either side of middle near end of carinae contiguous with margins
VOLUME 60, NUMBER 1
11
of mesial emargination; a shallow concavity subtending emargination; weak carina
connecting triangular teeth, below this another shallow concavity subtended by
a darkened tumescence (Fig. 4).
Parameres terete basally; abruptly flattened and flared apically; very apex ta¬
pering to digitiform process (Fig. 2). A finely shagreened, irregular tumescence
located anterolaterad of each paramere; tumescence stalked and its basal ligaments
attached to base of paramere. Aedeagus simple in structure; theca with prominent,
subapical, mesial tubercle dorsally; penisfilum elongate, strongly sinuately curved;
slightly flared apically (Fig. 3). Proctiger broad, inornate.
Female genitalia. — First gonocoxites small, widely separated. Triangulum
broadly, sinuately emarginate. Fused second gonocoxites shallowly transversely
strigose. Eighth paratergite with large marginal blotch either side of each ninth
paratergite (Fig. 5).
Holotype.—S ; labeled: mtns. N. Ajijic, Jalisco, Mexico. 5200-7600' 9 July 1964.
W. L. Nutting and sons. Deposited U.S. National Museum.
Allotype. — Labeled as holotype. Deposited U.S. National Museum.
Comments.— As in some other species of Dendrocoris, D. maculosus exhibits
a characteristic pattern of sexual dimorphism. The mesial abdominal tubercle is
well developed in the female, vestigial in the male. The second antennal segment
is proportionately longer in the female. The female is slightly larger.
D. maculosus has a pair of teeth on the posterior face of the pygophore, a
character shared with D. variegatus and D. reticulatus Barber. It also has the
hypopygial plates (the posterior visible portion of the pygophore) fused with the
sides of the pygophore as in D. variegatus to which the new species is obviously
related. D. maculosus differs from this latter species in having the head edentate,
the connexivum alternated, the spiracles ringed with black and the legs maculated.
Key to the Mexican Species of Dendrocoris
1. Femora and tibiae maculate (Jalisco) . maculosus, n. sp.
Femora and tibiae immaculate . 2
2. Dorsum ochraceous with anterior pronotum contrastingly dark brown (So¬
nora, Coahuila, Baja) . contaminatus Uhler
Anterior pronotum concolorous with dorsum . 3
3. Spiracles ringed with black (Chiapas, Morelos, Zacatecas, Guerrero) ....
. suffultus (Distant)
Spiracles pale (Oaxaca, Mexico, Guerrero, Chiapas, Sinaloa, Michoacan,
San Luis Potosi) . variegatus Nelson
Literature Cited
McDonald, F. J. D. 1966. The genitalia of North American Pentatomoidea (Hemiptera:Heteroptera).
Quaest. Entomol., 2:7-150.
Nelson, G. H. 1955. A revision of the genus Dendrocoris and it’s generic relationships. Proc. Entomol.
Soc. Wash., 57(2):49-67.
-. 1957. A new species of Dendrocoris and a new combination of Atizies (Hemiptera, Penta-
tomidae). Proc. Entomol. Soc. Wash., 59(4): 197-199.
Rolston, L. H. 1978. A revision of the genus Odmalea Bergroth (Hemiptera:Pentatomidae). N.Y.
Entomol. Soc., 86(l):20-36.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 12-15
Published 27 January 1984
Gastrodynerus, a New Genus of Eumenidae from Western
North America (Hymenoptera: Eumenidae)
Richard M. Bohart
Department of Entomology, University of California, Davis, California 95616.
The inclusion of “ Stenodynerus” vanduzeei in that genus has never been sat¬
isfactory because the faint dimples on the polished front face of the pronotum
are unlike the pair of pits so characteristic of Stenodynerus. Likewise, an assign¬
ment to Cephalodynerus is ruled out by the asulcate sternum II and unmarked
ocular emargination of vanduzeei. With the Ending of two additional species from
Mexico it now seems appropriate to establish a new genus.
Abbreviations used throughout the paper are T-I, T-II etc. for terga and S-I,
S-II etc. for sterna. Types of the new species are in the University of California
at Davis Entomology Museum (UCD).
Gastrodynerus Bohart, New Genus
(Gr.: gaster + Odynerus)
Generotype.— Stenodynerus vanduzeei Bohart, 1948.
Diagnosis. — Medium sized, 5-8 mm long, palpi 6-4, mandibles evenly 5-den-
tate in female, 5-dentate in male with subbasal one a little prominent, F-I 1.5-
2.0 times as long as broad, male antennal hook small and slender, clypeus apically
incised, vertex with strong tubercles or at least faint traces, female vertex pits
indistinct between postocellar tubercles or in a weak median postocellar depres¬
sion. Pronotum with a hardly visible pair of dimples on polished front face,
transverse carina weak and effaced toward middle, length of pronotal dorsum at
middle less than antennal socket diameter, no pretegular sulcus and following
carina, humeral angles moderate; notauli and parapsidal furrows weakly impressed
in posterior half, parategula slender and incurved, scutellum conspicuously flat¬
tened, metanotum flattened and not raised above level of scutellum, tegula slender
and with outer edge evenly curved to posterior point, omaulus undeveloped,
stemaulus absent, submarginal cell II narrowly truncate anteriorly, submarginal
III longer posteriorly than anteriorly, prestigmal radial stem short (Fig. 8), male
midleg relatively unmodified; propodeum without superior teeth or ridges, no
lateral tooth or carina, a sharp uptilted membranous projection laterally below,
valves connected but deeply incised (Fig. 1), median sulcus deep and sharp. T-I
moderately rounded at summit, about as long as broad, with a slightly raised and
smooth apical border; T-II sharply rounded (bent) in profile at basal fourth to
fifth (Fig. 1), apical margin convex and slightly to strongly reflexed, T-II basally
with a series of pits or ridges; S-II without a median sulcus, basal third nearly flat
and rather sharply bent out in profile (Fig. 1); S-III with massive punctures in
middle third (Fig. 9); genitalia relatively simple (Figs. 4, 5).
Discussion. — The combination of polished front pronotal surface, pitted base
of T-II, and asulcate as well as bent S-II distinguish Gastrodynerus from such
related genera as Stenodynerus and Cephalodynerus. The former has distinct
VOLUME 60, NUMBER 1
13
pronotal pits, and the latter has S-II with a median sulcus toward the base, and
teeth or carinae on the propodeum above. Further, Gastrodynerus differs in having
three lower propodeal projections (inferior membranous tooth and two valves)
instead of two (valves only). The genus is also rather unusual in having the ocular
emarginations dark. Three species are known, vanduzeei and two new ones de¬
scribed below.
Key to Species of Gastrodynerus
1. T-II hardly reflexed at apex (Fig. 1) hind margin of pronotum yellow at
least posteriorly (Fig. 6), T-I coarsely punctate and with a shallow groove
across summit (Fig. 1) vertex with only traces of polished tubercles (Fig.
6) . searsi Bohart
T-II reflexed at apex about 2-3 times median ocellus diameter, margin of
pronotum dark (Fig. 7), T-I rather evenly and moderately punctate across
summit, vertex various . 2
2. Vertex with large round polished postocellar tubercles (Fig. 7), interocellar
tubercles present, upper mesopleural spot well developed, usually a free
spot on T-II . vanduzeei (Bohart)
Vertex with insignificant or small and oblique tubercles, interocellar tu¬
bercles absent, upper mesopleural spot absent or rarely present and small,
no free spot on T-II . stangei Bohart
Gastrodynerus searsi Bohart, New Species
(Figs. 1-6)
Male holotype.— Length 6.5 mm, forewing 5.5 mm. Black with pale yellow as
follows: clypeus, inside of scape, interantennal dot, postocular dot, narrow and
irregular humeral margin, hind margin of pronotum narrowly in posterior two
thirds (Fig. 6), metanotal band, propodeum toward abdominal insertion, apical
dot on midfemur, T-I laterally and posteriorly, T-II posterior band, T-III-VII
banded, S-I mostly, S-II-V narrowly across apex; reddish yellow are: flagellomeres
inside but not including hook, legs partly; wings smoky, stigma brown. Pubescence
inconspicuous. Punctures of frons, vertex, and notum moderately coarse and close,
clypeus nearly impunctate, punctures of T-I coarse and close, those of T-II fine
to medium and mostly 2-3 puncture diameters apart, those of S-I coarse and
close, of S-II medium and well spaced. Face as in Figure 2, antennal apex Figure
3. traces of postocular and postocellar polished welts, T-I a little longer than
broad, and with a slight depression across summit, T-II as long as broad and
faintly reflexed apically (Fig. 1), genitalia relatively simple (Figs. 4, 5).
Female.— Length 7 mm, forewing 6 mm. Clypeus with lateral and subapical
dots, sometimes a tiny mesopleural dot, midfemur dark.
Holotype male. — Santo Nino, Chihuahua, Mexico, September 1, 1968 (T. A.
Sears, R. C. Gardner, C. S. Glaser, UCD). Paratypes, 19 males, 3 females, same
data as holotype, 4 females, 4-6 mi nw. Choix, Sinaloa, Mexico, August 6-
September 5, 1968 (T. A. Sears, R. C. Gardner, C. S. Glaser, UCD).
Discussion. — The weakly developed vertex tubercles, yellow hind pronotal mar¬
gin, absence of a free spot on T-II, somewhat flattened as well as coarsely punctate
T-I, and practically no apical reflex on T-II distinguish the species. It is named
for Terry Sears who collected a great many Mexican eumenids in 1968.
14
PAN-PACIFIC ENTOMOLOGIST
Figure 7. Gastrodynerus vanduzeei, male head and prothorax, dorsal. Figures 8, 9. Gastrodynerus
stangei. 8. Forewing apex. 9. Male sternum III.
VOLUME 60, NUMBER 1
15
Gastrodynerus stangei Bohart, New Species
(Figs. 8, 9)
Male holotype. —Length 7 mm, forewing 6 mm. Markings as given above for
searsi except: clypeus with median dark spot, pronotum narrowly banded in front
only, tegula with two dots, yellow streaks on tibiae, forewing stigma orange (not
brown), S-I half dark, S-III banded, other segments with faint apical bands. Punc¬
tures of frons, vertex and notum medium and close, clypeus nearly impunctate,
punctures of T-I medium and somewhat spaced across summit, those of T-II
medium and 1-2 puncture diameters apart except coarse and close before apical
reflex. Face and antennal apex much as in Figures 2, 3, postocellar polished
tubercles weak and punctate, no interocellar tubercles, T-I longer than broad,
more strongly convex than Figure 1, T-II as long as broad and with an apical
reflex of 2 MOD, genitalia about as in Figures 4, 5.
Female. — Length 8.5 mm, forewing 7.5 mm. About as in male, no markings
on abdominal segments after II.
Holotype male.— Three mi n. Petlalcingo, Puebla, Mexico, August 3, 1963 (F.
D. Parker, L. A. Stange, UCD). Paratypes, 6 males, same data as holotype, 1
male, Zimapan, Hidalgo, Mexico, June 11, 1951 (H. E. Evans, Cornell), 1 female,
Chapala, Jalisco, Mexico, July 6, 1952 (A. E. Michelbacher, UCB on loan to CAS).
Discussion.— As indicated in the key, stangei is most similar to vanduzeei. The
absence of a free spot on T-II distinguishes stangei from vanduzeei in southwestern
U.S. but not from all Mexican specimens. However, the weak postocellar tubercles
and absence of interocellar ones in stangei are adequate for separation. Another,
more subtle, distinction is the less strongly rounded T-I in side view. In this
respect stangei is halfway between vanduzeei and searsi. The species is named for
my friend, the well known Neuropterist and Hymenopterist, Lionel Stange.
Gastrodynerus vanduzeei (Bohart)
(Fig. 7)
Stenodynerus vanduzeei Bohart, 1948, Proc. Calif. Acad. Sci. (4)24:327.
Holotype male, Baboquivari Mts., Arizona (CAS).
Male.— About as described above for searsi except: length 8.0 mm, forewing
6.5 mm, a broad pale yellow band across pronotum anteriorly, tegula yellow with
a central reddish dot, parategula and a mesopleural spot yellow, legs more exten¬
sively yellow, forewing stigma orange, an attached lateral spot on T-I and usually
a free spot on T-II, S-I dark, abdomen dark beyond second segment, punctures
of T-I medium coarse and about a puncture diameter apart across evenly rounded
summit, postocellar polished tubercles large and round (Fig. 7), postocular ones
smaller, T-I and T-II about as long as broad, T-II with an apical reflex of 2-3
times median ocellus diameters, male genitalia about as in Figures 4, 5.
Female. — Length 9.0 mm, forewing 7.5 mm. About as in male.
Distribution.— Mostly mountainous areas in S Arizona, S New Mexico, W Tex¬
as, Baja California, Sonora, Nuevo Leon, Sinaloa, and Zacatecas.
Discussion. — The main structural features were illustrated by Bohart in Figures
9-12 of the original description. This is the only known Gastrodynerus in the
U.S., where its slender form, pronounced postocellar polished knobs, interocellar
swellings, asulcate and bent S-II, dark ocular emarginations, and polished pronotal
front margin make it easy to recognize. Most Mexican specimens lack the free
lateral spot on T-II but are quite similar otherwise.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 16-19
Published 27 January 1984
Is Drosophila pseudoobscura a Garbage Species?
Jerry A. Coyne, Ian A. Boussy, and Stephen Bryant
(JAC, IAB) Department of Genetics, The University of California at Davis,
Davis, California 95616; (SHB) Department of Biological Sciences, Western Il¬
linois University, Macomb, Illinois 61455. Present address: (JAC) Department
of Zoology, University of Maryland, College Park, Maryland 20742.
An enormous amount is known about the genetics of natural populations of
Drosophila pseudoobscura and its sibling species D. persimilis. Spatial and tem¬
poral patterns of the frequencies of allozymes, lethal genes, and third-chromosome
inversions have been studied in detail by Dobzhansky, his students, and colleagues
(Dobzhansky and Epling, 1944; Dobzhansky, 1970; Lewontin, 1974; Anderson
et al., 1975). In many instances, however, the interpretation of these patterns has
been hampered by our ignorance of the ecology of these flies. Very little is known
about where these species live, feed, and breed in nature. We report here findings
which suggest that they are at least partial human commensals and can be fre¬
quently found breeding in agricultural refuse.
There are only a few reports of immature stages of D. pseudoobscura and D.
persimilis in nature. Carson and his colleagues (1951, 1956) discovered a large
number of larvae and pupae in slime fluxes of the California black oak ( Quercus
kelloggii ) and one larva in a flux on white fir {Abies concolor) in the Sierra Nevada.
Additional larvae were found in black oak fluxes in the San Jacinto mountains
of California (Carson, 1978). Dobzhanksy and Epling (1944) describe two addi¬
tional slime flux breeding sites: one on the introduced deodar cedar {Cedrus
deodara) in Beverly Hills, California, and the other on an injured California
grapevine ( Vitis californica) in Sonoma, California. However, Carson (1978) re¬
ported that adult D. pseudoobscura and D. persimilis could be found in areas with
no apparent slime fluxes. This suggests that the fluxes may not be the major
breeding site—or even an important one.
Implicit in many studies, and in much written about these species, is the as¬
sumption that they breed only in natural substrates and are not associated with
human activity. However, there has been sporadic evidence that they may be
partial human commensals. Noting that D. pseudoobscura could be collected
around human habitation, Dobzhansky (1965) and Carson (1965) suggested that
the species had the potential to become a domestic colonizer. In addition, H.
Ikeda and H. Carson found young D. pseudoobscura adults in an isolated garbage
can in Malibu, California, although the flies were not observed hatching directly
from the garbage (Carson, 1971). Finally, Spencer (1944) reared 10 D. pseudoob¬
scura from 27,805 Drosophila pupae collected in a citrus dump near Azusa,
California. Despite these observations, the conventional wisdom appears to be
that these species are largely confined to natural habitats.
The possibility of human association and transport is nevertheless worth con¬
sidering since it may help explain certain genetical phenomena, especially the
synchronous rise and fall of inversion frequencies in many populations (Anderson
Table 1. Drosophila flies reared from agricultural refuse.
Location
Date collected
Substrate
Obscura group
flies reared
Obscura group
flies identified
Other species reared 1
Riverside, CA
Biocontrol citrus grove
May 1970
oranges
325
_2
me, si, im, re
grapefruit
503
_2
me, si, im, re
March 31, 1971
grapefruit
4781
_2
me, si, im
April 4, 1981
oranges
51
_2
me, si, im, re, bu
April 5, 1982
oranges
305
_2
me, si, im
Mill Valley, CA
Residence
Sept. 14, 1979
compost
7
1 pe, 6 az
me, si
Winters, CA
Fruit orchard
Nov. 1, 1979
pomegranates
1
1 pe
me, si
June 15, 1980
peaches
122
39 ps
me, si, im, re
cherries
83
26 ps
me, si, im, re
March-May 1982
oranges
139
52 ps
me, si, im
grapefruit
602
22 ps, 1 pe
me, si, im, re
tangerines
5
5 ps
si
lemons
246
75 ps
me, si, im, re
July 29, 1982
figs
63
33 ps
me, si
Camino, CA
Fruit orchard
Nov. 4, 1979
pears
47
47 ps
me, si, im
prunes
62
43 ps
me, si, im
apples
33
29 ps
me, si
Fruit warehouse
Jan.-Feb. 1980
apples
16
16 ps
Pompano State Beach, CA
Garbage can
March 8, 1980
orange peel
275
76 ps, 6 pe
Furnace Creek Ranch,
Death Valley, CA
Date grove
March 30, 1980
dates
2
2 ps
me, si
Jan. 17, 1981
dates
6
6 ps
me, si
April 2, 1982
dates
18
18 ps
me, re
1 Key to species of Drosophila : az,
azteca; bu, busckii\ im,
immigrant me, melanogaster,
pe, persimilis ; ps, pseudoobscura ; re,
species of the repleta group
(probably D. hydei ); si, simulans.
2 Probably both D. pseudoobscura and D. persimilis (both species present in the grove).
VOLUME 60, NUMBER 1
18
PAN-PACIFIC ENTOMOLOGIST
et al., 1975), the homogeneity of allozyme frequencies among populations (Le-
wontin, 1974), the reappearance of flies in desert oases following supposed summer
extinction (Jones et al., 1981), and the recent appearance of Drosophila pseu-
doobscura in New Zealand (Parsons and Stanley, 1982). Turner and Jeffrey (1980,
p. 777) find human transport an objectionable explanation since “it can be used
to explain almost any changes in chromosome arrangement patterns observed,”
but we do not think facts should be ignored because they suggest untestable
hypotheses.
Table 1 shows our breeding records of D. pseudoobscura and D. persimilis from
fruit and garbage. Rotting fruit or compost was collected from the ground in areas
where these species come to banana traps. The citrus fruit from the 1970 and
1971 collections in Riverside, California was artificially injured and placed on
the ground in a citrus orchard harboring fly populations. All material was brought
back to the laboratory and placed in jars or rearing cages containing damp sand.
Eclosing flies were counted, and samples of obscura group Drosophila flies were
identified by electrophoresis or karyotyping of larvae from females mated to
laboratory stocks (Anderson et al., 1977).
These records substantially increase the known breeding substrates of these
species, and show that they can indeed be reared from a variety of human¬
generated substrates from different localities. The association of these species with
agriculture appears to be a widespread and not merely a casual phenomenon. In
addition, D. pseudoobscura and D. persimilis are often reared out with the other
well-known “garbage” species D. melanogaster, D. simulans, D. immigrans, D.
busckii, and D. hydei. These species are among those drosophilids most easily
reared in the laboratory, probably because of their polyphagy in nature. The rearing
of D. azteca from compost appears to be the first breeding record of this species.
It is worth noting that a large number of flies can sometimes be bred from a
small amount of material: the piece of orange peel from Pompano State Beach
which produced 275 D. pseudoobscura and D. persimilis measured only 7 x
10 cm.
Drosophila pseudoobscura and D. persimilis thus appear to be facultative gar¬
bage species, partially associated with human activity. This association may play
some role in the genetic patterns described above, since agricultural cultivation
is widespread in the western United States. The remarkable appearance of D.
pseudoobscura in New Zealand could, for example, be the result of the importation
of citrus from the United States.
We should emphasize that we do not believe that all individuals of D. pseu¬
doobscura and D. persimilis derive from human refuse. They can often be collected
in areas far removed from human settlement, though not in the profusion found
in orchards. There may be some natural breeding sites besides slime fluxes, which
remain unknown despite the intensive work of Carson and his colleagues and our
own futile efforts to rear D. pseudoobscura and D. persimilis from “natural”
substrates (leaf litter, fungi, berries, animal feces, etc.). Further work must dem¬
onstrate what proportion of flies in the field originate from “domestic” versus
“natural” breeding sites.
Acknowledgments
We thank Timothy Prout, Phil Ward, and Hampton Carson for comments on
the manuscript. We are also grateful to Dr. Carson for permission to cite his
VOLUME 60, NUMBER 1
19
unpublished manuscript, to John Moore for the 1981 rearing record from Death
Valley, and to Lorraine Barr for doing the electrophoresis. This work was sup¬
ported by an NIH postdoctoral fellowship to JAC and NIH grant GM 22221 to
the UC Davis Department of Genetics.
Literature Cited
Anderson, W. W., F. J. Ayala, and R. E. Michod. 1977. Chromosomal and allozymic diagnosis of
three species of Drosophila. J. Heredity, 68:71-74.
-, Th. Dobzhansky, O. Pavlovsky, J. Powell, and D. Yardley. 1975. Genetics of natural pop¬
ulations. XLII. Three decades of genetic change in Drosophila pseudoobscura. Evolution, 29:
24-36.
Carson, H. L. 1951. Breeding sites of Drosophila pseudoobscura and Drosophila persimilis in the
transition zone of the Sierra Nevada. Evolution, 5:91-96.
-. 1965. Chromosomal morphism in geographically widespread species of Drosophila. Pp.
503-531 in H. G. Baker and G. L. Stebbins (eds.), The genetics of colonizing species. Academic
Press, New York.
-. 1971. The ecology of Drosophila breeding sites. Univ. Hawaii Arboretum Lecture #2, pp.
1-27.
-. 1978. Some thoughts on population structure in the obscura group of Drosophila. Unpub¬
lished ms.
—--E. P. Knapp, and H. J. Phaff. 1956. Studies on the ecology of Drosophila in the Yosemite
region of California. II. The yeast flora of some species of Drosophila. Ecology, 57:538-544.
Dobzhansky, Th. 1965. “Wild” and “domestic” species of Drosophila. Pp. 533-562 in H. G. Baker
and G. L. Stebbins (eds.), The genetics of colonizing species. Academic Press, , New York.
-. 1970. Genetics of the evolutionary process. Columbia University Press, New York.
-, and C. Epling. 1944. Contributions to the genetics, taxonomy, and ecology of Drosophila
pseudoobscura and its relatives. Carnegie Inst. Wash. Pub. 554, Washington, D.C.
Jones, J. S., S. H. Bryant, R. C. Lewontin, J. A. Moore, and T. Prout. 1981. Gene flow and the
geographical distribution of a molecular polymorphism in Drosophila pseudoobscura. Amer.
Nat., 98:157-178.
Lewontin, R. C. 1974. The genetic basis of evolutionary change. Columbia University Press, New
York.
Parsons, P. A., and S. M. Stanley. 1982. Special ecological studies—Domesticated and widespread
species. In M. Ashburner, H. Carson, and J. N. Thompson (eds.), The genetics and biology of
Drosophila. Volume 3a. Academic Press, New York.
Spencer, W. P. 1944. Iso-alleles of the bobbed locus in Drosophila hydei populations. Genetics, 29:
520-536.
Turner, M. E., and D. E. Jeffry. 1980. Drosophila pseudoobscura of the Great Basin. II. Third
chromosome arrangements of selected northern Utah populations. Amer. Nat., 115:771-779.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 20-22
Published 27 January 1984
Further Data on Mucrochernes hirsutus (Banks)
(Pseudoscorpionida, Chernetidae)
William B. Muchmore
Department of Biology, University of Rochester, Rochester, New York 14627.
Mucrochernes hirsutus was redescribed and placed in the new genus Mucrocher¬
nes on the basis of a single individual, the holotype female of Atemnus hirsutus
Banks, which was “found 10 ft. from ocean. No trees near.” at Laguna Beach,
Orange County, California (Muchmore, 1973). More recently I have received
from Kenneth W. Cooper, University of California, Riverside, additional material
which sheds more light on the nature of this very interesting species.
Mucrochernes hirsutus (Banks)
(Figs. 1-5)
Material.— Three males and 3 females “from between splits in intertidal rocks,”
at San Pedro, Los Angeles County, California, 14 October 1975, K. W. and R.
Cooper. Two males and 2 females have been mounted on slides and studied in
detail.
Supplemental description.— The males are generally similar to the females ex¬
cept for genital features and slightly deeper palpal chelal hands in some males
(Figs. 1-3). There is considerable variation in size and proportions in both sexes
and usually the larger specimens are the more robust.
As in the holotype, all setae long, thin, and completely simple or with a single
lateral spinule. Carapace with about 200 setae. Abdominal tergal chaetotaxy of
largest male 25:29:32:43:48:53:57:53:57:57:T43T:2. Sternal chaetotaxy of same
57:(2)[4-3]/42(3):(3)28(3):47:61:59:56:50:47:T39T:2; anterior genital operculum
with 10 long setae flanked by 47 shorter ones; posterior operculum with two
groups of very small setae, [4-3], at middle of anterior margin and 42 setae
scattered over face and along posterior margin. Internal genitalia of male rather
compact, but with no outstanding features. Spermathecae of female as shown in
Figure 4, somewhat longer and more convoluted than was noticed in the holotype.
Though not mentioned earlier, the anterior tracheal trunks are greatly expanded
anteriorly (Fig. 5). Cheliceral galea variously branched in different specimens, but
seems equally developed in males and females. Palps of the newly collected females
essentially like those of holotype (Muchmore, 1973: Figs. 4, 5); it appears, there¬
fore, that distortion of the holotype was not serious. For the 4 mounted specimens
(males and females), palpal femur 2.95-3.1, tibia 2.6-2.9, and chela 3.1-3.2 times
as long as broad; hand 1.1-1.35 times as long as deep; movable finger 1.15-1.25
times as long as hand. Chelal fingers with 50-60 marginal teeth and 4-8 internal
and external accessory teeth. Leg IV with entire femur 4.0-4.3 and tibia 4.7-5.5
times as long as deep.
Measurements (mm). — Body length 2.75-3.65. Carapace length 0.89-1.10. Che-
licera length 0.32-0.41. Palpal femur 0.77-1.04 by 0.25-0.34; tibia 0.755-1.00
by 0.28-0.36; chela (without pedicel) 1.42-1.89 by 0.45-?; hand (without pedicel)
VOLUME 60, NUMBER 1
21
Figures 1-5. Mucrochernes hirsutus (Banks). 1. Right palp of largest male, dorsal view. 2. Left
chela of largest male, lateral view. 3. Left chela of smallest male, lateral view. 4. Spermathecae of
female. 5. Spiracle and anterior tracheal trunk.
0.66-0.86 by 0.49-0.78; pedicel about 0.12 long; movable finger 0.77-1.0 long.
Leg IV: entire femur 0.67-0.92 by 0.17-0.215; tibia 0.53-0.74 by 0.11-0.135.
Remarks. — This species is apparently well adapted for life on the seashore,
where it may be periodically covered by water: 1) The general hairiness probably
serves to keep the surface of the animal dry by trapping air between the bases of
the setae. A similar, but not so great, increase in numbers of setae is seen in
Pachyolpium atlanticum Mahnert and Schuster (1981), a species which inhabits
the tidal zone in Bermuda. 2) The expanded anterior tracheal trunks appear
admirably suited to hold air during the time the pseudoscorpion is covered by
water. No other instance is known of this phenomenon. 3) The galea is very well
developed in both male and female, suggesting that both sexes of the adults spin
22
PAN-PACIFIC ENTOMOLOGIST
silken cocoons, possibly to protect them from the tides. A similar situation was
noted with respect to the littoral Epactiochernes tumidus (Banks) in southeastern
United States (Muchmore, 1974). 4) The genera Corosoma Karsch and Dasy-
chernes Chamberlin are also characterized by having very large numbers of ves-
titural setae (cf. Mahnert, 1982). These two genera are, however, apparently mod¬
ified for quite a different function from Mucrochernes, namely for protection (?)
in the hives of meliponine bees. Also, in these forms, while the vestitural setae
are long and thin, many are finely denticulate toward the tips (Mahnert, and
personal observation).
Acknowledgments
I am much indebted to Kenneth Cooper for sending me the specimens and to
C. H. Alteri for drawing the figures.
Literature Cited
Banks, N. 1914. A new pseudoscorpion from California. J. Entomol. Zool., Pomona Coll., 6:203.
Mahnert, V. 1982. The pseudoscorpion genus Corosoma Karsch, 1879, with remarks on Dasychernes
Chamberlin, 1929 (Pseudoscorpiones: Chemetidae). J. Arachnol., 10:11-14.
-, and R. Schuster. 1981. Pachyolpium atlanticum n. sp., ein Pseudoskorpion aus der Gezei-
tenzone der Bermudas—Morphologie und Okologie (Pseudoscorpiones: Olpiidae). Rev. Suisse
Zool., 88:265-273.
Muchmore, W. B. 1973. A new genus of pseudoscorpions based upon Atemnus hirsutus (Pseudo-
scorpionida, Chemetidae). Pan-Pacific Entomol., 49:43-48.
-. 1974. Pseudoscorpions from Florida. 3. Epactiochernes, a new genus based upon Chelanops
tumidus Banks (Chemetidae). Florida Entomol., 57:397-407.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 23-29
Published 27 January 1984
World Synopsis of the Riffle Beetle Genus Leptelmis
Sharp, 1888, with a Key to Asiatic Species and
Description of a New Species from India
(Coleoptera, Dryopoidea, Elmidae)
Harley P. Brown and M. P. Thobias
(HPB) Department of Zoology (and Stovall Museum of Science and History),
University of Oklahoma, Norman, Oklahoma 73019; (MPT) St. Thomas’ College
(University of Calicut), Trichur, Kerala 680 001, India.
Abstract.— The genus Leptelmis is close to Stenelmis, but more similar to Ste-
thelmis. Citations for each known species of Leptelmis (15 Asiatic, 8 African)
include reported distribution, all known illustrations, and location of holotype.
A key to Asiatic species is based primarily upon the literature. Leptelmis philom-
ina, n. sp. from the Western Ghats of southern India is described and figured
(habitus, aedeagus).
Sharp (1888) created the genus Leptelmis upon the basis of one mutilated
specimen from Tokyo, Japan. He wrote: “This insect, though allied to Stenelmis,
cannot be placed in that genus; it differs in several respects, more especially in
the great separation of the hind legs and the very dependent front of the proster¬
num. The form is different from that of Stenelmis, and the sculpture is not similar
to that of any other Elmid known; the eyes, too, are much smaller than in Ste¬
nelmis, and the terminal joint of the maxillary palpus more slender.” Sharp did
not mention that a major feature linking Leptelmis to Stenelmis was the lack of
a fringe of hairy tomentum on the inner margin of the anterior tibia, a character
that separates Stenelmis from virtually all other elmine riffle beetles. More sur¬
prising is the fact that he did not mention the most conspicuous feature separating
Leptelmis from Stenelmis : the anterior portion of the pronotum is set off from
the rest of the pronotum by a broad transverse impression, giving an appearance
reminiscent of a turtleneck sweater—in fact, the anterior portion of the pronotum
often appears at first glance to be part of the head (Fig. 1). Furthermore, the
pronotum lacks the median longitudinal impression so characteristic of Stenelmis,
the legs are quite long, and the tarsal claws are large and with basal teeth. All of
these latter characters not mentioned by Sharp are shared by Stethelmis (Hinton,
1945), which occurs in Chile and Argentina. The general aspect of Leptelmis is
also reminiscent of Ancyronyx in North America (see Brown, 1976 for illustration),
of Hintonelmis in South America (Spangler, 1966; Hinton, 1971), and of Ony-
chelmis in Central and South America (Hinton, 1941).
Bollow (1941) added to the characterization of the genus that the tarsi are 4-
segmented, but he had apparently seen specimens only of L.flavicollis. His Figure
2 shows this feature; however, it is not characteristic of the genus. Most, if not
all, other species have the usual 5 tarsal segments. In contrast with Sharp, Bollow
24
PAN-PACIFIC ENTOMOLOGIST
characterized the eyes as larger than those of Stenelmis. Actually, the eyes vary
considerably in size, depending upon the species.
Nothing is known about the immature stages of Leptelmis, and very little about
the habitat, habits, or behavior of the adults. The little that has been published
is entirely from Deleve (1966). Most specimens are presumably from streams,
but even this is seldom stated in the literature. The holotype of L. fragilis was
found on vegetation in a flowing stream. From the morphology of Leptelmis, this
is the habitat we would expect, especially in view of the nature of the legs and
tarsi. Other elmids with similar structure, such as Ancyronyx and Hintonelmis,
are typically found clinging tenaciously to waterlogged wood or other submerged
vegetation, and it is a good guess that Leptelmis exhibits similar behavior. Two
of the paratypes of L. major were taken from forest humus. We can only wonder
how they happened to be there and how far they were from water, e.g., it would
be of interest to know whether the site had been inundated a few weeks earlier.
Deleve (1945, 1966) reported that both macropterous and brachypterous indi¬
viduals occur within the same species (e.g., L. fragilis and L. orchymonti). Only
macropterous forms, of course, have been taken at lights, but the fact that some
come to lights informs us that Leptelmis can utilize flight as a means of dispersal—
a common ability among elmids just after emergence from pupation.
Of the 117 known genera of elmids, only Leptelmis and Stenelmis occur in
both Africa and the Orient (Brown, 1981). Leptelmis is also unusual in being the
largest elmid genus for which the larva is unknown. Perhaps our pointing out the
large gaps in our knowledge of this group will stimulate investigators to fill the
gaps.
While collecting in streams of the Western Ghats (Sahyadri Mountains) of South
India, one of us (M. P. Thobias) obtained two specimens of Leptelmis that ap¬
parently represent a new species, which we here describe.
Leptelmis philomina , New Species
(Figs. 1, 2)
Holotype male. —Length 2.15 mm, width 0.75 mm. Subparallel, convex. Cuticle
tuberculate, feebly shining; brown, with pronotum and head darker than elytra;
anterior margin of pronotum, antennae, palps and legs testaceous (somewhat
darker in paratype). Head retractable within pronotal collar; visible portion with¬
out distinct impressions; on vertex and in a band tapering on each side toward
but not reaching antennal base covered by ashen white plastron studded with
small granules separated by about their own diameters; frons, clypeus and labrum
black, rather finely punctate; circumocular area black. Eyes rather small, their
greatest dimension being less than interocular distance. Antennae barely reaching
elytral humeri, slender, with segments 3-11 progressively longer. Frontoclypeal
suture indistinct. Maxillary palp with apical segment elongate and somewhat larger
than apical antennal segment. Pronotum slightly longer than broad, 0.625: 0.575
mm; widest at basal 2 / 5 ; width at base 0.550 mm; width at transverse impression
0.465 mm; width at apex 0.50 mm. Sides conspicuously bisinuate; anterior angles
broadly rounded as seen from above, posterior angles subacute; apical margin
arcuate and projecting over the head; lateral borders feebly margined, most no¬
ticeably so in posterior third; lateral margins not perceptibly crenate. Surface
rather coarsely granulate or tuberculate, the flattened granules on disk separated
VOLUME 60, NUMBER 1
25
Figures 1, 2. Leptelmis philomina n. sp. 1. Dorsal aspect of adult. 2. Aedeagus, ventral aspect.
by about their own diameters. Without sublateral carinae. Transverse impression
at anterior third broad and deep, giving the appearance of a bottleneck. With
prominences as follows: a feeble, flattened, median longitudinal ridge in the basal
quarter suggests a rather long, narrow human nose with the scutellum representing
a small chin; continuing the comparison with a human face, each “cheek” is low
and composed of a doubled or divided very feeble prominence, and the “eyes,”
though small, bulge rather prominently; each “eye” is located half way between
the midline and lateral margin on the posterior edge of the major transverse
impression and anterolateral to the anterior end of the “nose”; between the “nose”
and each “eye” there is also a minor elevation. Scutellum small (0.075 mm across),
about as long as wide, with sides rounded. Elytra more than twice as long as
pronotum (1.5 mm); humeri not prominent, but distinctly broader than pronotum;
sides subparallel in anterior half, then tapering arcuately in posterior half to
rounded apex. Strial punctures on disk rather large, subcircular, and separated
from one another by slightly less than their own diameters; punctures smaller and
shallower on apical declivity. Strial intervals rather flat, none genuinely carinate,
but third somewhat elevated at base and intervals 6-8 fused basally and slightly
elevated basally; intervals 2 and 3 merge on apical declivity. Lateral borders very
feebly margined and finely serrate. Epipleura narrowed gradually toward apex.
Venter and legs (except tarsi) covered with flattened granules separated by less
than to about twice their own diameters and suggestive of a medieval coat of
mail; each granule bearing an inconspicuous, short, decumbent seta. Although we
have not observed living specimens under water, apparently the plastron covers
the entire ventral surface, entire pronotum, vertex of head, and all of the legs
except the tarsi. Prosternum rather long anterior to procoxae, deflected in such a
way as to permit retraction of head until only frons, clypeus, and labrum are
exposed, together with eyes and antennae; granules on deflected portion smaller
and sparser; prosternal process without carinae, about equal in width to diameter
26
PAN-PACIFIC ENTOMOLOGIST
of globular procoxa, slightly longer than wide, broadly rounded at apex, feebly
margined, coarsely granulate. Mesosternum short, coarsely granulate, with a shal¬
low anteromedian excavation to accommodate prosternal process. Metasternum
with a bare area between mesocoxae and anterior to median impressed line.
Abdomen without carinae or other conspicuous features; cuticular granules very
coarse on segment 1, progressively less coarse posteriorly; segment 1 longest, 2-
4 progressively shorter, but 5 almost as long as 1; posterior margins of segments
1-4 straight to broadly emarginate, but 5 sinuate laterally and rather narrowly
rounded at apex. Sternita without conspicuous teeth or other projections for
attachment to elytra. Legs long and slender, especially the tibiae and tarsi; tibiae
almost straight and cylindrical, without tomental brushes, spines, or other con¬
spicuous features but with small inner apical projections and inconspicuous rows
of small setae along inner margins distally. Tarsi 5-segmented, that of foreleg
shortest and that of hind leg longest, each with the segments progressively longer
from base to apex, the apical segment of each being longer than the other 4
combined; segments 1-4 each with a short apical tuft of pale setae on ventral
surface; claws large, each with a basal tooth and curved ca. 90 degrees. Genitalia
(Fig. 2) rather standard in general aspect, 0.50 mm in length; basal piece 0.26
mm; penis or median lobe 0.30 mm; parameres 0.23 mm. As viewed dorsally or
ventrally, the parameres are slender distally with bluntly rounded apices and the
relatively broad penis projects beyond them, tapering to a blunt point.
Female. — Unknown.
Type. — Male to be deposited in the British Museum (Natural History), London;
South India: Western Ghats, leg. M. P. Thobias. Genitalia mounted in Hoyer’s
medium on slide #Lpp-Agm-l, at BMNH.
Paratype.—OnQ male with same data as type, in Stovall Museum of Science
and History, University of Oklahoma, Norman, Okla., U.S.A.
Etymology. — This species is named in honor of the beloved mother of the junior
author.
Comments.— As shown in the list below, only one species of Leptelmis, L.
fracticollis Champion, has previously been reported from India, that being from
the Haldwani Division of Kumaon, far to the north (in the foothills of the Him¬
alaya Mountains north or northeast of Delhi. Much closer geographically is the
island of Sri Lanka (Ceylon), from which two species of Leptelmis have been
described: L. nietneri Champion and L. cederholmi Deleve. L. philomina n. sp.
differs from L. fracticollis in pronotal sculpture, in being less broad across the
humeri, and in having the cuticle granulate or tuberculate ventrally rather than
punctate. From L. nietneri, the new species differs in being considerably larger
(2.15:1.5 mm), in lacking elytral maculae, and in having a much deeper transverse
prothoracic impression. L. philomina differs from L. cederholmi in having the
venter granulate rather than punctate and in aedeagal structure: in L. cederholmi
the basal piece is proportionately shorter, the parameres broader, the penis longer
and parallel-sided with a subapical denticulate collarette. Differences from and
among other Asiatic species are indicated in the key below, which is based largely
upon characters extracted from the literature. It includes 13 of the 15 known
Asiatic species, L. parallela and L. formosana being omitted because we cannot
distinguish them from L. gracilis by the descriptions.
VOLUME 60, NUMBER 1
27
Key to Asiatic Species of Leptelmis (Except L. parallela
from Japan and L. formosana from Taiwan)
1. Elytra maculate . 2
Elytra essentially uniform in color or with the humeri lighter. 5
2. Each elytron with 5 yellow spots (Viet Nam) . L. signata
Elytron with fewer than 5 spots or markings. 3
3. Very small (1.5 mm); each elytron with an oblique testaceous streak (Sri
Lanka). L. nietneri
Larger than 1.5 mm; elytral markings not oblique . 4
4. Small (1.8 mm); elytral vitta in apical half on intervals 2 and 3 (Philippine
Islands) . L. tawitawiensis
Larger (2.3 mm); elytral vitta in apical half on intervals 3 and 4, and
expanded apically (Viet Nam) . L. basalis
5. Elytra expanded posteriorly, much wider than pronotum; 2.5-2.8 mm
(Japan) . L. gracilis
Elytra not expanded posteriorly . 6
6. Tarsi 4-segmented, head as broad as pronotum; 2.4 mm (South China)
. L. flavicollis
Tarsi 5-segmented . 7
7. Large (3 mm); pronotum with anterior transverse impression indistinct
medially; cuticle very shiny (Sumatra). L. stricticollis
Smaller; pronotum with transverse impression complete and distinct . . 8
8. Venter granulate; 2.15 mm (South India) . L. philomina, n. sp.
Venter punctate rather than granulate. 9
9. Antennae reddish. 10
Antennae yellow or yellowish at least at base . 11
10. Penis parallel-sided, with subapical collarette of denticles; parameres ta¬
pering from near base (Sri Lanka). L. cederholmi
Penis constricted near middle, without denticles; parameres subparallel
in apical half (Sumatra, Java) . L. sulcata
11. Smaller (2 mm) (North India) . L. fracticollis
Larger (2.2 mm or more) . 12
12. Elytra with strial interval 3 raised from base to apex; aedeagus with
parameres (in ventral aspect) tapering from base almost to apex (Viet
Nam) . L. obscura
Elytra with strial interval 3 feebly raised at base only; parameres sub¬
parallel in apical half (Viet Nam) . L. vietnamensis
World Checklist of the Species of Leptelmis
Zaitzev (1910) cited the type species and emended the generic name to Leph-
thelmis. Since this emendation was unjustified by the rules of zoological nomen¬
clature, it has been quietly dropped, except by Bollow (1941). Sato (1960) pub¬
lished a world checklist which at that time included a total of only 8 species,
compared with the present 23.
To be as helpful as possible to others, we indicate in the list below, after the
name and citation, reference to published illustrations, location of holotype, and
28
PAN-PACIFIC ENTOMOLOGIST
reported distribution. Location of holotype is placed in parentheses, with abbre¬
viations as follows: BMNH = British Museum (Natural History), London; FCM =
Frey Collection, Munich; LMS = Lund Museum, Sweden; MNHB = National
Hungarian Museum, Budapest; MPF = Museum of Paris, France; MPSA = Mu¬
seum of Pretoria, South Africa; MRAC = Musee Royal de l’Afrique Centrale;
MRHNB = Musee Royal d’Histoire Naturelle de Belgique, Brussels; NSMT =
National Science Museum, Tokyo; ZMUC = Zoological Museum, University of
Copenhagen, Denmark.
Leptelmis Sharp
Sharp, Ann. Mag. Nat. Hist. (6) II, 1888, p. 243.
Lephthelmis Zaitzev, Coleop. Cat., Pars 17, Dryopidae, etc., 1910, p. 21.
Type Species: L. gracilis Sharp, 1888.
amoena Deleve, 1966, p. 8, f. 15. (MRAC) AFRICA: COTE D’IVOIRE, GHANA.
basalis Deleve, 1968, p. 155, no figs. (MNHB) VIET NAM.
cederholmi Deleve, 1973a, p. 11, f. 23. (LMS) SRI LANKA (CEYLON).
collarti (as Stenelmis ) (Deleve), 1937, p. 152, f. 3. Transferred to Leptelmis by
Deleve, 1938, p. 364. Genitalia in Deleve, 1966, f. 2-4. (MPF) AFRICA:
CONGO (ZAIRE), GABON, CONGO (BRAZZAVILLE).
costulata Deleve, 1942, p. 1, f. 1 (habitus); genitalia in Deleve, 1966, f. 1. (MRHNB)
AFRICA: CONGO (ZAIRE), CONGO (BRAZZAVILLE).
flavicollis Bollow, 1941, p. 86, T. 7, f. 2 (habitus). (FCM) SOUTH CHINA.
formosana Nomura, 1962, p. 48, no figs. (NSMT?) TAIWAN.
fracticollis Champion, 1923, p. 167, f. 3 (habitus). (BMNH) NORTH INDIA.
fragihs Deleve, 1966, p. 7, f. 13, 14. (MPSA) SOUTH AFRICA: VAAL.
gracilis Sharp, 1888, p. 244; habitus, hind tarsus, maxillary palp, prosternal pro¬
cess, and male genitalia in Sato 1960, f. 1, 2. (BMNH) JAPAN: HONSHU,
SHIKOKU, KYUSHU.
major Deleve, 1966, p. 4, f. 8-10. (MRAC) AFRICA: CONGO: ELIZABETH-
VILLE, KIVU; TANGANYIKA.
nietneri Champion, 1923, p. 168, no figs. (BMNH) CEYLON (SRI LANKA).
obscura Deleve, 1968, p. 153, f. 6. (MNHB) VIET NAM.
orchymonti Deleve, 1942, p. 3; genitalia in Deleve, 1966, f. 5-7. (MRHNB)
AFRICA: CONGO: ELIZABETHVILLE; RUANDA; SOUTHERN RHO¬
DESIA.
parallela Nomura, 1962, p. 46, no figs. (NSMT?) JAPAN: HONSHU, SHIKOKU,
KYUSHU.
philomina Brown and Thobias, n. sp. (BMNH) SOUTH INDIA.
seydeli Deleve, 1966, p. 6, f. 11, 12. (MRAC) AFRICA: CONGO: ELIZABETH¬
VILLE.
signata Deleve, 1968, p. 152, f. 5. (MNHB) VIET NAM.
sobrina Deleve, 1974, p. 272, f. 1. (MNHB) AFRICA: GHANA.
stricticollis (as Stenelmis ) (Grouvelle), 1896, p. 45, no figs. Transferred to Lep¬
telmis by Hinton, 1941, p. 88. (syntypes in BMNH, MPF) SUMATRA, PA-
LEMBANG.
sulcata (as Stenelmis ) (Grouvelle), 1892, p. 188; genitalia in Deleve, 1970, f. 1,
2. Transferred to Leptelmis by Deleve, 1970, p. 236. (MPF) SUMATRA, JAVA.
VOLUME 60, NUMBER 1
29
tawitawiensis Deleve, 1973, p. 28, f. 20. (ZMUC) PHILIPPINES: TAWI TAWI.
vietnamensis Deleve, 1968, p. 154, f. 7. (MNHB) VIET NAM.
Literature Cited
Bollow, H. 1941. Monographic der palaearktischen Dryopidae mit Beriicksichtigung der eventuell
transgredierenden Arten. (Col.) (Fortsetzung). Mitteil. Miinchner Ent. Ges., 31(1): 1-89.
Brown, H. P. 1976. Aquatic dryopoid Beetles (Coleoptera) of the United States. U.S. Environmental
Protection Agency, Cincinnati, Ohio, 82 pp. (Paperback reprint of 1972 book.)
-. 1981. A distributional survey of the world genera of aquatic dryopoid beetles (Coleoptera:
Dryopidae, Elmidae, and Psephenidae sens. lat.). Pan-Pacific Ent., 57(1): 133—148.
Champion, G. C. 1923. Some Indian Coleoptera (11). Ent. Mo. Mag., 59:165-179.
Deleve, J. 1937. Dryopidae du Congo beige. Bull. Ann. Soc. Ent. Belg., 77:149-164.
-. 1938. Dryopidae du Congo beige. II. Dryopinae. III. Elmmae. Rev. Zool. Bot. Afr., 31(3-
4):351-375.
-. 1942. Contribution a l’etude des Dryopidae. I. Deux Leptelmis nouveaux du Congo beige.
II. Remarque sur le genre Microdinodes et description d’une espece nouvelle. Bull. Mus. Roy.
Hist. Nat. Belg., 18(59): 1-8.
-. 1945. Contribution a l’etude des Dryopidae. III. Le genre Pseudo macro nychus Grouvelle et
le dimorphisme alaire de ses especes. Bull. Mus. Hist. Belg., Brussels, 21(9): 1-12.
-. 1966. Contribution a l’etude des Dryopoidea. XVIII. Notes complementaires sur les especes
africaines du genre Leptelmis Sharp et descriptions d’especes nouvelles (Coleoptera Elminthi-
dae). Bull. Inst. Roy. Sci. Nat. Belg., 42(8): 1-10.
-. 1966a. Dryopidae et Elminthidae de 1’Angola. Separata de No. 76 das publicafoes culturais
da Companhia de Diamantes de Angola, Lisboa, pp. 39-62.
-. 1968. Dryopidae et Elminthidae (Coleoptera) du Vietnam. Ann. Hist.-Nat. Mus. Nat. Hung.,
Pars Zool., 60:149-181.
-. 1970. Contribution a l’etude des Dryopoidea. XXL Elminthidae (Coleoptera) peu ou mal
connus de l’lndonesie et du Vietnam. Bull. Ann. Soc. Roy. Ent. Belg., 106:235-273.
-. 1973. Limnichidae, Dryopidae et Elminthidae des lies Philippines et de l’Archipel Bismarck
(Insecta, Coleoptera, Dryopoidea). Steenstrupia (Zool. Mus. Univ. Copenhagen), 3(3): 17-30.
-. 1973a. Coleoptera: Dryopidae et Elminthidae. Ent. Scand. Suppl. 4:5-23. (=Report No. 25
from the Lund Univ. Ceylon Expedition in 1962.)
-. 1974. Elminthidae (Coleoptera) du Ghana. Acta Zool. Acad. Sci. Hung., 20(3-4):271-289.
Grouvelle, A. 1892. Trois especes nouvelles d’helmides des lies de la Sonde. Not. Leyden Mus., 14:
187-190.
-. 1896. Note VI. Descriptions de dryopides (Parnides) et helmides nouveaux. Not. Leyden
Mus., 18:33-52.
Hinton, H. E. 1941. New genera and species of Elmidae. Trans. Roy. Ent. Soc. London, 91 (3):65—
104.
-. 1945. Stethelmis chilensis, new genus and species of Elmidae from Chile (Coleoptera). Proc.
Roy. Ent. Soc. London (B), 14(5-6):73-76.
-. 1971. A revision of the genus Hintonelmis Spangler (Coleoptera: Elmidae). Trans. Roy. Ent.
Soc. London, 123(2): 189-208.
Nomura, S. 1962. Some new and remarkable species of the Coleoptera from Japan and its adjacent
regions. Toho-Gakuho, (12):35—51, pis. 1, 2.
Sato, M. 1960. On the genus Leptelmis Sharp from Japan with checklist of the world (Coleoptera:
Elmidae). Akitu, 9:43-46.
Sharp, D. 1888. XXXI. Descriptions of some new Coleoptera from Japan. Ann. Mag. Nat. Hist.,
(6)11:242-245.
Spangler, P. J. 1966. The Catherwood Foundation Peruvian-Amazon Expedition. XIII. Aquatic
Coleoptera (Dytiscidae; Noteridae; Gyrinidae; Hydrophilidae; Dascillidae; Helodidae; Pse¬
phenidae; Elmidae). Mon. Acad. Nat. Sci. Philadelphia, 14:377-443.
Zaitzev, P. A. 1910. Dryopidae, Cyathoceridae, Georyssidae, Heteroceridae. Coleopterorum Cata¬
logs, Pars 17, Vol. 14:1-68.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 30-32
Published 27 January 1984
Captiia hornigi , a New Winter Stonefly from the Western
Great Basin (Plecoptera: Capniidae)
Richard W. Baumann and Andrew L. Sheldon
(RWB) Department of Zoology, Brigham Young University, Provo, Utah 84602;
(ALS) Department of Zoology, University of Montana, Missoula, Montana 59812.
During a recent study on the distribution of stonefly species in the Great Basin
ranges of Nevada (Sheldon, 1979) a previously undescribed Capnia species was
discovered. The faunal affinities of this species are not with the Rocky Mountain
fauna (Baumann et al., 1977) but are with the Sierra Nevada fauna (Jewett, 1960).
Capnia hornigi Baumann and Sheldon, New Species
Male.— Wings macropterous. Length of forewing 4-5 mm; length of body 4-6
mm. Body and appendages dark brown; wings hyaline except for fumose area in
costal space beyond cord. Abdominal segments one to six without special mod¬
ifications. Seventh tergum with median posterior portion modified as large pos¬
terior directed process, apex of process darkly sclerotized and narrowly rounded.
Eighth tergum reduced to narrow sclerotized band, posterior portion of tergum
membranous. Ninth tergum produced dorsally into low bifurcate ridge upon which
epiproct rests. Tenth tergum with large reflexed epiproct which extends forward
to posterior margin of ninth tergum; base of epiproct stout laterally, slightly
narrowed at bend, becoming gradually broader anteriorly with rounded apex,
small membranous area visible at apex, bordered by narrow triangular pointed
process on each side dorsally; dorsal aspect of epiproct with narrow base, becoming
abruptly broader toward apex, base sclerotized but with membranous area ap¬
pearing slightly anterior to base which becomes increasingly larger until it covers
the complete apex, much like in the genus Paracapnia (Figs. 1, 2).
Female.— Wings macropterous. Length of forewing 6-7 mm; length of body
6-8 mm. General appearance similar to male. Dorsal broad membranous stripe
extends from abdominal terga one to eight, with sclerotized triangular patch at
base of eighth tergum. Subgenital plate well developed and very darkly sclerotized
encompassing entire eighth sternum, plate flat at base and broadly rounded at
apex, not extending over ninth sternum, joined broadly to seventh sternum with
no apparent separation except coloration. Oblong, sclerotized patches present on
lateral posterior margins of eighth segment, bordering subgenital plate (Fig. 3).
Material. — Holotype male, #76499, allotype and two male and seven female
paratypes, Middle Creek, White Mountains, Esmeralda Co., Nevada 10-11-1977,
A. L. Sheldon. Types deposited at the U.S. National Museum, Washington, D.C.
Additional paratypes: Nevada, Esmeralda Co., Middle Creek, White Mountains,
4-III-1979, C. E. Hornig, 14 males, 17 females.
Etymology. — This species is named for Evan Hornig who collected some of the
paratypes of this new species. Evan is an avid student of aquatic insects and has
collected many interesting specimens in Nevada and surrounding states.
Diagnosis. — Capnia hornigi is the sister species of Capnia barberi Claassen from
VOLUME 60, NUMBER 1
31
Figures 1-3. Capnia hornigi. 1. Male genitalia, dorsal. 2. Male genitalia, lateral. 3. Female genitalia,
ventral.
the Sierra Nevada. The projection on the male seventh tergum is much more
massive in C. hornigi. The epiproct is more rounded in lateral view and apically
in dorsal view in C. hornigi. Capnia barberi has an epiproct which is much more
angular and pointed at the apex. The processes on the dorsal lateral areas of the
epiproct near the apex are larger and much more distinctive in C. barberi.
The female subgenital plate is broadly rounded apically in C. hornigi where in
C. barberi it is flat with sharply angled corners.
Ecology.— Adults of Capnia hornigi were collected at three locations (2160-
2470 m) on Middle Creek. No adults were collected at the highest station sampled
(2590 m) but capniid nymphs were present. Capnia utahensis was the only capniid
emerging on nearby Chiatovitch Creek (2100 m) and no capniids were collected
on the open bajada (1950 m) below the confluence of Middle and Chiatovitch
creeks. Collections made at other times may, however, extend the altitudinal
range of C. hornigi.
32
PAN-PACIFIC ENTOMOLOGIST
Middle Creek originates at 3650 m on the east side of the White Mountains
on the slopes of the Jumpoff and Mt. Montgomery (4097 m). At the collecting
sites, the substrate is dominated by shifting granitic sands while stable gravels
and cobbles are rare. Channel structure is regulated to a considerable extent by
roots and limbs of cottonwood and water birch ( Betula occidentalis). Leaf packs
and debris dams are common.
Associated species (collected in Middle Creek) were Zapada cinctipes, Yoraperla
brevis, Frisonia picticeps, Sweltsa townesi and Triznaka diversa. Most of these
species are common in the Sierra Nevada and C. hornigi may occur there also.
Acknowledgments
This study was supported in part by Grant 7717 from the Penrose Fund of the
American Philosophical Society.
The drawings were made by Connie A. Bevan Bhagat.
Literature Cited
Baumann, R. W., A. R. Gaufin, and R. F. Surdick. 1977. The stoneflies (Plecoptera) of the Rocky
Mountains. Mem. Amer. Entomol. Soc., 31:1-208.
Jewett, S. G., Jr. 1960. The stoneflies (Plecoptera) of California. Bull. Calif. Insect Surv., 6:125-177.
Sheldon, A. L. 1979. Stonefly (Plecoptera) records from the basin ranges of Nevada and Utah. Great
Basin Natur., 39:289-292.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 33-36
Published 27 January 1984
A New Species of Dichaetocoris Knight from the
Western United States, with Notes on Other
Species (Hemiptera: Miridae)
Dan A. Polhemus
Department of Biology, University of Utah, Salt Lake City, Utah 84112.
Collections made in western Colorado and southeastern Utah over the past
several years have turned up many specimens of the poorly known genus Di¬
chaetocoris. Among these was a new species, D. anasazi, described herein. The
material reported here is held in the J. T. Polhemus collection (JTP) unless oth¬
erwise noted.
Measurements are given in units, with 40 units = 1 mm, unless otherwise noted.
Dichaetocoris anasazi, New Species
Description. — Male: Small, ovate; length 2.87 mm; maximum width 0.91 mm.
Coloration yellowish orange, occasionally tinged with green on venter and along
anterior angles of pronotum.
Head yellowish orange, eyes black; frons set with erect black setae intermixed
with fine recumbent silky pubescence; gula with a patch of pale erect setae; vertex
with erect black setae; width of vertex over twice the dorsal width of an eye.
Antennae amber brown, clothed with short stiff setae; segment I with three stout
setae; antennal proportions I-IV: 8; 27; 25; 9.
Pronotum yellowish orange, smooth, shining, width/length = 36/16; margins
set with dark pubescence; calli indistinct; posterior angles rounded. Scutellum
amber, smooth, wider than long, width/length = 23/17; mesoscutum broadly ex¬
posed.
Hemelytra yellowish orange, semi-transluscent, surface faintly rugulose; set with
black recumbent setae interspersed with fine, pale, recumbent silky pubescence;
margins nearly parallel; cuneus with anterior margin weakly sinuate; wing mem¬
brane dusky grey, veins gold.
Venter light gold, often shaded with green along femora and posterior margin
of genital segment; rostrum gold, tip infuscated, length 1.30 mm, reaching well
beyond tips of hind coxae. Legs golden brown, covered with short erect setae;
tibiae armed with stout brown spines; tips of tarsi infuscated. Abdomen sparsely
covered with fine pallid hairs; genital segment with bifurcate tergal process (see
Fig. 3); left clasper slender, curving, acuminate; right clasper broadly bifurcate,
multispinose, distinctive (see Fig. 4).
Female: Similar to male in structure and pubescence, coloration often lighter;
length 3.09 mm; maximum width 1.00 mm.
Material examined.— Holotype, male, and allotype: UTAH, San Juan Co., White
Canyon at Soldier’s Crossing, approx. 5 miles N of Fry Canyon Store, 1219 m
(4000'), V-29-78, D. A. & J. T. Polhemus (JTP). Paratypes: 7 6, 5 9, same data
as types, all taken on Pinus edulis (Pinyon Pine) (JTP); 15 6, 9 9, UTAH, San
Juan Co., Grand Flat nr. Collins Canyon, VI-1-82, D. A. & J. T. Polhemus (JTP);
34
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Dichaetocoris anasazi, n. sp. Male, dorsal view.
Figure 2. Dichaetocoris anasazi, n. sp. Male, lateral view.
VOLUME 60, NUMBER 1
35
Figures 3, 4. Dichaetocoris anasazi, n. sp. 3. Male genital segment, posterior view, showing po¬
sitions of male claspers and bifurcate tergal process. 4. Male right clasper.
1 S, COLO., Montrose Co., 10 miles W of Montrose, VII-8-81, D. A. & J. T.
Polhemus, on Pinus edulis (JTP).
Etymology .—The name anasazi is a Navajo word meaning “ancient ones” and
refers to the prehistoric cliff dwellers who once inhabited the canyons of the
Southwest.
Discussion
Dichaetocoris anasazi, n. sp. runs to couplet 11 in Knight’s (1968) key, where
it may be immediately distinguished by the bifurcate tergal process on the male
genital segment and by the distinctive male right clasper. The right clasper is
36
PAN-PACIFIC ENTOMOLOGIST
characterized by a deep bifurcation dividing two multispinose processes, and may
be easily viewed without dissection (see Figs. 3, 4). The left clasper is similar to
those of several other species, notably D. pinicola Knight, and is thus not diag¬
nostic. No other known species of Dichaetocoris possesses a bifurcate tergal pro¬
cess.
The following records are noted for other species in this genus:
Dichaetocoris utahensis Knight: UTAH, San Juan Co., head of Lake Canyon
along the Nokai Dome road, 4200' (1280 m), V-29-78, 1 3, D. A. & J. T. Polhemus;
Garfield Co., Hog Spring in North Wash, near Hite, 5300' (1615 m), V-30-78,
1 <3, D. A. & J. T. Polhemus (JTP).
Dichaetocoris nevadensis Knight: UTAH, San Juan Co., White Canyon at Sol¬
dier’s Crossing, approx. 5 miles north of Fry Canyon Store, 4000' (1219 m),
V-29-78, 5(3, 1 9, D. A. & J. T. Polhemus; Grand Co., 5 miles south of Moab
along U.S. 163, 4000' (1219 m), V-27-80, 2 <3, D. A. & J. T. Polhemus (JTP).
These records represent the first instances in which either of these species have
been recorded outside the Great Basin. It is likely that these species, and others
of the same genus, occur widely in the pinyon-juniper woodlands across southern
Utah, but have been generally overlooked due to the remote nature of the country
and the fact that these insects mature in late May and early June, well before most
collectors visit the region.
Acknowledgments
The author is indebted to J. T. Polhemus of Englewood, Colorado for helpful
comments and advice in the preparation of this paper.
Literature Cited
Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the Western United
States. Brigham Young Sci. Bull., IX(3): 1-282.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 37^46
Published 27 January 1984
Distribution and Seasonal Abundance of
Brine Flies (Diptera: Ephydridae) in a
San Francisco Bay Salt Marsh
Mark A. Barnby and Vincent H. Resh
Division of Entomology and Parasitology, University of California, Berkeley,
California 94720.
Abstract. — Spatial and temporal distribution patterns of adult brine flies (Dip¬
tera: Ephydridae) were examined in Petaluma Marsh, San Francisco Bay, Cali¬
fornia. Biweekly D-vac samples (November 1977-1978) were collected along three
strata located 1 m, 3 m, and 10 m from three sites, a 2- and a 5-yr-old mosquito
control recirculation ditch and a natural channel. Adults were also collected weekly
(March-December 1980) from ethylene glycol pan-traps designed to simulate
pond and pothole habitats. Of the 28 species collected, 22 occurred in D-vac
samples. Distribution patterns of five were analyzed statistically: three species
[Scatella ( Scatella) stagnalis Fallen, Scatophila sp., and Atissa litoralis (Cole)]
exhibited similar patterns at ditches and natural channels and two species exhib¬
ited dissimilar patterns [ Scatella ( Neoscatella ) setosa Coquillett and Psilopa ( Cer-
opsilopa) coquilletti (Cresson)]. Of 23 species collected in pan-traps, Ephydra
millbrae Jones accounted for 95.5% of the total catch.
As part of a study to evaluate the impact of mosquito control recirculation
ditches on San Francisco Bay marshlands, the arthropod fauna of selected salt
marshes has been examined in terms of community structure and biomass and
the response of selected populations to ditches (Resh and Balling, 1979). The
insect family Ephydridae has been of particular interest since it is one of the most
abundant and species-rich arthropod families collected in these marshlands. The
Ephydridae occur in a variety of aquatic and semi-aquatic environments, but are
most remarkable for their ability to exist in “unique” habitats (e.g., oil pools
[Crawford, 1912], inland saline lakes [Aldrich, 1912], and hot springs [Wirth and
Mathis, 1979]). The purpose of this study has been to examine the seasonality
and distribution of ephydrid adults associated with man-made and natural wa¬
tercourses in a hypersaline Pacific coast salt marsh.
Materials and Methods
Study site.— This study was conducted at Petaluma Marsh, Sonoma Co., Cal¬
ifornia, a 1145-ha salt marsh located in the Petaluma River basin. The marsh
flora is dominated by pickleweed, Salicornia virginica Linnaeus.
A variety of natural aquatic habitats occurs throughout the marsh, e.g., ponds
(10 m-100 m in diameter), potholes (~ 3 m), sloughs (~ 10m), and natural channels
(«3 m). In addition, man-made recirculation ditches have been constructed to
connect a variety of ponds and potholes with natural channels to allow for tidal
38
PAN-PACIFIC ENTOMOLOGIST
flushing, which results in effective mosquito control. Natural channels and recir¬
culation ditches examined in this study were similar in width (40 cm), although
natural channels in general are deeper than ditches (100 cm cf. 60 cm).
Experimental design .—The influence of mosquito control recirculation ditches
and natural watercourses on the distribution of adult brine flies was examined
from two approaches:
1) Are distributions influenced by distance from a watercourse?
2) Does watercourse type (recirculation ditches or natural channels) influence
distribution patterns?
Three watercourses were chosen as study sites, a 2-yr-old ditch, a 5-yr-old ditch,
and a natural channel. At each site, three parallel strata at distances of 1 m (stratum
1), 3 m (stratum 2), and 10 m (stratum 3) from the watercourse were designated
(Fig. 1). Stratum 3 served as a control for intra- and inter-site comparisons since
a variety of physical factors exhibited gradients that occur only within 3 m of
either watercourse type. For example, standpipes placed perpendicular to both
ditches and natural channels indicated that tidal depression and recharge of the
water table are limited to a 3-m distance. Also, groundwater salinity and soil
surface salinity increased with distance from the channel, both becoming asymp¬
totic before 10-m distance (Balling and Resh, 1982).
Collecting methods. — Adult brine flies were sampled at approximately biweekly
intervals during late morning hours from November 1977 through November
1978 using a gasoline-powered suction device, the D-vac (Dietrick et al., 1959),
which collected insects into a 0.25-mm nylon mesh bag. All 9 strata (3 sites, each
with 3 strata) were sampled over a distance of 30 m by placing the suction cone
onto the substrate for 5 sec at approximately 1-m intervals. Samples were then
placed in Berlese-Tullgren funnels for 24 hr with the brine flies being collected
directly into 70% ethanol.
Seasonal occurrence of brine fly adults was also determined from ethylene glycol
pan-traps that were designed to simulate small ponds or potholes in the marsh.
This artificial habitat consisted of 16 aluminum roasting pans (0.3 m x 0.6 m)
arranged in a rectangular 4x4 grid (Fig. 2). Samples were taken at weekly in¬
tervals from March 1980 (after the last heavy rain of the wet season) to December
1980 (prior to the first heavy rain of the subsequent wet season). Since periodic
examination of all 16 pans indicated no inter-pan difference in species or their
abundances, only flies from the four interior pans were identified and enumerated.
Data analysis. — Due to fluctuations in brine fly abundance throughout the
study, it was necessary to remove the seasonal variation in the D-vac samples to
equally weight all sampling dates. This was done by converting the number of
each species in a sample to a percentage of that species total collected at all sites
and distances on the same date. An arc-sine square-root transformation was used
to normalize the percentages for parametric statistical analysis. Comparisons be¬
tween distances and between sites for each species were made with one-way
ANOVA and Student-Neuman-Keuls multiple range tests (SNK).
Results
The fauna. — During this study 28 species of brine fly adults were collected in
Petaluma Marsh with 22 species recorded from D-vac and 23 species from pan-
VOLUME 60, NUMBER 1
39
Figure 1. Location of study sites (2-yr-old ditch, 5-yr-old ditch, and natural channel) and sampling
strata (1 m, 3 m, 10 m) in the Petaluma Marsh.
trap samples (Fig. 3). Of these, 11 species were not included in Simpson’s (1976)
list of marine species [Limnellia sejuncta (Loew), Napaea socia Cresson, Pelina
truncatula Loew, Hydrina nigrescens Cresson, Ilythea caniceps Cresson, Ilythea
spilota (Curtis), Nostima picta (Fallen), Psilopa ( Psilopa) olga Cresson, Psilopa
( Psilopa ) compta (Meigen), P. coquilletti, Lytogaster gravida (Loew)]. Although
several of these new records occur in low numbers and possibly reflect their
transient status in the marsh, the repeated occurrence and abundance of some
species ( H. nigrescens, N. picta, P. truncatula, P. compta, and P. coquilletti ) in
1977 and 1980 suggests the presence of established populations.
40
PAN-PACIFIC ENTOMOLOGIST
Figure 2. Ethylene glycol pan-traps used to collect adults associated with potholes in Petaluma
Marsh.
Total numbers of adult brine flies collected with both sampling methods was
43,271. D-vac samples contained 11,343 individuals with five species comprising
>98% of the total: S. stagnalis (11.5%), S. setosa (25.7%), Scatophila sp. (5.2%),
P. coquilletti (17.9%), and A. litoralis (38.1%). Pan-traps produced 31,928 indi¬
viduals with 95.5% (30,505) represented by one species, E. millbrae. Of the
remaining 22 species sampled by pan-traps, 6 species comprised 4%: S. stagnalis
(0.5%), Scatella ( Scatella ) paludum (Meigen) (0.7%), Lamproscatella muria Math¬
is (1.2%), P. coquilletti (0.5%), A. litoralis (0.7%), and Polytrichophora sp. (0.4%).
Seasonal occurrence and abundance.—Scatella stagnalis adults were common
from March to mid-June in D-vac samples, and to a lesser extent in pan-trap
samples (Fig. 3). Other workers have collected this species from a variety of
Holarctic locations throughout the warm periods of the year and in a variety of
freshwater and maritime habitats, usually along mud and sand shores (Dahl, 1959;
Williams, 1938; Deonier, 1965, 1972; Eastin and Foote, 1971; Scheiring and
Foote, 1973; Simpson, 1976; Miyagi, 1977; Zack, 1979). Foote (1979:65) con¬
sidered S. stagnalis to be a “fugitive, nonequilibrium species” common in unstable
habitats that are frequently flooded. Both larvae and adults are polyphagous,
feeding on algae, diatoms, and decaying material (Simpson, 1976; Scheiring and
Foote, 1973; Foote, 1977, 1979), although certain food items may be preferred
since development time appears to be sensitive to diet (Zack and Foote, 1978).
Scatella paludum adults were abundant from mid-March to early-June in pan¬
trap samples. D-vac samples contained fewer adults, but they were found during
approximately the same season (Fig. 3). Dahl (1959) collected adults from beds
VOLUME 60, NUMBER 1
41
D-VAC
PAN TRAP
1978
F
I
1980
A A
1 L.
Scatella stagnalis
Scatella paludum
Scatella setosa
Lamproscatella lattini
Lamproscatella muria
Limnellia sejuncta
Sea tophi la sp.
Ephydra mi librae
Ephydra cinerea
Paracoenia sp.
Napaea socia
Pelina truncatula
Brachydeutera sturtevanti
Hydrina nigrescens
llythea caniceps
llythea spilota
Nostima picta
Notiphila decoris
Notiphila aenigma
Hydrellia griseola
Psilopa olga
Psilopa compta
Psilopa coquilletti
Atissa litoral is
Polytrichophora sp.
Lytogaster gravida
Mosillus tibialis
Clanoneurum americana
232
128
4
3
Figure 3. Abundance (as log 10 ) of brine fly adults from 1977-1978 D-vac samples and 1980 pan¬
trap samples.
of decaying seaweed and from freshwater and maritime sand beaches in Scan¬
dinavia from May to October. Zack (1979) near Mt. Rainier, Washington, re¬
corded adults from sandy beaches along the Tahoma River from June through
August. Similar to their close relative S. stagnalis, they most likely feed on a
variety of algal species (Foote, 1979).
Scatella setosa adults were found throughout the year in D-vac samples with
their peak abundance occurring from spring through mid-summer (Fig. 3). Adults
were sampled in lower numbers in the pan-traps, primarily during the early spring.
Cameron (pers. comm.) found this species in the fall and winter months in a
nearby San Francisco Bay salt marsh. The habits of both larvae and adults are
reported as similar to S. stagnalis (Wirth and Stone, 1956).
Lamproscatella muria adults were abundant in pan-trap samples beginning in
late-March and extending into autumn; however, only one specimen was recorded
from the D-vac samples (Fig. 3). This species is known to occur primarily in
saline habitats along both eastern and western North American coasts, although
it has been found occasionally in freshwater environments (Sturtevant and Whee¬
ler, 1954) and in alkaline ponds (Mathis, 1979). Also, Scheiring and Foote (1973)
have reported adults from saline waters in Ohio during late summer.
Scatophila sp., possibly a complex containing more than a single species (W.
Mathis, pers. comm.), was found from early spring to autumn in D-vac samples
42
PAN-PACIFIC ENTOMOLOGIST
(Fig. 3) although only one specimen was found in pan-trap samples. Scatophila
adults have generally been found to occur from July to September in a variety of
freshwater and maritime habitats including mud and sand shores, glacial river
beds, manure heaps, salt and freshwater meadows, seeps, spoil banks, and in
greenhouses (Bolwig, 1940; Dahl, 1959; Deonier, 1965, 1974; Scheiring and Foote,
1973; Harris and Deonier, 1979; Zack, 1979). Their diet consists of blue-green
algae, diatoms, and gram-positive bacteria (Bolwig, 1940; Deonier, 1974).
Ephydra millbrae, collected only rarely in the D-vac samples, was by far the
most abundant species collected in the pan-traps (Fig. 3). All life stages commonly
occur throughout the year, with abundant numbers observed each year since 1977
in Petaluma Marsh. Adults of E. millbrae are known to frequent calm, saline
pools where they feed, mate, and oviposit into the larval habitat. Jones (1906)
first described this species from salt marshes along South San Francisco Bay,
although his illustration of the egg resembles that of the hemipteran waterboatman
Trichocorixa reticulata (Guerin-Meneville), a common coexisting species. De¬
pending on the availability of suitable habitat, Ephydra adults have been collected
year round (e.g., Ping, 1921; Dahl, 1959; Kelts, 1979) and Foote (1979) considered
species in this genus to be oligophagous in algal food selection.
Psilopa coquilletti adults occurred from early spring to autumn in both D-vac
and pan-trap samples (Fig. 3). The habits and seasonality of P. coquilletti have
not been previously described, although closely related species are known to be
scavengers (Simpson, 1976) and some freshwater Psilopa are leafminers (Hennig,
1943).
Atissa litoralis adults were found throughout most of the year (Fig. 3). A. litoralis
has been found along the eastern U.S. seashore (Sturtevant and Wheeler, 1954)
and in the midwestern U.S. where they occur in both freshwater and saline habitats
(Deonier, 1965; Scheiring and Foote, 1973; Steinly, 1978). Cresson (1942:110)
considered it a “scarce western species”, but in Petaluma Marsh it was the second
most abundant ephydrid collected. Its larval biology is unknown.
Polytrichophora sp. adults occurred from spring through the summer in the
pan-trap samples. In D-vac samples, adults occurred sporadically and in low
numbers during the spring and summer (Fig. 3). Adults of Polytrichophora sp.
have been collected along mud and sand shores in Washington (Zack, 1979), Ohio
(Scheiring and Foote, 1973), and Iowa (Deonier, 1965). Diatoms and other algae
are utilized as food items by species in this genus (Deonier, 1972).
Seasonal abundance of the other 19 ephydrid species in Petaluma Marsh are
presented in Figure 3.
Distribution of species in relation to watercourses. — Five of 23 species collected
in D-vac samples were sufficiently abundant to statistically analyze their distri¬
bution patterns in relation to recirculation ditches (2- and 5-yr-old ditches) and
natural channels (Table 1).
Psilopa coquilletti adults showed no consistent trends within sites. A significantly
higher number of adults occurred away (i.e., at the 10-m distance) from the 2-yr-
old ditch (P = 0.01), yet the natural channel displayed a different trend (P = 0.03),
with significantly fewer adults at the 10-m cf. the 3-m distance (P = 0.03). Com¬
paring sites, the 1-m and 3-m distances at the natural channel had significantly
higher numbers of adults than either ditch (P = 0.01), whereas for the 10-m
distance densities at the 2-yr-old ditch were significantly higher than at either the
VOLUME 60, NUMBER 1
43
Table 1. Distribution of adult brine flies in relation to man-made (2- and 5-yr-old ditches) and
natural (channel, NC) watercourses; cell values are mean percents of adults/sample and * and | refer
to horizontal rows and vertical columns, respectively. Within any row (e.g., NC), two cells marked
with * are significantly different from one another, as determined from SNK multiple range tests. If
only one cell is marked with *, that cell is significantly different from the other two cells. Vertical
columns are marked in an identical manner, except for the use of f.
P. coquilletti
S. setosa
1 m
3 m
10 m P
1 m
3 m
10 m
p
2-yr
13.3
14.6
23.3|* 0.01
2-yr
11.7
5.0
16.9
0.10
5-yr
10.7
11.9
12.2 0.88
5-yr
8.3
5.9
11.7
0.62
NC
20.7f
26.5|*
14.4* 0.03
NC
36.8f*
1.8
14.9
0.01
P
0.01
0.01
0.01
P
0.01
0.44
0.76
Scatophila sp.
5. stagnalis
1 m
3 m
10 m P
1 m
3 m
10 m
p
2-yr
*
-i-
r-
o
5.5
11.8* 0.04
2-yr
17.2
8.6
16.6
0.57
5-yr
25.9|
12.4
21.1 0.39
5-yr
21.0
9.6
13.2
0.18
NC
19.6
10.0
6.4 0.20
NC
31.8*
7.2*
13.4
0.04
P
0.04
0.45
0.10
P
0.36
0.93
0.86
A.
Utoralis
1 m
3 m
10 m P
2-yr
17.5
10.6
14.6 0.33
5-yr
13.3
6.0*
22.5* 0.02
NC
14.1
11.8
21.8 0.11
P
0.69
0.24
0.36
5-yr-old ditch or the natural channel (P = 0.01). Thus, even though the densities
at the natural channel appear to be significantly different from those at the 1 -m
and 3-m distances of either ditch, the significant differences between sites at the
10-m distance (i.e., the control—see Experimental design) indicate that factors
other than those associated with presence of watercourses may be affecting the
distribution of this species.
Scatophila sp. distribution similarly displayed no consistent trends within sites.
The 2-yr-old ditch had significantly fewer adults at the 1-m distance cf. the 10-m
distance (P = 0.04), but different patterns were evident at the other two sites.
Comparing sites, the only significant difference occurred between the 2-yr-old
ditch and the 5-yr-old ditch at the 1-m distance (P = 0.04). The 5-yr-old ditch
and the natural channel had similar abundances at both the 1 -m and 3-m distances.
Scatella setosa adults were more abundant at the 1-m distance than at the
3- and 10-m distances from the natural channel (P = 0.01), a pattern not repeated
at either ditch site. Likewise, a significant difference existed between the 1-m
distance strata at the natural channel and either ditch site (P = 0.01). Therefore,
S. setosa distribution patterns at ditches did not appear to be similar to those at
natural channels.
Scatella stagnalis adults at the natural channel were significantly more abundant
at 1-m distance cf. the 3-m distance (P = 0.04). While similar distribution patterns
occurred at both ditched sites, neither was statistically significant. Also, compar¬
isons between sites revealed no significant differences. From this we conclude that
44
PAN-PACIFIC ENTOMOLOGIST
ditches and the natural channel were not dissimilar with respect to the distribution
of this species.
Atissa litoralis adults occurred in significantly greater numbers at the 10-m
distance cf. the 3-m distance at the 5-yr-old ditch (P = 0.02). Although the same
pattern was present, no statistically significant trends were detected at either the
2-yr-old ditch or the natural channel. No significant differences were found be¬
tween sites at any distance. Thus, the ditches and the natural channel did not
appear to affect distribution patterns of A. litoralis dissimilarly.
Discussion
Of the five species which accounted for >98% of the total specimens collected
with the D-vac in 1977-1978, all were found to occur in the same season in the
1980 pan-trap samples. Scatophila sp., L. muria, and E. millbrae were sampled
predominantly by one method ( Scatophila sp. from the D-vac, the other two from
the pan-traps). Two possible reasons may account for this: 1) a difference in actual
population abundance between years, i.e., Scatophila sp. populations may have
declined drastically sometime between 1977-1978 and 1980 and L. muria and
E. millbrae densities increased; or 2) the two methods (D-vac/Berlese-Tullgren
funnels cf. pan-traps) sampled selectively. For E. millbrae and L. muria, this
second explanation is far more likely since the larvae are aquatic and the adults
occur almost exclusively on pond rather than ditch or natural channel surfaces.
Also, both species have been observed on potholes and ponds in rather large
numbers since 1977. In the case of Scatophila sp., insufficient information is
available to determine why only one individual was sampled in 1980 pan-traps.
The distribution of salt marsh arthropods has been observed to vary with
distances from natural channels, marsh ponds, and other water bodies (e.g., Hull
et al., 1934; Williams, 1938; Rockel, 1969; Evans et al., 1971; Garcia and Schlin-
ger, 1972; Freeman and Hansens, 1972; Foster and Treherne, 1975, 1976a, 1976b;
Magnarelli and Anderson, 1978) and results of this study clearly indicate that
population distribution patterns of some species can be related to distance from
man-made ditches and/or natural channels. Whether the addition of recirculation
ditches produces the same results as increasing the number of natural channels
also depends on the species involved. At man-made and natural watercourses
different distribution patterns were found for two species, S. setosa and P. co-
quilletti. For P. coquilletti, factors other than those associated with presence of a
watercourse are apparently affecting its distribution, since abundances at the con¬
trol distance (10-m) of the sites were significantly different. Of the remaining three
species, no significant differences were noted between the natural and man-made
watercourses at any distance.
Differences in natural and man-made watercourses may have varied and com¬
plex effects on the adult brine fly populations. For example, greater woody (i.e.,
perennial) biomass of Salicornia in Petaluma Marsh occurs at 1 -m distance from
the natural channels than at the same distance from ditches (Balling and Resh,
1983). This increase in woody material may afford adult brine flies better pro¬
tection against inundating tides, predators, and extreme temperature fluctua¬
tions. These factors may be more important to some species than others and thus
produce the different distribution patterns observed for the Petaluma Marsh brine
flies.
VOLUME 60, NUMBER 1
45
Acknowledgments
We thank S. S. Balling and J. N. Collins for field and laboratory assistance and
the Marin/Sonoma Mosquito Abatement District for their continued support and
cooperation. W. N. Mathis confirmed brine fly identifications. Support for this
project was provided by University of California Mosquito Research Funds.
Literature Cited
Aldrich, J. M. 1912. The biology of some western species of the dipterous genus Ephydra. J. N.Y.
Entomol. Soc., 20:77-99.
Balling, S. S., and V. H. Resh. 1982. Arthropod community responses to mosquito control recir¬
culation ditches in San Francisco Bay salt marshes. Environ. Entomol., 11:801-808.
-, and-. 1983. The influence of mosquito control recirculation ditches on plant biomass,
production, and composition in two San Francisco Bay salt marshes. Estuarine Coastal Shelf
Sci., 16:151-160.
Bolwig, N. 1940. The description of Scatophila unicornis Czerny, 1900 (Ephydridae, Diptera). Proc.
Roy. Entomol. Soc. London (B), 9:129-137.
Crawford, D. L. 1912. The petroleum fly in California Psilopapetrolei Coq. Pomona Coll. J. Entomol.,
4:687-697.
Cresson, E. T., Jr. 1942. Synopses of North American Ephydridae. I. The subfamily Psilopinae, with
descriptions of new species. Trans. Amer. Entomol. Soc., 68:101-128.
Dahl, R. G. 1959. Studies on Scandinavian Ephydridae. Opusc. Entomol., Suppl. 15:224 pp.
Deonier, D. L. 1965. Ecological observations on Iowa shore flies. Proc. Iowa Acad. Sci., 71:496-
510.
-. 1972. Observations on mating, oviposition, and food habits of certain shore flies. Ohio J.
Sci., 72:22-29.
-. 1974. Biology and descriptions of immature stages of the shore fly Scatophila iowana. Iowa
State J. Res., 49:17-22.
Dietrick, E. J., E. I. Schlinger, and R. van den Bosch. 1959. A new method for sampling arthropods
using a suction collecting machine and modified Berlese funnel. J. Econ. Entomol., 52:1085-
1091.
Eastin, W. C., and B. A. Foote. 1971. Biology and immature stages of Dichaeta caudata. Ann.
Entomol. Soc. Amer., 64:271-279.
Evans, P. D., C. N. E. Ruscoe, and J. E. Treherne. 1971. Observations on the biology and submergence
behavior of some littoral beetles. J. Mar. Biol. Ass. U.K., 51:375-386.
Foote, B. A. 1977. Utilization of blue-green algae by larvae of shore flies. Environ. Entomol., 6:
812-814.
-. 1979. Utilization of algae by larvae of shore flies. Pp. 61-72 in D. L. Deonier (ed.), First
symposium on the systematics and ecology of Ephydridae (Diptera). North American Ben-
thological Society, Erie, Pennsylvania, 147 pp.
Foster, W. A., and J. E. Treherne. 1975. The distribution of an intertidal aphid, Pemphigus trehernei
Foster, on marine saltmarshes. Oecologia, 21:151-155.
-, and-. 1976a. The effects of tidal submergence on an intertidal aphid, Pemphigus
trehernei Foster. J. Anim. Ecol., 45:291-301.
-, and-. 1976b. Insects of marine salt-marshes: Problems and adaptations. Pp. 5-42 in
L. Cheng (ed.). Marine insects. North-Holland Publishing Co., Amsterdam, 581 pp.
Freeman, J. V., and E. J. Flansens. 1972. Collecting larvae of the salt marsh greenhead Tabanus
nigrovittatus and related species in New Jersey: Comparison of methods. Environ. Entomol.,
2:653-658.
Garcia, R., and E. I. Schlinger. 1972. Studies of spider predation on Aedes dorsalis (Meigen) in a
salt marsh. Proc. Calif. Mosq. Contr. Ass., 40: 117-118.
Harris, S. C., and D. L. Deonier. 1979. Ecological observations of immature Ephydridae in North
Dakota springbrooks. Pp. 111-122 in D. L. Deonier (ed.), First symposium on systematics and
ecology of Ephydridae (Diptera). North American Benthological Society, Erie, Pennsylvania,
147 pp.
Hennig, W. 1943. Ubersicht fiber bisher bekannten Metamorphosestadien des Ephydriden, mit
46
PAN-PACIFIC ENTOMOLOGIST
Neobeschreibungen nach dem Material der Deutschen Limnologischen Sundaexpedition (Dip-
tera: Ephydridae). Arb. Morphol. Taxonom. Entomol., 10:105-138.
Hull, J. B., W. E. Dove, and F. M. Prince. 1934. Seasonal incidence and concentrations of sand fly
larvae, Culicoides dovei Hall, in salt marshes (Ceratopogonidae: Diptera). J. Parasitol., 20:162-
172.
Jones, B. J. 1906. Catalogue of Ephydridae, with bibliography and descriptions of new species. Univ.
Calif. Publ. Entomol., 1:153-198.
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37-57.
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Tabanidae (Diptera). J. Med. Entomol., 14:573-578.
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zoogeography of Nearctic Lamproscatella Hendel. Smith. Contrib. Zool., 295:1-41.
Miyagi, I. 1977. Fauna Japonica. Ephydridae (Insecta: Diptera). Keigaku Publishing Co., Tokyo,
113 pp., 49 plates.
Ping, C. 1921. The biology of Ephydra subopaca Loew. Mem. Cornell Univ. Agr. Exp. Sta., 49:557-
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San Francisco Bay marshlands: Preliminary considerations and experimental design. Proc. Calif.
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Proc. NJ. Mosq. Exterm. Ass., 56:102-116.
Scheiring, J. F., and B. A. Foote. 1973. Habitat distribution of the shore flies of Northeastern Ohio.
Ohio J. Sci., 73:152-155.
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Steinly, B. A. 1978. New records of Atissa in Ohio. Entomol. News, 89:107-108.
Sturtevant, A. H., and M. R. Wheeler. 1954. Synopsis of Nearctic Ephydridae (Diptera). Trans.
Amer. Entomol. Soc., 79:151-257.
Williams, F. X. 1938. Biological studies in Hawaiian water-loving insects. Part III. Diptera. A:
Ephydridae and Anthomyiidae. Proc. Hawaii. Entomol. Soc., 10:85-119.
Wirth, W. W., and W. N. Mathis. 1979. A review of Ephydridae living in thermal springs. Pp. 21-
46 in D. L. Deonier (ed.), First symposium on systematics and ecology of Ephydridae (Diptera).
North American Benthological Society, Erie, Pennsylvania, 147 pp.
-, and A. Stone. 1956. Aquatic Diptera. Pp. 372-482 in R. L. Usinger (ed.), Aquatic insects
of California. Univ. Calif. Press, Berkeley, 508 pp.
Zack, R. S. 1979. Habitat distribution of Ephydridae (Diptera) of Mount Rainier National Park
(Washington State). Pp. 81-98 in D. L. Deonier (ed.), First symposium on systematics and
ecology of Ephydridae (Diptera). North American Benthological Society, Erie, Pennsylvania,
147 pp.
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Environ. Entomol., 7:509-511.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 47-52
Published 27 January 1984
Two New Conifer-Inhabiting Phytocoris from Western
North America (Hemiptera: Miridae)
Gary M. Stonedahl
Systematic Entomology Laboratory, Department of Entomology, Oregon State
University, Corvallis, Oregon 97331.
Abstract. — The new plant bug species Phytocoris nobilis and P. occidentalis are
described from western North America. Phytocoris nobilis was collected on Abies
spp. and P. occidentalis was taken on several species of Pinus and also collected
at light. The adult male of P. nobilis and male genitalia of both species are
illustrated.
Relatively little is known about the plant bug fauna of the northwestern United
States. A recent survey of the Phytocoris species that occur in this region has
revealed the presence of several undescribed species. Phytocoris nobilis and P.
occidentalis are described to provide names for a forthcoming study of the Miridae
of Oregon. Phytocoris nobilis was taken at several localities in the Coast and
Cascade Ranges of Oregon on Abies spp. Phytocoris occidentalis occurs throughout
much of coastal western North America on pines and also has been taken at light.
The following abbreviations are used for specimen depositories cited in the
locality data: American Museum of Natural History (AMNH); California Acad¬
emy of Sciences (CAS); California State Department of Food and Agriculture
(F&A); Los Angeles County Museum (LACM); Oregon State University (OSU);
San Diego Natural History Museum (SDNH); University of British Columbia
(UBC); University of California, Berkeley (UCB); University of California, Riv¬
erside (UCR); National Museum of Natural History, Smithsonian Institution
(USNM).
Phytocoris nobilis, New Species
(Figs. 1, 2-4)
Male (Fig. 1).—Length 7.68-8.85 mm, width 2.50-2.74. Head: width across
eyes 1.20-1.27, vertex 0.36-0.39; dark reddish brown to nearly black; frons, gula,
and ventral margin of lorum marked with white or pale yellow; base of jugum
with conspicuous pale spot; vertex and frons with scattered, erect, black bristles.
Rostrum: length 3.36-3.69, extending to 6th or 7th abdominal segment. Antennae:
dark brown or black; with closely appressed, recumbent, light setae; I, length 1.50-
1.71, with scattered, pale, often indistinct spots, and stout erect black bristles; II,
length 3.46-3.93, narrowly pale at base; III, length 1.86-2.18; IV, length 1.14-
1.27. Pronotum: mesal length 1.12-1.18, posterior width 1.98-2.05, white or pale
yellow, lateral margins broadly fuscous. Scutellum: moderately convex, white or
pale yellow, pale midline bordered by broad fuscous band. Hemelytra: opaque
white, densely marked with small to large fuscous patches, particularly along
48
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Phytocoris nobilis, n. sp., adult male.
claval, cubital and radial veins; apices of clavus and cuneus, posterolateral angle
of corium, and posterior margin of paracuneus fuscous; membrane conspurcate,
veins dark, becoming pale posteriorly. Venter: dark brown or black, shining, with
scattered pale setae. Legs: coxae white or pale yellow; femora fuscous, apical half
VOLUME 60, NUMBER 1
49
Figures 2-4. Phytocoris nobilis. 2. Left paramere. 3. Right paramere. 4. Sclerotized process of
vesica. 5-7. Phytocoris occidentalis. 5. Left paramere. 6. Right paramere. 7. Sclerotized process of
vesica.
50
PAN-PACIFIC ENTOMOLOGIST
marked with scattered pale spots, metafemora with preapical, transverse, pale
band; tibiae fuscous, triannulate with white; tarsi fuscous; tarsal claws yellowish
brown. Vestiture: pronotal disk, scutellum, and hemelytra with semierect, black
setae intermixed with recumbent, silvery setae. Genitalia: Figures 2-4. Genital
segment with large, broad tubercle at base of left paramere, and similar but smaller
tubercle at base of right paramere. The male description is based on the exami¬
nation of nine specimens.
Female. — Similar to male in color and vestiture; hemelytra with fewer fuscous
patches, often nearly entirely white. Length 6.78-7.26 mm, width 2.32-2.60.
Head: width across eyes 1.12-1.20, vertex 0.44-0.47. Rostrum: length 3.63-3.82,
extending to base of ovipositor. Antennae: I, 1.46-1.63; II, 3.19-3.57; III, 1.67-
2.05; IV, 0.95-1.18. Pronotum: mesal length 1.01-1.12, posterior width 1.79-
1.98. The female description is based on the examination of 12 specimens.
Type data. — HOLOTYPE male, Oregon, Lane Co., H. J. Andrews Experimental
Forest, T15S, R6E, Sec. 29, NW l A, 1450 meters, July 31, 1979 (G. M. Cooper;
USNM). PARATYPES: OREGON. Benton Co.: 2 males, Marys Peak, T12S,
R7W, Sec. 21, SW l A, 1097 meters, Aug. 6, 1975 (J. D. Lattin; OSU); 1 male and
3 females, Marys Peak, T12S, R7W, Sec. 20, SW l A, 1070 meters, July 25-Aug.
17, 1979 (G. M. Cooper; OSU); 1 male and 2 females, Marys Peak, T12S, R7W,
Sec. 21, SW 1 A , 1097 meters, Aug. 3, 1979 (G. M. Cooper; OSU). Lane Co.: 1
female, same data as holotype (G. M. Stonedahl; USNM); 2 males and 5 females,
H. J. Andrews Experimental Forest, T15S, R6E, Sec. 29, SW X A and SE l A, 1480
meters, July 20 & 31, 1979 (G. M. Cooper, G. M. Stonedahl; AMNH, OSU);
1 male and 1 female, same data as above but July 13, 1980 (G. M. Cooper; CAS);
1 male, H. J. Andrews Experimental Forest, T15S, R6E, Sec. 7, SW X A, 1329
meters, July 31, 1979 (G. M. Stonedahl; OSU).
Specimens were collected on noble fir, Abies procera Rehd. and silver fir, Abies
amabilis (Dougl.) Forbes.
Remarks.— This species belongs to the junceus species group formed by Knight
(1974) and discussed by Kelton (1979) and Henry (1979). Phytocoris nobilis keys
to P. rainieri Knight in Knight (1974:126) based on the predominantly white
pronotal disk and hemelytra, and the relatively short first antennal segment. The
two species can be separated on the basis of color and vestiture of the dorsum,
legs, and antennae. The dorsum of P. nobilis has a lighter, grayish white cast
compared to the reddish brown cast of P. rainieri. The hind femora of P. nobilis
are fuscous with scattered pale spots that are largely restricted to the apical half
of the femur. In P. rainieri, the hind femora are pale yellow with a reticulate,
reddish brown pattern that extends to the base of the femur. The tibiae of P.
nobilis are clearly triannulate with white, but banding is indistinct or lacking in
P. rainieri. The femora and first antennal segments of P. rainieri are more densely
set with erect, black, bristle-like setae than are those of P. nobilis. Male genitalia
were not compared since the male of P. rainieri is unknown.
Phytocoris occidentalis, New Species
(Figs. 5-7)
Male. — Length 4.43-5.83 mm, width 1.53-1.71. Head: width across eyes 0.90-
0.94, vertex 0.29-0.32; pale yellow to testaceous; buccula, lorum, jugum, and
clypeus marked with red or reddish brown; frons with 6-8 fuscous striae each
VOLUME 60, NUMBER 1
51
side of midline. Rostrum: length 1.89-2.30, extending to 5th or 6th abdominal
segment. Antennae: dark reddish brown to fuscous; I, length 0.68-0.79, with
recumbent, black setae and 10-12 erect, pale bristles; II, length 1.78-2.03, nar¬
rowly pale at base; III, length 1.03-1.30, narrowly pale at base; IV, length 0.68-
0.90. Pronotum: mesal length 0.71-0.81, posterior width 1.28-1.42; testaceous
or grayish yellow, collar and calli marked with red; posterior submargin with 4-
6 slightly elevated tubercles outlined posteriorly by a wavy, fuscous band. Scu-
tellum: testaceous or grayish yellow; frequently marked or tinged with red, es¬
pecially along midline. Hemelytra: testaceous or grayish yellow with distinct red¬
dish cast, areas bordering claval suture darker brown; cuneus deeper red; membrane
translucent, confusedly sprinkled with fuscous spots which often coalesce to form
larger fuscous patches, veins pale, usually tinged with red and sometimes infus-
cated basally. Venter: testaceous or grayish yellow, extensively mottled with red
or reddish brown; base of 9th abdominal segment broadly fuscous. Legs: coxae
pale yellow with scattered, reddish patches; femora pale yellow, reticulated with
red or reddish brown so that pale areas appear as spots, metafemora more exten¬
sively infuscated apically; tibiae reddish brown to fuscous with scattered, pale
spots, pale markings on front tibiae often appearing as three distinct bands; tarsi
testaceous to fuscous; tarsal claws testaceous. Vestiture: pronotal disk, scutellum,
and hemelytra with semierect, black setae intermixed with brown or black, scale¬
like setae and white, scale-like setae. Genitalia: Figures 5-7. Genital segment with
a finger-shaped tubercle at bases of left and right parameres. The male description
is based on the examination of 46 specimens.
Female.—Very similar to male in color, vestiture, and size. Length 4.37-5.29
mm, width 1.51-1.71. Head: width across eyes 0.80-0.95, vertex 0.33-0.41. Ros¬
trum: length 1.98-2.47, extending to 5th or 6th abdominal segment. Antennae:
I, 0.68-0.80; II, 1.76-2.11; III, 1.12-1.30; IV, 0.76-0.94. Pronotum: mesal length
0.70-0.86, posterior width 1.19-1.49. The female description is based on the
examination of 29 specimens.
Type data. — HOLOTYPE male, Oregon, Josephine Co., 4 mi NE Cave Junction
on US 199, Aug. 8, 1979, ex Pinus ponderosa Dougl. (G. M. Stonedahl; USNM).
PARATYPES: OREGON. 3 males and 1 female, same data as holotype (OSU);
9 males and 3 females, same data as holotype but Aug. 24, 1979 (AMNH, CAS,
OSU).
Additional specimens. — CANADA. BRITISH COLUMBIA. Goldstream, Aug.
4, 1927 and Aug. 16, 1929 (W. Downes; UBC). UNITED STATES. CALIFOR¬
NIA. Alameda Co.: Berkeley, Aug. 28 and Oct. 1, 1965, at light (R. L. Langston;
UCB). Butte Co.: Oroville, July 13, 1926 (H. H. Keifer; CAS). Contra Costa Co.:
Moraga, July 24 and Sept. 23, 1976 (D. G. Denning; AMNH); San Pablo Dam
Res., June 29, 1978, ex Pinus radiata D. Don (C. P. Ohmart; OSU); Walnut Crk.,
Aug. 20, 1969, at blacklight (J. A. Powell; UCB). Mariposa Co.: Feliciana Mtn.,
July 26, 1946, ex Pinus attenuata Lemmon (R. L. Usinger; UCB). Plumas Co.:
Johnsville, Sept. 2, 1967, light trap (H. Pini; F&A). Sacramento Co.: Sacramento,
June 15-26, 1968 (R. A. Belmont; F&A). San Bernardino Co.: San Bernadino
Mts., Mill Crk., 1829 meters, Sept. 2, 1950, at light (Timberlake; UCR); San
Bernardino Mts., South Fork Camp, 1890 meters, Sept. 10, 1944, ex Pinus pon¬
derosa (Timberlake; UCR); San Bernardino Mts., Camp O-Ongo nr. Running
Springs, 1920 meters, Aug. 31, 1971 (C. L. Hogue; LACM); 4 mi S Twentynine
52
PAN-PACIFIC ENTOMOLOGIST
Palms, March 30, 1970, ex Yucca schidigera Roezl (UCB). San Diego Co.: Laguna,
Aug. 21, 1927 (SDNH). Shasta Co.: Lakehead, 350 meters, June 26, 1981, ex
Pinus sabiniana Dougl. (J. D. Lattin; OSU). Siskiyou Co.: McCloud, 1073 meters,
Aug. 9 & 20, 1980, ex Pinus attenuata (G. M. Stonedahl; OSU). Yolo Co.: 5 mi
N Rumsey, May 2, 1973 (J. C. Miller; OSU). OREGON. Grant Co.: Malheur
Nat. For., Dixie Campground, Aug. 24, 1981, ex Pinus ponderosa (G. M. Stone¬
dahl; OSU). Hood River Co.: Mt. Hood, Cloud Cap, 1829 meters. Sept. 5, 1979,
ex Pinus contorta Dougl. (G. M. Stonedahl; OSU); T1S, R10E, Sec. 31, Sept. 13,
1979, ex Pinus contorta (G. M. Stonedahl; OSU); T3S, R9E, Sec. 18, 1311 meters,
Sept. 13, 1979, ex Pinus contorta (J. D. Lattin; OSU). Josephine Co.: O’Brien,
Aug. 2, 1968 (P. Oman; OSU); 2 mi W O’Brien, Aug. 10, 1979, ex Pinus ponderosa
(G. M. Stonedahl; OSU). Linn Co.: 1 mi W jet. State Hwy. 22 on US 20, Sept.
19, 1979, ex Pinus contorta (G. M. Stonedahl; OSU); 0.5 mi N jet. US 20 on
State Hwy. 22, 1158 meters, Sept. 1, 1977, ex Pinus contorta (J. D. Lattin; OSU).
Remarks.—Phytocoris occidentalis will key to P. miniatus Knight in Knight
(1968:219) but is easily separated from miniatus by the submarginal, fuscous band
on the pronotal disk and the black, scale-like setae on the dorsum. Phytocoris
occidentalis is much larger than P. miniatus which has an average size of only
3.4 mm. The geographic ranges of these two species also are different. Phytocoris
miniatus is known only from Arizona and Utah.
Phytocoris occidentalis is superficially similar to another pine-inhabiting Phy¬
tocoris species, P. stellatus Van Duzee. Phytocoris stellatus is easily separated
from occidentalis by the absence of black, scale-like setae on the dorsum and the
structure of the male genitalia, particularly the absence of the genital tubercles.
Acknowledgments
I wish to thank John D. Lattin (OSU), Paul W. Oman (OSU), and Thomas J.
Henry (USDA, SEL, % USNM) for reviewing the manuscript; and special thanks
to Richard C. Froeschner (USNM) for loaning type specimens of P. abiesi Knight
and P. rainieri Knight for comparison. I also thank Bonnie B. Hall for the fine
illustration of P. nobilis and Julie A. Stonedahl for help in preparing the manu¬
script. The Theodore Roosevelt Memorial Fund (AMNH) and the Oregon State
University Foundation provided funds to study additional type material at the
USNM.
Literature Cited
Henry, T. J. 1979. Descriptions and notes on five new species of Miridae from North America
(Hemiptera). Melsheimer Entomol. Ser., 27:1-10.
Kelton, L. A. 1979. Two new species of Phytocoris from western Canada (Heteroptera:Miridae).
Can. Entomol., 111:689-692.
Knight, H. H. 1968. Taxonomic review: Miridae of the Nevada Test Site and the western United
States. Brigham Young Univ. Sci. Bull., Biol. Ser., 9(3): 1-282.
-. 1974. A key to the species of Phytocoris Fallen belonging to the Phytocoris junceus Kngt.
group of species (Hemiptera, Miridae). Iowa State. J. Res., 49:123-135.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 53-54
Published 27 January 1984
Description of the First Central American Species of
Stenopogon Loew (Diptera: Asilidae)
Larry G. Bezark
California Department of Food and Agriculture, Biological Control Services
Program, Division of Environmental Monitoring and Pest Management, Sacra¬
mento, California.
Abstract.—A new species of Stenopogon Loew from La Pacifica in Costa Rica
is described. This represents the first Central American record for this genus.
Stenopogon setosus , New Species
Male .—Length 12 mm. Head black; face silvery tomentose, frons and occiput
golden tomentose. Mystax composed of strong white bristles extending halfway
to antennae; white hairs subequal in length to antennal segment 1, extending %
the distance from upper edge of mystax to antennal bases; occipital bristles brown;
bristles of frons and ocellar tubercle dark-brown. Antennae (Fig. 1) dark-brown,
bristles whitish; segments 11-6-20-11.
Mesonotum black, humeri dark-reddish, tomentum light-brownish, central
stripes and intermediate areas darker. Hairs numerous, brownish, anteriorly as
long as antennal segment 1, becoming longer and whitish posteriorly. Bristles
whitish; 3 humeral; 3-4 strong post-humeral; 4 strong, 1 weak presutural; 5
supraalar; 3 postalar; 3 strong dark-brown, 1 strong white anterior and 4 white
posterior dorsocentral. Pleurae and coxae black, golden-brown tomentose, hairs
and bristles white. Entire scutellum light-golden tomentose; 4 long white marginal
bristles.
Abdomen dark-brownish, silvery tomentose dorsally, becoming brownish to¬
mentose laterally. Hairs white, longer laterally on 1-3; 4 white lateral bristles on
1. Sternites dark-brownish, brown tomentose; hairs erect white on 1-7. Genitalia
dark-brownish; hairs white (Figs. 2-4).
Legs black; tibiae and tarsi dark-brownish. Hairs and bristles white; a few dark-
brown bristles on the tibiae and tarsi.
Halteres pale-brown. Wings very lightly infuscated; posterior cell 1 open; pos¬
terior cell 4 closed and petiolate; anal cell closed at wing margin.
Female. — Unknown.
Holotype. — Male, La Pacifica, 4 km NW of Canas, Gte Province (Guanacaste),
Costa Rica. Dec. -22/27-73, P. Opler. Deposited on indefinite loan in the collection
of the California Academy of Sciences.
Paratypes. — 3 males, same data, except: Jan. -23-74. Specimens deposited in
the collections of the Essig Museum of Entomology, Berkeley and the author.
Stenopogon setosus belongs to the Albibasis Group as defined by Wilcox (1971).
It differs from the other known species of this group by the presence of long hairs
above the mystax which extend nearly to the antennal bases, and by the closed
54
PAN-PACIFIC ENTOMOLOGIST
Figures 1-4. Stenopogon setosus, n. sp. 1. Antenna. 2-4. Male genitalia, lateral, dorsal, and ventral
views.
and petiolate 4 th posterior cell. The presence of long anterior dorsocentral bristles
will separate it from the remainder of the Albibasis Group species except for S.
lomae Wilcox, in which the anterior mesonotal hairs are yellowish and are only
as long as the second antennal segment. There is some variation in the color of
the anterior dorsocentrals and in the number of marginal scutellar bristles, ex¬
hibited by the paratypes.
Two additional specimens (one male, one female) of Stenopogon from Alajuela,
Costa Rica, are similar except the thorax and abdomen are reddish in ground
color. It is possible that they represent a second new species, but as they are in
poor condition, species determination will have to be deferred until more material
is accumulated.
The illustrations were prepared by Susan Kaiser. Thanks to L. Knutson (USNM)
for sending specimens in his care, and to C. S. Papp for technical assistance.
Special thanks go to John Chemsak who never lets me leave U.C. Berkeley without
unidentified asilids.
Literature Cited
Wilcox, J. 1971. The genera Stenopogon Loew and Scleropogon Loew in America North of Mexico
(DipteraiAsilidae). Occasional Papers of the California Academy of Sciences, No. 89, 134 pp.,
263 figs.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 55-62
Published 27 January 1984
Notes on Neotropical Veliidae (Hemiptera) VI.
Revision of the Genus Euvelia Drake
John T. Polhemus 1 and Dan A. Polhemus 2
3115 S. York St., Englewood, Colorado 80110.
The genus Euvelia was founded by Drake (1957) with E. advena Drake as the
only included species. Whereas Drake had but a single female before him, we
fortunately have series of E. advena , and of three new species described below.
A redescription of E. advena is provided along with a key to the known species.
We are indebted to the following individuals and institutions for the opportunity
to study specimens; abbreviations for the various collections are given in paren¬
theses and used in the text: Dr. R. T. Schuh, American Museum of Natural History,
New York (AMNH); Dr. P. H. Arnaud, California Academy of Sciences, San
Francisco (CAS); Dr. T. J. Henry, United States National Museum (Natural His¬
tory), Washington (USNM); Dr. R. C. Froeschner, Curator, Drake Collection,
Smithsonian Institution, Washington (CJD); Dr. H. H. Weber, personal collection
(HHW) and Max Planck Institut of Fimnologie, Plon (Plon); J. T. Polhemus
collection (JTP).
Unless otherwise noted, measurements are given in units, 60 units = 1 mm.
Euvelia Drake
Euvelia Drake, 1957. Proc. Biol. Soc. Wash., 70:116.
Type-species Euvelia advena Drake, monotypy and original designation.
This genus is characterized by four large blade-like pretarsal structures on each
middle tarsus, short fusiform antennal segments, and the middle and posterior
acetabula approximate and far removed from the front pair; other characteristics
are typical of the Microveliinae. The most closely related genus, Husseyella Her¬
ring (1955), has four much smaller blade-like pretarsal structures on each middle
tarsus and relatively much longer, more slender antennae, but shares the unusual
placement of the middle acetabula. The only other American member of the
Microveliinae with blade-like pretarsal structures is Xiphovelia lacunana (Drake
& Plaumann) with three blades on each middle tarsus and the acetabula about
equidistant from each other (see Polhemus, 1977).
The differences between Euvelia and Husseyella are not greater than say the
austrina group of Microvelia and other Microvelia species. We believe, however,
that a revision of the generic structure of the Microveliinae should result from a
cladistic analysis of all taxa of the subfamily, so this matter will be treated in a
later publication.
1 Contribution from University of Colorado Museum, Boulder, Colorado 80309.
2 Contribution from Department of Biology, University of Utah, Salt Lake City, Utah 84112.
56
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Euvelia species, anterior trochanter. A. E. advena Drake. B. E. lata, n. sp.
Key to Species of Euvelia
1. Body very broad; connexival segments 1 and 4 with silvery hair patches;
connexiva without pruinose areas (Fig. 3) . E. discala, n. sp.
Body moderately broad to broad; connexival segments without silvery hair
patches; connexival segments 2 and 4-7 (at least) with pruinose areas
(Fig. 4) . 2
2. Male fore trochanter produced distally into a long distinct spur (Fig. la);
width of females across humeri about 1.5 times width of head across
eyes; females without glabrous area on pleura . E. advena Drake
Male fore trochanter not or only slightly produced; females with a glabrous
area on pleura or width of humeri almost twice as wide as head across
eyes . 3
3. Females broad, with short to moderately long stout setae and no glabrous
area on pleura; males with a short anteriorly directed spur or tuft of stiff
hairs distally on fore trochanter (Fig. lb) . E. lata, n. sp.
Females narrower, with no stout setae on pleura, but with a depressed
glabrous area on pleura (Fig. 2); males without spur or tuft on fore
trochanter . E. concava, n. sp.
Euvelia advena Drake
Euvelia advena Drake, 1957. Proc. Biol. Soc. Wash., 70:116.
Description.— Apterous male: Elongate, broadest across mesonotal angles; ab¬
domen tapering evenly to apex; tergite seven trapezoidal. Length 1.37 mm, max¬
imum width 0.80 mm.
Ground color black, with scattered pruinose areas. Head black, narrow area
along base of vertex and adjoining eyes reddish brown; frons with scattered stout
VOLUME 60, NUMBER 1
57
h
1mm
Figure 2. Euvelia concava, n. sp., female, lateral view showing depressed glabrous area on pleura.
Figure 3. Euvelia discala, n. sp., female, dorsal view (legs omitted).
58
PAN-PACIFIC ENTOMOLOGIST
Figure 4. Euvelia concava, n. sp., female, dorsal view (legs omitted).
setae. Antennae chestnut brown, terminal segment black; proportions of segments
I-IV: 13; 11; 14; 14.
Pronotum black; anterior lobe with rectangular area medially near anterior
margin not reaching inner eye margins pink, surrounded by larger rectangular
black area covering medial part of anterior lobe and extending laterally beyond
inner eye margins, flanked by lateral pruinose areas; posterior lobe brownish to
black medially, black laterally. Posterolateral angles of metanotum pruinose. Pleu¬
ral region sparsely set with stout setae, without glabrous areas.
Abdomen black; connexiva moderately reflexed; tergites two, five, six, and
seven, and connexival segments two, four, five, and six with broad pruinose areas;
pruinose areas appearing lavender on more distal segments.
Ventral surface dark grey; rostrum chestnut brown, glabrous; margins of coxal
cavity and venter embrowned; narrow glabrous areas present medially on ab¬
dominal sternites one through six; usual small tuft of setae arising at end of scent
channel above and between hind and middle coxae. Legs pale yellow, infuscated
distally along femora and tibiae; male fore trochanter with distinct spur (Fig. la);
fore tibia sparsely set with stout setae; middle femora with sparse tuft of four
setae at base. Legs with following proportions:
VOLUME 60, NUMBER 1
59
Femur
Tibia
Tarsal I
Tarsal II
Anterior
22
22
9
Middle
44
46
11
22
Posterior
28
31
4
15
Genital segments small, inconspicuous; parameres tiny, symmetrical, not di¬
agnostic.
Apterous female: Similar to male but slightly larger, body shape more ovate;
tergite 8 broader than in male, nearly rectangular. Length 1.55 mm, maximum
width 0.89 mm.
Material examined. — BOLIVIA: 1 2 (Holotype), Cuarto Ojo, Piray River, IV-
25-1957 (CJD). BRAZIL: 1 <3, 2 2, Rio Madeira, Porto Velho, IX-1937, A. M.
Ollala (JTP); 2 6, 7 2, Goias, Santa Isabel, Ilha de Bananal, Rio Araguaia, B.
Malkin (JTP); 10 <5, 4 2, Source region, Rio Xingu, VIII-28 and 29-1965, E. J.
Fittkau (JTP, HHW, Plon); 1 6, 1 2, Rio Negro, 30 km below Barcelos, A333, II-
7-1962, E. J. Fittkau (HE1W, Plon); 17 6, 13 2, Rio Cuieiras, Igarape Cachoeira,
A428, XI-25-1962, E. J. Fittkau (JTP, HHW, Plon). PERU: 2 2, Estiron, Rio
Ampiacu, Loreto, rapid forest stream, XI-28 to XII-7-1961, B. Malkin (USNM).
Euvelia lata, New Species
Description .—Apterous male: Ovate, broadest across mesonotal angles, abdo¬
men curving broadly and evenly to apex. Length 1.34 mm, maximum width 0.95
mm.
Ground color black, with scattered pruinose areas; body thickly covered with
fine golden pubescence. Head black, narrow area along base of vertex and adjoining
inner margins of eyes reddish brown; frons with distinct medial cleft, set with
scattered stout setae, especially prominent at base of labrum. Antennae chestnut
brown, terminal segment black; proportions of segments I-IV: 11; 11; 16; 16.
Pronotum black; anterior lobe set with scattered stout setae, with ovate area
medially near anterior margin but not reaching inner eye margins pink, surrounded
by larger rectangular black area covering medial portion of anterior lobe and
extending laterally to inner eye margins, flanked by lateral pruinose areas; posterior
lobe black, with faint medial furrow. Posterolateral angles of metanotum pruinose.
Pleural region sparsely set with stout setae, lacking glabrous areas.
Abdomen black; connexiva weakly reflexed; tergite one with two small trian¬
gular pruinose areas flanking medial black area; tergites two, three, five, six, and
seven, and connexival segments two, four, five, and six with broad pruinose areas;
connexival margins set with stout setae.
Ventral surface dark grey; rostrum chestnut brown, glabrous; coxal cavity black,
margins embrowned; small glabrous areas present medially on abdominal sternites
four through six; usual small tuft of setae present at end of scent channel. Legs
chestnut brown, femora infuscated distally; fore trochanter with small, anteriorly
directed tuft of setae (Fig. 1 b); middle femora with tuft of four stout setae basally.
Legs with following proportions:
Femur
Tibia
Tarsal I
Tarsal II
Anterior
24
22
11
Middle
42
46
11
22
Posterior
28
31
4
15
60
PAN-PACIFIC ENTOMOLOGIST
Apterous female: Similar to male but slightly larger, body shape slightly broader.
Length 1.64 mm, maximum width 1.05 mm.
Material examined. —Holotype, male, BRAZIL: Rio Cuieiras, Igarape de Arara,
A410, XI-22-1962, E. J. Fittkau (Plon). Paratypes as follows: BRAZIL: 1 6, 1 9 ,
same data as holotype (JTP, Plon); 1 9 , Rio Cururu, Lago da Ziganea, A103,
1-24-1961, E. J. Fittkau (HHW); 1 9 , above Manaus, Cachoeira da Igarape Gigante,
A198, VII-3-1961, E. J. Fittkau (Plon); 1 6, above Manaus, St. Antonio, Igarape
da Bajaon, A276, XII-11-1961, E. J. Fittkau (HHW); 3 < 5 , 3 9 , Lago Catua, IX-
12-1961, E. J. Fittkau (JTP).
Discussion. — The key characters will separate this species from all other known
Euvelia. The body is relatively broad, but not nearly as broad as in E. discala,
n. sp. The name lata refers to the broad body.
Euvelia concava , New Species
Description .—Apterous male: Elongate, broadest across mesonotal angles; ab¬
domen tapering evenly to apex, connexival margins weakly sinuate. Length 1.25
mm, maximum width 0.75 mm.
Ground color black, with scattered pruinose areas. Head black, narrow area
along base of vertex adjoining eyes light brown; frons with scattered stout setae
along inner margins of eyes, lower portion pruinose. Antennae chestnut brown,
segment III infuscated, terminal segment black; proportions of segments I-IV:
11; 9; 11; 13.
Pronotum black; anterior lobe with roughly triangular area medially near an¬
terior margin not reaching inner eye margins pink, surrounded by larger rectan¬
gular black area covering medial part of anterior lobe and extending laterally to
inner eye margins, flanked by small lateral pruinose areas; posterior lobe black,
with distinct medial depression. Posterolateral angles of metanotum pruinose.
Pleural region sparsely set with stout setae, without glabrous areas.
Abdomen black; connexiva moderately reflexed; tergite one with two small
triangular pruinose areas flanking medial black area, bases adjoining posterior
lobe of pronotum; tergites two, three, five, six, and seven, and connexival segments
two, four, five, and six with broad pruinose areas; pruinose areas appearing blue
on more proximal segments fading to lavender distally.
Ventral surface dark grey; rostrum chestnut brown, glabrous; margins of coxal
cavity and venter embrowned; glabrous areas present medially on abdominal
sternites three through five; usual small tuft of setae arising at end of scent channel
between hind and middle coxae. Legs pale yellow, lightly infuscated distally along
femora and tibiae; fore and middle tibiae sparsely set with stout setae. Legs with
following proportions:
Femur
Tibia
Tarsal I
Tarsal II
Anterior 20
16
9
Middle 39
35
18
7
Posterior 29
22
4
11
Apterous female: Similar to male but slightly larger; pruinose areas of slightly
different shape (Fig. 4). Length 1.49 mm, maximum width 0.80 mm. Anterior
lobe of pronotum with stout setae; medial pink pruinose area more rectangular
than in male; pleural region with large glabrous area (Fig. 2). Abdomen with
VOLUME 60, NUMBER 1
61
connexiva strongly reflexed; pruinose areas more extensive than in male; triangular
pruinose patches on abdominal tergite 1 extending to posterior margin of tergite.
Material examined. — Holotype, male BRAZIL: Capitao Vasconselos, on Rio
Tuatuari, Upper Xingu Basin, Matto Grosso, VII-31-1957, B. Malkin (CAS).
Paratypes, BRAZIL: 21 S, 71 2, 3 nymphs, same data as holotype (CAS, JTP,
USNM).
Discussion.—Euvelia concava, n. sp. most closely resembles E. advena Drake,
but may be easily separated by the lack of a spur on the male anterior trochanter,
and the depressed glabrous area on the pleura of the female. The latter character
separates the females from all other known Euvelia species, and is apparently the
result of a modification to facilitate phoresy. The name concava refers to the
glabrous concavity on the female.
Euvelia discala, New Species
Description.— Apterous male: Ovate, broadest across mesonotal angles, shape
as in Figure 3. Length 1.46 mm, maximum width 1.05 mm.
Ground color black, with scattered silvery hair patches, body thickly clothed
in fine golden pubescence. Head reddish brown; frons black, with distinct medial
furrow, set with stout setae. Antennae chestnut brown, terminal segment lightly
infuscated, segments two and three each with three stout spines; proportions of
segments I-IV: 11; 11; 18; 16.
Pronotum black; anterior lobe with rectangular area medially near anterior
margin reaching inner eye margins reddish brown, sparsely set with stout setae;
posterior lobe black, with two round silvery hair patches flanking faint medial
furrow, set sparsely with stout setae. Pleural region moderately set with stout
setae, lacking glabrous areas.
Abdomen black; connexiva moderately reflexed, set with stout hairs; tergites
two, six, and seven, and connexival segments one and four with small round
patches of silvery hairs.
Ventral surface dusky brown; rostrum chestnut brown, glabrous; margins of
coxal cavity and venter embrowned. Legs chestnut brown; fore and middle tibiae
sparsely set with stout setae; distal comb on fore tibia prominent, black. Legs
with following proportions:
Femur
Tibia
Tarsal I
Tarsal II
Anterior
28
26
11
Middle
50
48
13
20
Posterior
31
35
4
15
Apterous female: Similar to male, but slightly larger and broader, shape and
markings as in Figure 3. Length 1.70 mm, maximum width 1.19 mm. Reddish
brown area on anterior lobe of pronotum divided medially by black line; margins
of pronotum somewhat more heavily set with stout setae than in male.
Material examined. — Holotype, male BRAZIL: Rio Cuieirias, Igarape da Arara,
A410, XI-22-1962, E. J. Fittkau (Plon). Paratypes as follows: BRAZIL: 4 6, 3 9,
same data as holotype (JTP, HHW, Plon); 1 9, Rio Negro, Furo de Paracuuba,
Si/Sa 7, IX-17-1959, Sioli & Sattler (Plon); 1 6, above Manaus, Reserva Duke,
Igarape Barro Branco, A584, XI-6-1965, E. J. Fittkau (Plon). PERU: 1 6, 1 9,
62
PAN-PACIFIC ENTOMOLOGIST
Estiron, Rio Ampiacu, Loreto, rapid forest stream, XI-28 to XII-7-61, B. Malkin
(USNM).
Discussion. — The broad body, long hairs on the dorsum, and silvery spots on
the connexiva will separate E. discala from other known species of the genus.
The name discala refers to the shape of the insect.
Literature Cited
Drake, C. J. 1957. New neotropical water-striders (Hemiptera). Proc. Biol. Soc. Wash., 70:111-118.
Herring, J. L. 1955. A new American genus of Veliidae (Hemiptera). Florida Entomol., 38:21-25.
Polhemus, J. T. 1977. Type-designations and other notes concerning Veliidae (Insecta: Hemiptera).
Proc. Entomol. Soc. Wash., 79:637-648.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 63-64
Published 27 January 1984
A New Species of Phymatodes from New Mexico
(Coleoptera: Cerambycidae)
James S. Cope
6689 Mt. Holly Dr., San Jose, California 95120.
Abstract.— Phymatodes shareeae, is described as new from the Nacimiento
Mountains, Rio Arriba County, New Mexico. Larval and pupal habits in Acer
glabrum are also reported.
Members of the relatively large genus, Phymatodes, are found throughout the
Holarctic Region. The species are mostly small, slender, with maculate elytra,
and are found mainly upon broad-leaved trees. In North America however, a
considerable number of the described species are immaculate and many utilize
coniferous hosts. A new species, closely related to Phymatodes vulneratus (Le-
Conte), was collected from northern New Mexico.
Phymatodes shareeae, New Species
Male. — Form subcylindrical; color piceous with coxae, bases of femora, tibiae,
and antennae rufous or rufotestaceous; elytra with a single pair of oblique white
fasciae just behind middle, expanded at the margin. Head with upper frons and
vertex coarsely, closely punctate; antennae nearly as long as body, scape sparsely
punctate, sparsely ciliate, third segment about as long as fourth. Pronotum a little
wider than long, sides broadly rounded, base moderately constricted; disk coarsely,
closely punctate, clothed with short, coarse, erect, dark pubescence, interspaces
shining. Elytra about twice as long as basal width, sides subparallel, apical half
often explanate, apices broadly rounded externally, narrowly rounded internally,
surface distinctly punctate at base, more finely and subrugosely punctate behind
fasciae; pubescence short, dark, suberect, shining, sparsely clothed with short,
erect hairs; posterior tarsi with first segment distinctly longer than following two
together. Abdomen with sternites polished, shining, very sparsely punctate and
subglabrous; fifth sternite shorter than fourth, apex broadly, feebly emarginate.
Length 6-9 mm.
Female. — Antennae surpassing elytral fasciae; femora less strongly clavate; fifth
sternite about as long as fourth, apex narrowly rounded. Length 6-10 mm.
Holotype male, allotype (California Academy of Sciences) from 9 miles SW
Coyote, Nacimiento Mountains, Rio Arriba County, New Mexico, beaten from
Acer glabrum Torrey, 21 June 1977 (J. S. Cope). Paratypes as follows: 37 males,
28 females, same locality data, reared from Acer glabrum, emerged April 2-19,
1978 (J. S. Cope). Paratypes are deposited in the following collections: University
of California at Berkeley, San Jose State University, J. S. Cope, E. Giesbert, F.
T. Hovore, J. Huether, R. Penrose, R. Surdick, R. H. Turnbow, Jr., W. H. Tyson,
J. E. Wappes.
64
PAN-PACIFIC ENTOMOLOGIST
This species resembles Phymatodes vulneratus but the two may be separated
by the uniform, dark, short, erect elytral pubescence, expanded elytral fasciae,
and rufous antennal coloration of Phymatodes shareeae. In contrast Phymatodes
vulneratus may be characterized by longer, erect, golden elytral pubescence, par¬
allel sided elytral fasciae, and typically dark antennae. Antennal coloration of
specimens from southern California often tends to be a light reddish-brown.
Phymatodes vulneratus is a Pacific Coast species ranging from Southern California
to British Columbia while Phymatodes shareeae is located 500 miles to the east
in northern New Mexico.
Biology. — Both of these closely related species utilize various species of Acer
as hosts. The entire type series was beaten from or reared from Acer glabrum.
Larval work is typical of other species in the genus, as summarized by Chemsak
and Powell (1964). Larvae work just under the bark of larger branches, lightly
scoring the sapwood. The meandering galleries are loosely packed with a light,
fine, granular frass mixed with darker bits of bark. Prior to pupation the larva
cuts an oblique tunnel into the wood, then turns parallel to the wood grain where
a pupal chamber is constructed. The exit hole is plugged with packed granular
frass. Adult emergence at the type locality apparently occurs in June.
The following key segment may be inserted into the “Key to North American
Species of Phymatodes ” found in Linsley’s (1964) monograph of the Cerambyci-
dae.
3(2). Elytra with 2 pairs of white fasciae or with a single pair just before apex
.(Linsley’s 4) 5
Elytra with a single pair of white fasciae just behind middle, pronotum
with disk distinctly, closely punctate; elytra distinctly punctate from
base to apex . 4
4(3). Elytra with single pair of oblique white fasciae, each parallel sided or at
most curving slightly forward just before margin; basal % of elytra
usually rufous or rufotestaceous, rarely black or piceous; antennae
usually dark, with long flying hairs on segments 2-7. Pacific Coast
species. vulneratus
Elytra with single pair of oblique white fasciae, expanded at margin;
elytra piceous; antennae rufotestaceous with short suberect hairs on
segments 2-7. New Mexico . shareeae
Acknowledgments
I wish to express my appreciation to John Chemsak for reviewing this manu¬
script and to my wife Sharee for assistance, patience, and encouragement under
circumstances often well above the call of duty.
Literature Cited
Chemsak, J. A., and J. A. Powell. 1964. Observations of the larval habits of some Callidiini with
special reference to Callidiellum cupressi (Van Dyke). J. Kansas Entomol. Soc., 37:119-122.
Linsley, E. G. 1964. The Cerambycidae of North America, Pt. V, Taxonomy and classification of
the subfamily Cerambycinae, Tribes Callichromini through Ancylocerini. Univ. Calif. Pubis.
Entomol., Vol. 22, 197 pp.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, pp. 65-66
Published 27 January 1984
Scientific Note
Feeding Behavior of Lacon profusa (Candeze) (Coleoptera: Elateridae) in
Carpenter Ant Attended Colonies of Cinara spp. (Homoptera: Aphididae) 12
Very little is known about the food habits of elaterid adults. Most reports suggest
that they feed on plant juices and young, tender leaves. Balduf (1935, The bionom¬
ics of entomophagous Coleoptera, John Swift Inc. Co., St. Louis), reported Cor-
ymbites virens Schr. feeding upon Vanessa sp. caterpillars (Lepidoptera: Nym-
phalidae). Bradley and Hinks (1968, Can. Ent., 100:40-50) noted adults of Lacon
brevicornis (LeConte) at rest in colonies of Cinara gracilis Wilson attended by
Dolichoderus taschenbergi (Mayr) and by Formica obscuripes Forel. Lacon pro-
fusus (Candeze) is treated as a subspecies of L. brevicornis by some workers (Arnett,
1952, Supplementary review of the Adelocerina, Wasmann J. Biol., 10:103-126).
Day (1971, The southern potato wireworm, U.S.D.A. Tech. Bull. 1443) observed
large numbers of adult Conoderus falli Lane (Elateridae) associated with honeydew
on plants but did not mention whether they were feeding on it. Thomas (1940,
Biology and control of wireworms, Bull. Pa. Agric. Exp. Stn., no. 392) reported
that baits containing sugar solution or molasses were attractive to some elaterid
adults. The literature appears devoid of reference to elaterids or any other Co¬
leoptera attending Homoptera except for Wheeler’s (1928, The social insects:
Their origin and evolution, Degan, Paul, Trench, Trubner and Co. Ltd., London,
378 pp.) report of silvanid beetles of the neotropical genera Coccidotrophus and
Eunausibius attending pseudococcids.
Adult L. profusa were occasionally found associated with Cinara occidentalis
Davidson on white fir {Abies concolor Gord. and Glen.) and Cinara moketa Hottes
on sugar pine ( Pinus labertiana Doug.). When disturbed by attending carpenter
ants, Camponotus modoc Wheeler, the beetle retracted its appendages into pronotal
grooves and remained motionless.
During 1977, a total of four females were observed, three at C. moketa and
one at C. occidentalis colonies. The first beetle was observed for 12 hours at the
same C. moketa colony on July 14, 1977, and it remained on the same tree until
it was collected on July 19. On July 15, this aphid colony, previously consisting
of 12 apterae and one alate, had disappeared and the beetle had moved to another
branch. As this beetle was not marked, it was not certain whether or not it was
the same individual on both occasions. The second beetle was observed on July
26 and marked with a spot of red paint on the right elytron. It remained on the
same tree until August 3 and at the same C. moketa colony for 4 days.
On July 14, 1977, another adult L. profusus was observed at one end of an ant-
attended C. moketa colony. The beetle remained motionless while using its an-
1 Supported, in part, by a contract (CX 8000-6-0016) to the Department of Entomological Sciences,
University of California, Berkeley, from the United States Department of Interior, National Park
Service.
2 From a thesis submitted to the University of California, Berkeley, in partial fulfillment of the
requirements for the degree of Doctor of Philosophy in Entomology, June 1979.
3 Present address: Division of Forest Entomology, Department of Plant and Forest Protection,
Swedish University of Agricultural Sciences, P.O. Box 7044, S-750-07 Uppsala, Sweden.
66
PAN-PACIFIC ENTOMOLOGIST
tennae and mouthparts to stroke an aphid directly beneath its head. This stroking
continued while carpenter ants crawled over the beetle’s dorsum. After about 15
minutes of such behavior, the attending ants were carefully removed. The beetle
continued its stroking behavior for another 10 minutes with no apparent effect
on the aphid. At this point, the beetle slowly picked up the aphid between its
mandibles and began consuming it. After 15 minutes, the aphid’s head was still
protruding from the beetle’s mandibles. A newly-arrived ant stroked the pro¬
truding head of the aphid and then walked away. After consuming the aphid, the
beetle began stroking another aphid which raised its abdomen and exuded a drop
of honeydew. Minutes later, the beetle was observed collecting a honeydew droplet
from another aphid. It continued stroking the same aphid for another 5 minutes
at which time it received more honeydew. This behavior contrasts with that of
ants which would move on to a new aphid immediately after collecting honeydew.
On July 17, an adult L. profusus was observed eating a C. moketa while under
attack simultaneously by several ants. One Camp, modoc worker grasped the
partially consumed aphid and bracing itself against the beetle’s head, tore off about
half of the aphid’s body. Although many ants exhibited alarm behavior (i.e., rapid
running about and greatly increased turning rate), the beetle remained absolutely
motionless with mouthparts and antennae retracted.
On July 19, 1977, the same beetle (marked earlier) was found at another C.
moketa colony where it consumed one aphid over a period of 9 minutes. Sub¬
sequently, it walked back to the trunk where it elicited no response from a minor
worker of Camp, modoc and was then attacked by a major worker. The attack
lasted about 5 minutes during which time the beetle remained motionless.
By collecting honeydew and consuming aphids, L. profusa obtains both car¬
bohydrates and protein. This rich food resource is both immobile and fairly
dependable. Thus the beetle may feed for prolonged periods with very little energy
expended in search of food, thereby increasing its fecundity and longevity. Al¬
though attending ants reduce the beetle’s foraging efficiency with their frequent
attacks, the ant’s presence is probably beneficial to the beetle by reducing com¬
petition for aphids from other predators and by reducing aphid dispersal (Tilles
and Wood, 1981, Can. Ent., in press). Although Camp, modoc competes directly
with L. profusus for both honeydew and aphids, the ants are much more efficient
honeydew collectors and are often successful in pulling aphids from the beetle’s
grasp. Protective behavior may be stimulated by aphid produced compounds to
which the ants respond. Camp, modoc becomes very excited when a crushed C.
moketa is suspended above it.
The armoured exoskeleton of the L. profusus appears to be highly resistant to
ant attacks. The beetle is apparently undisturbed by attacking ants as it may
continue stroking aphids while under attack, or alternatively, can maintain a
protective posture for varying periods and thus resume feeding within seconds
after cessation of an ant attack.
L. profusus appears to be an inefficient predator because: 1) it spends long
periods of time soliciting honeydew from a single aphid, often up to 15 minutes,
and 2) it often takes as long as 20 minutes to consume a single aphid.
D. A. Tilles, 3 Department of Entomological Sciences, University of California,
Berkeley.
PAN-PACIFIC ENTOMOLOGIST
60(1), 1984, p. 67
Published 27 January 1984
Scientific Note
New Records for Two Cerambycids from the Pacific Northwest
Specimens of Phymatodes oregonensis Chemsak are uncommon in collections
and the larval host has apparently not been recorded. The author reared two
female P. oregonensis from small branches of an Alnus sp. (A. rubra Bong, or A.
rhombifolia Nutt.) collected in Dabney St. Pk., Multnomah County, Oregon, in
December 1978. The infested wood was collected from the crown of a fallen tree.
The larvae mined beneath the bark and pupated in chambers constructed in the
sapwood. Both specimens emerged in February 1979.
A male of Obrium californicum Van Dyke was collected by the author on the
University of Idaho campus, Latah County, in July 1981 (NEW STATE REC¬
ORD). The specimen was taken from the foliage of ocean spray ( Holodiscus
discolor (Pursh) Maxim.) growing in a stand of western white and lodgepole pine.
Linsley (Univ. Calif. Publ. Entomol., 21:139) recorded O. californicum as occur¬
ring from central Oregon and northern California south through the Sierra Nevada
to the Tehachapi and Santa Cruz Mountains. It has also been collected in the San
Gabriel Mts. of southern California (F. T. Hovore, pers. comm.). This species is
known to use western yellow and Jeffrey pines for larval development (ibid.).
Robert H. Turnbow, Jr., Department of Entomology, Texas A&M University,
College Station, Texas 77843.
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Vol. 60 April 1984 No. 2
THE
Pan-Pacific Entomologist
CHANDLER, D. S.—A new species and records of Tanarthrus (Coleoptera: Anthicidae). 69
CHEMSAK, J. A. and E. G. LINSLEY—A new California species of Phymatodes Mulsant
(Coleoptera: Cerambycidae). 73
KORMILEV, N. A.—A new species of the genus Chinessa Usinger and Matsuda, 1959, from
Papua New Guinea (Hemiptera: Aradidae).... 76
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Emery (Hymenoptera: Formicidae). 79
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SMITH, D. R. and G. A. P. GIBSON— Filacus, a new genus for four species of sawflies
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CHEMSAK, J. A.—Description of a new purpuricenine genus, Linsleyella (Coleoptera: Cer¬
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RUST, R. W. — Synonymy in California Channel Island Epeolini bees (Hymenoptera: Antho-
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SMETANA, A. —Review of the Japanese species of the genus Gabrius Stephens (Coleoptera:
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linidae) . 163
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PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 69-72
Published 17 April 1984
A New Species and Records of Tanarthrus
(Coleoptera: Anthicidae ) 1
Donald S. Chandler
Department of Entomology, University of New Hampshire, Durham, New
Hampshire 03824.
Interest in the potential rare and endangered insects of public lands has led to
the granting of several federal contracts to determine the endemic insect species
which are associated with sand dunes. One such contract was given to the Cali¬
fornia Department of Food and Agriculture Insect Taxonomy Laboratory and
has dealt almost exclusively with the major dune formations of the Mojave Desert
in California. During the course of this study a number of rarely collected Tan¬
arthrus species were found, as well as a new species from the southern San Joaquin
Valley. Since many of these species possessed only a few distribution records in
my revision (Chandler 1975), new records are entered following the description
of the new species.
A portion of the contract report by Andrews et al. (1979) presented data on
Tanarthrus coruscus Chandler which was obtained from their year-long pitfall
trap sets in the Eureka Dunes of Inyo County. They found that this species occurred
only along the margin of an adjacent playa. This preference for the margins of
alkali lakes and saline soils has been detailed by Chandler (1974, 1979). The
majority of the specimens of coruscus were collected in May, with small numbers
being trapped from June until September. Individuals of Tanarthrus tartarus
Chandler and Vacusus confinis (LeConte) were discovered feeding on a dead
Phodaga alticeps LeConte (Meloidae), but it is uncertain whether they were at¬
tracted to the cantharidin in the beetle, or merely to the presence of a food source.
All measurements are in millimeters in the description of the new species. The
specimens from which the new records are taken are in the collections of the
California Department of Food and Agriculture and the author. I would like to
thank Dr. John F. Burger, University of New Hampshire, for reading the manu¬
script.
Tanarthrus ( Tanarthropsis) andrewsi, New Species
(Figs. 1, 2)
General description.— Length 3.08-3.95. Coloration varying from head, prono-
tum, and first third of elytra orange and remainder of elytra dark brown, to body
all dark brown with orange antennae and legs. Head basally with distinct median
impression, punctures on frons shallow and moderately dense, microreticulation
distinct between punctures, puncture diameter equal to that of eye facet; eleventh
antennomere distinctly constricted, portion before constriction equal in length to
tenth antennomere, portion after constriction about twice that of basal portion.
Pronotum with antero-lateral angles rounded; punctures dense on disc, similar in
1 Scientific Contribution Number 1144 from the New Hampshire Experiment Station.
70
PAN-PACIFIC ENTOMOLOGIST
1
depth and size to those on frons, areas near lateral margins feebly granulate,
microreticulation between punctures distinct. Elytra feebly ridged and lacking any
obvious microreticulation; pubescence directed posteriorly, all setae appressed.
Male genitalia with tegmen as long as phallobase; internal sac smooth, lacking
internal spines; primary gonopore lacking sclerotized spots or ribs.
Female similar to the male in external appearance.
Male holotype. —Soda Lake, San Luis Obispo County, California. Length 3.46.
Head 0.78 long, 0.79 wide behind eyes; tenth antennomere 0.08 long, eleventh
antennomere with portion before constriction 0.07 long, after constriction 0.15
long. Pronotum 0.71 long, greatest width at point 0.46 from base, collar 0.04
thick, 0.35 wide. Elytra 1.97 long, 0.87 wide across humeri; body dark brown
with orange antennae and legs.
Specimens examined. — HOLOTYPE male: California: San Luis Obispo Co.:
Soda Lake, VI-8-1979, F. G. Andrews and A. J. Gilbert, on salt encrustations at
lake edge. PARATYPES: 8 males, 35 females, eutopotypical. The holotype is to
be deposited in the California Academy of Sciences. Paratypes will be placed in
the collections of the author, the California Department of Food and Agriculture,
and the Floyd G. Werner Collection, Tucson, Arizona. The species is named for
Fred G. Andrews, who has collected many fine species of Anthicidae.
Distribution. — Only known from an alkali lake at the southern end of the San
Joaquin Valley.
Relationships. — Most similar to vafer Chandler by the proportions of the last
two antennomeres, and by both the male internal sac and primary gonopore
lacking spines or sclerotized spots. In andrewsi the base of the head is simply
depressed medially rather than cleft, and the punctures of the head and pronotum
VOLUME 60, NUMBER 2
71
are shallow, not really umbilicate as in vafer. Vafer may be a somewhat larger
species as the single specimen known (the holotype) is 4.23 mm long.
New Records of Tanarthrus
Tanarthrus {Tanarthrus ) salinus LeConte
California: Inyo Co.: SE end Saline Valley, III-30-1976, D. Giuliani, edge of
dry lake. Kern Co.: Koehn Lake, VI-14-1964, F. G. Andrews. San Bernardino
Co.: Saratoga Springs, VI-13-1964, F. G. Andrews.
Tanarthrus ( Tanarthropsis) alutaceus (LeConte)
California: Imperial Co.: 3 mi NW Glamis, IX-15/16-1972, M. Wasbauer &
A. Hardy, blacklight. Inyo Co.: Saline Valley, Artesian Well, 1100 ft, IV-26-1977,
D. Giuliani.
Tanarthrus ( Tanarthropsis ) brevipennis Casey
New Mexico: Chaves Co.: Bottomless Lakes State Park, VII-25-1975, K. Ste¬
phan.
Tanarthrus {Tanarthropsis) coruscus Chandler
California: Inyo Co.: S end Owens Lake, sand dunes, VII-15-1978, A. R. Hardy
& F. G. Andrews, at blacklight; Saline Valley dunes, III-30-1976, D. Giuliani,
cereal bowl pit trap; Eureka Valley Dunes, VIII-30-1976, D. Giuliani, blacklight.
Riverside Co.: Palen Dunes, IX-20-1977, A. R. Hardy & F. G. Andrews, at
blacklight. San Bernardino Co.: 3 mi W Rice, VI-1972, D. Giuliani, sand dunes;
Salt Lake, 5 mi N Renoville, IV-17-1974, F. G. Andrews & M. S. Wasbauer;
Saratoga Springs, Death Valley, V-6-1965, F. G. Andrews. Nevada: Lincoln Co.:
Game Range Dunes, VII-14-1975, F. G. Andrews & A. R. Hardy, at blacklight.
MEXICO: Baja California: Diablo Canyon Dry Lake, VII-16-1979, D. Giuliani,
blacklight.
Tanarthrus {Tanarthropsis) inhabilis Chandler
California: Inyo Co.: Eureka Valley Dunes, VII-13-1975, F. G. Andrews & A.
R. Hardy. San Bernardino Co.: Salt Creek, 5 mi N Renoville, IV-17-1974, F. G.
Andrews & M. S. Wasbauer.
Tanarthrus ( Tanarthropsis ) inyo Wickham
California: Inyo Co.: Deep Springs Valley, Buckhorn Springs, IV-1 1-1975, D.
Giuliani; Big Alkali Lake, VI-14-1979, F. G. Andrews, lake edge. Mono Co.:
Mono Lake, Paoha Island, VII-5-1980, D. Giuliani. Nevada: Churchill Co.: Sand
Mountain, VII-19-1977, D. Giuliani, UV light. Esmerelda Co.: Clayton Valley
dunes, IV-17-1974, F. G. Andrews & A. R. Hardy. Pershing Co.: Woolsey, VI-
27-1973, T. R. Haig.
Tanarthrus {Tanarthropsis ) iselini Chandler
New Mexico: Roosevelt Co.: Salt Lake near Arch, IX-20-1977, W. A. Iselin.
Tanarthrus {Tanarthropsis) tartarus Chandler
California: Riverside Co.: Palen dunes, IX-20-1977, A. R. Hardy & F. G.
Andrews, at blacklight; 7 mi SE Freda, IV-24/VII-26-1978, F. G. Andrews & A.
72
PAN-PACIFIC ENTOMOLOGIST
R. Hardy, small sand area. San Bernardino Co.: Cadiz Dunes, IX-18-1977, A. R.
Hardy & F. G. Andrews, at blacklight.
Literature Cited
Andrews, F. G., A. R. Hardy, and D. Giuliani. 1979. The coleopterous fauna of selected California
sand dunes. A report in fulfillment of Bureau of Land Management Contract CA-960-1285-
1225-DEOO. California Department of Food and Agriculture, Sacramento, i + 142 pp.
Chandler, D. S. 1975. A revision of Tanarthrus LeConte with a presentation of its Mid-Cenozoic
speciation (Coleoptera: Anthicidae). Transactions of the American Entomological Society, 101:
319-354.
-. 1979. A new species of Tanarthrus from California (Coleoptera: Anthicidae). Pan-Pacific
Entomologist, 55:147-148.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 73-75
Published 17 April 1984
A New California Species of Phymatodes Mulsant
(Coleoptera: Cerambycidae)
John A. Chemsak and E. G. Linsley
University of California, Berkeley, California 94720.
Specimens of Phymatodes are most commonly encountered by rearing. The
following new species was reared from branches of Juniperus californicus by J.
Cope and W. Tyson from the Mt. Hamilton area of California. The larval habits
are similar to those of P. nitidus and P. decussatus as reported by Chemsak and
Powell (1964) and Chemsak (1965). P. tysoni, n.sp. will key out to P. nitidus in
Linsley’s (1964) key to Phymatodes but may be separated by the characters in the
diagnosis below.
This study was carried out during a National Science Foundation sponsored
study on North American Cerambycidae (Grant DEB-8015523). We wish to thank
J. Cope and W. Tyson for making their specimens available for study and Carolyn
Tibbetts for preparing the illustration.
Phymatodes tysoni, New Species
Male. — Form small, a little depressed; integument shining, pale reddish brown,
legs darker, elytra black behind anterior white fasciae except along suture to a
little behind apices of posterior white fasciae, metasternum often infuscated, ab¬
domen black. Head finely, sparsely punctate, thinly clothed with long erect hairs;
antennae extending to about apical l h of elytra, segments from sixth subopaque,
clothed with very short pubescence, basal segments rather sparsely ciliate, second
segment less than Vi as long as third, third about as long as fourth. Pronotum as
broad as long, sides broadly rounded; disk finely, sparsely punctate, punctures
smaller than those at base of elytra, sides coarsely, confluently punctate; erect
hairs long, black; prosternum coarsely punctate at sides; meso- and metasternum
shallowly punctate, hind coxae clothed with pale, appressed pubescence. Elytra
less than 2 l h times as long as broad, sides vaguely expanded behind middle;
punctures sparse, moderately coarse, becoming finer toward apex; long, dark,
erect hairs rather sparse; fasciae ivory-white, subglabrous, anterior pair arcuate,
usually oblique on disk, posterior pair oblique, slanting posteriorly from near
suture at middle; apices rounded. Legs slender, femora strongly clavate, sparsely
punctate, long, erect hairs numerous. Abdomen shining, sparsely punctate and
pubescent; last sternite slightly emarginate at apex. Length, 5-7 mm.
Female. — Form similar. Antennae extending to a little beyond middle of elytra.
Pronotum with sides rather finely punctate; prosternum finely punctate at sides.
Legs with femora more slender. Abdomen with last sternite rounded at apex.
Length, 5-8 mm.
Holotype male, allotype (California Academy of Sciences) and 26 paratypes (11
males, 15 females) from Arroyo Mocho Canyon, Alameda Co., California, reared
from Juniperus californicus, 25, 30 March, 1972 (W. H. Tyson). Additional para¬
types: 20 males, 20 females, Arroyo Bayo, Mt. Hamilton, Santa Clara Co., Cal-
74
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Phymatodes tysoni Chemsak and Linsley. < 3 .
ifornia, reared from Juniperus californicus, 20 March, 1979, 14, 21 March, 1981
(J. Cope); 2 males, 7 mi NW Bitterwater, San Benito Co., California, 10 April,
1967, ex Juniperus (J. A. Chemsak); 8 males, 8 females, Isabelle Valley, Santa
Clara Co., California, 3 April, 1981 (J. Cope).
The type series is fairly uniform in coloration. Occasionally the pronotum is
partially infuscated and the legs are black.
P. tysoni may be separated from P. nitidus LeConte by the arcuate, oblique
VOLUME 60, NUMBER 2
75
anterior fasciae of the elytra. These are transverse in P. nitidus. Additionally the
posterior fasciae begin at the middle of the elytra and extend back obliquely in
P. tysoni. In P. nitidus this pair has its apices at the apical x h of the elytra and are
less oblique. The punctation of the pronotum in P. tysoni is finer than that at the
base of the elytra. The punctures in P. nitidus are subequal.
We are pleased to dedicate this species to W. H. Tyson for his continuing
cooperation on this project.
Literature Cited
Chemsak, John A. 1965. Habits of Phymatodes decussatus decussatus (LeConte). Pan-Pac. Entomol.,
41:71.
-, and J. A. Powell. 1964. Observations on the larval habits of some Callidiini with special
reference to Callidiellum cupressi (Van Dyke). Jour. Kansas Entomol. Soc., 37:119-122.
Linsley, E. G. 1964. The Cerambycidae of North America. Part V. Taxonomy and classification of
the subfamily Cerambycinae, Tribes Callichromini through Ancylocerini. Univ. California
Pubs. Entomol. 22, 197 pp., 60 figs.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 76-78
Published 17 April 1984
A New Species of the Genus Chinessa Usinger and Matsuda,
1959, from Papua New Guinea (Hemiptera: Aradidae)
Nicholas A. Kormilev
87-17 Myrtle Avenue, Glendale, New York 11385.
For the privilege of studying a small lot of Aradidae from Papua New Guinea
and the Bismark Archipelago, I am sincerely grateful to Paul H. Arnaud, Jr.,
California Academy of Sciences, San Francisco. The material included a new
species of Chinessa Usinger and Matsuda, the description of which follows.
The genus Chinessa is unusual in that it has a large number of endemic species
that are recorded from New Guinea. Of the 22 known species, all but two are
from this area, with one each otherwise known from New Britain and Ceram.
The genus Chinessa more than any other genus of Mezirinae shows specific di¬
versity—particularly in the size and the shape of genae, in the shape of the an¬
terolateral angles of the pronotum, and in the shape of the lobes of the sixth and
seventh abdominal segments.
All measurements in this paper were taken with an ocular micrometer, 25 units
equal 1 mm. For convenience, the length of the abdomen was taken from the
apex of the scutellum to the tip of segment IX and to the tips of the lobes of
segment VII.
Chinessa arnaudi Kormilev, New Species
(Figs. 1, 2)
Female.— Elongate, ovate, partially granulate; fore lobe of pronotum laterad of
inner tubercles smooth and shiny. Head (inclusive of genae) much longer than its
width across eyes (48:33); anterior process strongly forked; genae pointed and
divergent, much longer than clypeus and reaching 3 A of antennal segment II.
Antenniferous tubercles acute and divaricate; postocular consisting of two granules
placed one over another and directed backward. Eyes strongly protruding. Vertex
raised and granulate. Antennae 1.73 times as long as width of head across eyes;
relative lengths 15:13:16:13. Labium not reaching hind border of labial groove,
which is closed posteriorly. Pronotum less than half as long on median line as its
maximum width (32:72). Collar truncate anteriorly; anteriolateral angles produced
forward and inwardly, leaving a deep incisure between them and collar. Lateral
border rounded on fore lobe, then sinuate and subparallel on hind lobe; hind
border sinuate medially. Fore disc with a short median sulcus flanked by 2 (1 +1)
high tubercles; laterad of them smooth and shiny; 2(1 + 1) smaller tubercles placed
sublaterally. Hind disc roughly granulate. Scutellum shorter than its basal width
(35:40); lateral borders straight and carinate; basal border convex and carinate;
median carina thin and high, granulate; disc laterad of it sharply, transversely
rugose. Hemelytra reaching fore border of tergum VII; corium reaching basal l h
of connexivum III; its apical angle acute, apical border straight and only at inner
angle sinuate. Metathoracic scent gland openings small, moderately gaping. Legs
unarmed. Abdomen ovate, shorter on median line than its maximum width across
VOLUME 60, NUMBER 2
77
Figures 1, 2. Chinessa arnaudi, n. sp., 2. 1. Head and pronotum. 2. Tip of abdomen from above.
78
PAN-PACIFIC ENTOMOLOGIST
segment V (85:95), but longer to the tips of lobes of segment VII (100:95). Con-
nexivum wide, with uneven and punctured upper surface. Tergum VII raised
backward and granulate, its hind border truncate; tergum VIII short and wide,
but less than width of head across eyes (27:33). Lateral borders slightly convex
from II to V; posteroexterior angles III to V progressively protruding; VI forming
acute lobes, directed obliquely backward; VII forming long, acute lobes directed
backward and produced far beyond segment IX. Paratergites small, conical, as
long as the slightly tricuspidate segment IX. Spiracles II to VI ventral, placed far
from border; VII placed near the border and slightly visible from above; VIII
dorsolateral. Color black; hind lobe of pronotum and connexivum sepia brown;
labium and tarsi brown.
Total length. — 8.08 mm to tip of segment IX; 8.68 mm to tips of lobes of
segment VII; width of pronotum 2.88 mm; width of abdomen 3.80 mm.
Holotype. — 2 , Papua New Guinea, Morobe District, Forestry Road north of
Wau, 9.X. 1969 (James E. Tobler). Deposited in the collection of the Department
of Entomology, California Academy of Sciences, type no. 14029. It is a pleasure
to dedicate this curious species to Dr. Paul H. Arnaud, Jr. He has kindly made
available to me the Aradidae under his care for many years.
In my key to Chinessa (1971:117) C. arnaudi, n. sp. runs to C. forfex Kormilev,
1971, but may be separated from it by: smaller size; genae relatively shorter,
reaching only to 3 4 of antennal segment II (longer than segment II in forfex)-,
posteroexterior angles of connexivum VI produced into acute lobes (not produced
in forfex ); antennae relatively shorter, only 1.73 times as long as width of head
across eyes (1.8 in forfex).
Literature Cited
Kormilev, N. A. 1971. Mezirinae of the Oriental Region and South Pacific (Hemiptera-Heteroptera:
Aradidae). Pacific Ins. Mon., 29:1-165, 233 figs., 3 maps.
Usinger, R. L., and R. Matsuda. 1959. Classification of the Aradidae (Hemiptera-Heteroptera).
British Museum (Nat. Hist.), London, vii + 410 pp., 102 figs., pis. 1-4.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 79-87
Published 17 April 1984
Biology of the Thatching Ant Formica haemorrhoidalis
Emery (Hymenoptera: Formicidae)
Emma E. MacKay and William P. MacKay
Departamento de Entomologia, Colegio de Graduados, Escuela Superior de
Agricultura, Ciudad Juarez, Chih., Mexico.
The rufa group of the genus Formica has several species with high population
densities and extensive foraging areas, which makes them very important in
ecosystems. The six species present in Europe have been extensively studied. The
biologies of the more than 20 species that occur in North America are almost
unknown. Preliminary investigations have been done by McCook (1884), Cole
(1932), Weber (1935), Ayre (1957), and Talbot (1959, 1963, 1972).
Here we present the biology of Formica haemorrhoidalis Emery, a species of
the rufa group which occurs in southern California, USA. Formica haemorrhoi¬
dalis is a typical member of the rufa species group. It is distributed throughout
the higher great plains and mountains of western North America (Brown, 1965)
including North and South Dakota, Colorado, Utah, Washington, eastern Nevada
to the western slopes of the Sierra Nevada Mountains (Creighton, 1950) south to
the mountains of southern California (MacKay and MacKay, unpublished).
Methods and Materials
The study area was located in the San Bernardino Mountains of southern
California, USA, on the north side of Bluff Lake at 2100 meters. The vegetation
consists of yellow pine ( Pinus jejferyi Grev. and Balf.) and fir {Abies concolor
(Gord. and Glend.) Lindl.). The ants are absent on highly exposed sites and on
south facing slopes, but generally occur in clearings within the coniferous forest.
Nest populations were estimated by complete excavation of five nests located
outside of the study area. We were also able to determine the population distri¬
bution within each nest as well as the nest structure and collect the inquilines.
The process of complete excavation of each nest required approximately 100
hours. Each excavation was begun before sunrise to reduce the numbers of foragers
not counted. All of the individuals seen were captured and counted, as well as
the brood and inquilines. The populations of nests within our study area were
visually estimated, based on comparisons of activity, size, etc. with the five nests
we excavated.
Nest density was determined by delimiting a six hectare area and labeling all
of the nests with numbered aluminum tags. The area was examined weekly to
determine the movement of nests and/or formation of new nests, which were also
marked.
The movement of winged females and males was observed by individually
marking them before the mating flights, with combinations of three different
colored dots on the thorax (using Testor’s model paint).
Foraging activity was determined by surrounding two nests with sheet metal
buried to a depth of about 10 cm, with about 20 cm above ground. Few ants
80
PAN-PACIFIC ENTOMOLOGIST
attempted to cross the enclosure so it was not coated with tanglefoot or similar
materials. Two plastic tubes (6 cm length, 2 cm diameter) were placed in the sheet
metal at ground level for the entrance and exit of the ants. The longer end of each
tube was about one centimeter above the ground to prevent the foragers from
using the wrong tube to enter or exit the nest. Every hour (day and night) and for
one day each week throughout the foraging season of 1978, we collected samples
of foragers by placing a glass jar under the entrance tube for one to five minutes,
depending on the level of activity. The ants were counted and returned to the
nest and the numbers of foragers per hour calculated. All hours are expressed in
Pacific Standard Time.
Nest temperatures were measured using thermisters implanted in three different
levels of the nests. Air temperatures at the soil surface were also recorded.
Results and Discussion
External nest structure. —The nesting sites consisted of domes made of “thatch”
or pine needles, pieces of bark and other such materials, usually constructed
adjacent to stumps or logs. This type of nest is characteristic of ants of the rufa
species group (Ayre, 1957). Examination of 108 nests demonstrated that 59.1%
consisted of thatch on logs, 37.3% of thatch on stumps, and 3.6% were constructed
on the surface of the ground (MacKay and MacKay, 1982). As most of the nests
were associated with logs and stumps of Pinus jejfreyi, the distribution of the ants
may be limited by the distribution of the pines. The ant is more common in
disturbed areas due to the greater availability of nesting sites.
Observations of the 108 nests show that the thatching was more commonly
placed on the east side (Fig. 1). Nest construction occurred primarily in the
morning on the side which was first exposed to the sun. Scherba (1958) suggested
that either the ants sense temperature and build on the warmest side, or the
warmer temperatures facilitate nest building over longer periods of time and at
a faster rate.
Internal nest structure.— The ant population was distributed in the thatching
and in chambers in logs and stumps of P. jejfreyi, made by termites, wood boring
beetles or ants ( Camponotus spp.). Formica haemorrhoidalis was able to displace
the original inhabitants (MacKay and MacKay, 1982). The brood were usually
placed inside the chambers and protected with thatching. The larger chambers
were packed with thatching. The dryer areas under the bark and upper tunnels
contained primarily adult ants and pupae. The eggs, larvae, and queen(s) were
found in the lower chambers. No ants were found in the hard interior of the log
or stump. Areas under the bark along the roots of stumps often contained large
numbers of ants. The ants were rarely found in the soil and then only in the top
5 cm. The internal nest structure is typical of the rufa group (Weber, 1935; Talbot,
1971; Kloftetal., 1973).
Nest distribution and density.— The nests of F. haemorrhoidalis were located in
clearings in the forest (Fig. 2). The only exception was nest #1678 in the lower
right hand corner of Figure 2. This nest’s population was very small and located
in an extremely rotten stump. It may have been a remnant of an area that pre¬
viously had a higher nest density which had become overgrown with trees. The
colony died in the summer of 1979. The clumped nests in the clearings exchanged
workers (MacKay and MacKay, in prep.).
VOLUME 60, NUMBER 2
81
COMPASS DIRECTION
Figure 1. Distribution of thatching on the nest sites of Formica haemorrhoidalis.
Many authors conclude that the nests of species of the rufa group are spaced
“fairly uniformly” or “regular” ( subnitens: Ayre, 1957, 1959; opaciventris: Scher-
ba, 1964). Scherba (1964) concluded that the regular spacing of nests is due to
the spacing of suitable or preferred nesting sites or a tendency to locate budded
Figure 2. The 6 ha study area. Horizontal lines indicate dense woods; vertical lines, semidense
woods. Stippled areas indicate rocky areas with few trees. Clear areas represent clearings. Squares
represent nests with populations above 20,000 adult ants; circles, above 10,000 adult ants; triangles,
less than 10,000 adult ants. Encircled nests: nests which produced sexuals in 1978. The numbers refer
to the individual nests.
82
PAN-PACIFIC ENTOMOLOGIST
Table 1. A comparison of the nest populations of five Formica haemorrhoidalis nests.
#
Date
Type of
nesting site
Adult
ants
Brood
1
July 1977
Stump
27,006
10,570
2
July 1977
Log
16,419
347
3
August 1977
Log
29,681
447
4
September 1977
Log
2024
0
5
April-May 1978
Thatched dome
61,000
*
* Brood not counted.
nests at a distance from the parental nest, an expression of territoriality. Yasuno
(1964) found that colonies of F. japonica were overdispersed where they occurred
in uniform low-density stands and aggregated where they occurred in species-rich
areas.
The nest density within the six hectare study area was 5.7 nests/ha. In other
areas in the San Bernardino Mountains the nest density ranged from 1 to 15
nests/ha.
Populations and nest phenologies.—The mean adult worker population was
33,529 ± 9595 SE (Table 1). The data for nest #4 were disregarded in the cal¬
culation of the mean as it was apparently an incipient nest. Each nest had several
queens.
Large numbers of eggs and small larvae were found in a nest excavation begun
on 20 April 1978. Apparently, the queen began laying sometime before then. The
last snowfall occurred on 17 April and much of the area was still covered with
snow at the time excavation was begun. By the first part of July the brood pop¬
ulation reached its peak (Table 1). In late summer and fall there were few or no
brood in the nest.
Reproduction and nuptial flights.— The nuptial flights of Formica spp. are sel¬
dom seen (Donisthorpe, 1927). In F. haemorrhoidalis, the first winged females
were seen and captured at 0900 on June 4, 1978. Others were captured on June
6 between 0800 and 0900 and July 9, 1978 at 1400. During these days the females
briefly exited the nest and then reentered it. The nuptial flights began on July 17
and ended on July 23 during both summers. The first sexuals appeared at 0700.
Certain conditions of temperature and sunlight act as stimuli to initiate mating
flights (Ayre, 1957). On rainy days or after 1000 when the air temperature at the
soil surface was above 24°C the sexuals remained inside the nests.
The same individuals made several trips in and out of the nest before the flight,
appearing each morning until they finally flew. Workers usually ignored them,
but occasionally a worker would attempt to follow a female, capture and wrestle
with her, but she always escaped and continued moving away from the nest. The
males were more cautious than the females. When they exited from the nest, they
either immediately flew or rapidly returned to the nest.
Both sexes climbed plants or other objects before flying. They usually flew
upward in a spiral path until they were above the pine trees and then flew north.
The females completed more spirals to reach the top of the trees, possibly because
they were larger and heavier than the males. The workers encouraged the sexuals
to climb vegetation and fly by biting their legs and gaster.
VOLUME 60, NUMBER 2
83
Only a few sexuals left the nests each day. We could not find a swarming area.
One pair was observed to copulate in the air: A male on a nest flew directly to a
female which was flying nearby and captured her at an altitude of three meters.
Both fell to the ground and continued copulation.
After mating, the females landed on objects such as trees and vegetation near
established nests (occasionally they landed on the investigators). After a few sec¬
onds they flew to the nest surface and entered the nest with no interaction from
the workers of the nest.
During the nuptial flight, nest queens also emerged and took up positions on
high objects, presumably waiting for males to reinseminate them, as was reported
by Marikovsky (1961). We did not observe mating in such females.
Apparently only a small portion of the nests in the six hectare study area
produced sexuals. In 1978, sexuals were observed leaving only nests #1644, 1786,
1795, and 1829 (Fig. 2, encircled nests), 12% of the nests in the study area. All
of the other nests in the study area were checked each day. Surprisingly it was
the smaller and not the larger nests that produced sexuals (Fig. 2). Two of the
nests (#1795 and #1829) were less than a year old. Nest #1644 produced sexuals
during both 1977 and 1978. Sexuals emerged from each of the nests in the study
area at the same time.
Each nest apparently produced only a few reproductives. In 1978, we captured
and marked 11 females and 12 males from nest #1644; 46 females and 4 males
from nest #1786; 15 females and 11 males from nest #1795, and 71 females and
one male from nest #1829. Certainly we did not capture all of the reproductives,
especially the males, but our evidence indicates that few are produced in this
species. Most ant species produce large numbers of reproductives (MacKay, 1981).
As the mated queens of F. haemorrhoidalis are allowed to enter established nests,
the mortality rate would be much lower than that which occurs in other species
where the females must individually found new nests. In 1978, at least 143 females
were produced in the 6 ha study area and 11 were seen to enter established nests.
There were undoubtedly more females that entered nests which we did not observe,
because we could not observe more than two nests at any one time.
Nest formation .—Ant species which practice secondary pleometrosis (mated
females enter established nests) usually practice a form of nest formation referred
to as “budding.” It is simply an emigration of part of a nest to a new nesting site.
When workers locate a suitable site, they begin to carry other individuals to the
new site. Within a few days there is considerable movement of carried individuals
from the “mother” nest to the “daughter” nest. The winged females of F. hae¬
morrhoidalis are carried in the same manner as the workers, but are heavier and
more difficult for the workers to carry. The males refuse to be carried and have
to be dragged to the new nesting site.
We observed the formation of two permanent nests by budding. The first one
began on September 10, 1977, and was completed by October 10, 1977, at a
distance of 27 meters from the mother nest. The second one occurred around
July 17, 1978, at a distance of 23 meters from the mother nest.
The process creates some confusion or conflict as individuals are carried in
both directions. During two periods of observation (15 minutes each) on 10
September 1977, 62 and 19 workers, respectively, were carried to the “daughter”
nest, while 14 and 12 were carried back to the “mother” nest. During a 15 minute
84
PAN-PACIFIC ENTOMOLOGIST
period on 21 September 1977, 26 were carried to the “daughter” nest and 3 were
carried back to the “mother” nest.
There is considerable interest in secondary pleometrosis and budding of nests
in ants as there may be a conflict among the workers as to which shall stay with
the parental queen and which shall go with the new queen (Macevicz, 1979). It
would be interesting to use electrophoresis to determine the relationships of the
individuals involved. Of the 11 mated females observed entering established nests,
apparently only one was reentering the nest she left. The other 10 females were
not marked and therefore presumably came from nests which were outside of the
6 ha study area. They were entering established nests other than those they left
and thus were not “sisters” of the workers in the nest.
Movement of colonies. — Ants move the entire colony and even the nest material
when conditions become unfavorable (Donisthorpe, 1927). We witnessed the
movement of one nest to at least three different locations in the 1978 season.
Apparently, the first location was too small; the second one was a recently cut
stump which did not have chambers produced by other insects. The third location
was in a shaded area. The fourth location was not found; the nest may have died.
The time between the movements varied between one and two weeks. The process
of moving was similar to budding. The worker ants carried all the brood and
many of the coworkers. When the nest queens were moving, the ants became
alert and assumed an attacking position. The process of moving lasted up to three
to four days; during the process the ants did not forage. No other nests were seen
to change locations in the study area in the two years of observations.
Interactions between nests.— There was considerable movement of workers be¬
tween nests, especially in the spring (MacKay and MacKay, in prep.). This species
showed no aggressiveness between individuals, even when we mixed workers that
were collected from areas several kilometers apart. Three winged females from
three different nests were placed together in a large glass jar. There was no aggres¬
sion between them and one was observed cleaning the antennae and thorax of
the other two individuals. They commonly engaged in trophallaxis.
Foraging. — Foraging began early in the spring when there was still snow on the
ground and continued until the first heavy snowfall in the winter (May to October).
The ants foraged 24 hours a day throughout most of the foraging season, with a
small peak in activity about mid-morning and a much larger peak in late afternoon
(Fig. 3). The number of foragers was low between 0200 and 0500 and between
1000 and 1600. Foraging did not stop during rain. When there was a full moon
the ants increased their nocturnal activity, possibly due to better visibility. The
radius of the foraging area extended up to 23 meters from the nest. The prey and
food consisted primarily of insects and honey dew from aphids (Vielma et al., in
prep.).
Daily foraging activity was apparently influenced by temperature (Fig. 3). Dur¬
ing the hottest times of the day, most foragers were inside the nest. Ants outside
the nest took refuge beneath pine cones and other objects or waited in the shade
of pine trees until the afternoon when the temperature dropped.
Inquilines. — Several species of guests occurred near or inside the nests of F.
haemorrhoidalis. Many were accidentals, but others could have relationships with
the ants or other animals in the mound involving commensalism, mutualism or
predation. Formica haemorrhoidalis is insectivorous and destroys large numbers
of insects (Vielma et al., in prep.), but apparently does not harm the numerous
VOLUME 60, NUMBER 2 85
17,700
Figure 3. Daily foraging activity of a nest of Formica haemorrhoidalis and changes in temperature
of the nest on 15 July 1978.
species of inquilines. The most common guests include various species of beetles
listed by MacKay (1980). In addition we collected many Goniusa alperti Kistner
(Coleoptera: Staphylinidae) in the nests, which also occur in the nests of F. ob-
scuripes in Washington (Kistner, pers. comm.). Myrmecophila manni Schimmer
(Orthoptera: Gryllidae) is very common within the nests. Other ant species found
nesting in the domes include Leptothorax hirticornis Emery, L. andrei Emery and
L. muscorum (Nylander), the latter being very common. A colony of Manica
bradleyi (Wheeler) was found within one nest. Numerous unidentified spiders
were also collected in the nests.
Aggressive behavior.— The workers exhibit aggressive behavior when the nest
is directly attacked or disturbed, when they are collecting prey, and when they
are moving the queens to a new nesting site. The adult workers lift the anterior
part of their body and direct their antennae forward when approached at a distance
of 30 cm. Following this, they double the gaster under the thorax and squirt a
liquid containing formic acid up to 30 cm. Individuals can squirt consecutively
at least twice. Repeated attack causes irritation of the skin ultimately leading to
the formation of painful blisters and peeling of the skin.
Summary
Studies of the biology of Formica haemorrhoidalis Emery were conducted in
the San Bernardino Mountains of southern California, USA. The nesting sites
86
PAN-PACIFIC ENTOMOLOGIST
were stumps or logs covered with pine needles and other plant materials. Colonies
contained a mean of 33,529 adult ants. Oviposition began in early April, peak
populations of brood occurred in July. Nest density ranged from 1 to 15 nests/
ha. Nuptial flights occurred in July; relatively few reproductives were produced.
The mated females entered established nests, but usually not the same nest they
were raised in. New nests were formed by emigration of part of an established
nest. Foraging occurred from May to October. The ants foraged 24 hours a day
with a small peak in activity in midmorning and a larger peak in later afternoon.
Numerous inquilines were found in the nests, especially beetles, crickets, and
other species of ants.
Acknowledgments
The United States Forest Service generously granted permission to conduct the
investigation on property under their jurisdiction. A. Fran§oeur verified the iden¬
tification of Formica haemorrhoidalis. D. Kistner, F. Andrews, and K. Cooper
identified the beetles. An anonymous reviewer made many valuable suggestions
in the manuscript. The research was partially supported by Sigma Xi, the Scientific
Research Society of North America.
Literature Cited
Ayre, G. L. 1957. Ecological notes on Formica subnitens Creighton (Hymenoptera: Formicidae).
Ins. Sociaux, 4(3): 173-176.
-. 1959. Food habits of Formica subnitens Creighton (Hymenoptera: Formicidae) at Westbank
British Columbia. Ins. Sociaux, 6(2): 105-114.
Brown, W. M. 1965. Studies on North American ants. I. The Formica integra subgroup. Entomol.
News, 76:181-186.
Cole, A. C. 1932. The thatching ant Formica obscuripes Forel. Psyche, 39(l-2):30-33.
Creighton, W. S. 1950. The ants of North America. Bull. Mus. Comp. Zool., 104:1-585 + 57 plates.
Donisthorpe, H. 1927. British ants. 2nd ed. Toutledge and Sons, London, xiii 4 - 244 pp.
Kloft, W. J., R. C. Wilkinson, W. H. Whitcomb, and E. S. Kloft. 1973. Formica integra (Hyme¬
noptera: Formicidae); I. Habitat, nest construction, polygyny, and biometry. Fla. Entomol.
56(2):67-76.
Macevicz, S. 1979. Some consequences of Fisher’s sex ratio principle for social hymenoptera that
reproduce by colony fission. Amer. Nat., 113:363-371.
MacKay, W. P. 1980. Ptomophagus californicus (Leconte) (Coleoptera: Leiodidae), an inquiline in
the nests of the ant Formica haemorrhoidalis Emery. Coleopt. Bull., 34(l):79-83.
-. 1981. A comparison of the nest phenologies of three species of Pogonomyrmex harvester
ants (Hymenoptera: Formicidae). Psyche, 88(l-2):25-74.
-, and E. MacKay. 1982. Coexistence and competitive displacement involving two native ant
species (Hymenoptera: Formicidae). Southwest. Nat., 27(2): 135-142.
Marikovsky, P. I. 1961. Material on sexual biology of the ant Formica rufa L. Ins. Sociaux, 8(1):
23-30.
McCook, H. C. 1884. The rufous thatching ant of Dakota and Colorado. Proc. Acad. Natur. Sci.
Philad., Part I, PP- 57-65.
Scherba, G. 1958. Reproduction, nest orientation and population structure of an aggregation of
mound nests of Formica ulkei Emery (Formicidae). Ins. Sociaux, 5(2):201—213.
-. 1964. Species replacement as a factor affecting distribution of Formica opaciventris Emery.
J. N.Y. Ent. Soc., 72:231-237.
Talbot, M. 1959. Flight activities of two species of ants of the genus Formica. Am. Midi. Nat., 61(1):
124-132.
-. 1963. Nest structure and flights of the ant Formica obscuriventris. Anim. Behav., 12:154-
158.
-. 1971. Flights of the ant Formica dakotensis Emery. Psyche, 78(3): 169—179.
VOLUME 60, NUMBER 2
87
-. 1972. Flights and swarms of the ant Formica obscuripes Forel. J. Kans. Entomol. Soc., 45(2):
254-258.
Weber, N. 1935. The biology of the thatching ant, Formica rufa obscuripes Forel, in North Dakota.
Ecol. Monographs, 5(2): 166-206.
Yasuno, M. 1964. The study of the ant population in the grassland at Mt. Flakkoda II. The distribution
pattern of ant nests at the Kayano grassland. Science Reports of the Tohoku University Sendai,
Japan. Ser. 4(Biol), 30(l):43-55.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 88-93
Published 17 April 1984
Descriptions of the Nearctic Larvae of Pseudosmittia gracilis,
Mesocricotopus thienemanni and Heleniella nr. ornaticollis
(Diptera: Chironomidae: Orthocladiinae)
John W. Steiner
U.S. Geological Survey, National Water Quality Laboratory, 6481-H Peachtree
Industrial Boulevard, Doraville, Georgia 30340.
The works of European chironomid taxonomists have been used extensively
by their North American counterparts (Roback, 1957; Simpson and Bode, 1980;
and many others). The review compilations by Pankratova (1970) and Strenzke
(1950) have proved to be especially useful for the identification of the larvae of
the subfamily Orthocladiinae. Because many species of this subfamily are holarctic
in their distribution (Beck, 1980), specific identifications of nearctic larvae can
often be made based on European descriptions. This is especially true when the
measurements and morphological characters of North American larvae are con¬
gruent with those of European specimens, as in Pseudosmittia gracilis Goetghe-
buer. Specific identifications can also be made when the larvae are extremely
distinctive and the genus is monotypic as in Mesocricotopus thienemanni Brundin.
The specimens for this study were collected in conjunction with U.S. Geological
Survey water resource projects. The descriptions that follow are the first that are
complete for the larval stages and also the first that are based on nearctic speci¬
mens. The diagnoses should be useful in water quality investigations. Unless
otherwise noted, the measurements in the descriptions are means expressed in
microns. The terminology follows Saether’s (1980a) glossary.
Pseudosmittia gracilis Goetghebuer
Diagnosis. — The larvae of the genus Pseudosmittia Strenzke can be distin¬
guished from those of all other nearctic Orthocladiinae by the following combi¬
nation of characters: procerci absent; seta interna of mandible absent; antennae
reduced with segment I short and square; SI and SII bifid with equal rami; semi¬
terrestrial habitat. The larva of P. gracilis differs from the other known species
of the genus in these characters: anal tubules hemispherical; mandible with 4
subequal inner teeth; antennal blade as long as segment I; premandible with 2
inner lobes and 2 sharp teeth (Fig. 1). Examined larvae (n = 4) were collected
from Bitter Creek near Fort LaClede, Wyoming on November 4, 1976.
4th instar larva. — Head capsule brown with darker mouth parts and occipital
margin. Body gray-white. Total length of larva about 3.3 mm.
Head. — Length about 330, width about 265; eyespots large, single. Mentum
with 11 teeth in shallow arc; median tooth with center peaked; width: 103. Ven-
tromental plates thin, short and curving to the base of mentum. Mandible with
4 short lateral teeth and a peg-like seta subdentalis; length: 113. Premandible bent
at a right angle with 2 blunt inner lobes and 2 long sharp teeth; apical tooth light
colored; length: 63. Antenna reduced, 3-segmented with blade about as long as
VOLUME 60, NUMBER 2
89
Figure 1. Pseudosmittia gracilis fourth-instar larva. Head, ventral aspect. Inset: antenna.
first segment; length of segments: 10:3:1; 1st segment about as wide as long with
a large ring organ. Epipharynx with SI and SII bifid with equal rami; pecten
epipharyngis consists of 3 small teeth.
Body. — Procerci absent. Posterior parapods reduced, each with 6-8 small yellow
claws. Anal tubules hemispherical. Body segments without setae.
The larva of this species has been described and partially illustrated by Thie-
nemann (1944), Strenzke (1950) and Pankratova (1970). It has not previously
been reported from North America. Because they are usually terrestrial, the larvae
of Pseudosmittia gracilis are seldom found in streams except after torrential rains
or cave-ins. Although at least nine species of this genus are described in European
literature, P. setavena (Saether, 1969) is the only described (adult stages) nearctic
species. A larva designated as “Pseudosmittia group” was keyed and pictured by
Oliver et al. (1978).
90
PAN-PACIFIC ENTOMOLOGIST
Mesocricotopus thienemanni Brundin
Diagnosis. — This is a monotypic genus that can be differentiated from other
nearctic Orthocladiinae by these characters: Antennae 5-segmented with segment
III very short and square; antennal blade exceeds antennal apex; ring organ at
about 0.4 of segment I; premandible simple with a clear inner lobe; mandible
with 3 small inner teeth and a smooth subdental margin; mentum with 12 dark
teeth with median teeth very large and laterally indented (Fig. 2). Examined larva
was collected from Sikolik Lake, Alaska on July 7, 1977.
4th instar larva. — Head capsule light brown with dark brown mouthparts. Body
light tan. Total length of larva 5.9 mm.
Head. — Length about 335, width about 260. Mentum dark brown with 12 teeth;
median teeth very large with lateral indentations; sides of mentum nearly parallel;
width: 110. Ventromental plates very thin and curving to basal lateral corners of
the mentum. Mandible with 3 small subequal inner teeth and a short clear seta
subdentalis; length: 93. Premandible simple and slightly bent; length: 64. Antenna
with blade exceeding apex; length of segments: 30:12:2:13:4. Antennal tubercles
prominent. Epipharynx with SI bifid and SII simple; pecten epipharyngis consists
of 3 narrow sharp teeth.
Body. — Posterior parapods each with 14 variable light yellow claws. Procerci
sclerotized each with 7-8 anal setae. Anal tubules pointed and digitate. Abdominal
segments with scattered simple setae.
Chernovskii (1949) keyed this larva as “Orthocladiinae gen.? karelica ” and
stated that it was found in “lakes and rivers of Karelia, [USSR] rare.” Pankratova
(1970) placed this species in the genus Limnophyes Eaton. Saether (1980b) stated
that the larva corresponds to adults described by Brundin (1956) as Mesocrico¬
topus. Later in a personal communication, Dr. Saether stated that the species is
M. thienemanni and that the larvae are “. . . found in the littoral to the profundal
zones of oligo- and mesotrophic lakes.” The species is apparently holarctic, but
confined to boreal regions.
Heleniella nr. ornaticollis Edwards
Diagnosis. — The larvae of Heleniella Gowin are distinctive in the following set
of characters: antenna 6-segmented with blade exceeding apex and segment II
interrupted; premandible bifid with an inner lobe; SI plumose; mentum with 12
teeth with last lateral teeth larger than penultimates; mandible with one sharp
and two blunt inner teeth (Fig. 3). The species described below has not been
associated with adults and no other larva of this genus has been described from
North America. Therefore a specific diagnosis is not possible. Examined larvae
(n = 7) were collected from an unnamed creek near Huntingdon, Utah on June
7, 1977.
4th instar larvae. — Head capsule yellow with cinnamon brown mouth parts.
Body gray-white. Total length of larva about 5.2 mm.
Head. — Length about 340, width about 250. Mentum with 12 unicolorous teeth;
median teeth tallest with a distinctive U-shaped notch between them; 4th lateral
teeth recessed between 3rd and 5th laterals; width: 90. Ventromental plates with
anterior margins overlying the bases of the 2nd and 3rd mental teeth. Mandible
with apical tooth and first inner tooth sharply pointed; subdental seta small; length:
80. Premandible bifid with a blunt inner lobe; length: 67. Antenna 6-segmented
VOLUME 60, NUMBER 2
91
Figure 2. Mesocricotopus thienemanni fourth-instar larva. Head, ventral aspect.
with blade exceeding apex; segment II interrupted at about 0.2; segment VI thread¬
like and difficult to see; length of segments: 48:23:6:5:3:2. Epipharynx with SI
plumose with 6-8 short points; SII simple; pecten epipharyngis consisting of 3-
5 blunt lobes.
Body. — Procerci short, yellow and sclerotized, each with 8-10 anal setae. Pos¬
terior parapods each with 16 clear simple claws. Anterior parapods each with 60-
90 mostly pectinate yellow claws.
Epilithic diatoms such as Synedra and Cymbella were found in the guts which
92
PAN-PACIFIC ENTOMOLOGIST
Figure 3. Heleniella nr. ornaticollis fourth-instar larva. Head, ventral aspect.
explains the worn condition of the mouthparts of most larvae. Saether (1969)
described Heleniella curtistyla and H. hirta from adults and pupae collected in
Canada. The larvae described here may correspond to either of these species or
VOLUME 60, NUMBER 2
93
to an unknown species. The specimens appear to be very similar to H. ornaticollis
as keyed and illustrated by Cranston (1979). Heleniella larvae are found primarily
in boreal or montane streams.
All specimens are deposited in the U.S. Geological Survey National Water
Quality Laboratory permanent collection in Doraville, Georgia.
Acknowledgments
My good friends, Broughton Caldwell and Craig Moore, reviewed the manu¬
script and offered many comments and suggestions. Their patient help is greatly
appreciated. I thank Dr. O. A. Saether who identified to species my drawing of
Mesocricotopus thienemanni.
Literature Cited
Beck, W. M. 1980. Interesting new chironomid records for the southern United States (Diptera:
Chironomidae). J. Georgia Entomol. Soc., 15( 1):64—73.
Brundin, L. 1956. Zur Systematik der Orthocladiinae (Dipt. Chironomidae). Rep. Inst. Freshwat.
Res. Drottningholm, 37:5-185.
Chemovskii, A. A. 1949. Identification of larvae of the midge family Tendipedidae. Inst. Akad.
Nauk. SSSR. 31. English translation of Russian by E. Lees, Freshwater Biological Association,
K. E. Marshall (ed.). National Lending Library of Science and Technology, Boston Spa, York¬
shire, England (1961), pp. 1-293.
Cranston, P. S. 1979. The biosystematics of British aquatic larval Orthocladiinae. Ph.D. thesis,
Queen Mary College, University of London.
Oliver, D. R., D. McClymont, and M. E. Roussel. 1978. A key to some larvae of Chironomidae
(Diptera) from the Mackenzie and Porcupine River watersheds. Can. Fish. Mar. Serv. Tech.
Rep., 791:1-73.
Pankratova, V. YA. 1970. Larvae and pupae of midges of the subfamily Orthocladiinae of the fauna
of the U.S.S.R. (Diptera, Chironomidae = Tendipedidae). Inst. Akad. Nauk. SSSR, 102:1-343.
(In Russian)
Roback, S. S. 1957. The immature tendipedids of the Philadelphia area. Monogr. Acad. Natur. Sci.
Philadelphia, 9:1-152 + 28 pis.
Saether, O. A. 1969. Some Nearctic Podonominae, Diamesinae, and Orthocladiinae (Diptera: Chi¬
ronomidae). Bull. Fish. Res. Bd. Can., 170:1-154.
-. 1980. Glossary of chironomid morphology terminology (Diptera: Chironomidae). Entomol.
Scand. Suppl., 14:1-51.
-. 1980b. The females and immatures of Paracricotopus Thienemann and Hamisch, 1932,
with the description of a new species (Diptera: Chironomidae). Aquatic Insects, 2(3): 129-145.
Simpson, K. W., and R. W. Bode. 1980. Common larvae of Chironomidae (Diptera) from New
York State streams and rivers with particular reference to the fauna of artificial substrates. New
York State Mus. Bull., 439, 105:1-105.
Strenzke, K. 1950. Systematik, Morphologie und Okologie der terrestrischen Chironomiden. Arch.
Hydrobiol. Suppl., 18:207-414.
Thienemann, A. 1944. Bestimmungstabellen fur die bis jetzt bekannten Larven und Puppen der
Orthocladiinen (Diptera Chironomidae). Arch. Hydrobiol., 39:551-664.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 94-96
Published 17 April 1984
Meridic Diet for Rearing of the Host Specific
Tropical Wood-borer Plagithmysus bilineatus
(Coleoptera: Cerambycidae)
John D. Stein and Janis E. Haraguchi
Pacific Southwest Forest and Range Experiment Station, Forest Service, U.S.
Department of Agriculture, Berkeley, California, and Honolulu, Hawaii.
Abstract.— A synthetic diet was successfully developed to rear Plagithmysus
bilineatus, a host specific tropical wood-borer. Host material incorporated into
the diet stimulated 1 st instar larval feeding immediately after eclosion. The basic
diet was also used to rear Plagithmysus funebris, Plagithmysus varians, Phora-
cantha semipunctata, Curtomerus flavus, Placosternus crinicornis, and Sybra al¬
ter nans.
The tropical wood-borer Plagithmysus bilineatus Sharp is an endemic pest
associated with the decline of ohia-lehua ( Metrosideros polymorpha Gaudichaud)
on the island of Hawaii (Papp et al., 1979). Concern for potential impact on the
ohia ecosystem prompted biological studies that necessitated rearing Plagithmysus
on artificial diet.
Several modifications of Adkisson’s et al. (1960) diet for the pink bollworm,
Pectinophora gossypiella (Saunders), have been used to rear wood-borers. Harley
and Willson’s (1968) modification was developed for rearing the lantana borer,
Plagiohammus spinipennis Thomson, and Gardiner (1970) incorporated pulver¬
ized plant material into his version to successfully rear temperate species of
cerambycid borers. We used both diets in Hawaii to rear adult beetles from larvae
collected in the field up to 40 days prior to pupation. However, our efforts to rear
the 1st instar larva on these diets were unsuccessful.
This paper reports yet another modification of the diet developed by Adkisson
et al. (1960) for the pink bollworm. The modified diet, which is easier to prepare,
differs from Adkisson’s diet in several ingredients. It has proven useful in labo¬
ratory cultures of Plagithmysus bilineatus and several other endemic and intro¬
duced cerambycids.
Preparation
To prepare 1500 ml of the diet (Table 1), first add agar to 900 ml of distilled
water in a 1000-ml beaker and bring to a boil. After the agar dissolves, pour the
solution into a large Waring Commercial Blender (CB-6) and slowly add peptone,
Vanderzant Adkisson Wheat Germ Diet (ICN Pharmaceuticals, Inc.) and 15 g of
ohia-lehua sawdust. 1 Blend for 2 minutes at low speed. Then pour the mixture
1 Trade names and commercial enterprises or products are mentioned solely for information. No
endorsement by the U.S. Department of Agriculture is implied.
VOLUME 60, NUMBER 2
95
Table 1. Composition of the Meridic diet used to rear Plagithmysus bilineatus with quantity of
ingredients per 1500 ml of media.
Ingredients
Quantity
Water
1260 ml
Agar
70 g
Vanderzant Adkisson Wheat Germ Diet
187.5 g
Sawdust
15 g
Peptone
3g
Inhibitor:
Ethyl alcohol (95%)
16.6 ml
Sorbic acid
2g
Methyl p-hydroxybenzoate
1.4 g
Potassium hydroxide (KOH)
3 g
Vanderzant Modification Vitamin Mixture
75 g
from the blender into three 1000-ml beakers, cover with foil, and autoclave for
15 minutes at 15 psi. While the wheat germ and sawdust mixture is in the au¬
toclave, mix the alcohol, sorbic acid, methyl p-hydroxybenzoate, potassium hy¬
droxide, and Yanderzant Modification Vitamin Mixture (ICN Pharmaceuticals,
Inc.) into 360 ml of distilled water until they dissolve. Remove the agar mixture
from the autoclave and cool to 50°C, then add the antimicrobial-vitamin solution
to the agar and mix in the blender for 2 minutes at low speed. Pour liquified diet
into 9 x 50 mm petri dishes with snap-tight lids or 16 x 100 mm disposable
petri dishes until three-fourths full. The petri dishes can be stored in the refrigerator
until needed.
The Plagithmysus diet, compared with that of Adkisson, deletes the antimicro¬
bial butyl p-hydroxybenzoate and adds linseed oil, cholesterol, and ascorbic acid—
which are in the wheat germ diet—and peptone, potassium hydroxide, and ohia-
lehua sawdust. The incorporated sawdust stimulates feeding and improves larval
acceptance of the diet immediately after hatching.
Insect Culture
Eggs were obtained by caging wild-type males and females in the laboratory at
21°C. Small pieces (4x10 cm) of ohia-lehua bark placed on the cage floor were
examined daily for eggs, which were collected and placed in small petri dishes.
Immediately after eclosion, 1 st instar larvae were transferred to the diet in petri
dishes with snap-tight lids. As the larvae matured they were transferred to the
larger petri dishes.
Plagithmysus bilineatus development time from the egg to the adult stage dif¬
fered significantly between the wild-type and diet-reared specimens (P < 0.01).
Wild-type specimens developed in an average of 269.6 days (SE 4.71) compared
with 171.4 days (SE 2.31) for diet-reared specimens. Size of adults reared on the
diet did not differ significantly from that of wild-type specimens {P < 0.05). Av¬
erage dry weight of laboratory-reared adults (28.7 mg, SE 1.01) and wild-type
adults (27.0 mg, SE 0.88) was similar.
Of the adults reared from the egg stage, 17% had deformed metatarsal segments.
This deformity was nonexistent for those field collected larvae placed on the diet
96
PAN-PACIFIC ENTOMOLOGIST
after the 2nd instar and may be indicative of some nutrient deficiency in the first
two instars. Larval mortality was 37% and was attributed primarily to injury of
1 st instar larva during transfer to the media.
Discussion
Use of this synthetic diet will reduce the development time of P. bilineatus and
produce laboratory-reared adults equivalent to the wild-type in size. This media
has not been evaluated for effects such as fecundity, loss of host preference, or
other criteria necessary for continuous laboratory rearing, but may serve as a basis
for modified diets of numerous tropical woodborers.
The synthetic diet proved successful for rearing other endemic beetles when
sawdust from host trees was substituted for ohia-lehua sawdust. We reared Pla-
githmysus funebris Sharp on diet containing mamane, Sophora chrysophylla (Salis¬
bury), sawdust and Plagithmysus varians Sharp on diet containing koa, Acacia
koa Gray, sawdust.
We also reared late instar larvae of the introduced beetles Phoracantha semi-
punctata (Fabricius) and Curtomerus flavus (Fabricius) on diet with Eucalyptus
robusta Smith sawdust, and Placosternus crinicornis (Chevrolet) and Sybra alter-
nans (Wiedeman) on diet with koa sawdust added. Late instar larvae of Plagith¬
mysus bilineatus, which is host specific on ohia-lehua, also developed on the
modified diet to which Eucalyptus robusta or koa sawdust had been added. The
development of Curtomerus flavus, Placosternus crinicornis, Sybra alternans, and
Plagithmysus bilineatus on diet with sawdust of nonhost tree species indicates
that acceptance of the diet was not a critical factor after the 2nd instar. This agrees
with observations made by Gardiner (1970).
Literature Cited
Adkisson, P. L., E. S. Vanderzant, D. K. Bull, and W. E. Allison. 1960. A wheat germ medium for
rearing the pink bollworm. J. Econ. Entomol., 53:759-762.
Gardiner, L. M. 1970. Rearing wood-boring beetles (Cerambycidae) on artificial diet. Canad. Ento¬
mol., 102:113-117.
Harley, K. L. S., and B. W. Willson. 1968. Propagation of a cerambycid borer on a meridic diet.
Canad. J. Zool., 46:1265-1266.
Papp, R. P., J. T. Kliejunas, R. S. Smith, Jr., and R. F. Scharpf. 1979. Association of Plagithmysus
bilineatus (Coleoptera: Cerambycidae) and Phytophthora cinnamomi with the decline of’ohi’a-
lehua forests on the island of Hawaii. For. Sci., 25:187-196.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 97-100
Published 17 April 1984
The Genus Acanthodoryctes Turner (Hymenoptera, Braconidae):
Redescription and New Generic Synonymy
Donald L. J. Quicke
Department of Zoology, University of Nottingham, Nottingham, NG7 2RD
England.
The classification of many tropical genera of Braconinae has been in a state of
considerable confusion for many years and is not yet fully sorted out. The Aus¬
tralian genus Acanthodoryctes Turner which belongs to the subfamily Doryctinae
(see Shenefelt and Marsh, 1976) was based on Iphiaulax morleyi Froggatt; Iphiau¬
lax itself belongs to the subfamily Braconinae. Recently, the author had the op¬
portunity of examining the type of the Indo-Australian genus Bispinariopsis Fah-
ringer which is currently placed among the Braconinae (Shenefelt, 1978) and found
it to be a doryctine agreeing well with the description of Acanthodoryctes and all
the specimens under Iphiaulax morleyi in the British Museum (Natural History)
collections; the type specimen of Acanthodoryctes has not been located. Below,
features of Acanthodoryctes are described and illustrated for the first time, and
the type species of Bispinariopsis is transferred to this genus.
Genus Acanthodoryctes Turner
Acanthodoryctes Turner, 1918:55. Type species: Iphiaulax morleyi Froggatt, lo¬
cation of type not known. Original designation.
Bispinariopsis Fahringer, 1942:37. Type species: Atanycolus tomentosus Szepli-
geti, in Hungarian Natural History Museum, Budapest. Monobasic and original
designation. NEW SYNONYMY.
Generic diagnosis. — Both sexes: Terminal flagellomere formed into a point at
the apex; basal flagellomere nearly twice as long as broad; all flagellomeres lon¬
gitudinally striate and longer than broad. Scapus and pedicellus short and shiny.
Maxillary palp six segmented. Hypoclypeal depression deep and dorsally rounded;
clypeus rugose; face in the centre smooth and raised, forming a ridge between the
antennal sockets, this central area also narrowing ventrally; face irregularly, coarse¬
ly punctured laterally. Face and clypeus with long pale hairs. Frontal depression
deep. Head cuboid without an occipital margin.
Pronotum produced anteriorly, margined anteriorly and dorsolaterally, within
these margins (on the pronope) there are a pair of mid-lateral, longitudinal ridges
which are formed into strong dorsal pointing spines, and with a short mid-posterior
ridge (Figs. 1 and 2). Middle lobe of the mesonotum smooth and shiny, notauli
shallow and wide with a few weak striae; posteriorly the notauli are margined on
the medial side, these margins enclosing a sunken area with a median carina.
Scutellar sulcus deep with approximately six crenulations. Scutellum smooth but
axillae crenulate. Metanotum with three prominent medial, longitudinal carinae.
Propodeum reticulate with a pair of more prominent carinae running anteriorly
98
PAN-PACIFIC ENTOMOLOGIST
Figures 1-4. $ Acanthodoryctes tomentosus (Szepligeti) comb. n. 1. Pronope and anterior of
mesonotum dorsal view. 2. Pronotum and anterior of mesothorax lateral view. 3. Propodeum and
first metasomal tergite dorsal view. 4. Right hind wing. Scale bar: Figures 1-3, 1 mm; Figure 4, 2
mm.
from its hind margin. Most of the propodeal sculpture is obscured by dense, long,
white, flattened hairs (Fig. 3).
Metasoma: Tergite 1 about IV 2 X longer than apically broad (Fig. 3), with a
raised central area bordered laterally by the dorsal carinae. The lateral areas are
obscured by long white hairs, the median area is generally smooth with some
longitudinal striations apically. Tergites 2 to 5 with coarse foveate sculpture and
with the hind margins densely covered with medially pointing white hairs. Tergite
2 with a very large, slightly raised medial area covering most of the tergum.
Wings: Forewing about % the length of the thorax and abdomen combined;
nervus recurrens interstitial, nervulus postfurcal, parastigma large and continuous
with the basalis. Radial cell reaching the wing margin % the way between the apex
of the stigma and the wing-tip. Second abscissa of the radius more than 2 x length
of the first. Subdiscoideus quite well developed, nearly reaching the wing margin.
First brachial cell long; second abscissa of discoideus approximately 0.6 x media.
Hindwing (Fig. 4); submediellan cell large, nervellus running distally towards wing
margin, basella postfurcal, postnervellus absent.
Legs: Fore femur broad, of equal length to tibia. Fore tibia with a longitudinal
row of pegs (“spines” or chaetobothria), tibial spur flattened and densely setose.
Middle legs with a row of weaker pegs along the tibiae. Hind femur only 3 x
longer than broad; tibia slightly longer than femur; tibial spurs setose and ap¬
proximately 0.3 x length of basitarsus; hind basitarsus without spines. All claws
unidentate, narrow with a poorly developed basal lobe.
VOLUME 60, NUMBER 2
99
Acanthodoryctes tomentosus (Szepligeti), New Combination
Acanthodoryctes tomentosus (Szepligeti), comb. nov. ( =Atanycolus tomentosus
Szepligeti; =Bispinariopsis tomentosus : Fahringer).
Since A. tomentosus, comb. nov. is the type species of Bispinariopsis, the latter
becomes a junior synonym of Acanthodoryctes.
Discussion
The initial placement of both Iphiaulax morleyi and Atanycolus tomentosus in
the Braconinae was probably largely due to the following combination of char¬
acters:
i) Occipital carina absent (apomorphous),
ii) Second cubital cell long (plesiomorphous),
iii) Postnervellus absent (apomorphous),
iv) Tergite 1 with a raised central area,
v) Hind coxae with neither teeth nor tubercles (doubtful state)
However, Acanthodoryctes shows great affinities to the Doryctinae:
i) Fore and middle tibiae with a row of pegs (apomorphous),
ii) Maxillary palp six segmented (plesiomorphous),
iii) Submediellan cell long (plesiomorphous),
iv) Crenulate axillae (? plesiomorphous),
v) Head cubicoid (plesiomorphous; both cubicoid and transverse heads are found
in the Braconinae),
vi) Tergite 1 of metasoma with well developed dorsal carinae (plesiomorphous).
A number of genera of Braconinae possess stout bristles or pegs on the fore tibiae
(e.g. Merinotus Szepligeti, Glyptomorpha Holmgren), however in these the bristles
form a large patch rather than a distinct row. The tibial pegs of the tribe Hister-
omerini also form a patch; this tribe has in the past been regarded as belonging
to both the Braconinae and the Doryctinae (Achterberg, 1976). The present author
prefers to regard the Histeromerini as doryctines on the basis of venation and the
six segmented maxillary palps. The presence of an occipital carina cannot be taken
as a diagnostic feature of the Doryctinae (Marsh, 1970) since it is also absent in
Liobracon Szepligeti, Liodoryctes Szepligeti and Binarea Brulle; its loss is a wide¬
spread apomorphous feature in the Braconidae (Tobias, 1967; Papp, 1974; Ach¬
terberg, 1976). The raised medial area on the first tergite of Acanthodoryctes is
significantly different from the area which partially characterises the Braconinae
in that it is bordered by the dorsal carinae and therefore widens towards the base
of the segment.
Acanthodoryctes appears close to Liobracon with which it shares the following
characters:
i) Occipital carina absent,
ii) Head cubicoid,
iii) Pronotum with margins enclosing a deep transverse excavation (apomor¬
phous),
iv) Converging carinae at junction of notauli (? apomorphous),
v) Hind coxae simple,
100
PAN-PACIFIC ENTOMOLOGIST
vi) Hind femora short and broad,
vii) Dense silvery hairs on side of thorax and propodeum (apomorphous).
Acanthobracon may be separated from Liobracon by the pronotal teeth, absence
of a postnervellus and by the radiellan cell not being divided by a spurious vein.
Acknowledgments
I wish to thank Dr. Jeno Papp (Budapest) for loan of the type specimen of
Atanycolus tomentosus and Mr. Tom Huddleston for allowing me access to British
Museum specimens of Iphiaulax morleyi.
Literature Cited
Achterberg, C. van. 1976. A preliminary key to the subfamilies of the Braconidae (Hymenoptera).
Tijdschr. Ent.,119(3):33—78.
Marsh, P. M. 1970. The Nearctic Doryctinae, VIII. The genera Liobracon and Pedinotus, with notes
on the definition of the subfamily (Hymenoptera, Braconidae). Proc. Ent. Soc. Wash., 72(3):
313-317.
Papp, J. 1974. A study on the systematics of Braconidae (Hymenoptera). Folia Entomol. Hung.,
27(2): 109-133.
Shenefelt, R. D. 1978. Hymenopterorum catalogus (nov. ed.). Part 15. Braconidae, 10:1459.
-, and Marsh, P. M. 1976. Hymenopterorum catalogus (nov. ed.). Part 15. Braconidae, 9:
1264-1265.
Tobias, V. I. 1967. A review of the classification, phylogeny and evolution of the family Braconidae
(Hymenoptera). Ent. Obozr., 46(3):645-669. (In Russian)
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 101-113
Published 17 April 1984
Filacus , A New Genus for Four Species of Sawflies
Previously Placed in Macrophya or Zaschizonyx
(Hymenoptera: Tenthredinidae)
David R. Smith and Gary A. P. Gibson
(DRS) Systematic Entomology Laboratory, IIBIII, Agricultural Research Ser¬
vice, USDA, % U.S. National Museum of Natural History, Washington, D.C.
20560; (GAPG) Department of Entomology, University of Alberta, Edmonton,
Alberta T6G 2E3, Canada.
Abstract.—Filacus, a new genus, is proposed for four species of sawflies related
to Macrophya Dahlbom and Zaschizonyx Ashmead and is placed in the tribe
Sciapterygini of the Tenthredininae. The included species, doanei (Rohwer), pro-
vancheri (Rohwer), pluricinctellus (Rohwer), and pluricinctus (Norton), all new
combinations, were previously considered as one variable species and were placed
in either Macrophya or Zaschizonyx. Representatives of all species occur in Cal¬
ifornia, with two also in Arizona and northern Mexico. Adults have been collected
from a great variety of flowers and plants, but the only definite larval feeding
records are on Phacelia (Hydrophyllaceae), Amsinckia (Boraginaceae), and Ra¬
nunculus (Ranunculaceae). Keys are provided for identification of adults of the
genera of Sciapterygini and for adults and larvae of Filacus. Larvae of two species,
pluricinctus and pluricinctellus, are described.
For over 30 years, a small group of sawfly species that occur in California,
Arizona, and adjacent Mexico have been treated as a single taxon, pluricincta
Norton, which was assigned either to the genus Macrophya Dahlbom or Zaschi¬
zonyx Ashmead of the subfamily Tenthredininae. Ross (1951) synonymized the
11 species existing at that time, treating them as one variable species, Macrophya
pluricincta Norton. Benson (1959) was the first to recognize that pluricincta was
not a typical Macrophya (tribe Macrophyini) and placed it in the genus Zaschi¬
zonyx of the tribe Sciapterygini. Smith (1979) followed Benson’s decision for lack
of a better placement, and Gibson (1980) also excluded the species from Macro¬
phya.
After studying many specimens of “pluricincta,” we discovered that it consisted
of four species, separable by characters of the lancet, male genitalia, hindtibial
spurs, and relatively stable color patterns. Host data and larvae, though not known
for all species, help to support our conclusion that four species exist. We also
believe that these species cannot be assigned to any described genus, and therefore,
we propose a new genus for them.
Adults of the four species separated below are rather commonly collected in
California, mostly from flowers in early spring. Plants from which adults have
been collected may not represent their true host plant, but larvae have been found
feeding on plants of the genera Phacelia, Amsinckia, and Ranunculus. Because
102
PAN-PACIFIC ENTOMOLOGIST
adults commonly visit flowers, they may play a role in pollination. Many speci¬
mens examined have parts of the body covered with pollen.
Filacus Smith and Gibson, New Genus
Type-species. —Macrophya pluricincta Norton.
Description. — Head evenly convex, without deep frontal or antennal furrows
and without antennal tubercles. Clypeus flat, anterior margin shallowly, broadly,
circularly emarginate, without acute or rounded lateral lobes (Fig. 1); labrum
rectangular with front margin truncate to slightly emarginated and depressed (Fig.
1); width of malar space slightly less than l /i diameter of front ocellus; occipital
carina present laterally, obsolete on upper lateral and dorsal margins of head; eyes
converging below, distance between eyes below slightly less than maximum eye
length; each mandible bidentate (Fig. 1). Antenna stout, length less than 1 Vi x
head width; 3rd segment nearly as long as segments 4 + 5; 1st and 2nd segments
each longer than broad. Metepimeral appendage rounded, smooth; epicnemium
present. Basal plates contiguous on meson, not separated nor with broad mem¬
branous area. Hindtibia longer than hindfemur; hindcoxa slightly enlarged but
smaller than mesepisternum in lateral view, therefore apex of hindfemur reaches
nearly to apex of abdomen; foretibial spurs slightly to distinctly curved, not
straight; hindtibial spurs less than or slightly more than apical width of hindtibia
(Figs. 2, 3); tarsal claw of female with inner tooth as long as and broader than
outer tooth, teeth not appressed to each other, that of male with inner tooth
slightly longer and broader than outer tooth and teeth appressed. Forewing with
vein 2A + 3A joined to 1A basad to center of anal cell, sometimes with very
short anal crossvein.
Adults of all species are rather stout sawflies, 6-8 mm long, black with various
yellow markings, commonly with yellow transverse stripes on the abdomen, and
with fine, uniform punctation on the head and thorax, the texture, however,
subshining and not with dull surface sculpture.
Matsuda (1957) described the head morphology of “ Macrophya pluricincta
but we cannot determine the species of Filacus to which this description refers.
Larva.— Antenna conical, 5-segmented. Clypeus with 4 setae; labrum with 6
setae; mandible with 2 setae on outer surface. Maxillary palpus 4-segmented; 2nd
segment with 1 seta; palpifer with 3 setae; stipes with 1 seta; lacinia with 12-13
short spines (Fig. 12). Labial palpus 3-segmented; prementum with 4 setae. Right
mandible with 3 ventral teeth, 3 medial teeth, and 1 broad dorsal tooth (Fig. 10);
left mandible with 3 ventral teeth, 3 dorsal teeth, and medial ridge extending
basally from outer dorsal tooth (Fig. 11). Clypeus with median depression. Each
thoracic leg with femur longer than trochanter. Abdominal segments 1-8 each
7-annulate (Fig. 9); tubercles and setae on annulets 2 and 4 and on each posts-
piracular lobe, subspiracular lobe, and surpedal lobe; 7th annulet with only short
setae; prolegs without setae. Hairs present on head and setae present on thoracic
legs, 10th tergum, and subanal area.
The larvae are typically caterpillar-like and feed externally.
Remarks.— Although species of Filacus show some affinities with members of
the tribe Macrophyini, most characters support its placement in the Sciapterygini,
where it was placed by Benson (1959). The evenly convex head, lack of furrows
on the head, lack of antennal tubercles, presence of a metepimeral appendage,
VOLUME 60, NUMBER 2
103
Figures 1-12. Filacus spp. 1. Clypeus, labrum, mandibles of F. pluricinctus. 2. Hindtibial spurs of
F. pluricinctellus. 3. Hindtibial spurs of F. pluricinctus. 4. Male genital capsule, ventral, of F. pluri¬
cinctus. 5. Male valve of F. doanei. 6. Male valve of F. pluricinctus. 7. Male valve of F. pluricinctellus.
8. Male valve of F. provancheri. 9-12. Larva of F. pluricinctellus. 9. 3rd abdominal segment, anterior
to right. 10. Right mandible, ventral. 11. Left mandible, ventral. 12. Maxilla, adl = anterodorsal lobe.
and slightly enlarged hindcoxa are characters shared with the Macrophyini, but
the following place Filacus in the Sciapterygini: Hindtibia longer than hindfemur,
hindtibial spurs shorter than or not much longer than apical width of hindtibia,
anterior margin of labrum truncate or emarginate and depressed, and occipital
104
PAN-PACIFIC ENTOMOLOGIST
carina present only laterally. Four genera are currently recognized in the Sciap-
terygini, and Filacus can be separated from them by the key presented here.
Filacus is separated from Zaschizonyx, the only other Nearctic genus of Sciap-
terygini, by the following: Distance between eyes, below less than eye length (sub¬
equal to or greater in Zaschizonyx ); basal plates contiguous on meson (separated
and with broad membranous area in Zaschizonyx ); malar space less than Vi
diameter of front ocellus (slightly less than diameter of front ocellus in Zaschi¬
zonyx :); each mandible bidentate and with single cutting margin (each mandible
with 5 or more teeth and with dorsal and ventral cutting edges in Zaschizonyx)-,
epicnemium present (absent in Zaschizonyx)-, metepimeral appendage broad and
apically rounded (narrow and acutely pointed apically in Zaschizonyx)-, and head
and thorax finely punctate (impunctate in Zaschizonyx). Deda Gibson (1980), a
genus in the Macrophyini, is superficially like Filacus and is also distributed
primarily in California; however, Deda has quadridentate mandibles, a setiferous
metepimeral appendage, an emarginate clypeus with rounded lateral lobes, a
convex labrum, and small eyes, with the distance between the eyes below greater
than the maximum eye length.
Larvae are known for two species of Filacus, but they cannot be separated from
other genera because generic characters for most North American larval Ten-
thredininae have not been determined. They are separated from larvae of Aglao-
stigma Kirby by the presence of two setae on each mandible (one in Aglaostigma),
and from Sciapteryx Stephens and Rhogogaster Konow by the lack of setae on
the prolegs (present in those two genera). Larvae of Macrophya and Tenthredo
Linnaeus may also be separated from the above three genera by the same char¬
acters, but larvae of Macrophya and Tenthredo are inadequately known, and the
larva of the single species of Zaschizonyx is unknown. Use of characters in the
above description and known host plant information may help to identify larvae
of Filacus.
The genus name is based on the abbreviation of California spelled backwards;
the gender is masculine.
Adults of the four species of Filacus may be identified by comparing the lancet
and male valves with figures, by certain color patterns, and with the help of the
key to species.
Key to Genera of the Tribe Sciapterygini
Adults
1. Antenna with furrow on outer side of segments 4-9 (antennal tubercles
high; malar space greater than diameter of front ocellus; clypeus with
central emargination and rounded lateral lobes; eyes parallel, not con¬
verging below; inner tooth of tarsal claw longer and stouter than outer
tooth) (China) . Tyloceridius Malaise
Antenna without furrow . 2
2. Eyes, scarcely converging below, far apart, lower interocular distance great¬
er than eye length; malar space usually 2 x or more diameter of front
ocellus; head and thorax densely punctate and dull (antennal tubercles
low) (Palearctic). Sciapteryx Stephens
Eyes converging below, lower interocular distance equal to or less than eye
VOLUME 60, NUMBER 2
105
length; malar space at most equal to diameter of front ocellus; head and
thorax finely punctate and subshining or impunctate . 3
3. Basal plates widely separated on meson, leaving large membranous area
(distance between eyes below, subequal to eye length; malar space slightly
less than diameter of front ocellus) (Nearctic) .... Zaschizonyx Ashmead
Basal plate contiguous on meson, without membranous area. 4
4. Clypeus emarginate for Vi or more of medial length, with narrow rounded
lateral lobes; anterior margin of labrum not emarginate or depressed
(Palearctic) . Elinor a Benson
Clypeus shallowly, circularly emarginated for less than V 6 its medial length,
without lateral lobes (Fig. 1); anterior margin of labrum emarginate and
depressed (Fig. 1) (Nearctic) . Filacus, new genus
Key to Species of Filacus
Adults
1. Female . 2
Male . 5
2. Abdominal terga with apical edges yellow dorsally and laterally; hindtibial
spurs longer than apical width of hindtibia; hindcoxa with apical V3-V2
yellow . 3
Abdomen black (if some terga have yellow apical edges, then only dorsally
and hindtibial spurs stout and about % of apical width of hindtibia);
hindtibial spurs shorter or longer than apical width of hindtibia; hindcoxa
black or with about apical x h yellow. 4
3. Serrulae of lancet each with anteriorly projecting subbasal teeth well sep¬
arated from ventrally projecting lobe (Fig. 16); each abdominal tergum
broadly black along entire basal margin dorsally and laterally, thus the
abdomen appearing transversely striped; mesepisternum always entirely
black . pluricinctus (Norton)
Serrulae of lancet each without anteriorly projecting subbasal teeth, ventral
margin flat (Fig. 14); each abdominal tergum generally broadly black
only dorsally, with lateral downturned margins yellow, thus the abdomen
with a broad, longitudinal yellow stripe on each side; mesepisternum
sometimes with small central yellow spot . doanei (Rohwer)
4. Hindcoxa entirely black and/or abdomen with dorsum of some terga yellow
apically; hindtibial spurs stout, about % apical width of hindtibia; ser¬
rulae of lancet each flattened ventrally, intersegmental setae continuous
from dorsal to ventral margin of lancet (Fig. 18) .
.. pluricinctellus (Rohwer)
Hindcoxa with apical edge yellow and/or abdomen black with yellow con¬
fined to basal plates and 9th tergum; hindtibial spurs slender, slightly
longer than apical width of hindtibia; serrulae of lancet each rounded
apically, with 1 or 2 indistinct anterior subbasal teeth, intersegmental
setae of lancet indistinct (Fig. 20) . provancheri (Rohwer)
5. Abdomen beyond basal plates entirely black . 6
Abdominal terga with apical edges yellow usually dorsally and laterally .. 7
6. Hindtibial spurs stout, about % apical width of hindtibia; hindcoxa black
106
PAN-PACIFIC ENTOMOLOGIST
or only extreme apical edge indistinctly yellow; valve elongate, with
anterodorsal lobe (Fig. 7). pluricinctellus (Rohwer)
Hindtibial spurs slender, longer than apical width of hindtibia; apical edge
of hindcoxa broadly yellow; valve ovate, without anterodorsal lobe (Fig.
8) . provancheri (Rohwer)
7. Lateral downturned margins of terga usually entirely yellow, thus the ab¬
domen with a broad longitudinal yellow stripe on each side; outer surface
of hindfemur commonly entirely yellow; valve with anterodorsal lobe
(Fig. 5). doanei (Rohwer)
Each abdominal tergum black basally and yellow apically both dorsally
and laterally, thus the abdomen with a transversely striped appearance;
outer surface of hindfemur usually with black stripe for entire length;
valve without anterodorsal lobe (Fig. 6) .. . pluricinctus (Norton)
Known Larvae
1. Body with lateral dark spots resulting in broken supraspiracular and sub-
spiracular longitudinal lines; usually with black stripe extending from
ocellus to hindmargin of head; on Phacelia . pluricinctus (Norton)
Head and body unicolorous; on Amsinckia . pluricinctellus (Rohwer)
Filacus doanei (Rohwer), New Combination
(Figs. 5, 13, 14)
Labidia doanei Rohwer, 1909a:91.
Macrophya truncata Rohwer, 1909b:331.
Female. —Antenna and head black; clypeus and labrum sometimes yellowish
anteriorly and laterally and palpus sometimes yellowish except for apical segment.
Thorax black with following yellow: Broad posterior margin of pronotum, tegula,
mesoscutellum, and sometimes small central spot on mesepisternum. Abdomen
with basal plates yellow, posterior margin black; remaining terga mostly black
basally and yellow apically on dorsum, usually solidly yellow laterally; apical
yellow margins broader toward apex of abdomen; basal 3 or 4 sterna usually black,
remaining sterna yellow; sheath black. Legs yellow with following black: Coxae
except for extreme apical margin of fore- and midcoxae and apical x h or more of
hindcoxa; 1 st segment of fore- and midtrochanters and inner surface of 1 st segment
of hindtrochanter; basal l h to Vi of fore- and midfemora, mid- and hindfemora
sometimes with inner surfaces completely black; apex of each tibia, usually only
on inner surface of fore- and midtibiae; inner surface of each tarsus black, hind-
tarsus more extensively so, sometimes apical segments of each tarsus entirely
black. Wings hyaline; veins dark brown; stigma pale brown.
Hindtibial spurs 1 x h x as long or longer than apical width of hindtibia; hind-
basitarsus equal in length to following V/i segments combined. Each serrula of
lancet with apical margin flat, with 6 or 7 coarse posterior subbasal teeth, without
anterior subbasal teeth (Fig. 14).
Male. — Coloration similar to female, except without yellow spot on mesepi¬
sternum, mesoscutellum sometimes black, and femora sometimes with less yellow.
Valve as in Figure 5, with anterodorsal lobe.
Larva.—Unknown.
Types. — Rohwer described the female of doanei and stated “Specimens from
VOLUME 60, NUMBER 2
107
Figures 13-20. Female lancets of Filacus spp. 13-14. F. doanei. 15-16. F. pluricinctus. 17-18. F.
pluricinctellus. 19-20. F. provancheri. Entire lancet on left, central portion of lancet on right. Photo¬
graphs by GAPG.
Stanford University, March 9, 1905; Palo Alto, March 31, 1893, and April 27,
1895.” We located one specimen from the type-series in the USNM labeled “9
Mar. 1905, Stn. U., Cal.,” “Macrophya doanei Roh., type 2,” “2 type No. 14270
U.S.N.M.” We designate this specimen the lectotype of doanei.
Macrophya truncata was described from 5 S and 8 2. We found 5 <5 and 3 2 in
the USNM labeled by Rohwer as cotypes, and all are labeled “Claremont Cal.,
108
PAN-PACIFIC ENTOMOLOGIST
Baker.” We designate a female lectotype with the additional labels “9 cotype No.
14269 U.S.N.M.,” “Macrophya truncata Roh., cotype 9,” and we have labeled it
so; the other 5 6 and 2 9 are paralectotypes.
Distribution.— California: Lake Co. and Bay region southward, most records
from coastal counties but occurring inland in the south; Lake Co., San Francisco
Co., San Mateo Co., Stanislaus Co., San Benito Co., Monterey Co., Tulare Co.,
San Luis Obispo Co., Santa Barbara Co., Ventura Co., Los Angeles Co., San
Bernardino Co., Riverside Co., San Diego Co. Arizona: Yuma, 7.IV. 1979. Mexico:
Baja California, Norte, 10 mi N San Vincente, 25.III. 1973. Two specimens are
labeled “Victoria, B.C., Mch. 10, ’05.” We consider these specimens mislabeled
since this locality is considerably disjunct from the most northern record of this
species in California.
Collection data .—Adults have been collected from February 17 (Riverside,
California) to May 15 (Wildwood Canyon, San Bernardino County, California),
with most records in March and April. We do not have data for reared material,
but the following collection data were found on adult specimens: Cryptantha
muricata, Cryptantha intermedia, Gilia achillaefolia, Sisyrinchium sp., Nemoph-
ila, on Nemophila menziesii, Phacelia, Umbelliferae, and ex Boraginaceae.
Remarks. — Of the four species in Filacus, adults of doanei have the most ex¬
tensive abdominal yellow. The sides of the abdomen are almost entirely yellow,
thus it appears partly black above and below with lateral, longitudinal yellow
stripes. This is also the only species in which a yellow spot may be present on
the female mesepisternum. Other distinguishing characteristics are the female
lancet and male valve (see Figs. 5-8, 13-20).
Filacus pluricinctus (Norton), New Combination
(Figs. 1, 3, 4, 6, 15, 16)
Macrophya pluricinctus Norton, 1862:118.
Macrophya multicincta Rohwer, 1909b:333.
Macrophya napensis Rohwer, 1911:410.
Female .—Antennae and head black; anterior margin of clypeus and anterior
margin of labrum sometimes yellow. Thorax black with broad posterior margin
of pronotum, tegula, and usually small or large spot on mesoscutellum yellow.
Abdomen with basal plates mostly yellow with basal margin black, rest of ab¬
domen with banded appearance, terga dorsally and laterally black on basal portion,
yellow apically, the yellow margins increasing in width toward apex of abdomen;
sterna also banded. Legs yellow with following black: Coxae except apical margins
of fore- and midcoxae and about apical Vs of hindcoxa; trochanters except most
of 2nd segment of fore- and midtrochanters and usually all of hindtrochanters;
inner surface of femora and usually basal V3 to V2 of outer surface; inner surface
of tibiae, or sometimes all surfaces at base and apex with black on hindtibia
broadest, sometimes hindtibia all yellow except for extreme base and apex; inner
surface of tarsi sometimes all blackish with part of basitarsi or all of hindbasitarsus
yellowish. Wings subhyaline; veins dark brown; stigma pale brown, especially on
ventral V2.
Hindtibial spurs 1 Vt x as long or longer than apical width of hindtibia; hind¬
basitarsus subequal in length to following 3 segments combined. Each serrula of
VOLUME 60, NUMBER 2
109
lancet with 2 anteriorly projecting subbasal teeth, well separated from ventrally
projecting lobe, and 10-11 posterior subbasal teeth (Fig. 16).
Male. — Coloration similar to female, except clypeus and mesoscutellum usually
black, basal plates black with posterior margin yellow, femora and tibiae usually
mostly yellow on outer surfaces, but hindfemur commonly with longitudinal black
stripe on outer surface, and hindtarsus mostly black. Genitalia as in Figures 4, 6;
valve without anterodorsal lobe.
Larva. — Characters as for genus. Head amber with black stripe from each ocellus
to hindmargin of head. Body with broken subspiracular and supraspiracular lon¬
gitudinal blackish stripes. (Larvae from Salinas, Monterey Co., Calif., May 11,
1978, on Phacelia sp.; in the Illinois Natural History Survey.)
Types.— The syntypes of pluricinctus Norton, a male and a female labeled “San
Mat.,” “pluricincta,” “type 13991” are in the Museum of Comparative Zoology,
Harvard University. The female is hereby designated lectotype, the male is a
paralectotype. Rohwer described multicincta from 3 2 from Claremont, California;
2 2 were found in the USNM, and one is hereby designated lectotype. The lectotype
is labeled “Claremont, Cal., Baker,” “2 cotype No. 14264, U.S.N.M.,” “Macro-
phya multicincta Roh., cotype 2.” The second specimen is a paralectotype. Rohwer
described napensis from one female; the holotype is in the USNM, and is labeled
“Napa Co., Cal.,” “Hym. slide 205,” “antenna mounted,” “2 type No. 14616
U.S.N.M.,” “Macrophya napensis Roh., type 2.”
Distribution. — California: Yolo and Napa counties southwards; Napa Co., Yolo
Co., Marin Co., Contra Costa Co., San Francisco Co., San Mateo Co., Stanislaus
Co., Mariposa Co., Monterey Co., Fresno Co., Tulare Co., Inyo Co., San Luis
Obispo Co., Santa Barbara Co., Kern Co., Los Angeles Co., San Bernardino Co.,
Riverside Co., San Diego Co., Imperial Co. Arizona: Globe, 14.IV.33; Catalina
Mts., 19.IV.1957; “A” Mtn., Tucson, 19.III.1962; Ragged Top, Silverbell Mts.,
Pima Co., 24.III.78, ex Phacelia sp.; N. side Tortolita Mts., 5.IV. 1966, on Phacelia
(?); Baboquivari Camp, Babq. Cn., 17.IV. 1966, on Hydrophyllum occidental;
Pima Co., Silverbell Bajada; N. ent. Tucson Mts., 16.III.63, on flower; Pima Co.,
Catalina Mts., 3000', Sabino Canyon Sta., 15.IV.65. Mexico: nr. La Zapopita
Valle de Trinidad, Baja Calif., 17.IV.62; Baja Calif. Norte, 7 mi SE Maneadero,
100' el., 25.III. 1973.
Collection data.— Specimens have been collected from February 24 (Agua Ca-
liente Ind. Res., Palm Canyon, Riverside Co., and Bakersfield, Kern Co., Cali¬
fornia) to May 28 (San Francisco, California), with most records in March and
April. The species has been reared from larvae feeding on Phacelia distans (San
Mateo Co., Pescadero, San Gregorio site, 6.V.71, A. R. Moldenke) and possibly
Phacelia sp. (Corona, Calif., 12.III.62, Brawner), of the family Hydrophyllaceae.
Data from labels on adult specimens are as follows: Phacelia, Phacelia distans,
Phacelia flower, coll, on Phacelia, Phacelia crenulata, on flower, Nemophila,
Amsinckia, on Hydrophyllum occidentale, ex lupine, ex Encilia farinosa, and on
Phacelia tanasetifolia. Most of the collection records are from Phacelia in Cali¬
fornia and Arizona.
Remarks. — The black and yellow transversely banded appearance of the ab¬
domen of adult specimens is similar to those of doanei and pluricinctellus, but in
doanei the lateral downturned margins of the terga are yellow, while in pluri¬
cinctellus they are black laterally. Also, adults of pluricinctellus have short apical
110
PAN-PACIFIC ENTOMOLOGIST
spurs on the hindtibia, shorter than the apical width of the tibia, whereas those
of pluricinctus are longer than the apical width of the hindtibia. The lancet of
females of pluricinctus has two distinct anterior subbasal teeth on each serrulae,
well separated from the ventrally projecting basal lobe; the lancets of females of
other species do not have such anterior subbasal teeth.
Filacus pluricinctellus (Rohwer), New Combination
(Figs. 2, 7, 17, 18)
Macrophya pluricinctellus Rohwer, 1909b:332.
Macrophya occidentalis Rohwer, 1909b:330.
Macrophya bakeri Rohwer, 1909b:331.
Macrophya melanostoma Rohwer, 1909b:333.
Female.— Antenna and head black; anterior margin of clypeus and most of
labrum sometimes yellow. Thorax black with posterior margin of pronotum,
tegula, and spot on center of mesoscutellum yellow. Abdomen black with following
yellow: Basal plates and usually apical margins of remaining terga dorsally, basal
terga may be entirely black, and amount of yellow on apical margins of terga
increases posteriorly. Legs yellow with following black: Coxae except sometimes
apical margins; trochanters; femora except basal Vi to % of outer surface; line on
inner surface of fore- and midtibiae; apices of mid- and hindtibiae; line on inner
surface of each tarsus with apical tarsal segment mostly black. Wings hyaline;
veins dark brown; outer surface of costa and subcosta and sometimes basal l A of
these yellowish; stigma entirely or with ventral V2 yellowish.
Hindtibial spurs short and broad, nearly triangular, their length equal to about
% of apical width of hindtibia (Fig. 2). Hindbasitarsus subequal in length to
following 3 tarsal segments combined. Each serrula of lancet truncate apically,
without anterior subbasal teeth and with about 8 coarse posterior subbasal teeth
(Fig. 18).
Male. — Coloration similar to that of female, but abdomen black, inner surface
of all femora with complete black stripe, mesoscutellum black, tegula partly black,
hindtarsus sometimes all black, and extreme apical margin of 1st tergum some¬
times yellowish. Valve as in Figure 7, with anterodorsal lobe.
Larva. — Characters as given for genus description. Head and body unicolorous
greenish, without black markings; only eyespot black. (Larvae from Cuyama,
California, April 12, 1956, on Amsinckia, H. T. Osborn; LaGrange, California,
April 15, 1953, Amsinckia sp., R. P. Allen; Lindcove, Tulare Co., California,
April 4, 1956, ex Amsinckia, R. P. Allen; in the Illinois Natural History Survey
and USNM.)
Types. — Rohwer described Macrophya pluricinctella from an unspecified num¬
ber of females from “Palo Alto, California; Stanford University, California; Clare¬
mont, California.” We found 3 females in the USNM labeled as cotypes, 2 from
Claremont and 1 from Palo Alto. A specimen from Claremont, labeled “Clare¬
mont, Cal., Baker,” “9 cotype No. 14267, U.S.N.M.,” “Macrophya pluricinctella
Roh., cotype 9” is hereby designated lectotype; the other two specimens are para-
lectotypes.
Macrophya bakeri was described from one male; the holotype is labeled “Clare¬
mont, Cal., Baker,” “6 type No. 14262 U.S.N.M.,” “Macrophya bakeri Roh.,
VOLUME 60, NUMBER 2
111
type 3.” Macrophya melanostoma was described from one female; the holotype
is labeled “Claremont, Cal., Baker,” “2 type No. 14263 U.S.N.M.,” “Macrophya
melanostoma Roh., type 2.” Macrophya occidentalis was described from one male;
the holotype is labeled “Claremont, Cal., Baker,” “<3 type No. 14266 U.S.N.M.,”
“Macrophya occidentalis Roh., type 6.”
Distribution. — California: Napa, Yolo, and Yuba counties southward in coastal
and central areas to San Diego Co.; Sonoma Co., Napa Co., Yolo Co., Yuba Co.,
Marin Co., Solano Co., Sacramento Co., El Dorado Co., Contra Costa Co., Ala¬
meda Co., San Francisco Co., San Mateo Co., Santa Clara Co., Stanislaus Co.,
Santa Cruz Co., Monterey Co., San Benito Co., Fresno Co., Kings Co., Tulare
Co., San Luis Obispo Co., Santa Barbara Co., Kern Co., Fos Angeles Co., San
Bernardino Co., Riverside Co., San Diego Co. Arizona: Boyce-Thompson Ar¬
boretum, 2500', Vh mi W Superior, Pinal Co., Feb. 24, 1973, on flowers.
Collection data. — Representatives of this species have been collected from Feb¬
ruary 4 (Stanford University, California) to May 19 (Alviso, California), with
most records in March and April. Adults have been reared from larvae feeding
on Amsinckia tessellata (Riverside Co., Riverside, California, March 23, 1971)
and Amsinckia sp. (Santa Barbara Co., Ventucopa, California, March 28, 1960)
of the plant family Boraginaceae. Collection records from adult label data are as
follows: on barley, Nemophila, at flowers Brassica, Brassica, Amsinckia, Am¬
sinckia douglasiana, on Sambucus, Phacelia, Ranunculus californicus, on Cali¬
fornia poppy, Ranunculus, on flowers, Mimulus, on elderberry, ex sweeping grass¬
es, ex wheat fiddleneck, ex wheat, on Amsinckia (yellow fiddleneck), Phacelia
transectifolia, and ex Boraginaceae. Most records are from Amsinckia including
a series of over 200 specimens from Mojave, California that were swept from
Amsinckia sp.
Remarks. — The combination of the black abdomen, at least black laterally in
the female, the short hindtibial spurs which are shorter than the apical width of
the hindtibia, and the lack of anterior subbasal teeth on the serrulae of the lancet
will distinguish adults of this species. In some female specimens the dorsal trans¬
verse yellow bands on the abdomen are more extensive, even slightly extending
laterally, and these may be confused with those of pluricinctus. However, females
of the latter species possess long hindtibial spurs, longer than the apical width of
the hindtibia, and well separated anterior subbasal teeth on the serrulae of the
lancet. The hindtibial spurs and genitalia should be examined for accurate iden¬
tification of pluricinctus adults.
Filacus provancheri (Rohwer), New Combination
(Figs. 8, 19, 20)
Macrophya albipes Provancher, 1895:95 (preoccupied in Macrophya by Macro¬
phya albipes (Dahlbom), 1835).
Macrophya provancheri Rohwer, 1909b:328 (new name for albipes Provancher).
Macrophya nigricornis Rohwer, 1909b:329-330.
Female. — Antenna and head black; apical margin of labrum and subapical
palpal segments may be yellowish. Thorax black with posterior margin of prono-
tum, tegula, and small or large spot on mesoscutellum yellow. Abdomen black,
basal plates with 2 broad yellow spots, last tergum yellow. Legs yellow with
112
PAN-PACIFIC ENTOMOLOGIST
following black: Coxae except for apical margins; 1 st trochanteral segment of fore-
and midlegs and inner surface of 1 st trochanteral segment of hindleg; femora
except apical Vi to l h of outer surface; line on inner surface of fore- and midtibiae;
base and apex of inner surface of hindtibia entirely; line on inner surface of each
tarsus and most of apical tarsal segment of each tarsus. Wings subhyaline; veins
dark brown, stigma pale brown.
Hindtibial spurs subequal to or slightly longer than apical width of hindtibia.
Hindbasitarsus subequal in length to following 3 Vi tarsal segments combined.
Serrulae of lancet low, rounded, with 1 or 2 indistinct anterior subbasal teeth and
9-10 coarse posterior subbasal teeth; segmental hairs indistinct, not confined to
distinct vertical rows and not extending from dorsal to ventral edges of lancet
(Fig. 20).
Male.— Coloration similar to female but apical tergum black, mesoscutellum
black, hindtibia black with yellow streak on outer surface, and hindtarsus some¬
times entirely black. Genitalia as in Fig. 8; valve without anterodorsal lobe.
Larva. — Unknown.
Types.— The lectotype of Macrophya albipes Provancher is in the Provancher
Collection, Universite Laval, Sainte-Foy, Quebec (Smith, 1975); it is from Los
Angeles. Macrophya nigricornis Rohwer was described from both sexes from
“Mountains near Claremont, California.” The lectotype, by present designation,
is a female in the USNM labeled “Mts. near Claremont, Cal., Baker,” “Macrophya
nigricornis Rohwer, cotype 9.” The male, a paralectotype, has the same data except
for the sex and an additional label “<3 cotype No. 14265 U.S.N.M.”
Distribution.— California: Mendicino, Napa, Yolo, and El Dorado counties south
in coastal and central areas to San Diego Co.; Mendocino Co., Sonoma Co., Napa
Co., Yolo Co., El Dorado Co., Solano Co., Marin Co., San Francisco Co., Contra
Costa Co., San Mateo Co., Santa Cruz Co., Santa Clara Co., Alameda Co., Sta¬
nislaus Co., Tuolumne Co., Merced Co., San Benito Co., Fresno Co., San Luis
Obispo Co., Santa Barbara Co. (including Santa Cruz Is.), Ventura Co., Los
Angeles Co., San Bernardino Co., San Diego Co.
Collection data.— Collection records are from January 18 (Redwood City, San
Mateo Co., California) to May 20 (Davis, California), with most collected in
March and April. One rearing record is from larvae feeding on Ranunculus nys-
triculus (Tuolumne Co., Buck Meadows, Mather Site, reared, 16-III-71, A. R.
Moldenke) of the family Ranunculaceae. Larvae were not found from this rearing.
Collection data from adult specimens are as follows: on Sambucus, dandelion,
swept from Ranunculus ceanothus, Ranunculus californicus, Ranunculus, Montia,
Montia perfoliata, ca. Montia under Q. lobata, coll, on Rubus, coll, on Ceanothus,
visiting miner’s lettuce, Nemophila menziesii, Rhus trilobata, ex. misc. grass. One
long series was collected on Montia perfoliata (Portulacaceae) at Strawberry Can¬
yon, California, and another series from Almaden, Quicksilver Park, a mixed-
oak woodland south of San Jose. Pertaining to the latter series, L. Bezark (pers.
comm.) stated that they were abundant very low to the ground in association with
miner’s lettuce, Montia perfoliata, “Upon closer inspection the critters were ap¬
parently stuck to the very tiny flowers by their mandibles. Leaves were torn off
to place specimens in a killing jar.”
Remarks.— Adults of provancheri are the darkest colored of the species of Fi-
lacus, the abdomen being black with yellow markings only on the basal plates
VOLUME 60, NUMBER 2
113
and the ninth tergum of the female. The male of pluricinctellus also has a black
abdomen, but the hindcoxae are all black and the hindtibial spurs are stout. The
lancet of females of provancheri is very different from those of other species,
lacking well defined rows of segmental hairs and having rounded serrulae with
both anterior and posterior subbasal teeth.
Acknowledgments
We thank the following for allowing study of specimens: P. H. Arnaud, Jr.,
California Academy of Sciences, San Francisco; R. M. Bohart, University of
California, Davis; W. Ferguson, San Jose State University, San Jose, California;
S. Frommer, University of California, Riverside; H. Goulet, Biosystematics Re¬
search Institute, Agriculture Canada, Ottawa; F. F. Hasbrouck, Arizona State
University, Tempe; C. L. Hogue and R. R. Snelling, Natural History Museum of
Los Angeles County, Los Angeles, California; J. D. Lattin, Oregon State Univer¬
sity, Corvallis; W. W. Middlekauff, University of California, Berkeley; H. K.
Townes, American Entomological Institute, Ann Arbor, Michigan; M. S. Was-
bauer and L. Bezark, State of California Department of Food and Agriculture,
Sacramento; D. Webb, State Natural History Survey Division, Champaign, Illi¬
nois; and F. G. Werner, University of Arizona, Tucson. Other specimens are in
the National Museum of Natural History, Washington, D.C. (USNM).
Literature Cited
Benson, R. B. 1959. Tribes of the Tenthredininae and a new European genus (Hymenoptera: Ten-
thredinidae). Proc. R. Entomol. Soc. Lond. (B), 28:121-127.
Gibson, G. A. P. 1980. A revision of the genus Macrophya Dahlbom (Hymenoptera: Symphyta,
Tenthredinidae) of North America. Mem. Entomol. Soc. Can., No. 114, 167 pp.
-. 1980. Deda, a new genus of sawfiies from western North America (Hymenoptera: Symphyta,
Tenthredinidae). Can. Entomol., 112:249-258.
Matsuda, R. 1957. Morphology of the head of a sawfly, Macrophya pluricincta Norton (Hymenoptera:
Tenthredinidae). J. Kans. Entomol. Soc., 30:99-108.
Norton, E. 1862. Catalogue of American species of Tenthredo, as arranged by Hartig. Proc. Boston
Soc. Nat. Hist., 9:116-122.
Provancher, L. 1895. Les dernieres descriptions de l’Abbe Provancher. Nat. Can. (Que.), 22:79-80,
95-97.
Rohwer, S. A. 1909a. Notes on Tenthredinoidea, with descriptions of new species (Paper III). Can.
Entomol., 41:88-92.
-. 1909b. Notes on Tenthredinoidea, with descriptions of new species. Paper VI. Western
Macrophya. Can. Entomol., 41:327-334.
-. 1911. New sawfiies in the collections of the United States National Museum. Proc. U.S.
Natl. Mus., 41:377-411.
Ross, H. H. 1951. Symphyta. Pp. 4-89 in C. F. W. Muesebeck et al. (eds.), Hymenoptera of America
north of Mexico, synoptic catalog. U.S. Dep. Agric., Agric. Monogr. 2.
Smith, D. R. 1975. The sawfly types of Abbe Leon Provancher (Hymenoptera: Symphyta). Nat.
Can. (Que.), 102:293-304.
-. 1979. Symphyta. Pp. 3-137 in K. V. Krombein et al. (eds.), Catalog of Hymenoptera in
America north of Mexico, Vol. 1. Smithsonian Institution Press, Washington, D.C.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 114-118
Published 17 April 1984
Description of a New Purpuricenine Genus, Linsleyella
(Coleoptera: Cerambycidae)
John A. Chemsak
University of California, Berkeley, California 94720.
The genus Deltaspis was proposed by Audinet-Serville (1834) for the species
auromarginata described at the same time. Since then, the genus has been a
dumping ground for species of uncertain affinities. One of these, D. virens Bates
(1885) is clearly unrelated to auromarginata and is being placed into a new genus
described below. Included are two apparently undescribed species.
This study was conducted in conjunction with a National Science Foundation
sponsored study on North American Cerambycidae (Grant DEB-8015523). Most
of the material utilized, in the Essig Museum of Entomology, Berkeley, was col¬
lected with financial support from the National Science Foundation, the Univer¬
sity Research Expeditions Program, University of California, Berkeley, and A. &
M. Michelbacher. Additional specimens were loaned by Cornell University, Field
Museum of Natural History, Chicago, Los Angeles Natural History Museum,
University of Kansas and Texas A&M University. Marlin Rice made his Texas
material available. Carolyn Tibbetts prepared the illustration.
Linsleyella, New Genus
Form moderate sized, somewhat depressed; integument metallic. Head small,
front short; palpi unequal, truncate at apices; mandibles short, angulate at apical
one-third; eyes large, deeply emarginate, finely faceted, not embracing antennal
insertions; antennae slender in males, outer segments enlarging in females, 11-
segmented, longer than body in males, shorter than body in females. Pronotum
flattened, sides broadly rounded; disk abruptly delimited at sides, not callused
dorsally; prosternum shallowly impressed, intercoxal process not extending be¬
yond margins of coxae, apex not expanded, coxal cavities wide open behind,
rounded externally; mesosternum with intercoxal process gradually arcuate, lying
below margins of coxae; metepistemum moderately broad, sides subparallel, slightly
tapering posteriorly. Elytra subparallel, slightly tapering posteriorly. Elytra sub¬
parallel; apices rounded to subtruncate. Scutellum triangular, as broad as long,
apex acute. Legs slender; hind femora arcuate, not extending to apices of elytra;
tarsi slender, third segment cleft almost to base.
Type species.—Deltaspis virens Bates.
This genus is distinctive by the metallic coloration of the integument, apically
expanding antennae of the females, and by the rounded flattened, pronotum.
It is a pleasure to dedicate this genus to my friend and colleague, E. G. Linsley.
Three species are presently known.
Key to the Species of Linsleyella
1. Elytra with punctures coarse to moderately coarse, pubescence moderately
dense, erect, long and short
2
VOLUME 60, NUMBER 2
115
Elytra with punctures fine, very dense, pubescence very dense, subde¬
pressed. Length, 11-15 mm. Texas. ricei
2. Elytra with punctures very coarse, contiguous. Hind tibiae with suberect
black setae. Antennae of females with tenth segment as long as broad.
Length, 8-12 mm. Mexico . virens
Elytra with punctures moderately coarse, subconfluent. Hind tibiae lacking
black setae. Antennae of females with tenth segment distinctly longer
than broad. Length, 10-12 mm. Mexico . michelbacheri
Lins ley ella ricei, New Species
(Fig. 1)
Male. — Form moderate sized; integument dark metallic greenish, appendages
black; pubescence whitish, dense, subdepressed and erect. Head with front con-
fluently punctate, moderately densely clothed with long erect hairs; vertex con-
fluently punctate, long, erect hairs numerous; antennae extending beyond elytra
by one or two segments, basal segments rather densely clothed with short, de¬
pressed, black setae, outer segments opaque, very finely, densely pubescent, first
segment shorter than third, fourth slightly longer than first, shorter than third,
eleventh segment appendiculate. Pronotum broader than long, disk convex, mod¬
erately coarsely, subconfluently punctate, with a vague median callus behind
middle; pubescence long, erect; prosternum coarsely punctate on posterior one-
half, pubescence long, dense; meso- and metasternum finely, densely punctate,
densely clothed with long, subdepressed pubescence. Elytra about 2Vi times as
long as broad; punctures fine, very dense, finer toward apex; pubescence dense,
subdepressed, longer and erect over basal one half; apices subtruncate. Legs with
femora densely pubescent; tibiae with numerous short, black setae along inner
margins. Abdomen finely, densely punctate, densely pubescent; last sternite sub¬
truncate at apex. Length, 11-15 mm.
Female. — Form similar. Antennae shorter than body, outer segments slightly
enlarged, longer than broad. Abdomen with last sternite broadly truncate at apex.
Length, 13 mm.
Holotype male (California Academy of Sciences) from White River Lake, Cros¬
by Co., Texas, 19 September 1980, on Heterotheca (M. E. Rice). Sixteen paratypes
(10 males, 6 females), same locality, 17 September 1980 (M. Dicetto), 6 October
1980 (Rice), 12 October 1980 (Rice), 2-3 October 1982 (Rice, E. G. Riley, R.
Turnbow). One male paratype from 4 mi E Loco Hills, Otero Co., New Mexico,
4 October 1970 (O’Brien, Richardson).
The fine, very dense punctation and dense pubescence of the elytra will readily
separate this species.
This species is named in honor of Marlin E. Rice for his aid and cooperation
in the studies on Cerambycidae.
Linsleyella virens (Bates), New Combination
Deltaspis (?) virens Bates, 1885, Biol. Centrali-Americana, Coleop., 5:323,
Male. — Yorm moderate sized; integument bright metallic greenish, blueish, or
bronze, appendages metallic; pubescence rather sparse, mostly pale, erect, short
and long. Head confluently punctate, sparsely clothed with long, erect hairs; an-
116
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Linsleyella ricei Chemsak, female.
tennae slightly longer than body, basal segments shining, with numerous, short,
black, subdepressed setae, outer segments minutely pubescent, first segment short¬
er than third, fourth subequal to first, eleventh appendiculate. Pronotum broader
than long, somewhat depressed; disk vaguely impressed at sides, coarsely, closely
punctate; pubescence sparse, long, erect; prosternum deeply punctate over pos¬
terior two-thirds, moderately densely pubescent; meso- and metastemum densely
punctate at sides, sparsely at middle, sides densely pubescent. Elytra less than 2Vi
VOLUME 60, NUMBER 2
117
times as long as broad; punctures very coarse, contiguous, becoming slightly finer
toward apex; pubescence moderately sparse, erect, long and short, longer toward
base; apices rounded. Legs with femora moderately sparsely clothed with long,
flying hairs; tibiae with short, dark setae. Abdomen sparsely, indistinctly punctate,
moderately densely pubescent; last sternite shallowly emarginate-truncate. Length,
8-11 mm.
Female. — Form similar. Antennae shorter than body, outer segments expanded,
about as broad as long. Abdomen with last sternite broadly truncate at apex.
Length, 9-12 mm.
Type locality.— Mexico.
Range.— Vera Cruz, Puebla, Oaxaca.
Flight period.— June to September.
Adults are commonly encountered on flowers of Selloa glutinosa.
The very coarse punctures, strongly shining integument and rather sparse pu¬
bescence will separate this species.
Material examined. — Mexico: 79 males, 26 females, 7 km SE Morelos Canada,
Puebla, 4-10 July 1974, 4 October 1975, 20 September 1977, on flowers of Selloa
glutinosa (J. Chemsak, J. Powell, E. G. Linsley, A. & M. Michelbacher); 1 male,
1 female, Tehuacan, Puebla, 6 July 1941 (H. Dybas), 2 July 1955 (R. & J. Selander);
1 male, 1 female, 6 mi SW Techuacan, 9 August 1980, 8 July 1981 (Schaffner et
al.); 2 males, Tecamachalco, Puebla, 24 June 1951 (H. Evans), 2 July 1953 (U.
Kansas Mex. Exped.); 2 females, 2 mi E Tecamachalco, 27 June 1964 (C. Johnson);
1 male, 19 mi NW Calcaloapan, Puebla, 30 July 1963 (J. Doyen); 1 male, 10 mi
N Miltepec, Oaxaca, 4 August 1976 (Peigler et al.); 1 male, Jalapa Rd., K 341,
14 October 1945.
Linsleyella michelbacheri, New Species
Male.— Form moderate sized; integument metallic dark blueish; pubescence
pale, moderately dense, erect, long and short. Head confluently punctate, mod¬
erately densely clothed with long, erect pubescence; antennae extending about two
segments beyond elytra, basal segments with a few black setae, outer segments
opaque, minutely pubescent, first segment shorter than third, third slightly longer
than fourth, eleventh appendiculate. Pronotum broader than long; punctures coarse,
subconfluent; pubescence rather sparse, long, erect; prosternum with a delimited
area of deep punctures extending to sides but not across middle, pubescence
moderately dense, erect; meso- and metasternum finely, densely punctate at sides,
sparsely at middle, sides densely pubescent. Elytra about 2Vi times as long as
broad; punctures moderately coarse, subconfluent, becoming finer toward apex;
pubescence rather sparse, short and suberect and long and erect, erect hairs more
numerous toward base; apices rounded. Legs with femora rather sparsely clothed
with long flying hairs; tibiae clothed with pale pubescence. Abdomen rather sparse¬
ly punctate, pubescence long, suberect; last sternite emarginate at apex. Length,
11-12 mm.
Female.— Form similar. Antennae shorter than body, outer segments slightly
enlarged, longer than broad. Abdomen with last sternite broadly truncate. Length,
10-12 mm.
Holotype male, allotype (California Academy of Sciences) and 3 paratypes (2
males, 1 female) from 16 miles SE Saltillo, 6500 ft, Coahuila, Mexico, 25 Sep-
118
PAN-PACIFIC ENTOMOLOGIST
tember 1976 (J. Chemsak, J. Powell, A. & M. Michelbacher). Additional paratypes
as follows: 4 females, 17 males SE Saltillo, 14 September 1977 (J. Chemsak, A.
& M. Michelbacher); 1 female, 1 mi N El Tunal, Coahuila, Mexico, 11 August
1977 (E. I. Schlinger); 2 males, 87 mi N. Zacatecas, Zacatecas, Mexico, 7 August
1968 (J. Bigelow, M. Cazier). An additional female from Fraile, Coahuila, Mexico,
10-18 July 1941 (K. L. Retherford) is assignable to this species.
This species differs from L. virens by the less coarsely punctate elytra, absence
of dark setae on the tibiae and by the more slender distal segments of the female
antennae.
I am pleased to name this species for A. E. Michelbacher for his friendship and
assistance over the years.
Literature Cited
Audinet-Serville, J. G. 1834. Nouvelle classification de la famille des Longicornes. Ann. Soc. Ento-
mol. France, (1)1:5-110.
Bates, H. W. 1879-1885. Longicomia. Biologia Centrali Americana, Coleoptera, 5:1-436, 17 pis.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 119-121
Published 1 7 April 1984
Synonymy in California Channel Island Epeolini Bees
(Hymenoptera: Anthophoridae)
Richard W. Rust
Department of Biology, University of Nevada, Reno, Nevada 89557.
A recent review of the endemic insect fauna of the California Channel Islands
(Miller, 1984) indicated 16 taxa of bees (Apoidea). Earlier revisions of bee taxa
containing California Channel Island endemics have shown that the island en¬
demics are synonyms of mainland taxa: Anthidium Grigarick and Stange, 1968;
Dioxys Hurd, 1958; Melissodes LaBerge, 1961; Andrena LeBerge and Ribble,
1975, Ribble, 1968; Bombus Milliron, 1971; Coelioxys Mitchell, 1973; Agaposte-
mon Roberts, 1972; Osmia, Halictus Sandhouse, 1927, 1941; Hylaeus Snelling,
1970; and Colletes, Bombus Stephen, 1954, 1957. Thus, the number of both
endemic species and subspecies has been greatly reduced. Some island endemics
are recognized as good taxa, such as Perdita layiae layiae Cockerell (Timberlake,
1956).
Here I present two additional synonyms of California Channel Island Epeolini
bees in the genera Epeolus and Triepeolus.
Epeolus minimus (Robertson, 1902)
Epeolus eastwoodae Cockerell was described from San Miguel Island, Cuyler’s
Cove, near the shore, July 27, 1927 [sic 37] (Cockerell) (Cockerell 1937). Ex¬
amination of the holotype (California Academy of Sciences #4651) and compar¬
ison with the holotype of Epeolus minimus (Robertson) (Illinois Natural History
Survey #22953) indicates the two are conspecific. An additional 11 male and 7
female specimens from San Miguel Island were also compared to both types.
There was some difference in the overall size of the specimens, but no major
differences. E. minimus is a widespread species from the midwestern United
States: Illinois and Wisconsin, westward to California (Hurd, 1979). In Cockerell’s
original description of E. eastwoodae he mentions that the specimen was taken
at flower of Malacothrix implicata Eastwood; however there is no plant label on
the specimen, nor is there the information that the bee was taken at Cuyler’s Cove
and the date is July 27, 1937 not 1927 as indicated in the publication.
E. minimus females are recognized by the broad pygidium bordered apically
with a carina, mid-apical edge being truncate; pygidium is covered with red-brown
hairs; pseudopygidium is a semi-circular patch of short, silver hairs offset pos¬
teriorly by white pubescence. Male pygidium is one-and-one-half times as wide
as long and the carina is broadly U-shaped and covered with silver-white pubes¬
cence. Apical white hair bands of tergites 2-5 have narrow medium black stripes,
band of the sixth is complete.
Epeolus minimus is found on five of the California Channel Islands (San Miguel,
Santa Rosa, Santa Cruz, Anacapa and Santa Catalina) and is represented by over
100 specimens.
120
PAN-PACIFIC ENTOMOLOGIST
Triepeolus heterurus (Cockerell & Sandhouse, 1924)
Epeolus piscatoris Cockerell was described from Santa Catalina Island, Fish¬
erman’s Cove, at flower of Sinapis, June 9, 1933, W. P. Cockerell (Cockerell,
1939). Examination of the holotype (CAS #6652) and comparison with the ho-
lotype of Triepeolus heterurus (Cockerell & Sandhouse) (CAS #1611) showed the
two to be conspecific. An additional 4 females from Santa Catalina Island were
also similar to T. heterurus and an additional 11 specimens of T. heterurus from
other California Channel Islands are also similar.
Triepeolus heterurus is a very distinctive species. Female pygidium is covered
with red-brown hairs parted along the dorsoventral mid-line giving the appearance
of two distinct patches. White apical hair bands on tergites 2-5 are complete.
Male pygidium is twice as long as wide, carina is acutely rounded apically and
the segment is covered with black pubescence. White apical hair bands of tergites
2-5 are uninterrupted, sixth has a narrow median black stripe.
Trieleolus heterurus is also known from Santa Cruz Island and Anacapa Island.
Acknowledgments
Drs. W. E. LaBerge and W. J. Pulawski made type specimens available for
study and the late Dr. P. D. Hurd supplied information on type specimen location.
Dr. G. E. Bohart helped with the original identification of T. heterurus and R. L.
Brumley aided with E. minimus identification.
Literature Cited
Cockerell, T. D. A. 1937. Bees from San Miguel Island, California. Pan-Pacific Entomol., 13:148-
157.
-. 1939. The bees of the southern California Islands. Proc. California Acad. Sci. Fourth Series,
23:427-436.
-, and G. A. Sandhouse. 1924. Parasitic bees (Epeolinae and Melectinae) in the collection of
the California Academy of Sciences. Proc. California Acad. Sci. Fourth Series, 13:305-324.
Grigarick, A. A., and L. A. Stange. 1968. The pollen-collecting bees of the Anthidiini of California
(Hymenoptera: Megachilidae). Bull. California Insect Surv., 9:1-113.
Hurd, P. D. 1958. American bees of the genus Dioxys Lepeletier and Serville (Hymenoptera: Me¬
gachilidae). Univ. California Publ. Entomol., 14:275-302.
-. 1979. Superfamily Apoidea. Catalog of Hymenoptera in America north of Mexico. Smith¬
sonian Institution Press, Washington, D.C.
LaBerge, W. E. 1961. A revision of the bees of the genus Melissodes in North and Central America.
Part III (Hymenoptera: Apidae). Univ. Kansas Sci. Bull., 42:283-663.
-, and D. W. Ribble. 1975. A revision of the bees of the genus Andrena of the western
hemisphere. Part VII. Subgenus Euandrena. Trans. Amer. Entomol. Soc., 101:371-446.
Miller, S. E. 1984. Endemic insects of the California Channel Islands. In A. S. Menke and S. E.
Miller (eds.), Proc. Symp. Insects California Channel Islands. Santa Barbara Museum of Natural
History, Santa Barbara, California.
Milliron, H. E. 1971. A monograph of the western hemisphere bumblebees (Hymenoptera: Apidae;
Bombinae). I. The genera Bombus and Megabombus subgenus Bombias. Mem. Entomol. Soc.
Canada, 82:1-80.
Mitchell, T. B. 1973. A subgeneric revision of the bees of the genus Coelioxys of the western
hemisphere. North Carolina State Univ. Press, Raleigh, North Carolina.
Ribble, D. W. 1968. Revisions of two subgenera of Andrena: Micrandrena Ashmead and Der andrena,
new subgenus (Hymenoptera: Apoidea). Bull. Univ. Nebraska State Mus., 8:237-349.
Roberts, R. B. 1972. Revision of the bee genus Agapostemon (Hymenoptera: Halictidae). Univ.
Kansas Sci. Bull., 49:437-590.
VOLUME 60, NUMBER 2
121
Sandhouse, G. A. 1924. Bees of the genus Osmia in the collection of the California Academy of
Sciences. Proc. California Acad. Sci. Fourth Series, 13:341-372.
-. 1941. The American bees of the subgenus Halictus. Entomol. Amer. New Series, 21:23-39.
Snelling, R. R. 1970. Studies on North American bees of the genus Hylaeus. 5. The subgenera
Hylaeus, s. str. and Paraprosopis (Hymenoptera: Colletidae). Los Angeles Co. Mus. Contr. Sci.,
180:1-59.
Stephen, W. P. 1954. A revision of the bee genus Colletes in America north of Mexico (Hymenoptera:
Colletidae). Univ. Kansas Sci. Bull., 36:149-527.
-. 1957. Bumble bees of western America (Hymenoptera: Apoidea). Oregon St. Coll. Agric.
Exp. Sta. Tech. Bull., 40:1-63.
Timberlake, P. H. 1956. A revisional study of the bees of the genus Perdita F. Smith, with special
reference to the fauna of the Pacific Coast (Hymenoptera: Apoidea). Part II. Univ. California
Publ. Entomol., 11:247-350.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 122-150
Published 17 April 1984
Review of the Japanese Species of the Genus Gabrius
Stephens (Coleoptera, Staphylinidae)
(121st Contribution to the Knowledge of Staphylinidae)
Ales Smetana
Biosystematics Research Institute, Agriculture Canada, K.W. Neatby Bldg.,
Ottawa, Ontario K1A 0C6, Canada.
Abstract. — The Japanese species of the genus Gabrius Stephens are reviewed.
Fourteen species are recognized, nine of them are described as new. All species
are described and illustrated, and a key is presented to aid in the identification
of the species. All available records and biological data for the species are provided.
Lectotypes are designated for G. egens (Sharp), G. kuanshanensis (Bernhauer)
and G. conicus (Bernhauer).
Gabrius conicus (Bernhauer) is placed in synonymy with G. sharpianus (Cam¬
eron).
Only a few species of the genus Gabrius Stephens are at present known from
Japan. Sharp (1874, 1889) described three species of Philonthus Stephens which
were later assigned to Gabrius (Scheerpeltz, 1933), however, two of these, G.
prolatus (Sharp) and G. spadiceus (Sharp), in fact belong to Philonthus. Cameron
(1930) described G. septempunctatus and erected the name G. sharpianus for the
specimens originally mentioned by Sharp (1889:40) as Philonthus nigritulus
(Gravenhorst) and Tottenham (1940) redescribed the latter species under the name
G. demarcatus. Bernhauer (1939a, 1939b) added three more species ( G. subde-
pressus, G. conicus and G. unzenensis). No additional species of Gabrius have
since been described from Japan. The list of species of Gabrius previously recorded
from Japan includes therefore only the following names, chronologically arranged:
Gabrius egens (Sharp, 1874)
Gabrius sharpianus (Cameron, 1930)
Gabrius septempunctatus (Cameron, 1930)
Gabrius conicus (Bernhauer, 1939)
Gabrius subdepressus (Bernhauer, 1939)
Gabrius unzenensis (Bernhauer, 1939)
Gabrius demarcatus Tottenham, 1940
The names G. conicus and G. demarcatus are synonyms of G. sharpianus. The
original material of G. septempunctatus was not found in the Cameron collection
in the British Museum (Natural History), London (G. Kibby, pers. comm.), and
the species cannot therefore be interpreted at this moment. It could be conspecihc
with any of the species with 7 punctures in the dorsal rows on the pronotum
treated in this paper.
For many years I have known that the above list is inadequate since the genus
VOLUME 60, NUMBER 2
123
Gabrius is, in fact, very rich in species in Japan. Several species described as new
in this paper have been in my collection for years and were supposed to be
described in a joint paper with Mr. Y. Watanabe, Tokyo. For several reasons,
however, the paper was never published although a preliminary study of the type
material of the involved species was carried out by myself.
In 1980 I had the opportunity to collect in Japan for almost five weeks, together
with my friend Ivan Lobl from the Museum d’Histoire Naturelle in Geneve
(Switzerland) and my wife Zdena. During this time a large collection of the genus
Gabrius, containing several hundred specimens and many new species, was made
and the availability of this material prompted me to return to my previously
abandoned project.
In the following paper all new species are described and a review of the species
of Gabrius known to occur in Japan at present is given. A key to species is included.
There is no doubt that the species included in this paper represent only a portion
of the existing fauna; it is hoped that the paper will stimulate the interest in this
genus in Japan and bring our knowledge to a level comparable to that of the west
Palaearctic fauna.
The material studied is deposited in the Canadian National Collection, Ottawa,
Canada, and in the Museum d’Histoire Naturelle in Geneve, Switzerland (MHNG).
Key to Species
1(12). Dorsal rows of punctures on pronotum each with seven punctures.
(Occasionally one puncture missing or one additional puncture
present unilaterally.)
2(3). Paramere of aedoeagus with short, obtusely triangular median pro¬
jection (Figs. 7, 8). Length 6.1-7.3 mm .3. G. kobayashii, n. sp.
3(2). Paramere of aedoeagus at least slightly emarginate in middle of apical
margin, frequently divided into two variably long branches apically
(Figs. 5, 14, 17, 20, 23).
4(5). Median lobe of aedoeagus in front of paramere abruptly, angulately
narrowed into obtusely lanceolate apical portion (Fig. 13). Length
5.8-7.0 mm .5. G. yamanei, n. sp.
5(4). Median lobe of aedoeagus in front of paramere evenly and gradually
narrowed into more or less conical apical portion (Figs. 4, 16, 19,
22 ).
6(9). Paramere of aedoeagus apically divided into two very long branches;
branches longer than basal portion of paramere (basal apodemes
not included) (Figs. 5, 20).
7(8). Branches of paramere of aedoeagus each with inner margin obliquely
truncate apically, truncate portion with numerous sensory tuber¬
cles; branches separated by rather wide arc mediobasally (Fig. 5).
Length 6.0-7.5 mm.2. G. subdepressus (Berhnauer)
8(7). Branches of paramere of aedoeagus each with inner margin simple,
not obliquely truncate apically, with only a few sensory tubercles
near apex; branches separated by rather acute angle mediobasally
(Fig. 20). Length 5.0-5. 5 mm .7. G. damon, n. sp.
9(6). Paramere of aedoeagus apically divided into two short branches;
branches shorter than basal portion of paramere (basal apodemes
not included) (Figs. 17, 23).
124
PAN-PACIFIC ENTOMOLOGIST
10 ( 11 ).
11 ( 10 ).
12 ( 1 ).
13(14).
14(13).
15(16).
16(15).
17(18).
19(20).
20(19).
21 ( 22 ).
22 ( 21 ).
23(24).
24(23).
25(26).
26(25).
Branches of paramere of aedoeagus separated mediobasally by deep,
V-shaped emargination (Fig. 17). Length 5.9 mm . G. io, n. sp.
Branches of paramere of aedoeagus separated mediobasally by less
deep, U-shaped emargination (Fig. 23). Length 5.0-5.7 mm, ....
.8. G. abas, n. sp.
Dorsal rows of punctures on pronotum each with six punctures. (Rare¬
ly one additional puncture present unilaterally.) 1
Apical portion of median lobe of aedoeagus strongly asymmetrical,
with apex hook-like curved (Fig. 41). Length 4.0-4.8 mm .
.14. G. nepos, n. sp.
Apical portion of median lobe of aedoeagus symmetrical, apex never
hook-like curved (Figs. 1, 10, 25, 28, 31, 34, 38).
Paramere of aedoeagus very wide, not constricted in middle, sensory
tubercles situated partially away from apical margin, forming char¬
acteristic pattern (Fig. 11). Length 6.0-6.5 mm ... 4. G. egens (Sharp)
Paramere of aedoeagus more or less narrow, distinctly constricted in
middle, sensory tubercles situated strictly on apical margin (Figs.
2, 26, 29, 32, 36, 39).
Apical margin of paramere of aedoeagus with small median emar¬
gination and one deeper and much wider emargination on each
side (Fig. 39). Length 4.2-4.9 mm . 13. G. ophion, n. sp.
Dorsal face of apical portion of median lobe distinctly swollen (lateral
view) (Fig. 35). Length 5.2-6.2 mm . . 12. G. sharpianus (Cameron)
Dorsal face of apical portion of median lobe not swollen (lateral view).
Antenna long, 3rd segment longer than 2nd, 4th segment distinctly,
5th segment slightly longer than wide. Aedoeagus and paramere as
in Figures 1,2. Length 5.5-6.2 mm ... 1. G. unzenensis (Bernhauer)
Antenna moderately long, 3rd segment as long as 2nd, 4th segment
at most slightly longer than wide, 5th segment about as long as
wide. Aedoeagi and parameres different (Figs. 25, 26, 28, 29, 31,
32).
Median lobe of aedoeagus strongly, almost conically narrowed api-
cally, with subacute apex (Fig. 25). Length 4.0-4.5 mm .
.9. G. demades, n. sp.
Median lobe of aedoeagus less strongly and not conically narrowed
apically, with rounded apex (Figs. 28, 31).
Sternite of male pygidium shallowly emarginate apically (Fig. 30).
Notch in middle of apical margin of paramere shallower and less
wide; sensory tubercles on apical margin of paramere larger and
more crowded together; median lobe shorter and stouter (Figs. 28,
29). Length 4.8-5.2 mm . 10. G. philo, n. sp.
Sternite of male pygidium deeply emarginate apically (Fig. 33). Notch
in middle of apical margin of paramere deeper and wider, sensory
tubercles on apical margin of paramere smaller and not crowded
together; median lobe longer and narrower (Figs. 31, 32). Length
4.7-5.1 mm . 11. G. kuanshanensis (Bernhauer)
1 Rare specimens with an atypical number of punctures in the dorsal rows of pronotum should be
run through both halves of couplet 1(12).
VOLUME 60, NUMBER 2
125
1. Gabrius unzenensis (Bernhauer, 1939)
(Figs. 1-3)
Philonthus unzenensis Bernhauer, 1939b: 153.
Gabrius unzenensis; Smetana, 1973:129.
Material studied.— Japan: Honshu: Mie Pref., 20.VI.61, Y. Watanabe (2).
For the description and other information about this species see Smetana 1973:
129-130.
See Figures 1-3 for details of the sternite of the male pygidium and the ae-
doeagus. Note especially the arrangement of sensory tubercles on the paramere
of the aedoeagus.
Distribution. —Gabrius unzenensis is at present known only from Japan (Kyushu
and Honshu).
Discussion. —In my previous paper (Smetana, 1973:129) the year 1938 is given
as the publication date of this species. The date “December 1938” is printed at
the bottom of the page with the description of G. unzenensis ; however, the wrapper
of the “III/IV Heft” of the Entomologisches Nachrichtenblatt (pages 113-176)
gives May 1939 (“Ausgegeben im Mai 1939”) as the publication date. The correct
year for this species is therefore 1939 and not 1938.
2. Gabrius subdepressus (Bernhauer, 1939)
(Figs. 4-6)
Philonthus subdepressus Bernhauer, 1939a:97.
Gabrius subdepressus ; Smetana, 1973:130.
Material studied. Japan: Honshu: Asama Plateau, Nidoage, 22. VIII.60, Y. Wa¬
tanabe (4); Nikko Nat. Park, Ryuzu, 1400 m, 16.VII.80, A. & Z. Smetana (4);
Nikko Nat. Park, below Konsei Pass (W side), 1500-1600 m, 15.VII.80, A. & Z.
Smetana (4).—Additional localities can be found in Smetana 1973:130.
Piceous black to black, elytra usually somewhat paler; palpi and legs brownish
yellow, inner portions of middle and hind tibiae darkened. Head feebly longer
than wide (index 24:22), indistinctly widened posteriorly, temporal angles rather
rounded. Eyes rather small and flat, tempora more than twice as long as length
of eyes in dorsal view (index 21:9). Antenna in general similar to that of G. astutus
Er., outer segments hardly to slightly transverse. Pronotum rather narrow, longer
than wide (index 29:22), usually slightly narrowed posteriorly, dorsal rows of
punctures often somewhat irregular, each with seven (exceptionally eight) punc¬
tures. Elytra flat and moderately long, at sides slightly longer than pronotum
(index 32:29), with fine and dense punctation. Punctation of abdomen finer than
that of elytra, seventh (fifth visible) tergite with whitish apical seam.
Male. — Sternite of pygidium deeply and widely emarginate, small triangular
median area before emargination flattened and smooth; lateroapical margins of
emargination bearing rather strong and long setae becoming gradually shorter
both medially and laterally (Fig. 6). Aedoeagus rather large and elongate, apical
portion of median lobe long, strongly and evenly narrowed apically (Fig. 4).
Branches of paramere each with inner margin obliquely truncate apically, truncate
portion with numerous sensory tubercles; branches separated by rather wide arc
mediobasally (Fig. 5).
126
PAN-PACIFIC ENTOMOLOGIST
Figures 1-3. Gabrius unzenensis. 1. Aedoeagus. 2. Underside of paramere. 3. Apical portion of
stemite of male pygidium.
VOLUME 60, NUMBER 2
127
Figures 4-6. Gabrius subdepressus. 4. Aedoeagus. 5. Underside of paramere. 6. Apical portion of
stemite of male pygidium.
128
PAN-PACIFIC ENTOMOLOGIST
Length 6.0-7.5 mm.
Distribution. —Gabrius subdepressus is at present known from Manchuria in
northeastern China and from Japan (Honshu and Kyushu).
Discussion. — Gabrius subdepressus is similar to G. astutus, however, it differs,
in addition to the differences on the aedoeagus, by the smaller eyes, the different
number of punctures in the dorsal rows on the pronotum (there are only six
punctures in each row in G. astutus) and by the different male secondary sexual
characters.
In my previous paper (Smetana, 1973:130) the year 1938 is given as the pub¬
lication date of this species. The date”September 1938” is printed at the bottom
of the page with the description of G. subdepressus ; however, the wrapper of the
“II. Heft” of the Entomologisches Nachrichtenblatt (pages 65-112) gives February
1939 (“Ausgegeben im Feber 1939”) as the publication date. The correct year for
this species is therefore 1939 and not 1938.
See Smetana (1973:130) for the information on the type series of this species.
3. Gabrius kobayashii , New Species
(Figs. 7-9)
Holotype (male) and allotype (female). — “Japan Gumma Pr. 7 km E Usui Pass
850 m 24.VII.80 A. & Z. Smetana.” In the Canadian National Collection, Ottawa
(CNCNo. 16178).
Paratypes. — Same data as holotype (9); Gumma Pref., 6 km E Usui Pass, 750
m, 20.VII.80, A. & Z. Smetana (8); Gumma Pr., 5 km E Usui Pass, 900 m,
25.VII.80, A. & Z. Smetana (3); Gumma Pr., Usui Bypass, 700 m, 20.VII.80 A.
& Z. Smetana (1); Mie Pr. Hirakura, 21.VI.61, Y. Watanabe (5).
Externally very similar to G. subdepressus, but differing as follows: eyes slightly
larger, index length of eyes to length of temples in dorsal view 11:19 (same index
in G. subdepressus 9:21), punctures on temporal portion of head in general less
numerous, microsculpture on head and pronotum finer, with less appreciable
tendency to form irregular meshes on frontal portion of head and on middle
anterior portion of pronotum.
Male. — Sternite of pygidium broadly emarginate, emargination slightly bisin-
uate, its apical margin bearing numerous long and rather strong setae becoming
gradually shorter towards midline (Fig. 9). Aedoeagus rather large, with bulbus
and basal portion of median lobe wide, median lobe anteriorly suddenly narrowed
into long and narrow, basally attenuate and apically rounded apical portion.
Paramere wide, entirely covering middle portion of median lobe, its apical margin
broadly arcuate with small, obtusely triangular median projection. Sensory tu¬
bercles on underside of paramere arranged along apical margin, including median
projection, and becoming distinctly less dense toward middle (Figs. 7, 8).
Length 6.1-7.3 mm.
Distribution. —Gabrius kobayashii is at present known only from one locality
each in Mie and Gumma Prefectures in Central Honshu.
Bionomics. — The specimens from near Usui Pass were collected by sifting fallen
leaves and other debris along creeks.
Discussion. — The external characters, distinguishing this species from G. sub¬
depressus are inconspicuous, however, the differences in the size of the eyes and
in the development of the microsculpture on the head and pronotum seem to be
VOLUME 60, NUMBER 2 129
Figures 7-9. Gabrius kobayashii. 7. Aedoeagus. 8. Underside of paramere. 9. Apical portion of
sternite of male pygidium.
130
PAN-PACIFIC ENTOMOLOGIST
constant. Gabrius kobayashii is also very similar in all external characters to G.
yamanei, but differs from the latter by the same set of characters as from G.
subdepressus.
Etymology. —Patronymic, named in honour of Dr. F. Kobayashi, Forestry and
Forest Products Research Institute, Ushiku, Ibaraki, in appreciation of his enor¬
mous assistance during our field work in Japan in the summer of 1980.
4. Gabrius egens (Sharp, 1874)
(Figs. 10-12)
Philonthus egens Sharp, 1874:44.
Gabrius egens\ Smetana, 1960:303.
Material studied.—Japan: Honshu: Kyoto Pref., Kyoto, 7.-8.VIII.80, Cl. Be-
suchet (MHNG)2.
For description and other information about this species see Smetana (1960:
303-304).
See Figures 10-12 for details of the sternite of the male pygidium and the
aedoeagus. Note especially the arrangement of sensory tubercles on the paramere
of the aedoeagus.
Type material . — I was able to study three out of four specimens of the original
series (see Sharp, 1874:45) deposited in the British Museum (Natural History),
London. The specimens are labelled: Spec. No. 1: “Japan. G. Lewis.’’/“Sharp
Coll. 1905-313.’’/“Philonthus egens Type D.S.” Spec. No. 2: “Japan. G. Lewis
1910-230”/“Philonthus egens mihi D.S.” Spec. No. 3: “Japan. G. Lewis.’’/“Ja¬
pan” (oval yellow label)/“Sharp Coll. 1905-313.” The specimen No. 1 bearing
the label “Philonthus egens Type D.S.” was dissected and the aedoeagus and the
pygidium were mounted in Canada balsam. It is hereby designated as the lectotype
of G. egens ; the label “Lectotype Philonthus egens Sharp A. Smetana des. 1981”
has been attached to this specimen.
Bionomics. —No details are known about the habitat requirements of this species.
Distribution. — Gabrius egens is at present known only from Hyogo (Sharp, 1874:
45) and Kyoto Prefectures (Honshu).
Discussion.— The aedoeagus of G. egens is in general appearance similar to that
of G. yamanei, however, it is distinctive mainly by the differently shaped apical
portion of the paramere with the differently arranged sensory tubercles (Figs. 11,
14).
The specimen subsequently designated as lectotype was received in damaged
condition with almost no antennae (only six outer segments of right antenna
present), with extensive damage to the legs and with the apical portion of the
aedoeagus missing. The quite characteristic paramere, however, is present. The
apical portion of the aedoeagus on Figure 10 was reconstructed from my previous
illustration (Smetana, 1960:304).
5. Gabrius yamanei, New Species
(Figs. 13-15)
Holotype (male) and allotype (female). — “Japan Gumma Pr. 5 km E Usui Pass
900 m 25.VII.80 A. & Z. Smetana.” In the Canadian National Collection, Ottawa
(CNCNo. 17180).
VOLUME 60, NUMBER 2
131
Figures 10-12. Gabrius egens. 10. Aedoeagus. 11. Underside of paramere. 12. Apical portion of
stemite of male pygidium.
132 PAN-PACIFIC ENTOMOLOGIST
Figures 13-15. Gabrius yamanei. 13. Aedoeagus. 14. Underside of paramere. 15. Apical portion
of stemite of male pygidium.
VOLUME 60, NUMBER 2
133
Paratypes. — Same data as holotype (41); Gumma Pref., sous Usui Pass, 400 m,
25.VII.80,1. Lobl (MHNG) 1; Gumma Pref., 6 km E Usui Pass, 750 m, 20.VII.80,
A. & Z. Smetana (47); Gumma Pref., sous Usui Pass, 850 m, 24.VII.80, I. Lobl
(MHNG) 25; Gumma Pref., 7 km E Usui Pass, 850 m, 24.VII.80, A. & Z. Smetana
(29); Gumma Pref., sous Usui Pass, 850 m, 24.VII.80, I. Lobl (MHNG) 5; Oku-
tama near Tokyo, 29.IV.61, K. Mizusawa (2).
Externally extremely similar to G subdepressus and differing only in characters
on aedoeagus and in having slightly different male secondary sexual characters.
Male. — Stemite of pygidium deeply, triangularly emarginate, small triangular
median area before emargination flattened and smooth; lateroapical margins of
emargination bearing rather strong and long setae becoming gradually shorter
both medially and laterally (Fig. 15). Aedoeagus rather large and in general similar
to that of G. kobayashii, however, with median portion of median lobe narrower,
with apical portion somewhat wider and with differently shaped paramere. Par-
amere anteriorly very broadly, arcuately emarginate, with small and narrow me¬
dian notch in middle of apical margin. Sensory tubercles on underside of paramere
arranged densely along apical margin, one isolated tubercle on each lateral margin
situated distinctly below apical row of tubercles (Figs. 13, 14).
Length 5.8-7.0 mm.
Distribution. — Gabrius yamanei is at present known from around Usui Pass
near Karuizawa in Gumma Prefecture and from near Tokyo (Central Honshu).
Bionomics.— All specimens from near Usui Pass were collected by sifting wet
fallen leaves and other debris along creeks, sometimes together with specimens
of G. kobayashii.
Discussion.— Except for the smaller average size, G. yamanei does not appre¬
ciably differ externally from G. subdepressus. Also the emargination of the sternite
of male pygidium is almost identical in both species, however, the strong setae
in the emargination are longer and denser in G. yamanei. For a comparison with
G. kobayashii and G. egens see the discussion under the respective species.
Etymology. — Patronymic, named in honour of Dr. A. Yamane, Forestry and
Forest Products Research Institute, Ushiku, Ibaraki, in appreciation of his great
help during our held work in Japan in the summer of 1980.
6. Gabrius io> New Species
(Figs. 16-18)
Holotype (male).—“ Japan Kyoto Pr. Seryo-toge, 500-700 m, 6.VIII. 80 A. &
Z. Smetana.” Deposited in the Canadian National Collection, Ottawa (CNC No.
17181).
Paratypes.— Kyoto Pref., Seryo-Toge, 13 km N Kyoto, 500-600 m, 6.VIII. 80,
I. Lobl (MHNG) 2.
Externally extremely similar to G. subdepressus and differing only in characters
on the aedoeagus and in having slightly different male secondary sexual characters.
Male. — Stemite of pygidium deeply, triangularly emarginate, small triangular
median area before emargination flattened and smooth; lateroapical margins of
emargination bearing rather strong and long setae becoming gradually shorter
both medially and laterally (Fig. 18). Aedoeagus very similar in general shape to
that of G. subdepressus, however, differing in shape of paramere. Paramere with
branches shorter and wider, shorter than basal portion of paramere and separated
134
PAN-PACIFIC ENTOMOLOGIST
Figures 16-18. Gabrius io. 16. Aedoeagus. 17. Underside of paramere. 18. Apical portion of stemite
of male pygidium.
VOLUME 60, NUMBER 2
135
mediobasally by deep V-shaped emargination, with sensory tubercles more nu¬
merous (Figs. 16, 17).
Length 5.9 mm.
Distribution. — Gabrius io is at present known only from the type locality in the
Kyoto Prefecture (Honshu).
Bionomics. — The holotype was taken by sifting a small pile of rotting grass.
Discussion. —The holotype of this species is slightly smaller than most specimens
of G. subdepressus, however, this is probably insignificant; the range of size in G.
io can be expected to be comparable to that of G. subdepressus.
The male secondary sexual characters on the pygidium of G. io are very similar
to those of G. subdepressus, however, the emargination is narrower and the setae
on lateroapical margins of the emargination are shorter and less strong in G. io.
Etymology. — The specific name is the Latin interjection expressing joy.
7. Gabrius damort, New Species
(Figs. 19-21)
Holotype (male) and allotype (female). — “Tachiyzawa - Vill. (Yamagata Pref.)
Japan 30. VII.60 Coll. Y. Watanabe.” In the Canadian National Collection, Ottawa
(CNCNo. 17182).
Paratypes. — Same data as holotype (1); same data as holotype but date 2. VIII.60
( 1 ).
Externally very similar to G. subdepressus and differing only by its smaller size,
by the characters on the aedoeagus and in having slightly different male secondary
sexual characters.
Male. — Sternite of pygidium rather deeply triangularly emarginate, small tri¬
angular median area before emargination flattened and smooth; lateroapical mar¬
gins of emargination bearing rather strong and long setae becoming gradually
shorter both medially and laterally (Fig. 21). Aedoeagus very similar in general
shape to that of G. subdepressus, however, in general smaller and less robust.
Paramere narrower and more elongate, branches separated by rather acute angle
mediobasally, each with inner margin simple, not obliquely truncate apically and
with only a few sensory tubercles near apex (Figs. 19, 20).
Length 5.0-5.5 mm.
Distribution. — Gabrius damon is at present known only from the type locality
in the Yamagata Prefecture (northern Honshu).
Bionomics. —No details are known about the habitat requirements of this species.
Discussion. — The male secondary sexual characters on the pygidium of G. da¬
mon are very similar to those of G. subdepressus, however, the emargination is
narrower and less deep in G. damon. It resembles that of G. io, however, in the
latter the setae on lateroapical margins of the emargination are more numerous.
Etymology.— The specific name is the name of a Pythagorean celebrated on
account of the friendship between him and Phintias.
8. Gabrius abas, New Species
(Figs. 22-24)
Holotype (male) and allotype (female). — “Japan Gumma Pr. 5 km E Usui Pass
900 m 25.VII.80 A. & Z. Smetana.” In the Canadian National Collection, Ottawa
(CNC No. 17183).
136
PAN-PACIFIC ENTOMOLOGIST
Figures 19-21. Gabrius damon. 19. Aedoeagus. 20. Underside of paramere. 21. Apical portion of
sternite of male pygidium.
VOLUME 60, NUMBER 2
137
Figures 22-24. Gabrius abas. 22. Aedoeagus. 23. Underside of paramere. 24. Apical portion of
sternite of male pygidium.
138
PAN-PACIFIC ENTOMOLOGIST
Paratypes. — Same data as holotype (6); Gumma Pr., 6 km E Usui Pass, 750 m,
20.VII.80 A. & Z. Smetana (2); Gumma Pr., Usui Bypass, 700 m, 20.VII.81, A.
& Z. Smetana (1).
Externally very similar to G. subdepressus and differing only by its smaller size,
by the characters on the aedoeagus and in having slightly different male secondary
sexual characters.
Male. — Sternite of pygidium rather deeply triangularly emarginate, small tri¬
angular area before emargination flattened and smooth; lateroapical margins of
emargination bearing rather strong and long setae becoming gradually shorter
both medially and laterally (Fig. 24). Aedoeagus very similar in general shape to
that of G. io, however, different in some details in shape of median lobe and
paramere; branches of paramere separated mediobasally by moderately deep,
U-shaped emargination; shorter than basal portion of paramere and each with
sensory tubercles arranged in a similar way as those of G. io (Figs. 22, 23).
Length 5.0-5.7 mm.
Distribution. — Gabrius abas is at present known only from around Usui Pass
near Karuizawa in Gumma Prefecture (Central Honshu).
Bionomics. — All specimens of the original series were taken by sifting fallen
leaves and other debris along creeks.
Discussion.— The male secondary sexual characters on the pygidium of G. abas
are almost identical to those of G. damon, however, the setae on the lateroapical
margins of the emargination are in general shorter in G. abas.
One paratype in the original series of G. abas has only six punctures in one of
the dorsal rows on pronotum.
Etymology.— The specific name is the name of the twelfth king of Argos, son
of Lyncaeus and Hypermnestra, grandson of Danaus, father of Acrisius and grand¬
father of Perseus.
9. Gabrius demades, New Species
(Figs. 25-27)
Holotype (male) and allotype (female). — “Japan Toyama Pr. Tateyama Mts.
Bijodaira 1000 m 28.VII.80 A. & Z. Smetana.” In the Canadian National Col¬
lection (CNCNo. 17184).
Paratypes. — Same data as holotype (4); Nikko Nat. Park, below Konsei Pass
(W side), 1500-1600 m, 15.VII.80, A. & Z. Smetana (2); Toyama Pr., Arimine,
Kaminikawa, 1100 m, 29.VII.80, A. & Z. Smetana (1); Ehime Pr., Ishizuchi Nat.
Park, Mt. Ishizuchi, 12.VIII.80, Cl. Besuchet (MHNG) (4).
General habitus of a species from Nigritulus-group. Elytra and to lesser extent
also head and pronotum with slight metallic lustre. Antennae brunneous with two
basal segments testaceous or entirely testaceous, legs rufotestaceous with inner
portions of metatibiae occasionally darkened. Eyes rather small, index length of
eyes to length of temples in dorsal view equals 7:14. Dorsal rows of pronotum
with six punctures each, occasionally one additional puncture present unilaterally.
Male.— Sternite of pygidium moderately deeply triangularly emarginate, small
triangular median area before emargination flattened and smooth; lateroapical
margins of emargination bearing fairly strong but not long setae becoming grad¬
ually finer and shorter medially (Fig. 27). Aedoeagus elongate with apical portion
of median lobe evenly and almost conically tapered toward fairly acute apex.
VOLUME 60, NUMBER 2
139
Figures 25-27. Gabrius demades. 25. Aedoeagus. 26. Underside of paramere. 27. Apical portion
of stemite of male pygidium.
140
PAN-PACIFIC ENTOMOLOGIST
Paramere rather short, with two stout and very short branches apically, separated
from each other by almost semicircular notch; sensory tubercles on underside of
paramere arranged along apical margin of each branch (Figs. 25, 26).
Length 4.0-4.5 mm.
Distribution. — Gabrius demades is as present known from a few localities in the
Nikko National Park, in the mountains of the Toyama Prefecture (Honshu), and
in the Ishizuchi National Park (Shikoku).
Bionomics.— The specimens of the original series were taken by sifting wet leaf
litter and moss along creeks (Nikko N.P. and Kaminikawa) and by sifting debris
around bases of old trees in an undisturbed deciduous forest (Bijodaira).
Discussion. — Except for one specimen with seven punctures in one of the dorsal
rows on pronotum, all remaining specimens have six punctures in these rows.
Etymology. — The specific name is the name of the famous Athenian rhetorician
Demades, a contemporary of Demosthenes.
10. Gabrius philo, New Species
(Figs. 28-30)
Holotype (male) and allotype (female). — “Japan Nikko N.P. Ryuzu 16.VII.80
1400 m, A. & Z. Smetana.” In the Canadian National Collection, Ottawa (CNC
No. 17185).
Paratypes.—Same data as holotype (1); Nagano Pr., Shiga, 1500 m, 23.VII.80,
A. & Z. Smetana (1); Nagano Pr., J. E. Kogen N. Park, Shiga, 1500 m, 23.VII.80,
I. Lobl (MHNG) 14; Toyama Pr., Arimine, Kaminikawa, 1100 m, 29.VII.80, A.
& Z. Smetana (2); same I. Lobl (MHNG) 2; Kyoto Pref., Seryo-Toge, 13 km N
Kyoto, 500-600 m, 6.VIII.80, I. Lobl (MHNG) 2, Ehime Pr., via Mt. Ishizuchi,
1000 m, 14.VII.80, I. Lobl (MHNG) 4.
Externally very similar to G. demades but differing by slightly larger size, by
slightly larger eyes (index length of eyes to length of temples in dorsal view equals
7:18), by the characters on the aedoeagus and by the different male secondary
sexual characters.
Male.— Sternite of pygidium shallowly, almost arcuately emarginate apically,
small triangular median area before emargination flattened and smooth; margins
of emargination without accumulation of strong setae (Fig. 30). Aedoeagus, in¬
cluding paramere, very similar to that of G. demades, however, in general larger
with apical portion of median lobe more robust, not conically narrowed anteriorly
and with obtusely rounded apex (Fig. 28); paramere as in Figure 29.
Length 4.8-5.2 mm.
Distribution. — Gabrius philo is at present known from the Nikko National Park,
from the Nagano and Toyama Prefectures (Honshu), and from the Ehime Pre¬
fecture (Shikoku).
Bionomics. — The specimens of the type series were collected by sifting wet moss
and debris around seepages in an old hardwood forest (Nikko N.P.), in debris
along a creek (Shiga) and by sifting wet moss and various debris at a small creek
(Kaminikawa).
Discussion.— The paramere of G. philo is very similar to that of G. kuansha-
nensis; however, in G. philo the notch in the middle of the apical margin of the
paramere is shallower and less wide, and the sensory tubercles on the apical margin
of the paramere are larger and more crowded (Figs. 29, 32).
VOLUME 60, NUMBER 2
141
Figures 28-30. Gabrius philo. 28. Aedoeagus. 29. Underside of paramere. 30. Apical portion of
stemite of male pygidium.
142
PAN-PACIFIC ENTOMOLOGIST
Etymology.— The specific name is the name of an academic philosopher of
Athens, teacher of Cicero.
11. Gabrius kuanshanensis (Bernhauer, 1914)
(Figs. 31-33)
Philonthus kuanshanensis Bernhauer, 1914:66.
Gabrius kuanshanensis', Smetana, 1973:132.
Gabrius kuanshanensis', Coiffait, 1974:33, Fig. 12G, H, I; 79.
Material studied. — See Smetana, 1973:132. For the description and other in¬
formation about this species see Smetana, 1973:132-133 and Coiffait, 1974:33,
79-80.
See Figures 31-33 for details of the sternite of the male pygidium and the
aedoeagus. Note particularly the arrangement of the sensory tubercles on the
paramere of the aedoeagus.
Distribution. —Gabrius kuanshanensis is at present known only from “Chi-
Kuan-Shan” (=Jiguanshan?) and from Chinkiang (=Chenjiang?) in the Peoples
Republic of China. It has not yet been found in Japan, however, it may occur
there.
Discussion. —Gabrius kuanshanensis is, in all characters, including the aedoea¬
gus, very similar to G. sharpianus; however, the latter differs by having a swelling
on the dorsal face of the apical portion of the median lobe of the aedoeagus. As
a matter of fact, the seven specimens in the Bernhauer collection under the name
P. kuanshanensis described in detail by myself (Smetana, 1973:132) and men¬
tioned also by Coiffait (1974:79) belong to two different species. Of the three males
from “Chinkiang,” the aedoeagi of two of them agree with drawings of the ae¬
doeagus of G. kuanshanensis given both by myself (Smetana, 1973:131, Fig. 6)
and by Coiffait (1974:33, Fig. 12G, H, I). However, I recently dissected the third
male and found that the aedoeagus agrees with that of G. sharpianus.
Gabrius kuanshanensis is also similar to G. philo\ however, the latter differs by
having the sternite of the male pygidium shallowly emarginate and by some details
in the shape of the median lobe and of the paramere (see Figs. 29, 32, and the
discussion under G. philo).
I erroneously considered the only original specimen of G. kuanshanensis in the
Bernhauer collection (see Smetana, 1973:132) as the holotype of this species. The
specimen cannot be in fact considered as the holotype (see Bernhauer, 1914:67).
It is therefore hereby designated as the lectotype; the label “Lectotype Philonthus
kuanshanensis Bernh. Smetana des. 1981” has been attached to it.
12. Gabrius sharpianus (Cameron, 1930)
(Figs. 34-37)
Philonthus sharpianus Cameron, 1930:207.
Philonthus conicus Bernhauer, 1939:99 (n. syn.).
Gabrius demarcatus Tottenham, 1940:68.
Gabrius sharpianus', Smetana, 1960:305.
Material studied. — Japan: Gifu Prefi, 8 km SW Gero, Hwy 257,450 m, 31 .VII.80,
A. & Z. Smetana (16); Gifu Prefi, 9 km S Gero, Hwy 41, 475 m, 31.VII.80, A.
VOLUME 60, NUMBER 2
143
Figures 31-33. Gabrius kuanshanensis. 31. Aedoeagus. 32. Underside of paramere. 33. Apical
portion of sternite of male pygidium.
144
PAN-PACIFIC ENTOMOLOGIST
Figures 34-37. Gabrius sharpianus. 34. Aedoeagus. 35. Apical portion of median lobe in lateral
view. 36. Underside of paramere. 37. Apical portion of sternite of male pygidium.
VOLUME 60, NUMBER 2
145
& Z. Smetana (3); Gifu Pref., 8 km SE Osaka, 750 m, 1.VIII.80,1. Lobl (MHNG)
1; Gumma Pref., 5 km E Usui Pass, 900 m, 25.VII.80, A. & Z. Smetana (4);
Gumma Pref., 6 km E Usui Pass, 750 m, 20.VII.80, A. & Z. Smetana (3); same,
l. Lobl (MHNG) 2; Gumma Pref., 7 km E Usui Pass, 850 m, 24.VII.80, A. & Z.
Smetana (5); same, I.Lobl (MHNG) 3; Gumma Pref., 4 km SW Tsumagoi, 1050
m, 18.VII.80, A. & Z. Smetana (3); same, I. Lobl (MHNG) 4; Nagano Pref., Shiga,
1500 m, 23.VII.80, A. & Z. Smetana (1); Kyoto Pref., Kyoto, 18-20.VIII.80, Cl.
Besuchet (MHNG) 15; Kyoto Pref., Kyoto, Mt. Hiei, 800 m, 2.VIII.80, Cl. Be-
suchet (MHNG) 5; Kyoto Pref., Seryo-Toge, 13 km N Kyoto, 500-600 m, 6.VIII.80,
Cl. Besuchet (MHNG) 2; Nara Pref., Nara, 7, 10 and 11.VIII.80, A. & Z. Smetana
(24); same, 27-31.VII.80, Cl. Besuchet (MHNG) 12; Toyama Pref., 10 km SE
Unazuki Hot Spa, 400 m, 27.VII.80, A. & Z. Smetana (2). Peoples Republic of
China: “Nordwestl. China Chinkiang Col. Reitter” (1). Additional localities in
Japan can be found in Smetana, 1960:305.
For the description and other information about this species, including the
synonymy and type material of G. sharpianus and G. demarcatus, see Smetana,
1960:305-306.
See Figures 34-37 for details of the sternite of the male pygidium and the
aedoeagus. Note especially the arrangement of sensory tubercles on the paramere
of the aedoeagus.
Type material of G. conicus.— The Bernhauer collection in the Field Museum
of Natural History, Chicago, contains two conspecific females under the name of
Philonthus conicus. They are labelled: Spec. No. 1. “Tsushima Japan”/“conicus
Brnh. Typ.”/“conicus Bernh. Typus”/“Chicago NHMus M. Bernhauer Collec¬
tion.” Spec. No. 2: “Nordwestl. China Chinkiang Col. Reitter”/“conicus Bernh.
Cotypus”/“Chicago NHMus M. Bernhauer Collection.” Specimen No. 1 is hereby
designated as the lectotype of G. conicus; the label “Lectotype Philonthus conicus
Bernh. Smetana des. 1981” has been attached to this specimen. Both specimens
are considered indistinguishable from specimens of G. sharpianus and the name
G. conicus should be considered as a synonym of G. sharpianus; my corresponding
determination label was attached to each of the two original specimens.
Bionomics. — Most specimens collected in Japan in 1980 were taken by sifting
wet moss, fallen leaves and other debris, including old flood debris, along creeks;
some were also taken by sifting fallen leaves and other floor debris on wet areas
in an old mixed forest (Nara).
Distribution. —Gabrius sharpianus is known from “Chinkiang” in the Peoples
Republic of China and from numerous localities in Japan on Honshu (Gifu,
Gumma, Nagano, Nara and Toyama Prefectures) and on Kyushu (Nagasaki Pre¬
fecture).
Discussion. — Gabrius sharpianus is in all characters, including the aedoeagus,
very similar to G. kuanshanensis; however, the aedoeagus of the latter differs, in
addition to some minor differences, particularly by the absence of the globular
swelling of the dorsal face of the apex of the median lobe which is quite char¬
acteristic of G. sharpianus.
The specimen from “Chinkiang” mentioned above is one of the seven specimens
originally deposited in the Berhnauer collection under the name P. kuanshanensis
(see also the discussion under G. kuanshanensis).
146
PAN-PACIFIC ENTOMOLOGIST
13 . Gabrius ophion, New Species
(Figs. 38-40)
Holotype (male) and allotype (female). — “Japan Gumma Pr. 7 km E Usui Pass
850 m, 24.VII.80 A. & Z. Smetana.” In the Canadian National Collection, Ottawa
(CNCNo. 17186).
Paratypes. — Gifu Pr., 8 km SW Gero, Hwy. 257, 450 m, 31.VII.80, A. & Z.
Smetana (1); Nikko Nat. Park, Lake Chuzenjiko, Chisan-Shukuhakusho 1300 m,
15.VII.80, A. & Z. Smetana (2); Kyoto Pr., Seryo-Toge, 13 km N Kyoto, 500-
600 m, 6.VIII.80, Cl. Besuchet (MHNG) 2; Kuril Islands, Kunashir Island, Men-
deleevo, 9.IX.72, Pototskaya (1).
Externally very similar to G. demades and differing only by slightly larger eyes
(index length of eye to length of temple in dorsal view equals 10:17), by characters
on the aedoeagus and in having different male secondary sexual characters.
Male. — Sternite of pygidium with wide and rather shallow, obtusely traingular
emargination, apical margin with numerous setae gradually becoming shorter and
weaker towards middle (Fig. 40). Aedoeagus very long, median lobe very elongate
but relatively wide, gradually narrowed anteriorly, apically rather suddenly nar¬
rowed into obtuse apex. Paramere with apical margin with small median emar¬
gination and one deeper and much wider emargination on each side; sensory
tubercles on underside of paramere arranged along apical margin, leaving small
median emargination free (Figs. 38, 39).
Length 4.2-4.9 mm.
Distribution. —Gabrius ophion is at present known from a few localities in the
Nikko National Park, in the Gifu, Gumma and Kyoto Prefectures (Honshu), and
from one locality on the Kunashir Island in the Kuril Islands.
Bionomics. — Most specimens of the original series were collected in wet leaf
litter and other debris (including old flood-debris) along creeks; some were taken
by sifting a pile of decaying grass (Nikko N.P.). The specimen from the Kunashir
Island was reared from a pupa found on a dead Abies spec. tree.
Discussion. — The species is quite distinctive by the characteristic shape of the
paramere.
One paratype from Japan has one additional puncture in one of the dorsal rows
on the pronotum.
The specimen from the Kunashir Island is quite teneral.
Etymology. — The specific name is the name of the Centaur Ophion who was
the father of Amycus.
14 . Gabrius nepos, New Species
(Figs. 41-43)
Holotype (male) and allotype (female). — “Japan Gumma Pr. Usui Bypass 700
m 20.VII.80, A. & Z. Smetana.” In the Canadian National Collection, Ottawa
(CNCNo. 17187).
Paratypes. — Same data as holotype (5); Gifu Pr., 8 km SW Gero, Hwy 257,
450 m, 31.VII.80, A. & Z. Smetana (3); Gifu Pr., 9 km S Gero, Hwy 41, 475 m,
31.VII.80, A. & Z. Smetana (3); Gifu Pr., 8 km SE Osaka, 750 m, 1.VIII.80, A.
& Z. Smetana (1); Toyama Pr., Arimine, Kaminikawa, 1100 m, 29.VII.80, A. &
VOLUME 60, NUMBER 2
147
Figures 38-40. Gabrius ophion. 38. Aedoeagus. 39. Underside of paramere. 40. Apical portion of
stemite of male pygidium.
148
PAN-PACIFIC ENTOMOLOGIST
Figures 41-43. Gabrius nepos. 41. Aedoeagus. 42. Underside of paramere. 43. Apical portion of
stemite of male pygidium.
VOLUME 60, NUMBER 2
149
Z. Smetana (2); Mie Pr., Hirakura, 20. or 21.VI.61, Y. Watanabe (4); Ehime Pr.,
Ishizuchi N. Park, Omogo, 12.VIII.80, Cl. Besuchet (MHNG) 3.
Externally very similar to G. demades and differing only in characters on the
aedoeagus and in having different male secondary sexual characters.
Male. — Sternite of pygidium with rather narrow and not deep, obtusely arcuate
emargination, small triangular median area before emargination flattened and
smooth; with group of setae at each side of emargination (Fig. 43). Aedoeagus
with median lobe asymmetrical, apical portion of median lobe fairly narrow,
distinctly hook-like curved and obtuse apically. Paramere strongly widened an¬
teriorly, anterior margin broadly and shallowly arcuately emarginate, sensory
tubercles on underside of paramere situated at apical margin and gradually dis¬
appearing towards middle (Figs. 41-42).
Length 4.0-4.8 mm.
Distribution. — Gabrius nepos is at present known from several localities in Gifu,
Gumma and Toyama Prefectures (Honshu), and one locality in Ehime Prefecture
(Shikoku).
Bionomics.— The specimens of the original series were taken by sifting wet
moss, fallen leaves and other debris along creeks, those from around Gero were
sifted from old flood debris at a creek.
Discussion. — The species is quite distinctive by the characteristic shape of the
paramere and by the assymmetrical, hook-like curved apical portion of the median
lobe.
One paratype has one additional puncture in one of the dorsal rows on the
pronotum.
Etymology.— The specific name is the name of a Roman historian, the friend
of Cicero, Atticus and Catullus.
Acknowledgments
The original specimens of some species described by Bernhauer and by Sharp
have been made available to me through the kindness of Dr. L. Watrous, Field
Museum of Natural History, Chicago and Mr. G. Kibby, British Museum (Natural
History), London. Their assistance is gratefully acknowledged.
I thank Mr. Y. Watanabe, Tokyo, for allowing me to study, many years ago,
the Japanese species of Gabrius and letting me keep some specimens in my
collection. I also thank my colleagues, Drs. E.C. Becker and J.M. Campbell in the
Coleoptera Unit of the Biosystematics Research Institute for their suggestions and
criticisms of the manuscript, and Mr. G. Sato for carefully finishing all the drawings
in this paper.
Literature Cited
Bernhauer, M. 1914. Neue Staphyliniden der palaarktischen Fauna. Col. Rdsch., 3:65-68.
-. 1939a. Zur Staphylinidenfauna von China u. Japan. (10. Beitrag). Ent. Nachrbl., 12:97-109.
-. 1939b. Zur Staphylinidenfauna von China u. Japan (11. Beitrag). Ent. Nachrbl., 12:145-
158.
Cameron, M. 1930. New species of Staphylinidae from Japan. Ent. Mon. Mag., 67:181-208.
Coiffait, H. 1974. Coleopteres Staphylinidae de la region palearctique occidentale II. Sous famille
Staphylininae, Tribus Philonthini et Staphylinini. Suppl. a la Nouv. Rev. Ent., Tome IV, fas.
4. Toulouse, 593 pp.
150
PAN-PACIFIC ENTOMOLOGIST
Scheerpeltz, O. 1933. Staphylinidae VII. In Junk-Schenkling: coleopterorum catalogus, VI, pars 129.
Berlin, pp. 989-1500.
Sharp, D. 1874. The Staphylinidae of Japan. Trans. Ent. Soc. London: 1-103.
-. 1889. The Staphylinidae of Japan. Ann. Mag. Nat. Hist. Ser., 6(3):28-44, 108-121, 249-
267, 319-334, 406-419, 463-476.
Smetana, A. 1960. Monographische Bearbeitung der palaarktischen Arten der Gattung Gabrius Curt,
aus der nigritulus-Gruppe (Coleoptera). Dtsch. ent. Z., N.F., 7:295-356.
-. 1973. Ueber einige von Dr. M. Bemhauer beschriebene Gabrius-Arten (Coleoptera:Staphy-
linidae) (90. Beitrag zur Kenntnis der Staphyliniden). Nouv. Rev. Ent., 3:125-136.
Tottenham, C.E. 1940. A new species of Gabrius Stephens (Col., Staphylinidae) from Japan. Ent.
mon. Mag., 76:68-69.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 151-154
Published 17 April 1984
Hour of Mating Flight in Three Species of Ants
(Hymenoptera: Formicidae ) 1
Elwood S. McCluskey and Dale K. McCluskey
(ESM) Departments of Biology and Physiology, Loma Linda University, Loma
Linda, California 92350; (DKM) Colton, California 92324.
The mating flights of ants usually occur at a time of day characteristic for the
species (Holldobler, 1976; Kannowski, 1963; Talbot, 1946). Also there is appar¬
ently a genus likeness in time of flight (McCluskey, 1973, 1974); but for study at
this taxonomic level, records for more species are desirable. For one species
reported here, the hour is very different from most other records in its genus.
The flights all occurred in or near our yard 1 km up Reche Canyon (elevation
350 m), near Colton, San Bernardino County, California. The observations of the
first species below were made by both of us, those of the second by ESM, and the
third by Esther McCluskey.
Solenopsis Maniosa Wheeler
Mallis (1938) states that the mating flights are in the late afternoon of a warm
day, and we have often seen the winged castes out of the nest on summer evenings.
The purpose here is to make a more quantitative statement. In Figure 1A are
assembled the flight records for a number of nests and days. A few of these records
were the basis for the “1815” flight hour given in Figure 1 of McCluskey (1974)
(there called S. xyloni).
The hour of peak flight is shown by an “x” for each record; the mean was 1750.
The span of time from the earliest (1630) to the latest (1855) ant seen to fly is
shown by the short bar underneath. Flights occurred only in the shade, between
25 and 32°C (temperature at 1 m from surface). Any given flight often lasted only
a small fraction of an hour, but was preceded by a build-up of workers and alates.
Alates flying per minute varied from less than one to many.
For several nests in July 1979 both light intensity and ground-level temperature
were observed. At least for the limited number of nests and days compared, nest-
to-nest or day-to-day variability of these measurements did not appear to explain
the variability in time of flight (nearly an hour). Peak flight occurred at the same
time as peak number of alates out. Flowever, for some days or nests there were
no flights seen, even when the number out was 10 or more at once, as if flight
requires something beyond that which initiates exit from the nest. (As DKM says,
“the ants fly when they feel like it.”)
No winged ants were even seen outside the nest on any of the 10 mornings
briefly observed (see Fig. 1B for two of these days), even if unusually warm and
humid.
1 Species kindly determined by Roy Snelling.
152
PAN-PACIFIC ENTOMOLOGIST
©
I-1-1
0000 0400 0800
SUNSET
t
xx
X XXX
X XXX
X XXX
—1-1-1--—I
1200 1600 2000 2400
o
CO
UJ
_I
<
z:
LjJ
<*■ r
LU 5 —
CO
SUNRISE
+
A
/ \
i \
! K
P. HYATT I
/
I
\
i- / A \
* / \ \
7 V '
—p-o-<£pO-o—r-^
HOUR
0500
0600
Figure 1. A. Flight hours of S. maniosa. Each x represents hour (PST) of peak number flying/
minute for 1 nest for 1 day. Horizontal bar underneath runs from earliest to latest hour any ant was
seen to fly. The 14 flights are from 7 nests and 11 days between 20 June and 24 July during years
1973-1979. B. Hour of surface appearance and of flight for 1 nest each of 2 species (between 13 and
26 July 1978): • = P. hyatti, 0=5’. maniosa. Upper curve in each case represents number out of nest
(points are shown for other end of day, but are all zero and are not connected by OUT line). Lower
curve represents number flying/minute; ordinate scale there is 0.2 per division instead of 1 per division
as for number out.
Pheidole Hyatti Emery
Early one morning I happened to see winged females at a nest under a fruit tree
and, thinking them to be S. maniosa, was surprised at such a flight hour. A few
minutes later we saw ants hovering over a bare area of the yard 50 m from the
nest. Both these ants and those at the nest turned out to be P. hyatti.
The hovering or swarming area was observed for several days beginning just
before sunrise at about 0500. Only females were seen flying. They were from 1
to at least 3 m above the ground, and varied in number from zero to five at any
given time. The number was undiminished at the last observation about 0600.
The sun was then beginning to reach the area. On two occasions a male was seen
with a female on the ground beneath, one of the pairs in copulation. (The latter
as well as ants from the nest were used for species determination.) No ants were
seen flying there in the evenings.
VOLUME 60, NUMBER 2
153
In Figure IB is shown the nest flight activity for several days, and for com¬
parison, that of a nest of S. maniosa on two of the same days. The flight hour of
the P. hyatti colony (peak at 0520) was at the opposite end of the day from that
of S. maniosa. Only females were seen at either nest. For each species the number
of workers around the nest was far higher at the flight hour than at the opposite
end of the day, when there were no females out at all. Another contrast between
the flight hours was the temperature, 18-19°C for P. hyatti and 30-27° for S.
maniosa. Neither nest was in the sun at flight time.
The present record adds a second morning species for this genus. The compi¬
lation of flight hour records for Pheidole by McCluskey (1974) included seven
species. All of these flew in the late afternoon or evening except one in the early
morning. And even in that species, flight from the nest itself was not seen and
could have occurred in the late evening (Williams and Illingworth, 1935).
Pheidole Pilifera Pacifica Wheeler
About 1430 on 3 November 1959 Esther McCluskey discovered a mating swarm
of this species upon bicycling through it along an open lane. Again the next
afternoon there was swarming, lasting until after 1530. Twenty or 30 males were
flying around each female, and occasionally a copulating pair would fall to the
ground (several such pairs were collected for species determination). It was partly
cloudy and the air temperature was 19°C. These records were the basis for the
“1530” flight hour given in Figure 1 of McCluskey (1974).
Comparison of Mating Swarms in Pheidole
The mating aggregations reported here for two species of Pheidole resemble
those more completely described by Wilson (1957) for P. sitarches as well as one
briefly noted by Williams and Illingworth (1935) for P. megacephala. In all four
species the swarms were relatively low, and over bare areas. This is in contrast
to some other ants, where swarms occur on or above tall objects or high in the
air (Sudd, 1967).
Three of these four species of Pheidole, and another, pallidula (Goetsch, 1957),
were alike in that mating pairs dropped out of the swarms to the ground. In P.
megacephala, however, Williams saw the pairs continue to fly in the swarm.
The time relation between flight from the nest and swarming is not well known.
In none of the five species of Pheidole mentioned above was nest flight hour
reported except for P. hyatti described here. In this case flight at the mating area
continued from before until after the flight at the one nest seen.
Summary
For comparison at higher taxon levels there is need for records of more species.
The “evening” flight time of Solenopsis maniosa is documented by 14 different
day or nest flight records. The mean peak flight hour was 1750, the earliest flight
starting at 1630 and the latest ending at 1855 just before sunset; all were in the
shade.
Flights from one nest of Pheidole hyatti on several of the same days in July as
S. maniosa, were centered at 0520, just after sunrise but in the shade. A mating
aggregation of the same species hovered at a nearby spot on the same mornings.
Morning flight records are unusual for Pheidole.
154
PAN-PACIFIC ENTOMOLOGIST
A swarm of Pheidole pilifera pacifica was also seen but in November and at
about 1500. Like P. hyatti and certain previously-reported species in the genus,
the swarming was over a bare area, it was low, and mating pairs dropped to the
ground beneath.
Literature Cited
Goetsch, W. 1957. The ants. University of Michigan, Ann Arbor, 173 pp.
Holldobler, B. 1976. The behavioral ecology of mating in harvester ants (Hymenoptera: Formicidae:
Pogonomyrmex). Behav. Ecol. Sociobiol., 1:405-423.
Kannowski, P. B. 1963. The flight activities of formicine ants. Symp. Genet. Biol. Ital., 12:74-102.
Mallis, A. 1938. The California fire ant and its control. Pan-Pac. Entomol., 14:87-91.
McCluskey, E. S. 1973. Generic diversity in phase of rhythm in formicine ants. Psyche, 80:295-
304.
-. 1974. Generic diversity in phase of rhythm in myrmicine ants. J. N.Y. Entomol. Soc., 82:
93-102.
Sudd, J. H. 1967. An introduction to the behaviour of ants. Arnold, London, 200 pp.
Talbot, M. 1945. A comparison of flights of four species of ants. Amer. Midland Naturalist, 34:
504-510.
Williams and Illingworth. 1935. Proc. Hawaiian Entomol. Soc., 9:3.
Wilson, E. O. 1957. The organization of a nuptial flight of the ant Pheidole sitarches Wheeler. Psyche,
64:45-50.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, pp. 155-162
Published 17 April 1984
Observations on the Biology of Eremapis parvula Ogloblin
an Anthophorid Bee with a Metasomal Scopa
(Hymenoptera: Anthophoridae)
John L. Neff
7307 Running Rope, Austin, Texas 78731.
Abstract. — Observations are presented on nests, floral host preferences and phe¬
nology of Eremapis parvula Ogloblin, a monotypic exomalopsine bee from the
deserts of northern Argentina. Dense nest aggregations were located in fine alluvial
soil deposits along a sandy river bank. Individual nests are constructed by a single
female and show many features observed in the related genus Exomalopsis. A
distinctive feature of Eremapis nest cells is the ringlike deposition of fecal pellets.
Eremapis appears to be an oligolege of Prosopis (Fabaceae) and the phenology of
local bee populations is synchronized with the flowering of various taxa of that
genus. Pollen is transported on unusual keeled setae on the metasomal sterna as
well as in the scopa of the hind legs.
Eremapis is a monotypic genus of tiny (female body length 3.8-4.2 mm) bees
endemic to the arid Monte Desert and adjacent portions of the western Chaco of
Argentina (Ogloblin, 1956). It forms, along with the related monotypic genera
Teratognatha of northern Argentina and Chilimalopsis of northern Chile, a dis¬
tinctive group within the Exomalopsini which may represent the remnants of a
relatively primitive stock within that taxon (Michener and Moure, 1957; Toro,
1976). Nothing has been published on the biology of any of these three genera.
The information I present here on Eremapis parvula Ogloblin may be of impor¬
tance in examining evolutionary trends within the Exomalopsini, a group of
considerable biological diversity.
During the six month southern spring-summer (Oct.-Mar.) flowering seasons
of 1972-1973 and 1973-1974, I was able to study the phenology and floral
relationships of Eremapis parvula in the vicinity of Andalgala, Catamarca, Ar¬
gentina, as part of my larger study of the flower-visiting insects of this region. No
nests were discovered near Andalgala, but during a trip through the central and
southern portions of the Monte Desert, I discovered several large nesting aggre¬
gations and excavated numerous nests. The data on nesting habits of Eremapis
presented here are based on observations taken near this site as well as from
observations made on nest-containing soil examined in the laboratory. Philip
Torchio supplied additional notes based on material from the nest site. Eggs,
larvae, and pupae recovered from the nests were deposited with Mr. Torchio for
description.
Nesting Sites and Nest Architecture
On 17 November 1973, three nesting aggregations were discovered along the
banks of the Rio Huaco near the Termas Aguas Hedionados, approximately 30
156
PAN-PACIFIC ENTOMOLOGIST
km north of San Jose de Jachal, San Juan, Argentina. The area is covered with
desert scrub vegetation and the river itself was lined with bushes of Baccharis
spp. (Asteraceae) and small Prosopis (Fabaceae) trees intermixed with shrubs of
Schinus sp. (Anacardiaceae), Larrea divaricata (Cavanilles), Bulnesia retama
(Hooker and Arnott) Griseb., Plectrocarpa sp. (Zygophyllaceae), and Atamisquea
emarginata Hooker and Arnott (Capparaceae). The soil near the river was pre¬
dominantly bare, loose sand. The nests were restricted to encrusted areas in low
depressions which were filled with silt during periods of high water. The nest sites
proper were devoid of vegetation but the entire area was surrounded by low
Prosopis trees which would have caused partial shading. The crust of the soil
surface of the nest-containing depressions was between 6 and 8 mm thick and
underlain by a moist silt layer about 15 to 18 cm deep. Beneath this layer was
pure sand. The largest nesting aggregation occupied a depression that was 8 m
long and 1.5 m wide that was completely covered with numerous low conical
tumuli from the closely packed nests. No turrets were observed. Between 8 and
11 open nests were present per 100 cm area suggesting a population of over 8000
females in this one aggregation.
Nest openings were made directly into the crust or through pre-existing cracks
in the crust. The burrows were unlined, 0.9-1.0 mm in diameter, and descended
vertically for 12-15 cm. After reaching a depth of about 15 cm, the burrows turned
abruptly giving rise to horizontal laterals that extended about 3-4 cm before
terminating in single, oval, apparently unlined cells (Fig. 1). The short amount
of time available for excavating and the density of the nests prevented an accurate
determination of the number of cells per nest, but it is assumed that completed
nests are multicellular. No linear series or clusters of cells were found and, in fully
provisioned cells, the laterals were filled with soil. A few cells were found in the
upper 5 cm of the silty soil and several others were found at the silt-sand interface
or just slightly under the interface in the sand. Although the nest site was dis¬
covered at dusk, numerous males and females were visible, flying over the nest
site or resting on the soil surface. Only single females were found within open
nests. The presence of numerous old cells in the soil samples indicated that the
nest site had been used in previous years.
Individual cells were oval, approximately 3-4 mm long and 2—2.5 mm wide
with the long axis roughly parallel to the connecting lateral (Fig. 2). They were
apparently unlined and could not be extracted from the soil matrix. The cell cap
was thin (0.3-0.6 mm) and apparently also unlined. In each case, the provisions
were molded pollen masses with no free liquid. The smooth, slightly elongate
pollen masses were approximately 2.0 mm long and 1.5 mm tall and deep. The
ventral surface of the pollen masses was flattened and slightly concave. In some
cells, new pollen balls appeared to have the end nearest the cap slightly raised,
reminiscent of pollen balls made by some species of Exomalopsis (Raw, 1977;
Rozen, 1977), although this feature was not observed in all cases. Without ex¬
ception, the pollen masses consisted solely of Prosopis pollen. Cells containing
eggs as well as all stages of immatures were found. Dark pupae and adults were
absent in closed cells. Most cells excavated contained developing larvae still
feeding on the provisions.
The elongate egg (about 1 mm long) was placed on the dorsal portion of the
pollen mass, parallel to the long axis of the cell (Fig. 2). The egg contacted the
VOLUME 60, NUMBER 2
157
1
2
I mm
i-1
Figures 1-3. Nests of Eremapis parvula. 1. Diagram of a typical open nest. 2. Cell, pollen mass,
and egg. 3. Cell with ring of fecal smears.
pollen mass only at its tips with the middle portion arched away from the pro¬
visions. No distinct grooves or indentations in the pollen mass for egg placement
were noted. Early instars moved around the middle of the pollen mass resulting
in distinct feeding grooves which commonly resulted in a dumbbell-shaped pollen
mass as the feeding progressed. In the last developmental stages, larvae completely
encircled the reduced pollen mass with their body axes aligned perpendicular to
the long axes of the cells.
Several cells contained larvae that had initiated defecation but which still con¬
tained portions of the pollen ball. In most cells with fecal smears, the pollen mass
had been completely consumed suggesting that Eremapis, like many exomalop-
sines, may initiate defecation before completion of feeding. Feces were deposited
as narrow, flattened strips 0.42-0.64 mm long and 0.10- 0.12 mm wide and 0.02-
0.03 mm high. Approximately 90% of the feces were laid down around the center
of the cell and most strips were parallel to equator of the cell thus forming a
distinct ring that could be removed intact from the cell (Fig. 3).
Following defecation and deposition of a light yellow material (probably Mai-
158
PAN-PACIFIC ENTOMOLOGIST
Table 1. Host records for Eremapis parvula .'
Family
Taxon
No. of
females
No. of
males
Total
Fabaceae
Prosopis chilensis
128
59
187
Fabaceae
Prosopis flexuosa
1
35
36
Fabaceae
Prosopis torquata
18
25
43
Fabaceae
Mimozyganthus carinatus
1
1
Rhamnaceae
Zizyphus mistol
16
16
Capparaceae
Atamisquea emarginata
2
2
Olacaceae
Ximenia americana
4
4
Euphorbiaceae
Jatropha excisa
1
1
Zygophyllaceae
Larrea divaricata
2
2
Zygophyllaceae
Larrea cuneifolia
1
1
147
146
293
1 Records from the Bolson de Pipanaco, Catamarca, Argentina summed over two seasons, 1971—
1972, 1972-1973.
pighian excreta) on the bottom of the cell, the larva laid down a complete, trans¬
parent, single-layered cocoon. The cocoon followed the contours of the cell except
at the periphery of the cell cap where it curved into the center face of the cap
rather than precisely following the contours of the cell. Fecal material was not
incorporated into the cocoon, but did adhere firmly to it.
Phenology and Floral Associations
All available data indicate that Eremapis parvula collects only the pollen of
species of Prosopis. During the two seasons of collecting in the Bolson de Pipanaco
in which Andalgala is located, males of Eremapis were taken at the flowers of a
number of plant taxa, but females were observed only on Prosopis flowers (Table
1). Microscopic examination of both scopal loads and nest provisions revealed
only Prosopis pollen. In the vicinity of Andalgala, Eremapis was collected from
19 October to 6 January. This long flight period (79 days) for a desert species is
somewhat misleading as individual populations appear to be active for no more
than 30-40 days (based on collection records for individual sites). In fact, there
appear to be two flight periods, the first involving populations associated with
the synchronous early (Oct.-Nov.) flowering of the large phreatophytic trees of
Prosopis chilensis (Molina) Stunz and P. flexuosa de Candolle (both in section
Algarobia ) and the second involving populations that foraged on the later bloom¬
ing Prosopis torquata (Lagasca) de Candolle (section Strombocarpa). Flowering
of the phreatophytic species of Prosopis is largely independent of local rainfall,
but flowering of Prosopis torquata is closely tied to rainfall and local patterns of
runoff.
Female Eremapis showing minimal wing wear were found associated with both
the early blooming and the later blooming species. The relatively long flight period
could thus be due to multivoltinism (suggested by the presence of pupae in the
active nests at Rio Huaco) or, more likely, the staggered emergence of univoltine
populations responding to environmental cues which facilitated synchrony of
emergence with the different species of Prosopis growing in different habitats.
VOLUME 60, NUMBER 2
159
Figures 4, 5. Metasomal scopal hairs of female Eremapis parvula. 4. Rows of specialized setae on
sterna 2, 3, and 4 forming part of the metasomal scopa (x 175). 5. Tip of an individual scopal seta
with Prosopis pollen grain (x 350).
Mating
Copulation was not observed. Males were frequently observed hovering about
Prosopis inflorescences that were being visited by females. Occasionally males
pounced on foraging females, but their overtures were always rebuffed. These
observations suggest that at least some mating may take place at the flowers. Males
were also present at the Rio Huaco nest site although no mating activity was
noted.
Pollen Transport
Most pollen collected by female Eremapis is transported on the sparsely branched
scopal hairs of the hind tibia and basitarsi, a scopa very similar in structure to
that of various Ancyloscelis species. However, a significant amount of pollen is
also transported in an auxiliary metasomal scopa formed by sparse bands of
specialized, falcate setae on the second to the fifth sterna (Figs. 4 and 5). Exam¬
ination of pollen bearing females at the Rio Huaco site indicated that females
with metasomal pollen loads always had at least some pollen in the basitarsal
scopa although not all pollen bearing females had metasomal pollen loads. The
presence of females with metasomal pollen loads at the nest indicated that the
metasomal scopae are not just specialized structures that are used to gather pollen
or accumulate it before loading it into the scopa on the legs. Metasomal scopa
are also found in several paracolletine bees associated with Prosopis in Argentina
(Simpson et al., 1977) although in some of these cases, the metasomal scopa may
be used more for collecting pollen than for its transport.
Discussion
The nesting biology of the Exomalopsini has most recently been reviewed by
Torchio (1974). Subsequently, additional data have been provided by Michener
160
PAN-PACIFIC ENTOMOLOGIST
(1974), Vogel (1974), Rozen (1977), Raw (1977), and Parker (in press). My in¬
terpretation of Torchio’s analysis is that the only biological characteristic that
consistently separated the Exomalopsini from the other anthophorids was the
presence of oval shaped cells. Since oval-shaped cells are probably a primitive
character for the “higher” bees, there are no derived biological characters that
distinguish the group. Eremapis shows a number of characters that appear to
be primitive both within the Exomalopsini and the Anthophorinae in general.
These include: nests in soil; absence of turrets; unlined burrows; oval cells; molded
provisions with the egg placed on the surface opposite that attached to the walls;
and a complete cocoon. The apparent absence a secreted cell lining would be a
derived character but, as a loss character, is of limited phylogenetic significance.
Moreover, absence of a secreted lining was not rigorously established. The ini¬
tiation of defecation before completion of larval feeding is a derived trait for bees
(Stephen, Bohart and Torchio, 1969), but may be primitive for anthophorids. The
most distinctive and perhaps only derived character of the nests of Eremapis is
the ringlike arrangement of deposited feces.
Nest architecture of Eremapis appears to be very similar to that described for
Exomalopsis with a vertical main burrow and distinct laterals. Cell orientation
in different species of Exomalopsis varies from consistently horizontal (Raw,
1977), at a 40-45 degree angle (Raw, 1977; Rozen, 1977) from horizontal, or
may be variable (Rozen and MacNeill, 1957). Defecation may occur before the
completion of feeding (Rozen, 1977) and fecal matter may (Rozen and MacNeill,
1957; Rozen, 1977) or may not (Raw, 1977) be included in the cocoon. Biological
characters uniting Exomalopsis and separating it from Eremapis include the char¬
acteristically molded provisions with a foot-like projection and the secreted, wax¬
like lining. Additionally, most, but not all, Exomalopsis appear to be communal
(several females per nest) or even semi-social. Ancyloscelis nests lack the typical
vertical main burrow with a horizontal lateral arrangement, but may show com¬
plicated branching patterns. Ancyloscelis nests may (Michener, 1954; J. G. Rozen,
pers. comm.) or may not (Torchio, 1974; Michener, 1974) possess an obviously
secreted lining. Cocoon structure in Ancyloscelis appears to be very similar to that
of Eremapis in that it is one-layered and fragile (Torchio, 1974; Michener, 1974)
and contacts the central but not the lateral portions of the cell closure. In Ancy¬
loscelis, the feces are deposited as a complete, thin layer (Torchio, 1974; Michener,
1974) rather than in discrete strips characteristic of Eremapis or Exomalopsis
(Rozen and MacNeill, 1957; Raw, 1977; Rozen, 1977).
Paratetrapedia and Tapinotaspis differ from Eremapis, Ancyloscelis and Exo¬
malopsis in that their eggs are oriented perpendicular rather than parallel to the
long axis of the cells. Paratetrapedia and Tapinotaspis also differ in that their
provisions are molded into a ball-like form. According to Vogel (1974), but not
Claude-Joseph (1926), provisions in Tapinotaspis caerulea Friese are formed into
a ball-like mass with a distinct basal layer filling the bottom of the cell.
Data on the biology of other exomalopsine genera are largely fragmentary or
non-existent, but suggest considerable diversity (Torchio, 1974). Distinct wax¬
like cell linings are present in the cells of Tapinotaspis (Claude-Joseph, 1926;
Vogel, 1974) as well as in those of Monoeca and Paratetrapedia (Michener and
Lange, 1958). Data on the nesting biology of Eremapsis are entirely consistent
with its placement in the Exomalopsini and suggest a relatively close alignment
VOLUME 60, NUMBER 2
161
to Exomalopsis and more distantly, to Ancyloscelis, but the monophyletic, as
opposed to paraphyletic, nature of the Exomalopsini is open to question.
Although the host relationships of most exomalopsines are poorly known, most
appear to be polylectic. Oligolecty is well established and perhaps universal in
Ancyloscelis and is known in the subgenera Anthophorisca and Anthophorula of
Exomalopsis (Timberlake, 1980; pers. obs.) and may occur in some Tapinotaspis
(A. Moldenke, pers. comm.), but none of these groups include bees oligolectic on
the Fabaceae. An undescribed Isomalopsis is commonly associated with Prosopis
in northern Argentina, but this bee also collects pollen of Acacia (Fabaceae) and
Atamisquea. With its specialized mouth parts, one might also expect oligolecty
in Teratognatha although its host relationships are unknown. A character, pre¬
sumably primitive, allying Eremapis with Exomalopsis, Isomalopsis, Ancyloscelis
and probably Caenonomada and separating it from Paratetrapedia, Chalepogenus,
some Lanthanomellisa, Tapinotaspis and probably Monoeca is the absence of
floral oil collection (Vogel, 1974; Neff and Simpson, 1981).
The presence of a functional metasomal scopae in Eremapis is undoubtedly a
de novo specialization, but it is of interest as the absence of such a scopa is
considered to be an important character separating the Anthophoridae from the
Megachiliidae and Fedeliidae. Specialized metasomal arrays of branched setae
(Thorp, 1979; pers. obs.) are found in various Svastra and Xenoglossodes (Eu-
cerini) associated with the Asteraceae, but in these cases they appear to be pri¬
marily involved in pollen collection with pollen transport as a secondary function.
The distinctive metasomal setal arrays found in certain Tapinotaspis are believed
to be involved in floral oil collection rather than pollen collection or transport
(Neff and Simpson, 1981).
The combination of primitive and derived biological traits seen in Eremapis
appears to support Michener and Moure’s conclusion (Michener and Moure, 1957)
based on adult morphology, that Eremapis is a survivor of a primitive stock of
the Exomalopsini. No derived biological characters link Eremapis with the oil¬
collecting bees of the Paratetrapedia-Tapinotaspis complex although this group
is still poorly known biologically. Eremapis does show various similarities to
Exomalopsis and Ancyloscelis but these primarily involve primitive characters.
The relationship of the cocoon to the cell cap may be a derived character linking
Ancyloscelis and Eremapis, but characters such as this have not been sufficiently
surveyed to allow any firm conclusions to be drawn.
Acknowledgments
I thank A. R. Moldenke for making possible my wanderings through Argentina,
J. C. Schultz for assistance in nest excavation, Padre J. S. Moure for confirmation
of Eremapis identifications, P. Torchio for providing me with his own notes on
the nest material as well as ideas on exomalopsine biology, and B. B. Simpson
for typing the manuscript and helping prepare the illustrations. A. R. Moldenke,
P. Torchio, J. G. Rozen, and W. P. Stephen commented on an earlier draft of
this manuscript. Research in Argentina was part of the International Biological
Program, Structure of Ecosystems Desert Scrub Project.
Literature Cited
Claude-Joseph, F. 1926. Recherches biologiques sur hymenopteres du Chili (Melliferes). Ann. Sci.
Nat. Zool., (10)9:113-368.
162
PAN-PACIFIC ENTOMOLOGIST
Michener, C. D. 1954. Bees of Panama. Bull. Amer. Mus. Nat. Hist., 104:1-176.
-. 1974. Further notes on nests of Ancyloscelis (Hymenoptera: Anthophoridae). J. Kansas Ento-
mol. Soc., 47:19-22.
-, and R. B. Lange. 1958. Observations on the ethology of Neotropical anthophorine bees
(Hymenoptera: Apoidea). Univ. Kansas Sci. Bull., 39:69-96.
-, and J. S. Moure. 1957. A study of the classification of the more primitive non-parasitic
anthophorine bees (Hymenoptera, Apoidea). Bull. Amer. Mus. Nat. Hist., 112:395-452.
Neff, J. L., and B. B. Simpson. 1981. Oil-collecting structures in the Anthophoridae (Hymenoptera):
morphology, function, and use in systematics. J. Kansas Entomol. Soc., 54:95-123.
Ogloblin, A. A. 1956. Dos generos nuevos de la tribu Exomalopsini (Apidae, Hym.) de la Republic
Argentina. Dusenia, 7:149-158.
Parker, F. D. In press. Biological notes on the bee Exomalopsis crenulata Timberlake (Hymenoptera:
Anthophoridae). Pan-Pacific Entomologist.
Raw, A. 1977. The biology of two Exomalopsis species (Hymenoptera: Anthophoridae) with remarks
on sociality in bees. Rev. Biol. Trop., 25:1-11.
Rozen, J. G., Jr. 1977. Immature stages and ethological observations on the cleptoparasitic bee tribe
Nomadini (Apoidea: Anthophoridae). Amer. Mus. Novit., 2638:1-27.
-, and C. D. MacNeill. 1957. Biological observations on Exomalopsis ( Anthophorula ) chionura
Cockerell, including a comparison of the biology of Exomalopsis with that of other anthophorid
groups (Hymenoptera: Apoidea). Ann. Entomol. Soc. Amer., 50:522-529.
Simpson, B. B., J. L. Neff, and A. R. Moldenke. 1977. Prosopis flowers as a resource. Pp. 84-107
in B. B. Simpson (ed.), Mesquite. Its biology in two desert ecosystems. Dowden, Hutchinson
and Ross, Stroudsburg.
Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The biology and external morphology of
bees. Agr. Exp. Sta., Oregon State Univ., Corvallis, OR.
Thorp, R. W. 1979. Structural, behavioral, and physiological adaptations of bees (Apoidea) for
collecting pollen. Ann. Missouri Bot. Gard., 66:788-812.
Timberlake, P. H. 1980. Review of North American Exomalopsis (Hymenoptera, Anthophoridae).
Univ. Calif. Publ. Entomol., 90:1-158.
Torchio, P. F. 1974. Notes of the biology of Ancyloscelis armata Smith and comparisons with other
anthophorine bees (Hymenoptera: Anthophoridae). J. Kansas Entomol. Soc., 47:54-63.
Toro, H. 1976. Chilimalopsis, nuevo genero chileno de Exomalopsini (Hymenoptera: Apoidea). Ann.
Mus. Hist. Nat. Valparaiso, 9:73-76.
Vogel, S. 1974. Olblumen and olsammelnde Bienen. Akad. Wiss. Lit. Mainz. Tropische und sub-
tropische Pflanzenwelt, 7:285-547.
PAN-PACIFIC ENTOMOLOGIST
60(2), 1984, p. 163
Published 17 April 1984
Ambracyptus, A New Name for Paracyptus See vers
(Coleoptera: Staphylinidae)
Randall W. Lundgren
815 South Fell Avenue, Normal, Illinois 61761.
Seevers (1971) erected a monotypic genus, Paracyptus, for P. minutissima, a
new species of rove beetle found in amber from Chiapas, Mexico. This generic
name, however, was previously used by Cameron (1944) for another genus of
staphylinid beetle from India. Paracyptus Seevers, 1971, is, therefore, a junior
homonym of Paracyptus Cameron, 1944. I herein propose to replace Paracyptus
Seevers with Ambracyptus Lundgren, new name. The sole species affected becomes
Ambracyptus minutissimus (Seevers), new combination.
Literature Cited
Cameron, M. 1944. Descriptions of new Staphylinidae (Coleoptera). Proc. R. Entomol. Soc. Lond.,
Ser. B, Taxon, 13:49-52.
Seevers, C. H. 1971. Fossil Staphylinidae in Tertiary Mexican amber (Coleoptera). Univ. Calif. Publ.
Entomol., 63:77-86.
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Vol. 60
No. 3
July 1984
THE
Pan-Pacific Entomologist
ERWIN, T. L.—Studies of the Tribe Bembidiini (Coleoptera: Carabidae): Lectotype designa¬
tions and species group assignments for Bembidion species described by Thomas L.
Casey and others... 165
PARKER, F. D. — Biological notes on the bee Exomalopsis crenulata Timberlake (Hymenop-
tera: Anthophoridae) . 188
MOCKFORD, E. L.—A systematic study of the genus Camelopsocus with descriptions of three
new species (Psocoptera: Psocidae). 193
CAVENDER, G. L. and R. D. GOEDEN— The life history of Paracantha cultaris (Coquillett)
on wild sunflower, Helianthus annuus L. ssp. lenticularis (Douglas) Cockerell, in Southern
California (Diptera: Tephritidae). 213
MANLEY, D. G. —Predation upon Velvet Ants of the genus Dasymutilla in California (Hy-
menoptera: Mutillidae). 219
RENTZ, D. C. F. and D. B. WEISSMAN—Five new species of the band-winged grasshopper
genus Trimerotropis Sthl (Orthoptera: Oedipodinae). 227
WAGNER, R. —Contributions to Nearctic Psychodidae (Diptera, Nematocera). 238
ALLEN, R. K.—A new classification of the subfamily Ephemerellinae and the description of
a new genus. 245
ANDERSON, R. S. and S. B. PECK—Bionomics of Nearctic species of Aclypea Reitter: Phy¬
tophagous “Carrion” beetles (Coleoptera: Silphidae). 248
MUZZIO, S. F.—Neosminthurus bellingeri, a new species from California (Collembola: Smin-
thuridae). 258
SCIENTIFIC NOTES. 244, 256
EDITORIAL NOTICE. 264
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PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 165-187
Studies of the Tribe Bembidiini (Coleoptera: Carabidae):
Lectotype Designations and Species Group Assignments
for Bembidion Species Described by
Thomas L. Casey and Others
Terry L. Erwin
Department of Entomology, National Museum of Natural History, Smithsonian
Institution, Washington, D.C. 20560.
Many species of the genus Bembidion were dealt with by Lindroth (1963) in
his monumental “Ground Beetles of Canada and Alaska,” synonymies were given,
and species were described and assigned to species groups. Later, Lindroth (1975)
designated lectotypes for the Casey names which pertain to species of the northern
United States and Canada. However, this left more than 100 Casey names and
several others by LeConte, Hayward, Dejean, Chaudoir et al. not yet covered with
regard to taxonomy, nor species group assignment.
As in the case of Tachyina (Erwin, 1974), another group worked on by Casey,
little can be done before this “housekeeping” task is undertaken and a good
working list of names becomes available. Lindroth (1963) provided an excellent
beginning with his arrangement of species groups. Using this arrangement, I have
assigned all remaining species described from north of Panama to species groups,
provided synonymies of names not dealt with by Lindroth, and I note distribu¬
tions, where possible, extracted from USNM specimens.
These data then provide a working list for use in NABFP, the USD A/SI Co¬
leoptera Catalogue, and monographic studies now under way (Erwin and Kavan-
augh, 1980, 1981) and are compiled here as an aid to those projects.
Of Bembidion names for the Americas north of Colombia all are here dealt
with except some of the species of Cyclolopha and two Motschulsky names. G.
Perrault has dealt with Cyclolopha (MS) and designated lectotypes, etc. Two of
these are mentioned here, as is one other he did not deal with. The Motschulsky
names B. obscuromaculatum (Mots, 1859:130) and B. laterimaculatum Mots.
(1859:130) are undoubted members of the dorsale group. Lindroth (1963:211)
mentions the first species but does not give any clues as to its relationships. From
Motschusky’s descriptions and the probable locality (Fort Ross, California) I
suspect both names will apply to B. indistinctum Dejean, however the types must
be examined.
Methods
Methods are the same as mentioned before (Erwin, 1974). Most of the material
studied resides in the USNM (Smithsonian Institution, Washington, D.C. 20560),
however some other museum acronyms are used too. These are as follows:
BMNH—British Museum (Natural History); London; P. Hammond, and N. Stork.
CAS—California Academy of Sciences, San Francisco, California; D. H. Kavan-
augh.
166
PAN-PACIFIC ENTOMOLOGIST
MCZ—Museum of Comparative Zoology, Harvard, Cambridge, Massachusetts;
M. Thayer, A. Newton, S. Foster.
MNHP—Museum National d’Histoire Naturelle, Paris; H. Perrin.
Acknowledgments
I heartily thank all the above curators and their respective staffs for aid in
locating type material or providing data from type labels. I also thank Noreen
Connell and Linda Sims of my own staff for aid during preparation of this paper.
Bembidion acticola Casey
Bembidion acticola Casey, 1884:63. Lectotype: USNM #37042, male, here se¬
lected. Type-area: New Jersey, USA.
Bembidion argutum Casey, 1918:123. New Synonymy. Lectotype: USNM #37045,
male, here selected. Type-area: Rhode Island, USA.
Bembidion assensum Casey, 1924:41. New Synonymy. Lectotype: USNM #37406,
female, here selected. Type-area: Near Brooklyn, New York, USA.
Notes.— This species belongs in the contractum group; its members are found
along the Atlantic Coast from Long Island south to Maryland.
Bembidion actuosum Casey
Bembidion actuosum Casey, 1918:65. Lectotype: USNM #36899, female, here
selected. Type-area: Boulder County, Colorado, USA.
Bembidion debilicolle Casey, 1924:34. New Synonymy. Lectotype: USNM #36935,
female, selected and labelled by Lindroth in 1972 but unfortunately left out of
his paper (Lindroth, 1975), therefore credited here. Type-area: Lake County,
Oregon, USA.
Notes.— See Lindroth (1963:318) for details of distribution and note that Lin¬
droth used Casey’s second name for this species.
Bembidion adductum Casey
Bembidion adductum Casey, 1918:149. Lectotype: USNM #37062, male, here
selected. Type-locality: Paraiso Hot Springs, Monterey County, California, USA.
Bembidion relictum Casey, 1918:153. New Synonymy. Holotype: USNM #37067,
male. Type-locality: Truckee, California, USA.
Notes. — This species belongs in the quadrimaculatum group; its members are
found in central California.
[Bembidion aegrotum Casey, see B. quadrulum]
Bembidion alpinianum Casey
Bembidion alpinianum Casey, 1924:30. Holotype: USNM #36883, female. Type-
area: Nevada County, California, USA.
Notes.— This species belongs in the nigrum group and is found in California.
[Bembidion amnicum Casey, see B. nubiculosum\
VOLUME 60, NUMBER 3
167
[Lymneops angusticeps Casey, see B. laticeps\
[Bembidium apicale Jacquelin du Val, see B. spretum\
Bembidion approximatum (LeConte)
Ochthedromus approximate LeConte, 1852:187. Lectotype: MCZ #5526, male,
here selected. Type-locality: San Diego, California, USA.
Bembidion cernans Casey, 1918:100. New Synonymy. Lectotype: USNM #36997,
female, here selected. Type-locality: San Diego, California. USA.
Bembidion haustum Casey, 1924:38. New Synonymy. Holotype: USNM #37000,
female. Type-area: Alameda County, California, USA.
Notes. — There is no difference between Casey’s and LeConte’s types.
Bembidion aratum (LeConte)
Ochthedromus aratus LeConte, 1852:189. Lectotype: MCZ #3343, female, here
selected. Type-area: Gila River Valley, Arizona, USA.
Bembidium scintillans Bates, 1882:150. New Synonymy. Lectotype: BMNH, fe¬
male, here selected. Type-locality: Capulalpam, Mexico.
Bembidion vinnulum Casey, 1918:116. New Synonymy. Lectotype: USNM #37029,
female, here selected. Type-area: “Arizona (southern),” USA.
Bembidion definitum Casey, 1918:166. New Synonymy. Holotype: USNM #37030,
male. Type-locality: Tucson, Arizona, USA.
Notes. — This species belongs in the incrematum group and is found in the
southwestern USA south through Mexico to Honduras. The two Casey names are
based on color variants.
[Bembidion argutum Casey, see B. acticola ]
[Bembidion assensum Casey, see B. acticola]
[Bembidion augurale Casey, see B. consuetum]
Bembidion auxiliator Casey
Bembidion auxiliator Casey, 1924:38. Lectotype: USNM #36964, male, here se¬
lected. Type-area: San Joaquin County, California, USA.
Notes. — This “species” belongs in the patruele group which is in need of a
complete revision. Many forms are found in the southwestern USA and Mexico,
for example B. conspersum Chd. and their taxonomic limits are poorly understood.
[.Bembidion aversans Casey, see B. rupicola]
Bembidion avidum Casey
Bembidion avidum Casey, 1918:53. Lectotype: USNM #36886, male, here se¬
lected. Type-locality: Reno, Nevada, USA.
Notes. —This species belongs in the nigrum group and is found in the Basin
and Range Province of the western United States.
168
PAN-PACIFIC ENTOMOLOGIST
[Bembidion badiipenne Casey, see B. mexicanum ]
[.Bembidion bellulum Casey, see B. scopulinum\
Bembidion californicum Hayward
Bembidion californicum Hayward, 1897:84. Lectotype: MCZ #16290, female, here
selected. Type-locality: Pomona, California, USA.
Bembidion innocuum Casey, 1918:63. New Synonymy. Lectotype: USNM #36894,
male, here selected. Type-locality: Hoopa Valley, Humboldt County, California,
USA.
Veto.—This species was treated by Lindroth (1963) only in his key and he
placed it in the semistriatum group. Its range is now known to extend from
Humboldt County south to Riverside County in California.
[.Bembidion caliginosum Casey, see B. insulatum\
Bembidion callens Casey
Bembidion callens Casey, 1918:112. Lectotype: USNM #37033, female, here se¬
lected. Type-locality: Tucson, Arizona, USA.
Veto.—This species belongs in the obtusangulum group and is found in the
southwestern USA.
[.Bembidion callidum Casey, see B. quadrulum ]
[Bembidion canonicum Casey, see B. mexicanum ]
[Bembidion cernans Casey, see B. approximatum]
[Bembidicidium chevrolati Gemminger and Harold, see B. spretum]
Bembidion citulum Casey
Bembidion citulum Casey, 1918:66. Lectotype: USNM #36925, male, here se¬
lected. Type-locality: Colonia Garcia, Sierra Madre, Chihuahua, Mexico.
Notes .—This species belongs in the grapei group; its members are flightless,
densely microsculptured, and resemble members of B. texanum. It is probable
that its range is restricted to the Sierra Madre of Mexico.
[Bembidion civile Casey, see B. constrictum ]
Bembidion ciudadense Bates
Bembidion ciudadense Bates, 1891:263. Lectotype: BMNH, male, here selected.
Type-locality: Ciudad, Durango, Mexico.
Notes .—This species belongs in the transversale group and is found on the
Mexican altiplano.
Bembidion clemens Casey
Bembidion clemens Casey, 1918:159. Lectotype: USNM #37080, female, here
selected. Type-locality: Provo, Utah, USA.
VOLUME 60, NUMBER 3
169
Bembidion vapidum Casey, 1918:160. New Synonymy. Lectotype: USNM #37073,
male, here selected. Type-locality: Mt. Diablo, Contra Costa County, California,
USA.
Bembidion disparile Casey, 1918:161. New Synonymy. Lectotype: USNM #37074,
male, here selected. Type-locality: Santa Barbara, California, USA.
Bembidion invidiosum Casey, 1918:162. New Synonymy. Lectotype: USNM
#37081, male, here selected. Type-area: “Road between Fort Wingate and Je-
mez Springs, New Mexico,” USA.
Bembidion remotum Casey, 1918:162. New Synonymy. Lectotype: USNM #37086,
male, here selected. Type-locality: Paraiso Hot Springs, Monterey County, Cal¬
ifornia, USA.
Notes.— This species belongs in the anguliferum group and is found in the
western USA. Lindroth’s B. pseudocautum appears to be closely related and sub¬
sequent studies may show it to be conspecific.
[Bembidion cogitans Casey, see B. texanum ]
Bembidion cognatum Dejean
Bembidium cognatum Dejean, 1831:168. Holotype: MNHP, female. Type-area:
Mexico.
Notes.— This species, with convergent frontal furrows, forms its own group. Its
relationship to other groups is, at present, unclear.
Bembidion coloradense Hayward
Bembidium coloradense Hayward, 1897:98. Lectotype: MCZ #16295, female, here
selected. Type-area: Vicinity of Rico, Dolores County, Colorado, USA.
Notes.—Lindroth (1963) adequately discusses this species but did not designate
a lectotype.
Bembidion conspersum Chaudoir
Bembidion conspersum Chaudoir, 1868:244. (New name for B. tessellatum LeConte
1852:188, not Brulle, 1838:44.)
Bembidium tessellatum LeConte, 1852:188, not Brulle, 1838:44. Lectotype: MCZ
#5532, female, here selected. Type-locality: San Diego, California, USA.
Bembicidium xanthostictum Gemminger and Harold, 1868:424. (New name for
tessellatum LeConte.)
Notes. — This species belongs in the dor sale group; its members are found
throughout California.
Bembidion constrictum LeConte
Bembidium constrictum LeConte, 1848:462. (New name for contractum Dejean.)
Bembidium contractum Dejean, 1831:123, not Say, 1823:85.
Bembidion vernula Casey, 1884:62. New Synonymy. Holotype: USNM #37038,
female. Type-area: Cape May, New Jersey, USA.
Bembidion festinans Casey, 1918:121. New Synonymy. Lectotype: USNM #37043,
female, here selected. Type-area: Texas, USA.
170
PAN-PACIFIC ENTOMOLOGIST
Bembidion civile Casey, 1918:124. New Synonymy. Lectotype: USNM #37044,
female, here selected. Type-locality: Brownsville, Texas, USA.
Notes. — This species belongs in the contractum group as set forth by Lindroth
(1963). The species is an Atlantic and Caribbean coastal form which ranges from
Nova Scotia, Canada, south to Texas. Casey’s names are based on forms within
the variable species.
Bembidion consuetum Casey
Bembidion consuetum Casey, 1918:93. Lectotype: USNM #36955, female, here
selected. Type-locality: Gualala River, Mendocino County, California, USA.
Bembidion augurale Casey, 1924:38. New Synonymy. Lectotype: USNM #36954,
male, here selected. Type-locality: San Francisco, California, USA.
Notes. — This species belongs to the dor sale group and is found around the Bay
Area near San Francisco, California, USA.
[Bembidion continens Casey, see B. idoneum]
[Bembidium contractum Dejean, see B. constrictum\
Bembidion cordatum LeConte
Bembidium cordatus LeConte, 1848:457. Lectotype: MCZ #5522, female, here
selected. Type-area: New York, USA.
Bembidion placabile Casey, 1918:119. New Synonymy. Lectotype: USNM #37034,
male, here selected. Type-locality: Big Spring, Floward County, Texas, USA.
No to.—This species belongs to the dor sale group; its members are found be¬
tween the Rocky and Appalachian Mountain systems from Manitoba to Texas.
Bembidion cubanum Darlington
Bembidion cubanum Darlington, 1937:121. Holotype: MCZ #22491, male. Type-
locality: Sierra de Rangel, Pinar del Rio Province, Cuba, 1500'.
Notes.— This species belongs in the vernale group; its members are similar to
those of B. mexicanum and are found only on the island of Cuba.
Bembidion curtulatum Casey
Bembidion curtulatum Casey, 1918:39. Lectotype: USNM #36859, male, selected
by Lindroth, 1975. Type-locality: Hoopa Valley, Humboldt County, California,
USA.
Bembidion effetum Casey, 1918:40. New Synonymy. Lectotype: USNM #36861,
female, here selected. Type-locality: Soda Springs, Anderson Valley, Mendocino
County, California, USA.
Bembidion flebile Casey, 1918:41. New Synonymy. Lectotype: USNM #36867,
female, by Lindroth 1975:117. Type-locality: Santa Rosa, California, USA.
Notes. — This species belongs in the planiusculum group and was well described
by Lindroth (1963). Unfortunately, Lindroth did not study the type of B. effetum
which has page priority over B. flebile, and he did not decide definitely that the
teneral male type of curlulatum Casey was conspecific. I believe it to be and it
has page priority.
VOLUME 60, NUMBER 3
171
[Bembidion daphnis Casey, see B. nubiculosum ]
[Bembidion debilicolle Casey, see B. actuosum ]
[Bembidion definitum Casey, see B. aratum]
[.Bembidion delectum Casey, see B. wickhami\
[Bembidion demissum Casey, see B. egens\
[Bembidion denveranum Casey, see B. nebraskense ]
[Bembidion derisor Casey, see B. indistinctum\
[.Bembidion disparile Casey, see B. clemens\
[Bembidion docile Casey, see B. rapidum ]
Bembidion durangoense Bates
Bembidium durangoense Bates, 1891:263. Lectotype: BMNH, female, here se¬
lected. Type-locality: Villa Lerdo, Durango, Mexico.
Bembidion gilae Lindroth, 1963:246. New Synonymy. Holotype: MCZ #32536,
male. Type-locality: San Carlos, Gila River Valley, Arizona, USA.
Notes.— This species belongs in the coxendix group; its members are found in
the southwestern USA and northern Mexico.
[Bembidion editum Casey, see B. patruele ]
[Bembidion efficiens Casey, see B. idoneum\
Bembidion egens Casey
Bembidion egens Casey, 1918:132. Lectotype: USNM #37016, male, here selected.
Type-locality: Jemez Springs, New Mexico, USA.
Bembidion demissum Casey, 1918:133. New Synonymy. Lectotype: USNM
#37019, female, here selected. Type-locality: “near Benson, Arizona,” USA.
Notes.— This species belongs in the versicolor group which is badly in need of
study.
Bembidion evidens Casey
Bembidion evidens Casey, 1918:93. New Synonymy. Lectotype: USNM #36956,
male, here selected. Type-locality: Milford, Utah, USA.
Notes.— This species belongs in dorsale group and is known from Utah.
[Bembidion expositum Casey, see B. indistinctum ]
[Bembidion extensum Casey, see B. falsum ]
Bembidion falsum Blaisdell
Bembidion falsum Blaisdell, 1902:76. Lectotype: CAS #2661, male, here selected.
Type-area: Mendocino County, California, USA.
Bembidion extensum Casey, 1918:42. New Synonymy. Lectotype: USNM #36870,
172
PAN-PACIFIC ENTOMOLOGIST
female, here selected. Type-locality: Soda Springs, Anderson Valley, Mendocino
County, California, USA.
Bembidion kincaidi Hatch, 1950:100. New Synonymy. Locality: USNM #75671,
male. Type-locality: Mt. Baker, Oregon, USA.
Notes.— This species belongs in the planiusculum group; its members are found
in the Coastal and Cascade Mountains.
[.Bembidion fastidiosum Casey, see B. pedicellatum\
[Notaphus fastidiosus LaFerte, see B. spretum\
[Bembidion festinans Casey, see B. constrictum ]
Bembidion festivum Casey
Bembidion festivum Casey, 1918:45. Lectotype: USNM #36874, female, here
selected. Type-locality: Santa Barbara, California, USA.
Notes.— This species belongs in the mundum group; its members are found in
California and Oregon.
[Bembidion flebile Casey, see B. effetum ]
Bembidion flohri Bates
Bembidium flohri Bates, 1878:602. Lectotype: MNHP, male, here selected by G.
E. Ball and so labelled by him in 1972. Type-area: “Mexico, near the capitol.”
Bembidion henshawi Hayward, 1897:87. New Synonymy. Lectotype: MCZ #16292,
male, here selected. Type-area: Salt Lake, Utah, USA.
Notes. — This species belongs in the scudderi group; its members are halophilus
and occur from the Mexican transverse volcanic belt north to Manitoba.
[Bembidion formale Casey, see B. indistinctum]
[Bembidion franciscanum Casey, see B. indistinctum ]
[Bembidion frugale Casey, see B. impotens\
[Bembidion fugitans Casey, see B. rapidum]
[Bembidion gilae Lindroth, see B. durangoense ]
Bembidion grandiceps Hayward
Bembidium grandiceps Hayward, 1897:70. Holotype: Not located at MCZ. Type-
locality: Unknown.
Notes. — This species belongs in the planum group; its range extends from Mas¬
sachusetts to Kansas, south to Texas. For some reason, the type of this species
did not arrive at MCZ with the rest of the Horn collection, although Hayward
clearly states that is where he put the type.
Bembidion grandicolle (LeConte)
Ochthedromus grandicollis LeConte, 1852:189. Holotype: MCZ #5539, female.
Type-locality: San Diego, California, USA.
VOLUME 60, NUMBER 3
173
Notes. — This species belongs in the obtusangulum group; its members are known
from southern California.
Bembidion grapei Gyllenhal
Bembidion grapei Gyllenhal, 1827:403. See Lindroth, 1963:319.
Bembidion scrutatum Casey, 1918:64. New Synonymy. Lectotype: USNM #36987,
male, here selected. Type-locality: Eldora, Colorado, USA.
Notes. —This Holarctic species has many synonyms in the New and Old Worlds;
Lindroth (1963) neglected to mention this one.
Bembidion graphicum Casey
Bembidion graphicum Casey, 1918:108. See Lindroth (1963, 1975) for details.
Bembidion lassulum Casey, 1918:118. New Synonymy. Holotype: USNM #37040,
female. Type-locality: Tucson, Arizona, USA.
Notes.— Lindroth adequately discusses this species but neglected to mention
this Casey synonym.
[Bembidion gratuitum Casey, see B. impotens ]
Bembidion hageni Hayward
Bembidium hageni Hayward, 1897:87. (New name for sexpunctatum LeConte
1852:186.)
Bembidium sexpunctatum LeConte, 1852:186. Holotype: MCZ #99, female. Type-
area: Colorado River, California, USA.
Notes. — This species belongs in the obtusangulum group; its members are found
in southwestern USA.
[Benbidion haustum Casey, see B. approximatum]
[.Bembidion henshawi Hayward, see B. flohri\
Bembidion hilare Casey
Bembidion hilare Casey, 1918:44. Lectotype: USNM #36873, male, here selected.
Type-locality: Cloverdale, Sonoma, California, USA.
Notes. — This species belongs in the mundum group; its members are found in
California.
Bembidion idoneum Casey
Bembidion idoneum Casey, 1918:90. Lectotype: USNM #36946, male, here se¬
lected. Type-area: Mendocino County, California, USA.
Bembidion obsequens Casey, 1918:90. New Synonymy. Lectotype: USNM #36947,
male, here selected. Type-area: Lake County, California, USA.
Bembidion efficiens Casey, 1918:90. New Synonymy. Lectotype: USNM #36949,
female, here selected. Type-locality: Mokelumne Hill, Calaveras County, Cal¬
ifornia, USA.
Bembidion continens Casey, 1918:91. New Synonymy. Lectotype: USNM #36948,
female, here selected. Type-area: Siskiyou County, California, USA.
174
PAN-PACIFIC ENTOMOLOGIST
Notes .—This species belongs in the dor sale group; its range extends throughout
the western United States in the mountainous regions.
[Bembidion imbelle Casey, see B. impotens ]
Bembidion impotens Casey
Bembidion impotens Casey, 1918:129. (New name for B. pictum LeConte 1848:
461, not Duftschmid 1812:218). See Lindroth, 1963, 1975.
Bembidion frugale Casey, 1918:130. New Synonymy. Lectotype: USNM #37006,
male, here selected. Type-locality: Fort Yuma, California, USA.
Bembidion imbelle Casey, 1918:130. New Synonymy. Lectotype: USNM #37007,
male, here selected. Type-locality: Dallas, Texas, USA.
Bembidion gratuitum Casey, 1918:130. New Synonymy. Lectotype: USNM #37004,
male, here selected. Type-area: Arizona, USA.
Bembidion virgatulum Casey, 1918:131. New Synonymy. Lectotype: USNM
#37003, male, here selected. Type-locality: Reno, Nevada, USA.
Bembidion indigens Casey, 1918:133. New Synonymy. Lectotype: USNM #37017,
male, here selected. Type-area: Federal District, Mexico.
Bembidionpullulum Casey, 1918:133. New Synonymy. Lectotype: USNM #37018,
male, here selected. Type-locality: Amecameca, Mexico.
Notes .—This highly variable species belongs in the complex versicolor group;
its members are found in the southwestern USA and Mexico north to Canada.
Detailed study may reveal cryptic species in the complex.
Bembidion incertum Motschulsky
Bembidium incertum Motschulsky, 1845:350. See Lindroth (1963:272).
Bembidion testatum Casey, 1918:30. New Synonymy. Lectotype: USNM #36842,
male, here selected. Type-area: Lake Tahoe, California, USA.
Notes. —Lindroth (1963) adequately discusses this species however he neglected
to deal with this Casey synonym.
[Bembidion indigens Casey, see B. impotens ]
Bembidion indistinctum Dejean
Bembidium indistinctum Dejean, 1831:67. See Lindroth (1963:361).
Bembidion derisor Casey, 1918:99. New Synonymy. Lectotype: USNM #36986,
female, here selected. Type-area: Eastern shore of San Francisco Bay, California,
USA.
Bembidion expositum Casey, 1918:101. Lectotype: USNM #36999, male, here
selected. Type-locality: Tehachapi Pass, California, USA.
Bembidion formale Casey, 1918:101. New Synonymy. Lectotype: USNM #36995,
male, here selected. Type-locality: San Francisco, California, USA.
Bembidion reconditum Casey, 1918:102. New Synonymy. Lectotype: USNM
#36996, male, here selected. Type-locality: San Diego, California, USA.
Bembidion franciscanum Casey, 1918:102. New Synonymy. Lectotype: USNM
#36991, male, here selected. Type-locality: Alameda, California, USA.
Bembidion ornatellum Casey, 1918:102. New Synonymy. Lectotype: USNM
#36992, male, here selected. Type-locality: Alameda, California, USA.
VOLUME 60, NUMBER 3
175
Notes.— This species, a highly variable one, belongs in the dorsale group; its
members occur in the southern half of California in coastal localities and coastal
hills as far east as the Tehachapi.
[Bembidion innocuum Casey, see B. californicum\
[Bembidion inquietum Casey, see B. texanum ]
Bembidion insulatum (LeConte)
Ochthedromus insulatus LeConte, 1852:186. See Lindroth (1963:362) for details.
Bembidion caliginosum Casey, 1918:119. New Synonymy. Lectotype: USNM
#37035, male, here selected. Type-locality: Douglas, Arizona, USA.
Notes.— Lindroth (1963) adequately discusses this species but did not mention
this Casey synonym.
[.Bembidion invidiosum Casey, see B. clemens]
Bembidion jacobianum Casey
Bembidion jacobianum Casey, 1918:101. New Synonymy. Lectotype: USNM
#36993, female, here selected. Type-locality: San Diego, California, USA.
Bembidion procax Casey, 1918:103. New Synonymy. Lectotype: USNM #36994,
male, here selected. Type-locality: San Diego, California, USA.
Notes.— This species belongs in the dorsale group; its members are found in
southern California.
Bembidion jamaicense Darlington
Bembidion jamaicense Darlington, 1934b:76. Holotype: USNM #75334, male.
Type-area: Jamaica, Greater Antilles.
Notes.— This species belongs in the vernale group and is found only on the
island of Jamaica.
Bembidion jucundum Horn
Bembidium jucundum Horn, 1895:230. Lectotype: CAS #1, male, here selected.
Type-locality: San Jose del Cabo, Baja California, Mexico.
Notes. — This species belongs in the spaeroderum group; its members are known
only from the type locality.
[Bembidion lassulum Casey, see B. graphicum ]
Bembidion latebricola Casey
Bembidion latebricola Casey, 1918:100. Lectotype: USNM #36998, male, here
selected. Type-area: “Arizona (probably southern),” USA.
Notes. — This species belongs in the aeneicolle group which is badly in need of
revision; the distribution of this species is unknown, even the type area, as given
by Casey, is obscure.
176
PAN-PACIFIC ENTOMOLOGIST
Bembidion laticeps (LeConte)
Lymnaeum laticeps LeConte, 1858:61. Holotype: MCZ #5562, female. Type-
locality: San Diego, California, USA.
Lymneops angusticeps Casey, 1918:169. New Synonymy. Lectotype: USNM
#46903, male, here selected. Type-locality: San Pedro, California, USA.
Notes. — This species belongs in the nigropiceum group (Erwin and Kavanaugh,
1980a); its members are found on the seashore along the California coast.
[Ochthedromus laticollis LeConte, see B. nubiculosum ]
Bembidion luculentum Casey
Bembidion luculentum Casey, 1918:122. Lectotype: USNM #37050, male, here
selected. Type-area: Indian River, Florida, USA.
Bembidion prosperum Casey, 1918:122. New Synonymy. Lectotype: USNM
#37049, female, here selected. Type-locality: Lake Worth, Florida, USA.
Notes. — This species belongs in the contractum group; its members are found
in the southeastern USA.
[Bembidium lugubre LeConte, see B. mexicanum ]
Bembidion macrogonum Bates
Bembidium macrogonum Bates, 1891:262. Lectotype: BMNH, male, here select¬
ed. Type-locality: Cordova, Vera Cruz, Mexico.
Notes.— This species belongs in the transfer sale group; it is presently known
only from Mexico.
Bembidion marinianum Casey
Bembidion marinianum Casey, 1924:29. Lectotype: USNM #36893, female here
selected. Type-area: Marin County, California, USA.
Notes.— This species belongs in the semistriatum group and is found in central
California.
[Bembidion mediocre Casey, see B. patruele ]
Bembidion mexicanum Dejean
Bembidium mexicanum Dejean, 1831:126. Lectotype: MNHP, male, here selected
by G. E. Ball and so labelled by him in 1972. Type-area: Mexico.
Bembidium lugubre LeConte, 1857:6. Holotype: MCZ #5511, female. Type-area:
“Valley of the Rio Grande.”
Bembidium stabile LeConte, 1879:508. New Synonymy. Lectotype: MCZ #5512,
male, here selected. Type-locality: La Veta, Colorado, USA.
Bembidion badiipenne Casey, 1918:60. New Synonymy. Lectotype: USNM #36913,
female, here selected. Type-area: Road between Fort Wingate and Jemez Springs,
New Mexico, USA.
Bembidion vafrum Casey, 1918:60. New Synonymy. Lectotype: USNM #36921,
male, here selected. Type-area: “Arizona (probably southern),” USA.
VOLUME 60, NUMBER 3
177
Bembidion canonicum Casey, 1918:61. New Synonymy. Holotype: USNM #36920,
male. Type-area: Arizona, USA.
Bembidion retectum Casey, 1918:61. New Synonymy. Lectotype: USNM #36923,
male, here selected. Type-locality: St. George, Utah, USA.
Notes.-This highly variable species belongs in the vernale group and its range
extends from Utah to Central America.
Bembidion modocianum Casey
Bembidion modocianum Casey, 1924:29. Lectotype: USNM #36882, male, here
selected. Type-area: Modoc County, California, USA.
Notes.— This species belongs in the nebraskense group; its members are known
thus far only from the type locality.
Bembidion nebraskense LeConte
Bembidium nebraskense LeConte, 1863:19. Holotype: MCZ#5506, female. Type-
area: “Nebraska, near the Rocky Mountains,” USA.
Bembidion denveranum Casey, 1918:64. New Synonymy. Lectotype: USNM
#36898, male, here selected. Type-area: Boulder County, Colorado, USA.
Bembidion tractabile Casey, 1918:64. New Synonymy. Lectotype: USNM #36901,
female, here selected. Type-area: Utah, USA.
Notes.— This species was adequately discussed by Lindroth (1963) but he did
not cover these Casey synonyms.
[Bembidion negligens Casey, see B. rapidum ]
Bembidion nevadense Ulke
Bembidium nevadense Ulke, 1875:811. (See Lindroth, 1963, for details.)
Bembidion viaticum Casey, 1918:65. New Synonymy. Lectotype: USNM #36884,
female, here selected. Type-area: New Mexico, USA.
Notes. — Lindroth adequately discusses this species but did not cover this Casey
synonym.
Bembidion nogalesium Casey
Bembidion nogalesium Casey, 1924:42. Lectotype: USNM #37020, female, here
selected. Type-locality: Nogales, Santa Cruz County, Arizona, USA.
Notes.— This “species” belongs in the patruele group which is in need of a
complete revision (see B. auxiliator).
Bembidion nubiculosum Chaudoir
Bembidium nubiculosum Chaudoir, 1868:244. (New name for O. laticollis Le¬
Conte, not Duftschmid, 1812:19.)
Ochthedromus laticollis LeConte, 1852:187, not Duftschmid. Lectotype: MCZ
#5523, male, here selected. Type-area: Colorado River, California, USA.
Bembidion daphnis Casey, 1918:120. New Synonymy. Lectotype: USNM #37036,
male, here selected. Type-locality: El Paso, Texas, USA.
178
PAN-PACIFIC ENTOMOLOGIST
Bembidion amnicum Casey, 1918:121. New Synonymy. Lectotype: USNM #37037,
male, here selected. Type-locality: Brownsville, Texas, USA.
Notes. — This variable species belongs in the dorsale group; its members are
found throughout the southwestern USA and northern Mexico.
[Bembidion obsequens Casey, see B. idoneum]
Bembidion operosum Casey
Bembidion operosum Casey, 1918:103. Lectotype: USNM #36960, female, here
selected. Type-locality: Santa Cruz, California, USA.
Notes.— This species belongs in the patruele group and is found in California.
Bembidion oppressum Casey
Bembidion oppressum Casey, 1918:40. Lectotype: USNM #36866, male, here
selected. Type-locality: Duncan’s Mill, Sonoma County, California, USA.
Notes. — This species belongs in the planiusculum group; its members are known
from central California in the coastal range.
[Bembidion ornatellum Casey, see B. indistinctum]
[Bembidion particeps Casey, see B. viridicolle]
Bembidion patruele Dejean
Bembidium patruele Dejean, 1831:69. (See Lindroth, 1963, for details.)
Bembidion mediocre Casey, 1918:107. New Synonymy. Holotype: USNM #36967,
male. Type-area: “Atlantic regions,” USA.
Bembidion editum Casey, 1918:125. New Synonymy. Lectotype: USNM #37041,
male, here selected. Type-area: “New Jersey (near the Delaware River),” USA.
Notes. — This species was adequately covered by Lindroth (1963) but he did not
discuss these two Casey synonyms.
Bembidion pedicellatum LeConte
Bembidium pedicellatum LeConte, 1857:6. Holotype: MCZ #5551, male. Type-
area: Lancaster County, Pennsylvania, USA.
Bembidion strigulosum Casey, 1918:150. New Synonymy. Lectotype: USNM
#37068, male, here selected. Type-locality: Washington, D.C., USA.
Bembidion fastidiosum Casey, 1918:150. New Synonymy. Lectotype: USNM
#37069, male, here selected. Type-locality: St. Louis, Missouri, USA.
Notes. — This species was adequately discussed by Lindroth (1963:383) who also
hinted at the synonymy of Casey’s names.
Bembidion perbrevicolle Casey
Bembidion perbrevicolle Casey, 1924:25. Lectotype: USNM #36826, female, here
selected. Type-area: Placer County, California, USA.
Notes. — This species belongs in the nebraskense group; its members are known
from the foothills of the Sierra Nevada of California.
VOLUME 60, NUMBER 3
179
Bembidion pernotum Casey
Bembidion pernotum Casey, 1918:62. Lectotype: USNM #36922, female, here
selected. Type-locality: Jemez Springs, New Mexico, USA.
Notes. —This species belongs in the transversale group; its members are found
in the American southwest and will likely be found also in Mexico.
Bembidion pimanum Casey
Bembidion pimanum Casey, 1918:98. Lectotype: USNM #36982, female, here
selected. Type-area: “near Benson, Arizona,” USA.
Notes.—This “species” belongs in the patruele group which is in need of a
complete revision.
[Bembidion placabile Casey, see B. cordatum]
Bembidion placeranum Casey
Bembidion placeranum Casey, 1924:28. Holotype: USNM #36865, female. Type-
area: Placer County, California, USA.
Notes. — This species belongs in the incertum group and is found in the foothills
of the Sierra Nevada of California.
Bembidion placitum Bates
Bembidium placitum Bates, 1878:602. Type-area: “Mexico, near the capital.”
Notes. — I was unable to locate the type series of this species in MNHP where
it should be, however, I saw 3 specimens “ex Salle” in BMNH that Bates mentions
(1882:151). The species belongs in the scudderi group and is found in Mexico.
Bembidion platynoides Hayward
Bembidium platynoides Hayward, 1897:78. Lectotype: MCZ #16287, male, here
selected. Type-locality: Pomona, California, USA.
Bembidion sedulum Casey, 1918:70. New Synonymy. Lectotype: USNM #36931,
male, here selected. Type-area: “Southern California,” USA.
Notes.— This species was adequately discussed by Lindroth (1963) but he ne¬
glected to mention this Casey synonym.
Bembidion portoricense Darlington
Bembidion portoricense Darlington, 1939:86. Holotype: MCZ #23507, male. Type-
locality: El Yunque, Puerto Rico, Greater Antilles.
Notes. — This species belongs in the vernale group and is known only from the
island of Puerto Rico.
Bembidion praecinctum LeConte
Bembidium praecinctum LeConte, 1879:509. Holotype: MCZ #5547, female. Type-
locality: Alamosa, Colorado, USA.
Bembidion veridicum Casey, 1918:152. New Synonymy. Lectotype: USNM #37066,
male, here selected. Type-locality: Elko, Nevada, USA.
180
PAN-PACIFIC ENTOMOLOGIST
Notes. — This species was adequately discussed by Lindroth (1963) but he merely
hinted at Casey’s synonymy.
[Bembidion procax Casey, see B. jacobianum ]
[Bembidion prosperum Casey, see B. luculentum ]
[Bembidion provanum Casey, see B. semipunctatum ]
[.Bembidion pullulum Casey, see B. impotens]
Bembidion quadrimaculatum (Linne)
Cicindela quadrimaculata Linne, 1761:211. (See Lindroth, 1963, for details.)
Bembidion tenax Casey, 1918:152. New Synonymy. Lectotype: USNM #37065,
male, here selected. Type-locality: Fort Wingate, New Mexico, USA.
Notes. — This species was adequately discussed by Lindroth (1963) but he did
not cover this Casey synonym.
Bembidion quadrulum LeConte
Bembidium quadrulum LeConte, 1861:340. Lectotype: MCZ #5498, male, here
selected. Type-area: “East of Fort Colville,” Oregon, USA.
Bembidion callidum Casey, 1918:50. New Synonymy. Lectotype: USNM #36878,
male, here selected. Type-locality: Truckee, California, USA.
Bembidion tritum Casey, 1918:50. New Synonymy. Lectotype: USNM #36879,
female, here selected. Type-area: Boulder County, Colorado, USA.
Bembidion aegrotum Casey, 1918:51. New Synonymy. Lectotype: USNM #36880,
male, here selected. Type-area: Colorado, USA.
Notes. — This species was adequately discussed by Lindroth (1963) but he did
not cover these Casey synonyms.
Bembidion rapidum (LeConte)
Ochthedromus rapidus LeConte, 1848:453. Lectotype: MCZ #5533, female, here
selected. Type-area: “Rocky Mountains” as given by LeConte, restricted by
Lindroth (1963:363) to Colorado Springs, Colorado, USA.
Bembidion docile Casey, 1918:126. New Synonymy. Lectotype: USNM #37055,
female, here selected. Type-area: “Arizona (probably southern),” USA.
Bembidion negligens Casey, 1918:127. New Synonymy. Holotype: USNM #37052,
male. Type-locality: El Paso, Texas, USA.
Bembidion fugitans Casey, 1918:127. New Synonymy. Lectotype: USNM #37053,
male, here selected. Type-area: “Arizona (probably southern),” USA.
Notes.— This species was adequately discussed by Lindroth (1963) but he ne¬
glected to cover these Casey synonymys.
[Bembidion reconditum Casey, see B. indistinctum]
[Bembidion relictum Casey, see B. adductum ]
[Bembidion remotum Casey, see B. clemens ]
VOLUME 60, NUMBER 3
181
Bembidion renoanum Casey
Bembidion renoanum Casey, 1918:72. Lectotype: USNM #36932, male, here
selected. Type-locality: Reno, Nevada, USA.
Notes. — This species belongs in the bimaculatum group and is known only from
the type locality.
[Bembidion retectum Casey, see B. mexicanum]
Bembidion rubiginosum LeConte
Bembidium rubiginosum LeConte, 1879:508. Holotype: MCZ #5541, female. Type-
locality: Garland, Colorado, USA.
Notes. — This species belongs in the muscicola group and is known only from
the type locality.
Bembidion rucillum Darlington
Bembidion rucillum Darlington, 1939:85. Holotype: MCZ #23506, male. Type-
locality: Loma Rucilla, Dominican Republic, Greater Antilles (8000').
Notes.— This species belongs in the vernale group and is known only from the
Dominican Republic.
Bembidion rupicola (Kirby)
Peryphus rupicola Kirby, 1837:53. (See Lindroth, 1963, for details.)
Bembidion aversans Casey, 1924:35. New Synonymy. Lectotype: USNM #36941,
female, here selected. Type-area: Mexico.
Notes.— Lindroth (1963) adequately covered this species but did not discuss
this Casey synonym.
Bembidion satellites Bates
Bembidium satellites Bates, 1884:291. Lectotype: BMNH, female, here selected.
Type-locality: Pena Blanca, Panama.
Notes.— This species belongs in the vernale group; its members are found in
upland habitats from northern Costa Rica to western Panama.
Bembidion scenicum Casey
Bembidion scenicum Casey, 1918:159. Lectotype: USNM #37079, male, here
selected. Type-area: Lake Tahoe, California, USA.
Notes.— This species belongs in the anguliferum group and is known only from
the type-area.
[Bembidium scintillans Bates, see B. aratum ]
Bembidion scopulinum (Kirby)
Peryphus scopulinum Kirby, 1837:53. (See Lindroth, 1963, for details.)
Bembidion bellulum Casey, 1918:71. New Synonymy. Holotype: USNM #36902,
female. Type-locality: Las Vegas, New Mexico, USA.
182
PAN-PACIFIC ENTOMOLOGIST
Notes.— This species was adequately discussed by Lindroth (1963:342) but he
neglected to mention this Casey synonym. This species is widespread and enough
material is presently available in museums to provide a good basis for a geographic
study.
[Bembidion scrutatum Casey, see B. grapei\
[Bembidion sedulum Casey, see B. platynoides]
Bembidion semifasciatum Say
Bembidium semifasciatum Say, 1834:438. Type: Lost. Type-locality: Mexico.
Notes.— This rather well-described species belongs in the dorsale group. Des¬
ignation of a Neotype should await a revision of the Mexican Bembidion or the
dorsale group.
Bembidion semiopacum Casey
Bembidion semiopacum Casey, 1924:39. Holotype: USNM #36965, female. Type-
area: San Joaquin County, California, USA.
Notes.— This species belongs in the complex patruele group which badly needs
revision; members of B. semiopacum are known from California’s central valley.
Bembidion semipunctatum Donovan
Bembidium semipunctatum Donovan, 1806:22. (See Lindroth, 1963:367, for de¬
tails.)
Bembidionprovoanum Casey, 1918:105. Lectotype: USNM #36966, female, here
selected. Type-locality: Provo, Utah, USA.
Notes.— This species was discussed adequately by Lindroth (1963) but he did
not mention this Casey synonym.
[Bembidium sexpunctatum LeConte, see B. hageni ]
Bembidion sparsum Bates
Bembidium sparsum Bates, 1882:151. Lectotype: BMNH, male, here selected.
Type locality: Oaxaca, Mexico.
Notes. — This species belongs in the dorsale group; its members are found in
low lying habitats and coastal localities from Sonora, Mexico south to Nicaragua
on the west coast, to Vera Cruz on the east coast, Jamaica, Hispaniola, and in
Colombia.
Bembidion sphaeroderum Bates
Bembidium sphaeroderum Bates, 1882:147. Lectotype: BMNH, male, here se¬
lected and previously labelled by G. Perrault. Type-locality: Jalapa, Mexico.
Notes. — This species belongs in the sphaeroderum group and is known only
from Mexico.
VOLUME 60, NUMBER 3
183
Bembidion sphaerulifer Bates
Bembidium sphaerulifer Bates, 1891:261. Lectotype: MNHP, male, here selected
and previously labelled by G. Perrault. Type-locality: Yentanas, Durango, Mex¬
ico.
Notes. — This species belongs in the sphaeroderum group and is known only
from Mexico.
Bembidion spretum Dejean
Bembidium spretum Dejean, 1831:70. Holotype: MNHP, female. Type-area: Mex¬
ico.
Notaphusfastidiosus LaFerte-Senectere, 1841:49, not Jacquelin du Val, 1851:563.
Bembidium apicale Jacquelin du Val, 1856:23, not Menetries, 1832:137.
Bembidicidium chevrolati Gemminger and Harold, 1868:409, new name for B.
apicale Jacquelin du Val.
Notes.— This species belongs in the dor sale group and is widespread in Mexico
and the Caribbean area.
[Bembidium stabile LeConte, see B. mexicanum]
[Bembidion strigulosum Casey, see B. pedicellatum]
Bembidion subaerarium Casey
Bembidion subaerarium Casey, 1924:31. Lectotype: USNM #36896, male, here
selected. Type-locality: Blue Lakes, Alpine County, California, USA.
Notes.— -This species belongs in the nigrum group and is known only from the
type locality.
Bembidion subangustatum Hayward
Bembidium subangustatum Hayward, 1897:83. Type-area: Arizona and New
Mexico, USA.
Notes. — The type series of this species could not be located in MCZ where it
should be deposited. From the description I judge it to be a member of the grapei
group.
Bembidion submaculatum Bates
Bembidium submaculatum Bates, 1882:149. Lectotype: BMNH, male, here se¬
lected. Type-locality: Oaxaca, Mexico.
Notes. — This species belongs in the incrematum group and is known from
Mexico.
[Bembidion temperans Casey, see B. triviale]
[Bembidion tenax Casey, see B. quadrimaculatum]
[Bembidium tessellatum LeConte, see B. conspersum]
[Bembidion testatum Casey, see B. incertum]
184
PAN-PACIFIC ENTOMOLOGIST
Bembidion texanum Chaudoir
Bembidium texanum Chaudoir, 1868:240. Lectotype: MNHP, male, by Lindroth,
1963. Type-area: Texas, USA.
Bembidion cogitans Casey, 1918:69. New Synonymy. Lectotype: USNM #36928,
male, here selected. Type-area: “?Indiana,” USA.
Bembidion inquietum Casey, 1918:67. New Synonymy. Lectotype: USNM #36924,
here selected. Type-locality: Jemez Springs, New Mexico, USA.
Notes.— This species was adequately discussed by Lindroth (1963:325) but he
neglected to mention these synonyms.
Bembidion tigrinum LeConte
Bembidium tigrinum LeConte, 1879:509. Holotype: MCZ #5521, female. Type-
area: “Southern California,” USA.
Notes.— This species belongs in the tigrinum group and is found along the
California coast.
[Bembidion tractabile Casey, see B. nebraskense\
[Bembidion tritum Casey, see B. quadrulum ]
Bembidion trixiale Casey
Bembidion trixiale Casey, 1918:134. Lectotype: USNM #37008, male, here se¬
lected. Type-area: Lake County, California, USA.
Bembidion temperans Casey, 1918:135. New Synonymy. Lectotype: USNM
#37009, female, here selected. Type-area: Lake County, California, USA.
Notes. — This species belongs in the patruele group and is found in central
California.
Bembidion turquinum Darlington
Bembidion turquinum Darlington, 1937:122. Holotype: MCZ #22492, male. Type-
locality: Pico Turquino, north side, Cuba, 4500'-6000', Greater Antilles.
Notes.— This species belongs in the xernale group and is found solely on Cuba.
[Bembidion xafrum Casey, see B. mexicanum]
[Bembidion xapidum Casey, see B. clemens\
[Bembidion xeridicum Casey, see B. praecinctum ]
Bembidion xernale Bates
Bembidium xernale Bates, 1882:149. Lectotype: BMNH, male, here selected. Type-
locality: Pena Blanca, Panama.
Notes. — This species belongs in the xernale group; its members are found in
upland habitats from Guatemala to Panama. It probably also occurs in Mexico,
but I saw no specimens from there.
[Bembidion xernula Casey, see B. constrictum]
[Bembidion xiaticum Casey, see B. nexadense ]
VOLUME 60, NUMBER 3
185
[Bembidion vinnulum Casey, see B. aratum]
[Bembidion virgatulum Casey, see B. impotens]
Bembidion viridicolle (LaFerte)
Notaphus viridicollis LaFerte-Senectere, 1841:48. (See Lindroth, 1963:374, for
details.)
Bembidion particeps Casey, 1918:124. New Synonymy. Lectotype: USNM #37039,
female, here selected. Type-area: “Arizona (probably southern),” USA.
Notes.— Lindroth (1963) adequately discussed this species but he did not cover
this Casey synonym.
Bembidion vividum Casey
Bembidion vividum Casey. 1884:66. Lectotype: USNM #37048, female, here se¬
lected. Type-area: Cape May, New Jersey, USA.
Notes. —This species belongs in the contractum group and is found along the
coast of the mid-northern Atlantic.
Bembidion vulcanium Darlington
Bembidion vulcanium Darlington, 1934:157a. Holotype: MCZ #19625, male. Type-
locality: Volcan Irazu, 2800-3000 m, south slope, Costa Rica.
Notes. —This species belongs in the vernale group and is found in higher ele¬
vation throughout central Costa Rica.
Bembidion vulpecula Casey
Bembidion vulpecula Casey, 1918:126. Lectotype: USNM #37047, female, here
selected. Type-locality: Brownsville, Texas, USA.
Notes. — This species belongs in the dorsale group and is known from the Amer¬
ican southwest and Mexico.
Bembidion wickhami Hayward
Bembidium wickhami Hayward, 1897:112. Lectotype: MCZ #16299, male, here
selected. Type-locality: Dunsmuir, California, USA.
Bembidion delectum Casey, 1918:44. New Synonymy. Lectotype: USNM #36872,
male, here selected. Type-locality: Gilroy Hot Springs, Santa Clara County,
California, USA.
Bembidion carlhi Erwin and Kavanaugh, 1981:37. New Synonymy. Holotype:
CAS#13657. Type-locality: Steamboat Creek at Steamboat Falls, 410m, Doug¬
las County, Oregon, USA.
Notes.— This species belongs in its own group and is known from Oregon and
California. While this paper was in press, Erwin and Kavanaugh (1981) revised
the erasum group of species and in so doing described as new, Bembidion carlhi
Erwin and Kavanaugh, 1981. Unfortunately, this species had already been de¬
scribed by Hayward (1897) as B. wickhami, therefore carlhi is a junior synonym
of Hayward’s name. Kavanaugh and I thank Dr. Kenneth Cooper for pointing
out this error.
[Bembicidium xanthostictum Gemminger and Harold, see B. conspersum]
186
PAN-PACIFIC ENTOMOLOGIST
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Proceedings of the Zoological Society of London, 1878, pp. 587-609.
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-. 1891. Additions to the carabideous fauna of Mexico, with remarks on some of the species
previously recorded. The Transactions of the Entomological Society of London, 1891, pp. 223-
278.
Blaisdell, F. E. 1902. The frons in Bembidium, with descriptions of new species. Proceedings of the
Academy of Natural Sciences of Philadelphia, pp. 70-79.
Brulle, A. 1838. Insectes. Pp. 17-56 in d’Orbigny, Voyage d’Orbigny dans l’Amerique meridionale
(1837-43). Paris, 6(2): 17-56.
Casey, T. L. 1884. Contributions to the descriptive and systematic coleopterology of North America.
Part II. Collins Printing House, Philadelphia, pp. 61-198.
-. 1918. A review of the North American Bembidiinae. Memoirs of the Coleoptera, 8:1-223.
-. 1924. Additions to the known Coleoptera of North America. Memoirs of the Coleoptera,
11:1-347.
de Chaudoir, M. 1868. Observations synonymiques sur les carabiques de l’Amerique septentrionale
et descriptions d’especes nouvelles de ce pays. Revue et Magasin de Zoologie, ser. 2, 20:239-
245.
Darlington, P. J. 1934a. Four new Bembidiini (Coleoptera: Carabidae) from Costa Rica and Colom¬
bia. Occasional Papers of the Boston Society of Natural History, 8:157-162.
-. 1934b. New West Indian Carabidae, with a list of the Cuban species. Psyche, 41:61-131.
-. 1937. West Indian Carabidae III. New species and records from Cuba, with a brief discussion
of the mountain fauna. Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey.”,
11:115-136.
-. 1939. West Indian Carabidae V. New forms from the Dominican Republic and Puerto Rico.
Memorias de la Sociedad Cubana de Historia Natural “Felipe Poey.”, 13:79-101.
Dejean, P. F. M. A. 1831. Species generale des coleopteres de la collection de M. le comte Dejean,
5:1-883.
Donovan, E. 1806. The natural history of British insects, &c., XI. London, pp. 1-100.
Duftschmid, C. E. 1812. Fauna Austriaca. Oder Beschreiburg Osterreichischen Insecten, fur ange-
hende Freunde der entomologie, 2:1-311.
Erwin, T. L. 1974. Studies of the Subtribe Tachyina (Coleoptera: Carabidae: Bembidiini). Supplement
A: Lectotype designations for new world species, two new genera, and notes on generic concepts.
Proceedings of the Entomological Society of Washington, 76(2): 123-155.
-, and D. H. Kavanaugh. 1980. On the identity of Bembidion puritanum Hayward (Coleoptera:
Carabidae: Bembidiini). The Coleopterists Bulletin, 34(2):241-242.
-, and-. 1981. Systematics and Zoogeography of Bembidion Latreille: I. The carlhi and
erasum groups of Western North America (Coleoptera: Carabidae: Bembidiini). Entomol. Scand.
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Gemminger, M., and E. von Harold. 1868. Catalogus coleopterorum hucusque descriptorum syn-
onymicus et systematicus, 1:424.
Gyllenhal, L. 1827. Insecta Suecica descripta, Classis I, Coleoptera sive Eleutherata, 1 (4):761 p.
Lipsiae.
Hatch, M. H. 1950. Studies on the Coleoptera of the Pacific Northwest. II: Carabidae: Bembidiini.
The Pan-Pacific Entomologist, 26(3):97-106.
Hayward, R. 1897. On the species of Bembidium of America north of Mexico. Transactions of the
American Entomological Society, 24:32-143.
Horn, G. H. 1895. Coleoptera of Baja California (Supplement I). Proceedings of the California
Academy of Sciences, ser. 2, 5:225-259.
Jacquelin de Val, P. N. C. 1851. De Bembidiis Europaeis. Annales de la Societe Entomologique de
France, serie 2, T. 9, pp. 441-576.
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Kirby, W. 1837. Insects. Coleoptera. In Richardson, Fauna Boreali-Americana; or the zoology of
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R. N., Josiah Fletcher publisher, London, 249 pp.
VOLUME 60, NUMBER 3
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de LaFerte-Senectere, F. T. 1841. Description de dix carabiques nouveaux du Texas et d’une espece
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4:173-474.
-. 1852. Descriptions of new species of Coleoptera, from California. Annals of the Lyceum of
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Mexican Boundary Commission, under Major W. H. Emory, U.S.A. Proceedings of the Acad¬
emy of Natural Sciences of Philadelphia, [vol. 10], 1858, p. 59-89.
-. 1861. New species of Coleoptera inhabiting the Pacific district of the United States. Pro¬
ceedings of the Academy of Natural Sciences of Philadelphia, 13:338-359.
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Smithsonian Miscellaneous Collections, No. 167, pp. 1-86.
-. 1879. The Coleoptera of the alpine Rocky Mountain regions—Part II. Bulletin of the United
States Geological and Geographical Survey of the Territories, 5(3):499-520.
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Part 3. Opuscula Entomologica supplementum 24, pp. 201-408.
-. 1975. Designation of Holotypes and Lectotypes Among Ground Beetles (Coleoptera, Ca¬
rabidae) Described by Thomas L. Casey. The Coleopterists Bulletin, 29(2): 109-147.
Linne, C. 1761. Fauna suecica. &c. 2. ed. Stockholm, pp. 1-578.
Menetries, E. 1832. Catalogue raisonne des objets de Zoologie recuellis dans un voyage au Caucase
et jusqu’aux frontieres actuelles de la Perse. St. Petersbourg, 1832. 4. 271 pp.
von Motschulsky, V. 1845. Observations sur le Musee entomologique de l’Universite imperiale de
Moscou. Article I. Bulletin de la Societe Imperiale des Naturalistes de Moscou, 18(4):332-388.
von Motschulsky, V. 1859. Coleopteres nouveaux de la Californie. Bulletin de la Societe Imperiale
des Naturalistes de Moscou, 32(2): 122-185.
Say, T. 1823. Descriptions of insects of the families of Carabici and Hydrocanthari of Latreille,
inhabiting North America. Transactions of the American Philosophical Society, ser. 2, 2:1-
109.
Ulke, H. 1875. Report upon the collections of Coleoptera made in portions of Nevada, Utah,
California, Colorado, New Mexico, and Arizona during 1871, 1872, 1873, and 1874. In Report
upon geographical and geological explorations and surveys west of the one hundredth meridian,
in charge of First Lieut. Geog. M. Wheeler . . . , zoology, Washington, 5:811-827.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 188-192
Biological Notes on the Bee Exomalopsis crenulata
Timberlake (Hymenoptera: Anthophoridae)
Frank D. Parker
Bee Biology & Systematics Laboratory, Agricultural Research Service, USD A,
Utah State University, UMC 53, Logan, Utah 84322.
Abstract.— Nests of a ground nesting anthophorid bee, Exomalopsis crenulata
Timberlake, are described. Females nested gregariously and made jug-like cells
attached singly to lateral burrows. The nesting site had been used for more than
1 year. This bee used mainly Eriogonum pollens to provision its nests. A de¬
scription of the nesting site, arrangement, and description of the cells is presented.
A parasitic bee, Hypochrotaenia {Micronomada) gutierreziae (Cockerell), was
found in some Exomalopsis cells.
Exomalopsis is a moderately sized group of New World anthophorid bees that
are primarily Neotropical in their distribution. In the United States these small
bees (2.5 to 7.5 mm) are grouped into 4 subgenera and 47 species. The nesting
habits of Exomalopsis are known only for three species (E. ( Anthophorisca ) chio-
nura Cockerell, Rozen 1957; E. ( Pheanomalopsis ) solani Cockerell, Linsley, 1954;
E. ( Anthophorula ) torticornis Cockerell, Hicks, 1936): all nest in the ground in
gregarious or communal nests. Flower visitation records have been summarized
by Timberlake (1980) who reports that some species of Exomalopsis visit a wide
variety of flowers (e.g., E. solani, E. similis Cresson) but others are oligoleges of
composites {E. chionura) and mallows {E . sidae Cockerell).
A recently described species, E. (. Anthophorula) crenulata Timberlake (Tim¬
berlake, 1980) was commonly collected in southern Utah on Eriogonum flowers
in the summers of 1979 and 1980. Many nests of this bee were discovered in the
fall of 1980. This paper describes the nest architecture, cells, nest associates, and
host plants of E. crenulata.
Nesting Site
The location of the Exomalopsis nesting site was 2.1 km E of Little Gilson
Butte, approximately 40 km N of Hanksville, Emery Co., Utah. Here, the San
Rafael Desert is marked by numerous moving crescent dunes that are surrounded
by large areas of stabilized sandy soil. The vegetation is diverse, and perennial
shrubs {Eriogonum, Chrysothamnus, Gutierrezia ) and forbs {Oenothera, Helian-
thus, Senecio ) are abundant.
The nesting site was discovered on the sides and the base of a low mound of
soil which is the refuse of an attempt at mining (Fig. 1). The soil composing the
mound was sandy beneath but on the surface and 15 cm beneath it was hard and
baked. Many female bees were observed entering the numerous nesting holes
VOLUME 60, NUMBER 3
189
found in this mound; activity was greater in the afternoon and more than one
female entered and exited the same hole.
Nest Architecture
The descriptions of the nests were made from clusters of cells that were exca¬
vated and studied later in the laboratory. Our brief visit and a sandstorm prevented
detailed nest excavations at the site. However, many details of the nesting biology
were obtained from the nest samples and the limited measurements taken at the
nest site.
Entrance. — The nest entrances were small (3 mm wide), oval-shaped openings
clustered close together in the hard surface (Fig. 2). None of the entrances were
covered by a turret.
Burrows. — The burrows were larger (3.5 mm wide) just beneath the surface and
consisted of a maze of similarly sized tunnels that were interconnected at the cell
level approx. 30 cm beneath the surface. The burrows were lined by finely masoned
soil of about 0.5 mm thickness with a glazed appearance; the linings were easily
distinguished from the surrounding soil layers (Fig. 3). The composition of the
glaze is not known, but it was not waxy because applied water droplets easily
impregnated the lining.
There was no pattern in burrow orientation since burrows were constructed on
all planes and ran in many directions. It is of interest to note that the nesting site
had been used in previous years; many old, exited, and refilled cells were found
adjacent to newly provisioned cells and old and new cells were connected to the
same lateral burrow.
Cells. — The cells diverged from the lateral burrows at an angle of 90° to 180°
(Fig. 4) and were attached singly to the lateral burrows. Since the burrows were
multi-directional, cell orientation to the surface plane was also multi-directional.
Examination of several cells at different stages of construction revealed that a
“typical” cell was made by first “roughing-out” a short elongate chamber off a
lateral burrow and then lining the walls with a smooth, shiny paste of fine-grained
soil (Fig. 4, arrow). The walls of the chamber were 0.5 to 1.5 mm thick. The
finished cells were jug-shaped with the elongated neck region receiving the cell
cap. The oval chambers that held the pollen loaf averaged 6.0 ± 0.3 mm in length
and 3.7 ± 0.2 mm wide at the widest point (n = 20). The neck of this chamber
was constricted to 3 mm and capped by a spiral plug of soil (with 4 coils) (Fig.
5). The cap was 0.7 ± 0.4 mm thick, U-shaped and smooth on the outer surface
(Fig. 3, arrow) and was connected to the burrow by a short (0.8 to 3.4 mm) plug
of compact soil. The inner surface of the cell was polished and shiny, but when
water droplets were placed on it artificially, they were quickly absorbed.
Nest Biology
Provisions.— All cells with old provisions were moldy and the shape of the
pollen loaf could not be clearly seen, but the loaf was cylindrical and attached to
the cell wall by a small “foot” (Rozen, 1957). The free end of the provision
projected towards the cell base. Egg placement was not observed, but several dead
early instar larvae were found on the top of the loaf and near the “foot.”
The pollens comprising the loaf and samples of pollen loads taken from foraging
females were 99% Eriogonum.
190
PAN-PACIFIC ENTOMOLOGIST
Figures 1-8. 1. View of nesting site on the mound of soil N of Hanksville, Utah. Hills in the
background are the San Rafael Swell. 2. Soil surface of nesting site and several oval nest entrances.
3. Jug-like cells of Exomalopsis. Note the smooth cell cap and the thin paper-like cocoon. 4. Single
cells of Exomalopsis attached to lateral burrows. Note the thick soil plug connecting the cell to the
lateral burrow. 5. Cell cap (ventral view) showing spiral construction. 6. Cell with coiled overwintering
larva. 7. Mature larva; note the subapical ventral swelling. 8. Cell with cocoon and fecal pellets.
VOLUME 60, NUMBER 3
191
Larvae. — The mature overwintering larvae were creamy-white and coiled into
a tight C-shape (Fig. 6). The larvae rested on the dorsal surface with the head
oriented either towards the cell cap or bottom. The mature larva had prominent
subapical swellings on the venter (Fig. 7).
Fecal pellets.— The fecal pellets were smeared against the cell walls in longi¬
tudinal irregularly shaped stripes. At the top of the cell several pellets were loose.
Cocoon.— A thin light tan to brown layer of silk was spun against the cell walls
and over the smeared fecal pellets (Fig. 3). The thin, silk layer readily tore apart.
The dome-shaped top of the cocoon was formed just beneath the cell cap (Fig.
8). The cocoon appeared to be made from a single layer of silk. All live larvae
were found in cocoons and also, all the old exited cells contained traces of cocoon
material.
Nest Associates
A dozen females of the parasitic nomadine bee, Hypochrotaenia ( Microno-
mada) gutierreziae (Cockerell), were collected as they flew about the nesting site.
Several of them entered nests and remained for short intervals. In addition two
Exomalopsis cells contained dead but unemerged adults of this parasite. Like
other species of this group of parasites (Parker, 1981), the parasitic larva did not
spin a cocoon and its fecal pellets were small and evenly scattered on the cell
walls.
A female Exomalopsis was observed to attack one of these parasitic bees as it
tried to enter a nest. In their brief struggle a forewing of the parasite was torn off
by the host bee.
Many of the Exomalopsis cells had been robbed by small ants that frequented
the nesting site. The ants made small tunnels among the cells and removed all
the provisions, fecal pellets, and parts of the cocoon.
Discussion
Some of the nesting habits of E. crenulata are similar to other Exomalopsis.
All species make jug-like earthen cells, but the formation and the lining of the
walls differ. Cells of E. crenulata were easily removed from the surrounding soil,
the cell walls were easily distinguished by the fine-grained texture, and the inner
cell walls were polished but not waxed; cells of E. chionura could not be removed
intact nor be distinguished from the surrounding soil, and the inner walls were
lined with wax and were nonpermeable to water droplets (Rozen, 1957). Cells of
E. crenulata were attached singly to lateral burrows but other Exomalopsis cells
were found in short chains (Claude-Joseph, 1926; Rozen, 1957). Although more
than one female of E. crenulata used the same nest entrance, there were several
hundred openings being used concurrently at the dense nesting site. This contrasts
with E. torticornis and E. solani where several hundred females have been found
using the same entrance (Hurd and Linsley, 1975). Floral affinities differ among
Exomalopsis (Timberlake, 1980) and E. crenulata is another example of an oli-
golectic species ( Eriogonum ).
It is of interest to note that species of Exomalopsis that use a common nest
entrance are polylectic and one, E. solani, is known to practice cooperative pro¬
visioning. Michener (1966) found that some E. solani females that were gathering
pollen were not ready to lay eggs. Thus, the sharing of a common nest entrance,
192
PAN-PACIFIC ENTOMOLOGIST
cooperative cell provisioning (Michener, 1966), and polylectic foraging may be
steps in the evolution of a social organization among some species of Exomalopsis.
The host association for the parasitic bee, H. gutierreziae, is the first host record
for this group of nomadine bees (species that have a flattened clypeus). Snelling
(pers. comm.) reported that the specimens from Exomalopsis nests were the
smallest he had examined. Perhaps this nomadine bee also parasitizes larger and
related bees such as Diadasia.
Acknowledgments
I am indebted to D. Veirs for the photographs used in this paper and to T.
Griswold for field assistance. Thanks are due to those persons who made helpful
suggestions on the manuscript—N. Youssef, Utah State University; E. G. Linsley,
Univ. of California-Berkeley; R. R. Snelling, Los Angeles County Museum; and
V. J. Tepedino of this laboratory. R. R. Snelling also identified the nomadine bee.
Literature Cited
Claude-Joseph, F. 1926. Recherches biologiques sur les Hymenopteres du Chili. Ann. des Sciences
Naturelles, Zoologie, 9:113-268.
Hicks, C. H. 1936. Nesting habits of certain western bees. Canad. Entomol., 68:47-52.
Hurd, P. D., Jr., and E. G. Linsley. 1975. The principal Larrea bees of the southwestern United
States. Smithsonian Contr. Zool. 193, 74 pp.
Linsley, E. G., J. W. MacSwain, and R. F. Smith. 1954. A note on the nesting habits of Exomalopsis
solani Cockerell. Pan-Pac. Entomol., 30:263-264.
Michener, C. D. 1966. Evidence of cooperative provisioning of the cells in Exomalopsis. J. Kansas
Entomol. Soc., 39:315-317.
Parker, F. D., and G. E. Bohart. 1981. Notes on the biology of Andrena ( Callandrena ) helianthi
Robertson. Pan-Pac. Entomol., 58:111-116.
Rozen, J. G., Jr., and C. D. MacNeill. 1957. Biological observations on Exomalopsis ( Anthophorula )
chionura Cockerell, including a comparison of the biology of Exomalopsis with that of other
anthophorid groups. Ann. Entomol. Soc. Amer., 50:522-529.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 193-212
A Systematic Study of the Genus Camelopsocus with
Descriptions of Three New Species
(Psocoptera: Psocidae ) 1
Edward L. Mockford 2
Department of Biological Sciences, Illinois State University, Normal, Illinois
61761.
Abstract.— Five species of Camelopsocus are diagnosed and keyed. All known
distribution records are given for each species. C. bactrianus, n. sp. and C. hiem-
alis, n. sp. from southern California and C. tucsonensis, n. sp. from southern
Arizona are described. The sexes of C. bactrianus and C. hiemalis, species which
occur together, are associated by temporal and character concordance. Eggs, the
embryonic oviruptor and surrounding structures, and the nymphal instars of C.
bactrianus and C. hiemalis are described, but it is not yet possible to distinguish
the two species in these stages. The two species are univoltine in southern Cali¬
fornia. Nymphs and adults occur from November to May. This phenology con¬
trasts to that seen in C. monticolus and C. similis in the Rocky Mountains and
desert ranges of New Mexico, where nymphs and adults appear in the summer.
Camelopsocus is probably most closely related to Oreopsocus and Atlantopsocus.
Since the original discovery of Camelopsocus (Mockford, 1965), three additional
species have been found. New distribution records of the two described species
have been made, and observations of several characters not previously considered
allow more precise determination of taxonomic relationships of the genus.
It is the purpose of this paper to describe the new species, to describe the egg,
postembryonic stages, and phenology of two of the new species, to present new
distribution records of the previously known species, to present a key to the species,
and to reassess the taxonomic relationships of the genus.
The new species are from southern California and southern Arizona. They differ
phenologically from the previously known species in that adults are restricted in
their occurrence to the winter and spring. Adults of the two previously known
species, which are found at relatively high altitudes in the Rocky Mountains, the
desert ranges of New Mexico and Arizona, and the mountains and high plains of
Mexico, occur only in summer in the northern part of this distribution (i.e., the
United States), although they have been taken from late March through late
December in Mexico.
Materials and Methods
This study is based on observation of 236 adult specimens of which 187 rep¬
resent the new species. In addition, 136 nymphs of C. bactrianus, n. sp. and C.
hiemalis, n. sp. were observed. Whole specimens were placed for drawing in a
1 Contribution No. 518, Bureau of Entomology, Division of Plant Industry, Florida Department of
Agriculture and Consumer Services, Gainesville, FL 32602.
2 Research Associate, Florida State Collection of Arthropods.
194
PAN-PACIFIC ENTOMOLOGIST
Table 1. Measurements (A), counts, and ratios for new species of Amelopsocus. Abbreviations are
explained in text.
Species
+ sex
FW
HW
F
T
t.
^2
cten
f.
fa
f 3
IO/D
PO
bactiranus $
4971
3695
683
1462
303
204
18
693
702
595
1.90
0.95
bactrianus <?
5168
1435
738
1559
406
216
17
772
803
681
2.05
1.01
bactrianus 9
343
675
1278
301
208
488
520
459
2.85
0.84
bactrianus 9
249
674
1257
281
206
456
508
436
2.60
0.94
hiemalis <3
5129
3885
697
1382
318
194
9
691
661
571
2.07
0.91
hiemalis 6
5100
3848
697
1543
388
202
9
796
697
638
2.06
0.93
hiemalis 9
229
675
1163
261
204
434
390
354
2.79
0.86
hiemalis 9
257
686
1179
255
193
457
446
407
tucsonensis^
5347
4050
725
1552
388
172
15
796
730
656
1.88
1.03
tucsonensis $
5150
3948
695
1414
342
179
2
671
592
543
2.07
1.00
tucsonensis 9
223
617
1069
230
182
375
331
300
2.60
0.84
tucsonensis 9
190
637
1186
245
185
438
399
372
2.66
0.84
well slide filled with glycerine over cotton strands and were drawn with aid of a
drawing tube on a dissecting microscope. Morphological observations were made
on slide preparations in Hoyer’s medium under a compound microscope. Color
observations were made on whole specimens in alcohol with direct light under a
dissecting microscope. Measurements were made with a filar micrometer. The
micrometer unit was 0.987 p.
The following abbreviations are used in the measurements (Table 1): FW =
length of forewing; HW = length of hindwing; F = length of hind femur (this and
all other leg measurements are made condyle to condyle); T = length of hind tibia;
t l5 t 2 = length of hind tarsomeres in order from base; cten = number of ctenidia
(comblike scales at bases of setae) on first hind tarsomere; f l5 f 2 , f 3 = length of
first three flagellomeres in order from base; IO/D = least distance between com¬
pound eyes divided by greatest antero-posterior diameter of eye in dorsal view;
PO = transverse diameter of eye divided by its greatest antero-posterior diameter
in dorsal view.
Association of Sexes
In dealing with the taxonomy of species that are highly sexually dimorphic, one
is faced with the problem of association of the sexes. The solution generally
involves finding characters that are trans-sexually distributed. In Camelopsocus
the dorsal abdominal turret of segment 5 marks both sexes of the genus, but if
two or more species occur at one locality, the investigator must search for trans¬
sexual characters at the species level. Data on physical and temporal cooccurrence
of the forms involved are also useful. The sexes of C. monticolus Mockford and
C. similis Mockford were readily associated by several trans-sexual color markings
differing between the species, and by presence of both sexes of one species at
several localities where the other species was absent.
Adults of two species have been taken in Riverside County, California. Both
species are currently known only from that county. For empirical analysis of
associational data, we will consider the males as forms numbers 1 and 2, and the
females as forms numbers 3 and 4.
Space-time associations are considered first. The four forms are known from
VOLUME 60, NUMBER 3
195
O Form #1
• Form #2
A Form#3
▲ Form#4
Scores, mark of abdominal terga
Figure 1. Distribution of four forms of Camelopsocus scored for two trans-sexual characters. The
nodes at which data may fall are indicated by points; at those occupied by data the symbol for the
form and the number of individuals are indicated. The characters and scoring procedures are discussed
in the text.
three localities, but at two of the localities only one form is known, so that only
the third offers associative data. Here collections of Camelopsocus have been
made on 14 dates over a three-year period. Two forms, the trans-sexual pair 1
and 3, are known only from that locality. On six dates trans-sexual combinations
were taken. Four of the six trans-sexual combinations were 1 + 3; one was 1 +
3 + 4, and one was 1 + 2 + 3 + 4. Therefore, the combination 1 + 3 was three
times as frequent as 1 + 4 and six times as frequent as 2 + 3. Each of the dates
of collection of trans-sexual combinations was separated from each of the others
by at least ten days.
Material from Pima County, Arizona, representing a third species of Came¬
lopsocus, bears on this question. The males of the Arizona species closely resemble
form 2, and the females closely resemble form 4.
A search of the material for trans-sexual characters in which the species differ
(i.e., 1 from 2, 3 from 4) revealed two: (1) distinctness of a longitudinal dark band
along the dorsal midline of the abdominal terga from the turret of segment 5 to
the clunium, and (2) extent of doubling of the median lineation of the postclypeus.
Both of these characters were variable, and each was scored into four classes—
196
PAN-PACIFIC ENTOMOLOGIST
Figures 2-5. Camelopsocus bactrianus, n. sp. 2. <5, terminal abdominal segments in open position,
lateral view. E = epiproct. Scale of Figure 3. 3. Terminal abdominal segments in closed position,
lateral view. Scale = 0.2 mm. 4. <3, lateral view, antennae beyond first flagellomeres not shown. Scale =
1 mm. 5. 2, lateral view. Scale of Figure 4.
the abdominal mark ranging from absent (1) to very dark (4) and the postclypeal
mark ranging from completely divided (1) to completely solid (4). The results are
shown graphically in Figure 1. They support an hypothesis of association of forms
1 and 3 as one species, 2 and 4 as a second. No trans-sexual characters were found
which tended to support the opposite association.
All of the available data consistently support the same hypothesis of association.
Therefore, I have designated forms 1 and 3 as one species, here called C. bactri¬
anus, n. sp., and forms 2 and 4 as a second, here called C. hiemalis, n. sp.
Diagnosis of Genus Camelopsocus
To the existing diagnosis (Mockford, 1965:3) should be added the following:
male epiproct in form of a chair (Fig. 2, E), capable of rotation through about
VOLUME 60, NUMBER 3
197
90°, the extreme positions involving 1) paraprocts, seat of epiproct, and distal
end of hypandrium exposed (open position, Fig, 2), and 2) paraprocts, seat of
epiproct, and distal end of hypandrium hidden (closed position, Fig. 3).
Camelopsocus bact nanus, New Species
Diagnosis. — Distal segment of maxillary palpus uniformly dark; first tarsomeres
medium brown throughout. Male wings without or with extremely pale general
tawny wash; pterostigma uniformly medium brown. Hypandrium (Fig. 9) with
median strap broadest at base, tapering toward distal end; a small field of tubercle-
based setae basal to median strap. Process of distal end of phallosome (Fig. 10)
short, truncated apically, heavily sculptured, flanked by two heavily pigmented
lobes of low, rounded relief. Region between arms of subgenital plate parallel¬
sided, near apex containing a pigmented band (Fig. 11); distal unpigmented area
of plate variable in its extent.
Male.—Morphology. General facies as in Figure 4. Wing venation (Figs. 6, 7)
typical of the genus. Epiproct (Fig. 8, slightly flattened) with margin most heavily
sclerotized on posterolateral edges; dorsal edge of epiproct (uppermost in figure)
slightly lobed laterally (seen on only one side in figure, the other side folded under
in mounting). Paraproct (Fig. 8) with field of24-34 trichobothria in two specimens
counted.
Color (in alcohol). Compound eyes gray. Ground color of remainder of head
and body creamy white; markings various shades of brown. Vertex of head with
fields of medium brown spots bordering median ecdysial line nearly to ocelli,
bordering median margin of each compound eye, and bordering entire posterior
margin of head; ocelli with dark brown pigment cups on their inner margins; a
medium brown transverse spot between ocelli and upper margin of postclypeus;
postclypeal striations medium brown; maxillary palpi dark brown, first three
segments colorless on their distal ends. Antenna with scape, pedicel, and basal
third of first flagellomere pale brown; first flagellomere darkening to dark brown
in distal two-thirds; remainder of antenna dark brown. Thorax dorsally medium
to dark brown on notal lobes, creamy white on sutures; pleura variegated creamy
white (from muscles showing through colorless cuticle) and medium to dark
brown, creamy white above leg bases, along meso-metathoracic junction, on a
large spot in ventral half of mesepisternum and on another in middle of metepi-
sternum. Coxae medium brown, paling to colorless distally, except mesocoxa
medium brown throughout; femora medium brown dorsally, creamy white ven-
trally; tibiae medium brown on inner faces, creamy white on outer faces with
brown spots at bases of some setae. Tarsi medium brown. Forewings (Fig. 6) clear
except pterostigma medium brown throughout, stigmasaum medium brown in
basal two-thirds, white in distal third; some veins bordered in medium brown as
indicated in figure; hindwings (Fig. 7) clear except with pale brown wash along
anterior margin. Abdomen with preclunial segments largely dull white variegated,
especially along sides, on turret, and along dorsal midline with purplish brown.
Clunium and external genitalia medium to dark brown on heavily sclerotized
areas, creamy white on membrane.
Female. —Morphology. Of typical micropterous facies of females of the genus
(Fig. 5). Ocelli represented by two minute pigment spots. Tarsi lacking ctenidia.
Subgenital plate (Fig. 11) with pigmented arms directed forward basally, bending
198
PAN-PACIFIC ENTOMOLOGIST
Figures 6-12. Camelopsocus bactrianus, n. sp. 6. <3, forewing. Scale = 1 mm. 7. 6, hindwing. Scale
of Figure 6. 8. <3, epiproct and paraproct. Scale = 0.1 mm. 9. 6, hypandrium. Scale of Figure 11. 10.
<3, phallosome. Scale of Figure 11. 11.2, subgenital plate. Scale = 0.1 mm. 12. 2, ovipositor valvulae.
D = distal lobe of third valvula. Scale = 0.1 mm.
VOLUME 60, NUMBER 3
199
abruptly outward near their bases, abruptly narrowed and back-curved at their
ends; region between bases of arms as described in diagnosis. Ovipositor valvulae
as in Figure 12.
Color (in alcohol; for pattern in general, see Fig. 5). Comparable parts as de¬
scribed in male except ground color clear white; first flagellomere pale brown
throughout; thoracic terga with broad white band along dorsal midline occupying
about half width of each tergum, the terga dark brown laterally. Forewinglets
white with two medium brown lines running from base to half or more length of
winglet, these ending separately or converging; Anal angle of forewinglet brown;
hindwinglets white. Preclunial abdominal segments variegated medium brown
and white, mostly white ventrally.
Holotype 6, allotype, 3 6 and 7 9 paratypes.— California: Riverside Co.: Sec. 32,
T7S, R1E, 9 km south of Sage on Hwy. 3, 25 March 1976, beating jojoba ( Sim-
mondsia chinensis Link), collector not indicated. These types will be deposited
in the collection of the Entomology Department, University of California, Riv¬
erside.
Additional paratypes (collector not known where not indicated).—Type locality,
26 February 1976, beating jojoba, 3 6, 6 9, J. D. Pinto; 18 March 1976, beating
jojoba, 2 3, 23 9; 31 March 1976, beating jojoba, 6 9; 4 April 1976, beating jojoba,
3 6, 8 9; 9 April 1976, beating jojoba, 16, 13 9; 5 May 1976, beating jojoba, 2 9;
21 May 1976, beating jojoba, 2 9; 20 December 1976, beating jojoba, 1 6; 25
January 1977, beating jojoba, 3 9; 2 March 1977, beating jojoba, 1 6, 3 9; 23 March
1977, beating jojoba 4 9; 29 March 1977, beating jojoba, 16 9; 23 March 1978,
1 6; 1 April 1978, beating jojoba, 2 9; 14 March 1979, beating miscellaneous
chaparral plants, 10 6, 15 9, E. L. Mockford and J. D. Pinto. Of these, the majority
will be deposited in the collection of the Entomology Department, University of
California, Riverside; 23 specimens (9 6, 14 9) will be deposited in my collection,
and one pair will be deposited in the Florida State Collection of Arthropods,
Gainesville.
Note. — Numerous nymphs were taken in association with these adults but can¬
not yet be distinguished from nymphs of C. hiemalis.
Camelopsocus hiemalis, New Species
Diagnosis. — Distal segment of maxillary palpus uniformly dark brown; first
tarsomeres uniformly medium to dark brown throughout. Male wings (Figs. 13,
14) with general tawny wash; pterostigma uniformly medium brown, paler than
in C. bactrianus. Hypandrium (Fig. 16) with median strap broadest before middle,
tapering toward both ends; a field of tubercle-based setae basal to median strap,
more extensive than in C. bactrianus. Process of distal end of phallosome (Fig.
17) tapering to a blunt point, flanked by two heavily pigmented lobes of low,
rounded relief. Region between arms of subgenital plate (Fig. 18) tapering to apex;
distal unpigmented area of plate continuing forward nearly to level of bases of
arms.
Male.—Morphology. Wing venation (Figs. 13, 14) typical of the genus. Epiproct
(Fig. 15, slightly flattened) with margin most heavily sclerotized along entire
posterior edge; dorsal edge of epiproct (uppermost in figure) not lobed laterally.
Paraproct with field of 21-24 trichobothria in two specimens counted.
Color (in alcohol). As described for C. bactrianus except brown markings of
head dark brown; antennae dark brown throughout. Brown markings of thoracic
200
PAN-PACIFIC ENTOMOLOGIST
Figures 13-19. Camelopsocus hiemalis, n. sp. 13. <5, forewing. Scale = 1 mm. 14. 3, hindwing.
Scale of Figure 13. 15. 3, epiproct. Scale = 0.1 mm. 16. 3, hypandrium. Scale = 0.1 mm. 17. 3,
phallosome. Scale of Figure 16. 18. 2, subgenital plate. Scale = 0.1 mm. 19. 2, ovipositor valvulae.
Scale = 0.1 mm.
VOLUME 60, NUMBER 3
201
terga dark brown throughout; mesoparapsidal sutures medium brown (creamy
white in C. bactrianus). Coxae dark brown; tibiae medium brown throughout.
Wings (Figs. 13, 14) differing from C. bactrianus as described in diagnosis. Pre-
clunial abdominal coloration in general darker.
Female.—Morphology. As described for female of C. bactrianus except subgen¬
ital plate (Fig. 18) with pigmented arms directed anterolaterad from their bases;
other details of subgenital plate as described in diagnosis. Ovipositor valvulae as
in Fig. 19.
Color (in alcohol). As described for female of C. bactrianus except tarsi generally
darker; forewinglet white only on margin, its interior medium brown, hindwinglet
white but in some individuals with a central brown spot.
Holotype 6, allotype, 2 <3 and 3 2 paratypes. — California: Riverside Co.: Sec. 32
T7S, R1E, 9 km south of Sage on Hwy. 3, 14 March 1979, beating miscellaneous
chaparral plants, E. L. Mockford and J. D. Pinto. These types will be deposited
in my collection.
Additional paratypes (collector not known where not indicated).—Type locality,
25 March 1976, beating jojoba, 1 2; 9 April 1976, beating jojoba, 1 2; 25 January
1977, beating jojoba, 1 2. 23 March 1977, beating jojoba, 1 2, 29 March 1977,
beating jojoba, 2 2, California: Riverside Co.: Sec. 25, T6S, R5E, south of Black
Hill on west side of Hwy. 74, 23 March 1978, beating jojoba, 1 2. California:
Riverside Co.: Dripping Springs Campground on Hwy. 79, Cleveland National
Forest, 14 March 1979, beating miscellaneous chaparral plants 6 2, E. L. Mockford
and J. D. Pinto. Of these, six will be deposited in the collection of the Entomology
Department, University of California, Riverside; two will be placed in the Florida
State Collection of Arthropods, Gainesville, and four will be placed in my col¬
lection.
Additional records. — California: Riverside Co.: Riverside, 15 February 1940,
on Artemisia californica (Hall) W. Watson, 1 <3, P. H. Timberlake; 24 February
1950, on Salvia apiana Jepson, 1 <3, P. H. Timberlake; 3 March 1952, on Salvia
apiana, 1 <3, P. H. Timberlake.
Camelopsocus monticolus Mockford
Camelopsocus monticolus Mockford, 1965:4
Diagnosis .—Distal segment of maxillary palpus ranging from medium brown
on all but dull white anterior face to uniformly dark brown; first tarsomeres ranging
from pale brown with darker base to uniformly medium brown. Male wings with
only extremely pale general tawny wash. Pterostigma with median longitudinal
dark line on paler brown background. Median strap of hypandrium (Fig. 20)
broadest before middle, tapering toward both ends. Process of distal end of phal-
losome (Fig. 21) tapering distally to a point, with scaly sculpturing basally, lacking
flanking lobes. Region between arms of subgenital plate (Fig. 22) parallel-sided
in distal third, lacking a central pigmented band.
Records.— Mexico: Durango: Hwy. 40, 38.6 km west of Durango, 16 July 1963,
beating pines (Pinus sp.), 4 nymphs, E. L. Mockford and F. Hill; Nuevo Leon: 4
km south of San Roberto junction, Hwy. 57, el. 1900 m, 20 December 1978,
beating branches and foliage of shrubs, 2 2, 1 nymph, A. N. Garcia Aldrete;
Oaxaca: 8 km southeast ofNochixtlan, Hwy. 190, 21 August 1973, from persistent
dead leaves of Seloa sp., 2 2, A. N. Garcia Aldrete; Zacatecas: 150 km north of
202
PAN-PACIFIC ENTOMOLOGIST
Figures 20-25. Camelopsocus monticolus Mockford and C. similis Mockford. 20. C. monticolus
$, hypandrium (left side damaged in mounting). Scale of Figure 21. 21. C. monticolus 6, phallosome.
Scale = 0.1 mm. 22. C. monticolus $, subgenital plate. Scale = 0.1 mm. 23. C. similis <3, phallosome.
Scale of Figure 22. 24. C. similis 6, hypandrium. Scale = 0.1 mm. 25. C. similis $, subgenital plate.
Scale = 0.1 mm.
Zacatecas, 11 August 1969, beating grass under Cenizo ( Leucophyllum texanum
Bentham), 2 2, 3 nymphs, A. N. Garcia Aldrete. united states: Arizona: Pima
Co.: Catalina Hwy. at Bear Canyon Picnic Area, el. 1830 m, 19 July 1975, beating
dead branches of oaks ( Quercus sp.), 2 nymphs, E. L. Mockford; Colorado: Larimer
Co.: Pingree Park, 20 August 1924, 1 nymph, C. R. Crosby; New Mexico: Catron
Co.: 1 mile northwest of junction of Hwy. 180 and 12 (type locality), 6 July 1963,
beating scrub oak ( Quercus sp.), pine ( Pinus sp.) and juniper ( Juniperus sp.), 1 8
(holotype), 2 2 (allotype and paratype), 29 nymphs, E. L. Mockford and F. Hill.
The types are in my collection.
Note: The forewing figure, Figure 7, in Mockford (1965) is erroneously labelled
C. similis and is really this species.
Camelopsocus similis Mockford
Camelopsocus similis Mockford, 1965:6
Diagnosis. — Distal segment of maxillary palpus with round white spot covering
most of anterior surface, remainder medium brown; first tarsomeres white except
VOLUME 60, NUMBER 3
203
Figures 26-30. Camelopsocus tucsonensis, n. sp. 26. <5, epiproct. Scale = 0.1 mm. 27. <3, hypan-
drium. Scale = 0.1 mm. 28. S, phallosome. Scale of Figure 27. 29. 2, subgenital plate. Scale = 0.1 mm.
30. 2, ovipositor valvulae. Scale = 0.1 mm.
medium brown at both ends (uniformly medium brown in a few males). Male
wings with only extremely pale general tawny wash. Pterostigma uniform light to
medium brown. Median strap of hypandrium (Fig. 24) widest basally, tapering
toward tip. Process of distal end of phallosome (Fig. 23) broad basally, tapering
toward pointed distal end, lacking flanking lobes. Region between arms of subgen¬
ital plate (Fig. 25) abruptly tapering to acuminate point distally.
Records.— Mexico: Coahuila: 30 km south of Saltillo, Hwy. 57, el. 1900 m, 29
March 1979, beating branches and foliage of Juniperus sp. on mountain side, 1
9, 2 nymphs, A. N. Garcia Aldrete and M. C. Herrera; Durango: 38.6 km west
of Durango, Hwy. 40 (type locality) 16 July 1963, beating oaks and small erica-
ceous shrubs, 1 6 (holotype), 7 9 (allotype and paratypes), 12 nymphs, E. L.
Mockford and F. Hill; 48 km west of Durango, el. 2438 m, 6 May 1961, 1 S, 3
9, 1 nymph, H. F. Howden and J. E. H. Martin; Mexico, 1.6 km west of Continental
Divide, Hwy. 136, el. 3300 m, 3 July 1962, beating broad-leaved shrubs, 2 S, 8
9, 18 nymphs, E. L. Mockford and F. Hill; Nuevo Leon: 3 km north of San
Roberto junction, Hwy. 57, el. 1800 m, 29 March 1979, beating foliage of Larrea
sp., 4 9, A. N. Garcia Aldrete and M. C. Herrera; Oaxaca: 8 km south of No-
chixtlan, Hwy. 190, 18 August 1968, sweeping grasses (Aristida sp.), 6 9, 11
nymphs, E. L. Mockford and A. N. Garcia Aldrete; San Luis Potosi: 30.5 km
204
PAN-PACIFIC ENTOMOLOGIST
north of Huizache, Hwy. 57, 25 July 1963, beating desert shrubs, 1 nymph, E.
L. Mockford and F. Hill, united states: Colorado: Chaffee Co.: Agate Camp
Ground in Gunnison National Forest, el. 2682 m, 21 July 1969, beating sagebrush
{Artemisia sp.), 6 nymphs, E. L. Mockford; Gunnison Co.: Hwy 50, 9.7 km east
of Blue Mesa Dam, el. 2316 m, 23 July 1969, beating oaks and small heath plants
on hillside, 2 2, E. L. Mockford. Wyoming: Teton Co.: Grand Teton National
Park, Hwy. 26, 22.5 km north of Jackson, 5 August 1966, beating Douglas firs
{Pseudotsuga taxifolia Britton), 1 2, 1 nymph, E. L. Mockford; Sweetwater Co.:
14.5 km west of Point of Rocks, 26 July 1976, prey of robberfly, 1 2, R. J. Lavigne
et al. The types are in my collection.
Camelopsocus tucsonensis f New Species
Diagnosis. — Similar to C. hiemalis, differing in following details: male epiproct
(Fig. 26) with dorsal edge (uppermost in figure) somewhat narrower (compare to
Fig. 15); median strap of hypandrium only approximately 1.3 x as broad at its
greatest breadth (near middle, Fig. 27) as at base, vs. approximately 2x in C.
hiemalis\ frame of phallosome (Fig. 28) decidedly wider distally than basally vs.
nearly of same width in C. hiemalis (Fig. 17); pigmented arms of subgenital plate
(Fig. 29) each approximately 1.2 x as broad at greatest breadth as at base, vs.
approximately 2.1 x in C. hiemalis. Female ovipositor valvulae as in Figure 30.
Morphology and color otherwise as described for C. hiemalis except dorsal
abdominal turret of male not darkly marked.
Holotype $, allotype, 9 S and 7 2 paratypes, and 11 nymphs: Arizona: Pima
Co.: 6.4 km west of Tucson on road to Desert Museum, 1 February 1979, beating
jojoba, J. D. Pinto. The holotype, allotype, 7 male and 5 female paratypes will
be deposited in the collection of the Entomology Department, University of Cal¬
ifornia, Riverside. One pair of paratypes will be deposited in the Florida State
Collection of Arthropods, Gainesville, Florida, and another pair of paratypes will
be deposited in the collection of the author.
Key to the Species of Camelopsocus
1. Macropterous, forewings at rest exceeding tip of abdomen (as in Fig. 4)
. . . males. 2
Micropterous, wings reduced to minute scales, not reaching posterior end
of thorax (as in Fig. 5) . . . females . 6
2. Hypandrium with at least a few tubercle-based setae anterior to base of
median strap (Fig. 16); distal process of phallosome flanked by two
heavily pigmented lobes of low, rounded relief (Figs. 10, 17) . .. 3
Hypandrium lacking tubercle-based setae anterior to base of median strap;
distal process of phallosome not flanked by pair of low, rounded lobes
(Fig. 21). 5
3. Median strap of hypandrium approximately parallel-sided most of its length
(Fig. 9); distal process of phallosome short, not appreciably longer than
its basal width, truncated apically (Fig. 10); wing membrane, except along
anterior veins of forewing and anterior margin of hindwing nearly clear,
with only a trace of tawny wash (Figs. 6, 7) . C. bactrianus, n. sp.
Median strap of hypandrium widest in middle, tapering to each end (Figs.
16, 27); distal process of phallosome decidedly longer than its basal
VOLUME 60, NUMBER 3
205
width, tapering to a blunt point apically (Figs. 17, 28); wing membranes
with a decided tawny wash (Figs. 13, 14) . 4
4. Median strap of hypandrium approximately 1.3 x as wide at its greatest
width as at base (Fig. 27); epiproct decidedly tapering from middle to
anterior (upper in Fig. 26) end. C. tucsonensis, n. sp.
Median strap of hypandrium approximately 2 x as wide at its greatest
width as at base (Fig. 16); epiproct broadly truncated at anterior end,
the truncated area much broader than in C. tucsonensis (Fig. 15 vs. Fig.
26) . C. hiemalis, n. sp.
5. Hypandrium (Fig. 24) with median strap widest immediately beyond base,
gradually narrowing to distal end; at distal end expanded as leftward-
directed process. Edge of frame of phallosome broad to sides of distal
process (Fig. 23) . C. similis Mockford
Hypandrium (Fig. 20) with median strap widest before middle, tapering
from there toward base and toward distal end; at distal end expanded
as bulbous region and leftward-directed process. Edge of frame of phal¬
losome narrow to sides of distal process (Fig. 21).
. C. monticolus Mockford
6. Abdomen with a major turret on segment 5 and a minor one on segment
4 (lesser turrets visible on segments 3 and 2 in some specimens) . 7
Abdomen with a major turret on segment 5 and no other turrets. 9
7. Pigmented arms of subgenital plate diverging from their bases; region
between arms pointed or slightly truncated distally but never parallel¬
sided and never including a distinct pigmented band (Figs. 18, 29) ... 8
Pigmented arms of subgenital plate parallel at bases, then diverging abrupt¬
ly; region between arms parallel-sided where arms parallel, containing a
distinct pigmented band (Fig. 11) . C. bactrianus, n. sp.
8. Pigmented arms of subgenital plate relatively narrow (Fig. 29) with rela¬
tively broad bases; greatest width of an arm approximately 1.2 x basal
width . C. tucsonensis, n. sp.
Pigmented arms of subgenital plate relatively broad (Fig. 18) with relatively
narrow bases; greatest width of an arm approximately 2.1 x basal width
. C. hiemalis, n. sp.
9. Pigmented arms of subgenital plate with anterior ends tapering and curved
backward; region between arms truncated posteriorly (Fig. 22).
. C. monticolus Mockford
Pigmented arms of subgenital plate with anterior ends truncated and di¬
rected laterally; region between arms acuminate posteriorly (Fig. 25)
. C. similis Mockford
Eggs, Postembryonic Development, and Phenology of
Camelopsocus bactrianus and C. hiemalis
Eggs.—On 14 March 1979, I visited the site near Sage, Riverside County,
California (see records of C. bactrianus and C. hiemalis) and collected adults of
C. bactrianus and C. hiemalis. Some gravid females were kept alive on twigs of
chaparral shrubs in order to obtain eggs. It was not possible to distinguish the
species in the field, nor was it possible to examine the live material with a dissecting
microscope for several days after their capture. By that time, oviposition had
206
PAN-PACIFIC ENTOMOLOGIST
already started, and because the live material included both species, the eggs of
the two were not separated. The eggs appear quite uniform, and it is likely that
those of the two species are closely similar.
Ovipositing females were held during their first four days of captivity in cotton-
plugged shell vials 24 mm in diameter and 94 mm in height. The vials were held
during that time in a plastic bag containing a small amount of moist paper towel
to assure a higher than ambient relative humidity. After four days, the vials, then
at my laboratory, were transferred to a desiccator jar at 75% relative humidity,
24°C daytime and 19°C nighttime temperature, and 17 hr light, 7 hr dark period.
Eight separate ovipositions were found, one consisting of a single egg, one of
three, one of four, two of eight, one of 11, one of 24, and one of 44 (x = 13). All
eggs are covered with crusty particles of debris of a color similar to that of the
surrounding bark. The debris coating of the grouped eggs covers the entire group,
so that they appear as a continuous clump. The larger clumps are oriented length¬
wise of the rather small (ca. 2.5 mm in diameter) twigs on which they were placed.
The one egg laid singly shows a ridge in the debris coating of its upper surface
along each side in the anterior half (i.e., the half containing the hatching orifice).
This egg measured 681 y in length and 393 ^ in greatest width. It tapers slightly
toward the hatching end.
After oviposition had been completed, the eggs were stored at ambient relative
humidity but otherwise under the conditions in which the females had been held.
I intended to return them to high relative humidity in late October, shortly before
the presumed eclosion time; however, I forgot about them until late November,
at which time they had already started to eclose. Most of the nymphs were dead,
probably of desiccation and/or starvation. The fact that some remained alive
under conditions of ambient relative humidity and low food supply suggests that
eclosion had not been occurring for much more than a week.
Embryonic exuviae .—These structures, extending from the openings of eclosed
eggs, were mounted on a slide in Hoyer’s medium for examination. The oviruptor
(Fig. 31, O) is a pointed, toothlike structure arising from the slightly depressed
middle of a platform-like region probably consisting of the entire anterior surface
of the embryonic head. A fossa occupies the morphologically ventral surface of
the oviruptor. Around and on the base of the oviruptor are numerous minute
denticles. A pair of slender brachiae (Pearman, 1928) (Fig. 31, Br) arise below
the oviruptor and diverge strongly. The platform to the sides of the depressed
middle rises as a pair of lobes, each one surmounted by an anteriorly directed
process. One of the processes is knobbed at the tip and the other acuminate.
Immediately dorsal to the oviruptor lies a flap-like structure with its base posterior
to the edge of the platform.
Nymphal instars.— At the Sage site, insects were sampled on jojoba shrubs at
regular intervals throughout the year from February 1976 through April 1978
(Pinto, pers. comm.). The data for these two species of Camelopsocus are com¬
bined for phenological consideration for the years 1976, 1977, and 1978. Nymphs
of C. bactrianus and C. hiemalis cannot yet be distinguished. Therefore, obser¬
vations on the identification of instars and on their phenology presented below
probably pertain to both species.
From the literature, it now seems safe to make two generalizations about de¬
velopmental patterns of nymphs of suborder Psocomorpha. First, in general, there
VOLUME 60, NUMBER 3
207
Figures 31-42. Camelopsocus bactrianus, n. sp. or C. hiemalis, n. sp. 31. Anterior end of embryonic
exuviae. O = oviruptor; Br = brachiae. Scale = 0.05 mm. 32. Second instar, sex unknown, dorsal view
of head and thorax. Scale = 0.5 mm. 33. Second instar, outlines of turrets of abdominal segments 4
and 5, lateral view. Scale of Figure 32. 34. Third instar 3, structure and scale as in Figure 32. 35.
Third instar 3, structure and scale as in Figure 33. 36. Third instar 2, structure and scale as in Figure
32. 37. Third instar 2, structure and scale as in Figure 33. 38. Fourth instar 3, structure and scale as
in Figure 32. 39. Fourth instar 3, structure and scale as in Figure 33. 40. Fourth instar 2, structure
and scale as in Figure 32. 41 and 42. Fourth instar 2 (two variants), structure and scale as in Figure
33.
are six nymphal instars. The only exceptions appear to be small, very neotenic
males of some Archipsocids (Mockford, 1957). Secondly, first instars have six
flagellomeres, while subsequent instars have eleven. The following literature ci¬
tations serve to establish both generalizations: Weber, 1931; Sommerman, 1943a,
1943b, 1943c, 1944; Mockford, 1957; Eertmoed, 1966; Dunham, 1972; Fahy,
1972; Garcia Aldrete, 1973; Heilbronn, 1975. Nymphs of Camelopsocus appear
208
PAN-PACIFIC ENTOMOLOGIST
to conform with both of these generalizations. Nymphs of six size classes, probably
representing all instars, were found in material from the Sage site.
The smallest nymphs were taken in the earliest sample during the period of
occurrence of these species. Because they have six flagellomeres, they are un¬
doubtedly first instars. They have no trace of wing pads, and they lack abdominal
turrets. These characters were also observed in nymphs newly emerged from eggs
held in the laboratory.
The second instar (Figs. 32, 33), in addition to its larger size and possession of
a minute turret of the fifth abdominal tergum, has eleven flagellomeres and small
wing pads. The first flagellomere is about equal in length with the pedicel. Because
only four specimens of this instar are present in the samples, it is not known if
the sexes can be distinguished at this stage.
Third instars (Figs. 34-37), in addition to larger size, show a more prominent
turret of the fifth abdominal tergum, a trace in some individuals of a turret of the
fourth, relatively larger wing pads, and the first flagellomere very slightly longer
than the pedicel. In this instar there is marked sexual dimorphism in size of wing
pads, those of the male being slightly more than twice the length of those of the
female.
Fourth instars (Figs. 38-42), in addition to being larger, have the abdominal
turrets more prominent, wing pads of the female very slightly larger but those of
the male more than twice as long as in the previous instar. The relative length of
the first flagellomere has increased to about 1.5 x the length of the pedicel.
In the fifth instar (Figs. 43-46), the abdominal turrets have changed very little
in prominence. Wing pads of both sexes have increased in relative size, the increase
in the male being about 1.8 x. The relative length of the first flagellomere has
increased to about 2 x that of the pedicel in the female and nearly 3 x that of the
pedicel in the male. Primordia of external genitalia have become visible in both
sexes.
Sixth instars (Figs. 47, 48, 49, 50) show a marked increase in prominence of
the abdominal turrets. Wing pads of the female have increased very little in relative
size but have oriented somewhat more downward. Wing pads of the male have
more than doubled in length. The relative length of the first flagellomere has
increased to slightly less than 3 x that of the pedicel in the female and slightly
more than 4 x that of the pedicel in the male. In the male, the mesonotal lobes
have become more prominent, and ocellar primordia are visible. The latter are
represented in the female by two small pigment spots which do not increase in
size in the adult. Primordia of the external genitalia are more distinct in both
sexes.
Phenology. — Table 2 shows seasonal occurrence of the nymphal instars and
adults at the Sage site. It would appear that eclosion starts to occur in early
November. Nymphal development is largely complete by mid-March. Adults,
which started to appear in mid-December, persist in small numbers until late
May.
There can be no doubt that these two species are univoltine. Most of the eggs
have been deposited by mid-April, and apparently no eclosion occurs until late
fall, after the return of cool weather. Such a regimen apparently permits the
nymphs and adults of these species to exist under approximately the same climatic
conditions as are experienced by those stages of C. monticolus and C. similis
VOLUME 60, NUMBER 3
209
Figures 43-50. Camelopsocus bactrianus, n. sp. or C. hiemalis, n. sp. 43. Fifth instar <3, dorsal
view of head and thorax. Scale = 0.5 mm. 44. Fifth instar <5, outlines of turrets of abdominal segments
4 and 5, lateral view. Scale of Figure 43. 45. Fifth instar 2, structure and scale as in Figure 43. 46.
Fifth instar 2, structure and scale as in Figure 44. 47. Sixth instar 8, structure and scale as in Figure
43. 48. Sixth instar 8, structure and scale as in Figure 44. 49. Sixth instar 2, structure and scale as in
Figure 43. 50. Sixth instar 2, structure and scale as in Figure 44.
210
PAN-PACIFIC ENTOMOLOGIST
Table 2. Seasonal distribution of life history stages of Camelopsocus bactrianus, n. sp. and C.
hiemalis, n. sp. at a site near Sage, Riverside County, California.
Number of individuals
1st _
in- 2nd 3rd 4th 5th 6th Adult
Date star instar instar instar instar instar (3b, 9b, 3h, $h)
16 November
5 3
1 9
10 December
1 3, 8 9
9 3, 1 9
2 3, 3 9
16-20 December
1 1 3, 3 9
8 3, 10 9
7 3, 6 9
3 3
1 3b
25 January
3 9
Of
oo
16 3, 9 9
3 3b, 1 9h
26 February
1 3, 1 9
3 3b, 6 9b
2 March
4 3, 2 9
1 3b, 3 9b
18 March
1 3, 2 9
2 3b, 33 9b
23-25 March
1 9
5 3b, 13 9b, 2 9h
29 March-
1 3
3 3b, 32 9b, 3 9h
1 April
4 April
3 3b, 8 9b
9 April
1 3b, 13 9b
5 May
2 9b
21 May
2 9b
during the summer in the mountains of Wyoming, Colorado, New Mexico, and
Arizona, and perhaps throughout much of the year in the mountains and high
plateaus of Mexico. Only the eggs of the southern Californian species are exposed
to the potentially desiccating conditions of summer and early fall.
Taxonomic Relationships of Camelopsocus
This genus clearly belongs to Subfamily Psocinae (sensu Badonnel, 1951).
Smithers (1972) made a cladistic analysis of subfamily Psocinae but based it only
on certain hypandrial characters and one venational character. The information
of other venational characters, the male epiproct, the phallosome, female genitalic
features, and wing marking characters was not used. Furthermore, several genera
could be interpreted as being out of place in Smithers’ scheme. Thus, Oreopsocus
and several species of Trichadenotecnum can be considered to have a median
strap-like structure on the hypandrium. An intriguing partial review of the prob¬
lems of relationships in the Psocinae is provided by Thornton (1961). It is evident
from these works that we are still groping in the dark with problems of generic
limits and general relationships within this subfamily.
The information-rich male epiproct in the Psocinae has not been adequately
studied, and I hope to deal with it in a separate work. The chair-shaped male
epiproct of Camelopsocus (Fig. 2) also occurs in Loensia (pers. observ.), Oreop¬
socus (Roesler, 1939, Fig. 10A), and several species of Ptycta (pers. observ.). In
each case, it shows concordance with parameres converging basally. I am unable
to assess the structure of the male epiproct in Atlantopsocus from the existing
literature, although Meinander’s figure (1973, Fig. 7C) suggests that it is not chair¬
shaped.
Other features suggest within the assemblage of genera Oreopsocus-Loensia-
Ptycta (at least in par t)-Atlantopsocus-Camelopsocus a closer proximity among
VOLUME 60, NUMBER 3
211
Oreopsocus, Atlantopsocus, and Camelopsocus. The forewing venation of Atlan-
topsocus and Camelopsocus shows in common a shallow pterostigma, a shallow,
elongate stigmasaum, and shape of the areola postica with the distal segment of
Cula directed distally. In Oreopsocus the pterostigma is not quite so shallow, the
stigmasaum not quite so long, and the distal segment of Cula directed not quite
so distally (Badonnel 1943, Fig. 93). The base of the phallosome in the form of
an apodeme is common to Camelopsocus and Atlantopsocus, but not to Oreop¬
socus. All three genera share a relatively unpigmented distal end of the subgenital
plate, a long, slender distal process of the second valvula, and a relatively large
distal (or ‘inner’) lobe of the third valvula (Fig. 12, D). Oreopsocus and Came¬
lopsocus share a markedly similar distal end of the phallosome: bilaterally sym¬
metrical with a pointed median process and lateral shoulders. In both genera the
median sclerotized strap of the hypandrium is bent abruptly to the left at its distal
end.
Acknowledgments
Most of the material of the new species described above was borrowed from
the Research and Teaching collection of the Department of Entomology, Uni¬
versity of California, Riverside. Mr. Saul Frommer, curator of the collection,
arranged the loans. Mr. Frommer also very kindly aided me in numerous ways
during my visit to Riverside, primarily to collect these insects, in March 1979.
Dr. John D. Pinto, also of the Riverside institution, collected the type series of
C. tucsonensis as well as some of the material of the other two new species. Dr.
Pinto accompanied me in the field during part of my 1979 visit to Riverside. My
collecting in Mexico in 1968 was supported by a National Science Foundation
grant, NSF GB7729, to Illinois State University. Dr. Alfonso N. Garcia Aldrete,
then a graduate student, accompanied me on the 1968 trip and did part of the
collecting. Dr. Garcia Aldrete, now a member of the research staff of the Instituto
de Biologia, UNAM, has collected additional Mexican material of Camelopsocus
subsequent to my 1968 trip and very kindly put it all at my disposal during my
visit to Mexico City in June 1979. The manuscript was reviewed by the publi¬
cations committee, Division of Plant Industry, Florida Department of Agriculture
and Consumer Services. To the above individuals and institutions I extend my
sincere thanks.
Literature Cited
B adonnel, A. 1943. Psocopteres. Faune de France, 42:1-164.
-. 1951. Ordre des Psocopteres. In P. Grasse (ed.), Traite de Zoologie, Paris, Vol. 10(2): 1301—
1340.
Dunham, R. S. 1972. A life history study of Caecilius aurantiacus (Hagen) (Psocoptera: Caeciliidae).
Great Lakes Entomol., 5:17-27.
Eertmoed, G. E. 1966. The life history of Peripsocus quadrifasciatus (Psocoptera: Peripsocidae). J.
Kansas Entomol. Soc., 39:54-65.
Fahy, E. D. 1972. The life history and immature stages of an exotic bark frequenting psocid from
southern Ireland (Psocoptera). Entomol. Gazette, 24:319-323.
Garcia Aldrete, A. N. 1973. The life history and developmental rates of Lachesilla pacifica Chapman
(parthenogenetic form) at four levels of temperature (Psocopt. Lachesillidae). Ciencia, 28:73-
77.
Heilbronn, T. D. 1975. Some aspects of the biology of the bark louse Graphopsocus cruciatus (L.)
(Psocoptera: Stenopsocidae). Entomol. Record, 87:132-136.
212
PAN-PACIFIC ENTOMOLOGIST
Meinander, M. 1973. The Psocoptera of the Canary Islands. Notulae Entomol., 53:141-158.
Mockford, E. L. 1957. Life history studies of some Florida insects of the genus Archipsocus (Pso¬
coptera). Bull. Florida St. Mus., 1:253-274.
-. 1965. A new genus of hump-backed psocids from Mexico and southwestern United States
(Psocoptera: Psocidae). Folia Entomol. Mex., 11:1-15.
Pearman, J. V. 1928. Biological observations on British Psocoptera. Entomol. Mon. Mag., 64:239-
243.
Roesler, R. 1939. Beitrage zur Kenntnis der Copeognathenfauna Deutschlands. Zool. Anz., 125:
157-176.
Smithers, C. N. 1972. The classification and phylogeny of the Psocoptera. Austral. Mus., Mem., 14:
1-349.
Sommerman, K. M. 1943a. Description and bionomics of Caecilius manteri, n. sp. (Corrodentia).
Proc. Entomol. Soc. Washington, 45:29-39.
-. 1943b. Bionomics of Lachesilla nubilis (Aaron) (Corrodentia, Caeciliidae). Canad. Entomol.,
75:99-105.
-. 1943c. Bionomics of Ectopsocuspumilis (Banks) (Corrodentia, Caeciliidae). Psyche, 50:53-
64.
-. 1944. Bionomics of Amapsocus amabilis (Walsh) (Corrodentia, Psocidae). Ann. Entomol.
Soc. America, 37:359-364.
Thornton, I. W. B. 1961. The Trichadenotecnum group (Psocoptera: Psocidae) in Hong Kong with
descriptions of new species. Trans. Royal Entomol. Soc. London, 113:1-24.
Weber, H. 1931. Die Lebensgeschichte von Ectopsocus parvulus (Kolbe 1882). Zeitschr. Wiss. Zool.,
138:457-486.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 213-218
The Life History of Paracantha cult avis (Coquillett) on
Wild Sunflower, Helianthus annuus L. ssp. lenticularis
(Douglas) Cockerell, in Southern California
(Diptera: Tephritidae)
G. L. C A VENDER AND R. D. GOEDEN
Department of Entomology, University of California, Riverside, California
92521.
Paracantha cultaris is one of several phytophagous insect species infesting the
flowerheads of wild sunflower, Helianthus annuus ssp. lenticularis, in southern
California. Heretofore, little was known about the life history of this tephritid.
The immatures and adults described herein were field-collected or reared in the
insectary of the Division of Biological Control, University of California, Riverside,
from flowerheads of wild sunflower collected in western Riverside and south¬
western San Bernardino counties during 1979-1980. A few specimens were in¬
sectary reared from the capitula of commercial sunflower, H. annuus var. ma-
crocarpus (deCandolle) Cockerell, collected in Riverside in 1980. Insectary
conditions were 26 ± 1°C, 30-60% R.H., and a 12/12-hr (light/dark) photoperiod.
Distribution and host plants. — Foote and Blanc (1963) described the range of
P. cultaris as Arizona, California, Idaho, Nebraska, Nevada, New Mexico, Oregon,
Texas, Washington, and Mexico, south to Costa Rica. Table 1 records rearings
of P. cultaris from flowerheads of wild and cultivated sunflowers from different
locations in Riverside and San Bernardino counties in southern California, coun¬
ties from which this tephritid was unreported in Foote and Blanc (1963). Wasbauer
(1972) listed Borrichia frutescens (L.) Decandolle, Cirsium occidentale (Nuttal)
Jepson, and “ Helianthus spp.” as other host plants of P. cultaris besides H. annus.
The second author (RDG) and D. W. Ricker have reared many P. gentilis Hering
from C. occidentale capitula collected in southern California in recent years (un-
publ. data), but never P. cultaris. Therefore, this rearing record for P. cultaris
from C. occidentale in southern California, first published in Foote and Blanc
(1963), is suspect.
Biology.—Egg. Newly laid eggs (Fig. la) were smooth, shiny-white, elongate-
ellipsoidal, and had a long thread-like pedicel bearing the micropyle at its apex.
Measurements of 7 eggs yielded the following means (±SE): body length = 0.93 ±
0.02 mm, body greatest width = 0.29 ± 0.007 mm, pedicel length = 1.41 ± 0.03
mm, pedicel width near middle = 0.043 ± 0.002 mm, pedicel width at the distal
end = 0.068 ± 0.002 mm. The total lengths of the eggs including their pedicels
averaged 2.30 ± 0.02 mm. The egg of Aciurina ferruginea (Doane) pictured by
Tauber and Tauber (1967) is very similar to P. cultaris, as is the egg of P. culta
(Wiedemann) described by Benjamin (1934).
Only very small (2-4 mm in diameter), immature capitula were chosen for
oviposition by P. cultaris in the field. When ovipositing, the females pierced the
phyllaries. The body of the egg was inserted between the developing disk florets
214
PAN-PACIFIC ENTOMOLOGIST
Table 1. Field collections of P. cultaris in southern California during 1979 and 1980.
County
Locality
Host
plant 3
Months
collected
Stage(s)
collected 11
Riverside
Moreno
1
II
a
Riverside
Riverside
m
II
p
Riverside
nr. Highgrove
1
III
p
Riverside
Moreno
1
III
a
Riverside
Riverside
m
III
P
Riverside
Riverside
1
III
P
San Bernardino
Colton
1
IV
P
Riverside
Moreno
1
IV
P
Riverside
Riverside
1
IV
P
Riverside
Rubidoux
1
IV
P
Riverside
Moreno
1
V
P
Riverside
Riverside
1
V
1
Riverside
Rubidoux
1
V
P
Riverside
Moreno
1
VI
P
Riverside
Riverside
1
VI
a
Riverside
Beaumont
1
VII
P
San Bernardino
Chino
1
VII
1, P
San Bernardino
Colton
1
VII
1, P
San Bernardino
nr. Fontana
1
VII
P
Riverside
Moreno
1
VII
P
Riverside
nr. Murrieta
1
VII
1, P
Riverside
Perris
1
VII
P
Riverside
Riverside
1
VII
P
Riverside
Rubidoux
1
VII
1, P
Riverside
Temecula
1
VII
P
San Bernardino
Colton
1
VIII
1, P
Riverside
Riverside
1
VIII
1
Riverside
Temecula
1
VIII
1, P
Riverside
Riverside
1
IX
P
Riverside
Riverside
1
X
1
Riverside
Moreno
1
XII
a
a 1 = H. annuus ssp. lenticularis, m = H. annuus var. macrocarpus.
b 1 = larvae, p = puparia, a = adults.
with the long axis of the egg perpendicular to the long axes of the florets. The egg
stalk projected from the oviposition scar in an outer phyllary (Fig. 2a). In mature,
infested heads, oviposition by P. cultaris usually was evidenced by long, narrow,
slitlike scars through the involucral bracts. The scars always ran lengthwise along
the bracts. The empty white chorions usually were found within the scars, with
their stalks projecting from the outermost or middle phyllaries (Fig. 2a). The
lengths of the oviposition scars in the inner surfaces of the phyllaries of 20, mature,
wild sunflower heads averaged 0.95 ± 0.04 mm, 1.60 ±0.14 mm, 1.91 ± 0.16
mm, in the outermost, middle, and inner bracts, respectively.
Females rarely oviposted in caged flowerheads in the insectary, and when they
did, oviposition was not observed. Paracantha cultaris females deposited an av¬
erage of 1.2 ± 0.11 (range: 1-3) eggs in 23 small, immature capitula collected in
the field. Although up to 3 eggs were deposited in a single capitulum, dissected
field-collected flowerheads rarely yielded more than 1 larva or puparium.
VOLUME 60, NUMBER 3
215
Figure 1. Paracantha cultaris: (a) egg, (b) 3rd instar, (c) 3rd instar posterior spiracles, (d) 3rd instar
cephalopharyngeal skeleton, (e) puparium.
Larva. — There were 3 instars, but the first 2 rarely were collected. The 3rd instar
(Figs, lb, 2b) is glossy, cream-colored, barrel-shaped, anteriorly conical, and pos¬
teriorly truncate. Five 3rd instars averaged 3.47 ± 0.33 mm in length and 2.30 ±
0.12 mm in greatest width. The posterior spiracles (Fig. lc) of the 3rd instars
have 3 pairs of well-developed stigmatic slits. The stigmatic area was depressed.
Eighteen stigmatic slits from 3 fully developed 3rd instars averaged 0.077 ± 0.001
mm in length and 0.022 ± 0.0005 mm in width. The cephalopharyngeal skeleton
(Fig. Id) of the 3rd instar was heavily sclerotized and darkened. The cephalo¬
pharyngeal skeletons of 3, 3rd instars yielded the following average measurements:
total length from tip of mouth-hook to end of dorsal wing plate = 0.71 ± 0.13
mm, width between the dorsal wing plate and pharyngeal plate = 0.32 ± 0.02
mm, length of neck region between dorsal bridge and the mouth-hook = 0.15 ±
0.007 mm, width of neck region = 0.07 ± 0.005 mm, length of mouth-hook =
0.11 ± 0.002 mm, length of elliptical holes in pharyngeal plates = 0.13 ± 0.006
mm, and width of holes - 0.06 ± 0.004 mm.
The young larva tunneled across the top of the receptacle through the bases of
numerous immature disk florets within an immature capitulum. The damaged
areas soon darkened. Mature capitula of wild sunflower infested with P. cultaris
usually were deformed, asymmetrical, and stunted and occasionally had clorotic
bracts and lighter yellow petals.
The older larva continued mining the upper part of the receptacle. In mature
216
PAN-PACIFIC ENTOMOLOGIST
Figure 2. Life stages of Paracantha cultaris on wild sunflower: (a) egg chorion projecting from an
oviposition scar in a phyllary, 20 x, (b) larva-infested head, 5 x, (c) puparium inside a malformed
head, 2 x, (d) adult female on flower petal, 6 x.
flowerheads, larval damage was evidenced as a premature browning of the nor¬
mally whitish, disk florets. The larvae produced much frass, which darkened and
usually was mixed with a sticky sap or resin. Cavities were excavated among the
ovaries by the last instars (Fig. 2b). The larvae pupated in these cells surrounded
by tightly packed frass. The disk florets became darkened and matted just above
the feeding areas.
Both P. cultaris and Trupanea bisetosa (Coquillett) commonly attack wild sun¬
flowers in southern California. Infestations of these 2 tephritids could be distin¬
guished in the field by the size of the infested heads, the deformity of P. cultaris-
infested capitula, and by the numbers of larvae or puparia in each flowerhead.
Trupanea bisetosa 3rd instars and puparia were found in wild sunflower heads
with disks averaging 21.8 ± 0.47 (range 11-29) mm in diameter (n = 47). Puparia
and 3rd instars of P. cultaris were found in wild sunflower heads with disks
averaging 18.0 ± 0.57 (range: 11-30) mm in diameter (n = 56). These means
were highly significantly different (P < 0.001, Mest). Trupanea bisetosa larvae
did not deform the flowerheads like P. cultaris. Paracantha cultaris- infested flow¬
erheads contained only 1.02 ± 0.01 (range: 1-2) larvae or puparia per flowerhead
( n = 96); whereas, T. bisetosa averaged 4.6 ± 0.33 (range: 1-15) larvae or puparia
VOLUME 60, NUMBER 3
217
per infested head (n = 83). These larval means also were significantly different.
Trupanea bisetosa and P. cultaris commonly were reared from flowerheads col¬
lected from the same stand of host plants, but only rarely were found together in
the same capitulum.
Pupa. — The puparia (Figs, le, 2c) are barrel-shaped, black, and shiny. The
cephalic end of the puparium is narrower than the rounded posterior end. Twenty
P. cultaris puparia dissected from sunflower heads averaged 4.65 ± 0.08 mm in
length and 2.44 ± 0.05 mm in width. Field-collected puparia were held in the
insectary for 11 to 13 days before the adults emerged. Emergence of the adults
from puparia usually occurred between 5 (lights-on) and 8 a.m. in the insectary.
Adult. — The adult (Fig. 2d) has a characteristic wing pattern that easily separates
it from other tephritid genera in California (Foote and Blanc, 1963). The longev¬
ities of 18 adult males fed a yeast hydrolyzate (2 parts) + honey (5 parts) mixture
averaged 51.1 ± 6.39 (range: 20-115) days; whereas, 16 females lived 57.3 ±
8.09 (range: 19-126) days. These means were not significantly different (Utest).
Paracantha cultaris presumably is multivoltine, as adults and puparia were
collected in southern California during most months of the year (Table 1 and
Foote and Blanc, 1963). This species probably mainly overwinters as adults but
also as immature stages in flowerheads on the occasional host-plant individuals
that escape killing frosts and continue to bloom during mild winters in southern
California.
Plant damage.— Tephritid infestations of wild sunflower heads can influence
the seed set and number of viable seeds per head. Twenty capitula infested with
P. cultaris had significantly lower numbers of seeds than 31 uninfested capitula
(106.3 ± 11.47 vs. 226.0 ± 6.83, respectively) as well as significantly smaller
diameters (19.8 ± 1.23 mm vs. 28.7 ± 0.92 mm, respectively). Twenty larvae
each destroyed an average of 34.14 ± 2.85 (range: 14-58) disk florets during their
development. About half the number of florets normally contained in uninfested
flowerheads from healthy plants were directly and indirectly lost to each P. cultaris
larva.
Adult behavior.— When caged singly, P. cultaris adults mostly remained mo¬
tionless except for their sponging mouthparts, which constantly moved in a pump¬
ing manner. The wings usually were held motionless slightly overlapped above
and near the abdomen. Occasionally, a female alternately arched each wing for¬
ward, twisting it when nearly perpendicular such that its ventral surface faced
anteriorly.
A behavioral pattern involving both males and females repeatedly was observed
in insectary cagings. When a male approached within ca. 3 cm of a female, he
began to sway back and forth and inch forward. With his mouthparts fully ex¬
tended towards the female, the male leaned his body and sidestepped as he zig¬
zagged towards the female. With each forward movement, he thrust both wings
forward in unison, so that they formed a straight line with their ventral surfaces
facing anteriorly. The male stopped when he neared the female. The female largely
remained motionless during this time with her rostrum pulsating. The male folded
his wings along his dorsum and kept his mouthparts fully extended. Sometimes
the female would turn and walk away, or turn away, then back again, just before
they next joined mouthparts. Occasionally, a female touched the extended mouth¬
parts of the male with her prolegs before extending her proboscis and joining his.
218
PAN-PACIFIC ENTOMOLOGIST
The ventral surfaces of their labella were placed flat together in close contact.
Both flies stood motionless for a few minutes with their mouthparts in contact.
The wings of the female were overlapped and at rest; those of the male, only
slightly outstretched. One pair of flies stood with their mouthparts joined for 6
minutes and 47 seconds. They then broke apart and shortly rejoined for an ad¬
ditional 6 minutes and 50 seconds.
Some tephritids are aggressive and use wing displays and other means to ward
off intruders (Bateman, 1972). Pritchard (1967) observed that aggressive behavior
had to be subdued in Rioxa pornia (Walker) in order for mating to take place.
He suggested that an elaborate courtship behavior was necessary to overcome the
female’s normal tendency to fight or take flight. Male R. pornia produced a mound
of foam on which the females fed during copulation. Perhaps the P. cultaris male
similarly transfers a pacifying substance or recognition token to the female when
their mouthparts are joined. This “kissing” behavior often was observed in in¬
sectary cagings but mating never was observed to follow. Batra (1979) reported
“kissing” between Euaresta festiva (Loew) adults during courtship as well as
territorial behavior.
Mortality factors.— Pteromalus (Habrocytus ) sp. (Hymenoptera: Pteromalidae)
was reared from P. cultaris puparia. Bracon nuperus Cresson (Hymenoptera:
Braconidae) was reared from P. cultaris larvae.
Acknowledgments
The parasites were identified by E. E. Grissell and P. M. Marsh, Systematic
Entomology Lab., Insect Identification and Beneficial Insect Introduction Insti¬
tute, % U.S. National Museum, Washington, D.C. The technical assistance of D.
W. Ricker is gratefully acknowledged. This report was derived in part from a
thesis submitted by the first author in partial satisfaction of the requirements of
the M.S. degree in Entomology, at the University of California, Riverside.
Literature Cited
Bateman, M. A. 1972. The ecology of fruit flies. Ann. Rev. Entomol., 17:493-518.
Batra, S. W. T. 1979. Reproductive behavior of Euaresta bella and E. festiva (Diptera: Tephritidae),
potential agents for the biological control of adventive North American ragweeds (Ambrosia
spp.) in Eurasia. J. New York Entomol. Soc., 87:118-125.
Benjamin, F. H. 1934. Descriptions of some native trypetid flies with notes on their habits. U.S.
Dept. Agric. Tech. Bull. No. 401:1-95.
Foote, R. H., and F. L. Blanc. 1963. The fruit flies or Tephritidae of California. Bull. Calif. Insect
Surv., 7:1-117.
Pritchard, G. 1967. Laboratory observations on the mating behavior of the island fruitfly, Rioxa
pornia. J. Aust. Entomol. Soc., 6:127-132.
Tauber, M. J., and C. A. Tauber. 1967. Reproductive behavior and biology of the gall-former,
Aciurina ferruginea. Can. J. Zool., 45:907-913.
Wasbauer, M. W. 1972. An annotated host catalog of the fruit flies of America north of Mexico
(Diptera: Tephritidae). Calif. Dept, of Agric., Bur. Entomol., Occas. Papers, 19:1-172.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 219-226
Predation upon Velvet Ants of the Genus Dasymutilla in
California (Hymenoptera: Mutillidae)
Donald G. Manley
Department of Entomology, Fisheries and Wildlife, Clemson University, Clem-
son, South Carolina 29631. Mailing address: Pee Dee Experiment Station, P.O.
Box 5809, Florence, South Carolina 29502.
Abstract. — Most diurnal velvet ants appear to be warningly colored, though
some appear to be cryptically colored. Field observations failed to implicate any
likely predators upon these organisms. In the laboratory, velvet ants of the genus
Dasymutilla Ashmead were exposed to predation by grasshopper mice, two species
of lizards, a mantid, and a scorpion. Results indicated that lizards may provide
one source of predation upon diurnal velvet ants.
Insects make up a large part of the diet of many animals, both vertebrates and
invertebrates. Since most velvet ants are brightly colored, conspicuous creatures,
many of which can be found running around on the ground in daylight, it seems
unlikely that they could exist without a means of defense. Furthermore, since
velvet ants are parasitic on other wasps and bees, they frequently are exposed to
the bites and stings of the latter. But the velvet ants are well adapted to their
ecological niche, as noted by Schmidt and Blum (1977) in their account of the
defensive adaptations of the velvet ant Dasymutilla occidentalis (Linnaeus).
As a first line of defense, these wingless wasps have a very heavy, deeply pitted
integument. The integument functions as a suit of armor, and can be penetrated
by the stings and bites of other insects only with great difficulty, if at all (Evans
and Eberhard, 1970).
The most significant defense mechanism exhibited by velvet ants is their sting.
The stinger is a modified ovipositor, and may be half as long as the entire body.
The sting of the velvet ant is very painful (Kirkpatrick, 1957). And, while the
sting is rarely used in an offensive role, it is used with remarkable effectiveness
as a defensive weapon.
Since most velvet ants are brightly colored, presumably they would draw the
attention of potential predators. However, since the females are such vicious
stingers, it is supposed that the bright coloration assumes the role of a warning
signal to prospective enemies. Due to warning coloration, it is assumed that natural
enemies learn more quickly to avoid these painful stingers (Metcalf and Flint,
1932).
The velvet ants exhibit another defensive characteristic, a file on the base of
the third visible abdominal tergite. When rubbed against a scraper (a sharp ridge
on the underside of the second visible abdominal tergite), a squeaking sound is
produced (Hinton et al., 1969). There has been speculation that this sound serves
both as a sexual signal and in a defensive role. Kirkpatrick (1957) stated that it
220
PAN-PACIFIC ENTOMOLOGIST
is reasonably certain that this sound increases the effectiveness of the warning
coloration.
Evans and Eberhard (1970:221) sum up this combination of defense mecha¬
nisms with the statement that . . an animal that discovers that the mutillid is
hard-bodied and a powerful stinger is likely to remember its brilliant and unusual
color pattern and to avoid it in the future.”
It is interesting to note that while the males of the family do not possess any
means of stinging, they too have the warning coloration. Through the use of
pseudoaposematic coloration, the males are receiving benefits from the females’
ability to sting (Hinton et al., 1969).
Another defensive measure apparently used in at least one species is protective
coloration, or mimicry. D. gloriosa (Saussure), a species which inhabits the deserts
of New Mexico, Arizona, and southern California, is covered with long, white
hairs which make it appear remarkably like thistle down. Unless moving, velvet
ants of this species are easily overlooked. Since most related forms are brightly
colored, and are not known to be heavily preyed upon by any other organism,
the reason for this mimicry is not known (Cockerell, 1895).
While D. gloriosa provides the most obvious example of mimicry within the
genus, there are other species that also may exhibit this defensive mechanism. It
seems odd that some species would exhibit protective coloration if there were no
predators to produce a selective pressure for this phenomenon. This study was
conducted to determine what organisms might be applying a selective pressure
for the protective coloration exhibited by a few velvet ants of the genus Dasy-
mutilla.
Materials and Methods
After location of specimens in numbers suitable for study (Manley, 1975), velvet
ants were observed under natural conditions. Field observations were made
throughout the summer of 1974. Due to their greater mobility, males were more
difficult to observe. Most observations of males were made in conjunction with
observations of females, the latter frequently luring males into the open. Once
females were located, individual specimens were followed and their activities
observed. Care was taken to remain at distances sufficient to insure that normal
behavior would not be altered.
A primary concern in this project was to subject the velvet ants to predation
by various animals which might prey upon them under natural conditions. In
order to do this, specimens were collected and kept alive in the laboratory. After
specimens were transported to the laboratory, they were transferred to plastic
aquariums approximately 10 cm by 18 cm at the bottom. Fine mesh screen was
placed over the containers to prevent escape. Sand was placed in the bottom and
water was administered through glass vials plugged with cotton. Specimens were
kept in the containers at room temperature (ca. 22°C) until used in the experiments.
Five potential predator species were utilized: two southern grasshopper mice
(Onychomys torridus (Coues)); a scorpion (Hadrurus arizonensis (Ewing)); two
lizards ( Sceloperus magister Hallowell and Dipsosaurus dorsalis (Baird and Gi¬
rard)); and a mantid of undetermined species.
All experiments were conducted by introducing the velvet ants to containers
with the “predator.” In the first experiment, the container was a glass jar and the
VOLUME 60, NUMBER 3
221
predator was a desert mantid. In the remaining experiments the container was a
20 cm by 30 cm glass aquarium covered by a coarse mesh wire screen. In each
experiment the predator was presented with a velvet ant. In all but the trials with
the lizards, the predators then were presented with other arthropods to determine
if a predatory drive existed.
The first laboratory experiment was undertaken on August 23, 1974. A female
D. clytemnestra (Fox) was placed in a glass jar with a desert mantid captured at
light the previous night. Following removal of the velvet ant, two moths and a
fly were introduced into the jar (one at a time) to test the predatory response of
the mantid.
The second experiment was undertaken on September 6, 1974, using two lab¬
oratory-born southern grasshopper mice. Since the mice had been fed only stan¬
dard commercial rat pellets, they were conditioned to live food for one week prior
to the experiment. They were fed two live crickets daily. After one week their
predatory response to live food seemed well-developed.
The mouse that had shown the keenest predatory response was transferred to
the glass aquarium and allowed to become acclimated to the new surroundings
for one hour. A female D. magnified Mickel then was introduced into the aquar¬
ium. In order to test the mouse’s predatory drive, a bumblebee then was placed
in the aquarium with the mouse. The first mouse then was removed and the
second mouse was placed in the aquarium with the velvet ant. A milkweed bug
(Oncopeltus fasciatus (Dallas)) was later introduced into the aquarium in order
to test predatory response. About an hour later, the first mouse again was intro¬
duced into the aquarium and confronted by a female D. magnifica, followed by
a female D. satanas Mickel.
The third experiment was conducted December 13-20, 1974. On December
13, a female D. magnifica was placed in the aquarium with a scorpion ( H. ari-
zonensis) that had been captured in August 1974. Observations were made, pe¬
riodically, from December 13 through December 20. Cockroaches and mealworms
subsequently were placed in the aquarium with the scorpion in order to test
predatory response.
The final experiment took place May 14-19, 1975. In that experiment, two
lizards were used which had been caught May 11, 1975 and which had not been
fed since capture. On May 14 the first lizard ( D. dorsalis ) was placed in the
aquarium and allowed one hour to acclimate to its new surroundings. At that
time a thread was tied around a female of D. nocturna Mickel in such a way that
its movements were not restricted. It was then lowered in front of the lizard. The
second lizard ( S. magistef) was subsequently placed in the aquarium and the
procedure was repeated.
On May 16, the procedures were repeated, again using both lizards. This time
each lizard was presented first with a female of D. nocturna, then with a female
of D. clytemnestra. A final test was conducted on May 19 by lowering another
D. nocturna on a thread in front of the Sceloperus lizard.
Results
While most taxa of terrestrial animals include at least some insectivorous mem¬
bers, at no time was an attempted capture of a velvet ant observed under natural
conditions. The opportunity for predation by rodents or reptiles existed frequently.
222
PAN-PACIFIC ENTOMOLOGIST
It was not uncommon to see female velvet ants enter a rodent or reptile burrow
during the course of their explorations. On a number of occasions in which a
velvet ant was seen entering a small burrow, a lizard was seen exiting the burrow
after the velvet ant’s entry. Thus, it would seem that the lizards had no great
desire to confront the velvet ants under natural conditions. Lizards involved in
those observations were not identified.
There were instances in which velvet ants became entangled in spider webs
that guarded the entrance to burrows. On those occasions, the velvet ants quickly
and easily freed themselves without any interference from the occupants of the
webs (which appeared to be recent and presumably occupied).
Possible enemies of the velvet ants under field conditions appeared to be true
ants. Columns of ants going to or from their nest were frequently encountered by
the velvet ants. On every occasion, they would cross the column with great haste.
On one occasion a female D. gloriosa was dropped near the entrance to one of
these ant nests and was quickly killed by the ants. It seems ironic that the velvet
ant, whose integument is impenetrable to the stings of other wasps and bees, was
so easily overcome by ants.
When placed with the mantid, the D. clytemnestra was not attacked. Soon after
introduction of the velvet ant into the jar, the mantid appeared ready to strike,
but then backed off and showed no further interest in the mutillid. The mantid
eagerly devoured two moths and a fly that were then placed in the jar.
When the velvet ant was placed in the aquarium with the first mouse, the latter
immediately attacked the velvet ant. It picked the velvet ant up with its front feet
and attempted to decapitate its prey in the manner characteristic of this species
(Cyr, 1970). For its efforts, the mouse received a sting in the mouth. Instinct to
attack movement caused the mouse to repeatedly attack the velvet ant. Each time
the attack was terminated as the mouse was stung in the mouth. This lasted for
approximately five minutes with the mouse becoming increasingly hesitant to
attack. Pain apparently overcame instinct and attacks ceased. At that time the
mouse appeared groggy and glassy-eyed. The struggle between instinct and pain
resulted in considerable displacement behavior on the part of the mouse, including
rolling and digging in the sand. When the bumblebee was confined with the mouse,
the bee was captured, decapitated, and eaten.
When the second mouse was placed in the aquarium, it immediately attacked
the velvet ant that was again placed in the aquarium. It, too, was stung in the
mouth. The predatory drive was apparently not as strong in the second mouse
and, after being stung once, it ceased to attack. Again, displacement behavior was
observed. Females of D. satanas and D. gloriosa were subsequently introduced
into the aquarium. Though different colors from D. magnifica, these two latter
species apparently looked similar enough to D. magnifica that no attacks upon
them were observed. When a milkweed bug was introduced into the aquarium,
it was quickly captured, decapitated, and eaten.
When the first mouse again was placed in the aquarium, it seemed to have
regained its original vigor and predatory drive. Again, it was stung repeatedly in
the mouth. The mouse finally assumed what appeared to be a “do or die” attitude
and, in spite of numerous stings, kept the velvet ant in its mouth long enough to
cause injury. Once the velvet ant had been injured, it was decapitated, after which
the mouse attempted to eat its prey. For whatever reason, the mouse was not
VOLUME 60, NUMBER 3
223
successful in consuming its subdued victim. The dead velvet ant was left with the
mouse for a period of one week following the experiment and was not consumed
during that time.
After the mouse had attempted, and failed, to eat the velvet ant, the dead
specimen was replaced by another live velvet ant, a female D. satanas. The mouse
again attacked repeatedly and was stung repeatedly. The mouse soon appeared
groggy and all predatory activity ceased. Displacement behavior was again ob¬
served, but no further advances were made toward the velvet ant during the next
fifteen minutes.
While the velvet ant was confined with the scorpion, whenever it passed nearby,
the scorpion would attempt to grab it. Whenever captured, the velvet ant was
released almost immediately. It appeared that the scorpion was stung upon the
pedipalp in order to cause release. The scorpion, however, showed no reaction
other than release of the velvet ant. This sequence of events occurred repeatedly
throughout the day.
When next observed, on December 16, the velvet ant was still alive. Similar
displays continued to take place through December 20, after which time the velvet
ant had spent one week with the scorpion. From observation, it appeared that a
key to the scorpion’s inability to capture the velvet ant was that pressure applied
by the pedipalp of the scorpion seemed unable to injure the velvet ant. Cock¬
roaches and mealworms placed in the aquarium with the scorpion were quickly
killed and consumed.
When the velvet ant was lowered in front of the first lizard ( D. dorsalis ), the
lizard made no movement toward the velvet ant until it had been annoyed (by
the presence of the velvet ant on and in front of its face) for about five minutes.
After apparently being stung, the lizard made two or three snaps at the velvet ant,
but did no apparent damage. When the procedure was repeated with the second
lizard (S. magister), it did not attack the velvet ant even after five minutes of
annoyance. When repeated on May 16, the results were the same. The D. dorsalis
finally snapped at the velvet ant, but did no damage. The S. magister did not
attack the velvet ant.
On the final trial, the Sceloperus first showed no inclination to attack the D.
nocturna, even after considerable annoyance. Finally, however, it snapped at the
D. clytemnestra, nearly severing the velvet ant between the head and thorax. The
velvet ant was not eaten by the lizard, but died shortly thereafter from injuries
that it had sustained. The D. nocturna was left in the aquarium with the Dipso-
saurus until May 19 and was not injured. Later that day, another D. nocturna
was lowered on a thread in front of the Sceloperus. After only a few seconds the
lizard crushed and consumed the velvet ant in one quick movement. The lizard
exhibited no difficulty in consuming the velvet ant, and the sounds of crushing
of the tough integument were easily heard.
Discussion
This study was stimulated by the apparent fact that a few velvet ants are
cryptically colored while the majority are brightly colored. The most striking
example of this is D. gloriosa, which Cockerell (1895) described as resembling
thistle down. Cockerell related that, unless moving, these velvet ants were easily
overlooked.
224
PAN-PACIFIC ENTOMOLOGIST
In this study, it was noted that all specimens of D. gloriosa that were observed
were found in the vicinity of creosote (Larrea divaricata Cav.), and that these
velvet ants showed a remarkable resemblance to the fruit of the creosote. While
this made the velvet ants easy to overlook when at rest, it was discovered that
their movements made them equally difficult to observe. These velvet ants moved
in a jerky manner which resembled a creosote fruit being blown in the wind. More
than once a blowing creosote fruit was mistaken for one of these velvet ants. It
is not known how many velvet ants were overlooked as they sat motionless.
Another example of this cryptic coloration is seen in D. clytemnestra. This
small, pale yellow velvet ant was found most frequently in the vicinity of dry,
pale yellow grass. Unless moving, it was most difficult to see. On one occasion,
a specimen that was being observed stopped in the grass. Though the position of
this individual was known, it was almost impossible to distinguish the velvet ant
from its surroundings at a distance of more than a few centimeters.
Still another example is D. nocturna, a predominately pale species that is found
upon sand dunes. This species is very inconspicuous upon the light-colored sand
and, like D. gloriosa, is very likely to be overlooked unless moving.
Since cryptic coloration apparently exists in some velvet ants, the obvious
question is why it exists. In order for cryptic coloration to be an advantage, the
potential predator must possess color vision. Using this as a criterion, it may be
possible to reduce the number of potential predators. For example, most mammals
do not have the ability to see color. This is particularly true of nocturnal mammals
for which the light necessary for color vision is usually not available. And, since
most desert mammals are nocturnal, it seems unlikely that they would provide
the predatory pressure necessary to favor cryptic coloration by velvet ants.
Grasshopper mice were chosen as an example of mammalian predators because
they are small, voracious, and highly insectivorous predators (Cyr, 1970). Like
most desert mammals, grasshopper mice are nocturnal. Therefore, it is quite
possible that they would rarely, if ever, encounter these diurnal velvet ants. And
even though they may dig in the sand for prey, the habits of the velvet ants are
not well enough known to establish whether they might be encountered by this
potential predator.
The laboratory experiment with the two grasshopper mice was repeated four
times. On three trials the mice were subdued by stings of the velvet ants. On one
trial the velvet ant was killed. It seemed very doubtful, however, that under natural
conditions, the grasshopper mice would have taken as much punishment as they
did before retreating. Under conditions other than the confined laboratory situ¬
ation, the grasshopper mice probably would seek an easier meal.
One observation that remains unexplained at this point is that the grasshopper
mouse failed to eat the velvet ant even after it had been killed and decapitated.
Velvet ants have a very tough integument. Grasshopper mice, however, are ac¬
customed to eating beetles of the family Tenebrionidae (including Eleodes sp.),
and these beetles also have very tough integuments. It is possible that the dense
pubescence may have made the velvet ant unpalatable to the mouse.
Perhaps other arthropods prey upon the velvet ants. It is known that many
arthropods have color vision. Their vision, however, is presumably quite different
from our own. Therefore, it is not known if cryptic coloration (as we see it) is
cryptic to potential arthropod predators. The fact that many of the potential
VOLUME 60, NUMBER 3
225
predators are nocturnal (including scorpions and solpugids) raises further doubts
as to their predatory potential upon diurnal velvet ants.
The fact that the mantid started to move toward the velvet ant and then stopped
is a very interesting point. It seems very unlikely that this nocturnal predator had
ever before seen a velvet ant. Yet it seemed to recognize it as a source of danger.
Perhaps stridulation by the velvet ant served to warn this potential predator that
persistence would result in a sting. At any rate, the mantid seems an unlikely
predator of velvet ants under natural conditions.
The scorpion, too, had little success preying upon the velvet ant, even under
the confining laboratory conditions. And it, too, is nocturnal.
It would have been interesting to have placed a velvet ant in confined conditions
with a large solpugid (had one been available). Whether it would have been any
more successful than the scorpion is not known. But the solpugid is also nocturnal,
and even if it had been successful, it would not have explained cryptic coloration.
An untested possibility of predation upon the velvet ants comes from birds.
The fact that so many birds are diurnal, that they possess color vision, and that
some are insectivorous all make them suspect of such predatory activity. Dis¬
cussions with Dr. Charles T. Collins and Dr. Stuart L. Warter (ornithologists at
California State University, Long Beach) have led to the opinion that likely can¬
didates for predation upon the velvet ants might be some members of the family
Mimidae, the thrashers. These birds are diurnal, insectivorous and occupy much
of the same habitat as the velvet ants. However, no birds were seen harassing
velvet ants in the field, even after many hours of observation of velvet ants under
natural conditions.
A possible source of predation of velvet ants that emerged from this study is
insectivorous lizards. While no predatory activity was observed in the field, lab¬
oratory experiments showed that lizards have the capability of eating velvet ants.
This, like all of the laboratory experiments, occurred under confined conditions,
and with hungry potential predators. The fact that lizards can eat a velvet ant
does not establish that they ordinarily do so.
At this point it seems that the only two potential predators that might pursue
velvet ants would be certain birds and lizards. And, based upon observations
which have been made to this point, it seems unlikely that even these organisms
would encounter or pursue velvet ants on any regular basis.
If the color exhibited by some of the velvet ants is, indeed, cryptic coloration,
then it seems most likely that some selective pressure exists, whether it be by one
of the organisms already mentioned, or by something entirely different.
Another possibility is that the colors are not designed to be cryptic at all, and
that it is only to our eyes that they appear so. The lighter colors may be ther¬
moregulatory in function. But both of these possibilities present irregularities.
Most importantly, why are other velvet ants, both larger and smaller, brightly
colored? Whether the colors are defensive in nature, or physiological, there should
be some consistency. The brightly colored and cryptically colored individuals
occupy the same areas.
In conclusion, it seems that the colors exhibited by some velvet ants probably
are cryptic in function. What selection pressure exists is not known. Why other
velvet ants are not also cryptically colored but are, instead, wamingly colored is
also not known. Whatever the answers, it seems probable that velvet ants do not
226
PAN-PACIFIC ENTOMOLOGIST
make up a large part of the diet of any other animal. It seems that there must be
an easier meal than the hard-bodied, powerfully stinging, velvet ant.
Acknowledgments
I thank Dr. John DuRant and Dr. Albert Johnson for their comments and
suggestions on this manuscript.
Literature Cited
Cockerell, T. D. A. 1895. A mutillid which resembles thistle-down. Psyche, suppl., 7(233):6.
Cyr, M. A. 1970. Predatory behavior of the Grasshopper Mouse ( Onychomys ). Ph.D. thesis, Uni¬
versity of California, Los Angeles.
Evans, H. E., and M. J. West Eberhard. 1970. The Wasps. Univ. of Mich. Press, Ann Arbor.
Hinton, H. E., D. F. Gibbs, and R. Silberglied. 1969. Stridulatory files as diffraction gratings in
mutillid wasps. J. Insect Physiol., 15(4): 549—5 52.
Kirkpatrick, T. W. 1957. Insect life in the Tropics. Longmans, Green & Co., London.
Manley, D. G. 1975. The effects of temperature and light on the daily activity patterns in Velvet
Ants of the Genus Dasymutilla Ashmead (Hymenoptera: Mutillidae) in California. M.A. thesis,
California State University, Long Beach.
Metcalf, C. L., and W. P. Flint. 1932. Fundamentals of insect life. McGraw-Hill Book Co., New
York.
Schmidt, J. O., and M. S. Blum. 1977. Adaptations and responses of Dasymutilla occidentalis
(Hymenoptera: Mutillidae) to predators. Ent. Exp. & Appl., 21:99-111.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 227-237
Five New Species of the Band-winged Grasshopper Genus
Trimerotropis Stal (Orthoptera: Oedipodinae)
D. C. F. Rentz and David B. Weissman
(DCFR) CSIRO, Division of Entomology, P.O. Box 1700, Canberra City, A.C.T.
2601, Australia; (DBW) Research Associate, Department of Entomology, Cali¬
fornia Academy of Sciences, Golden Gate Park, San Francisco, California 94118.
Abstract.— Five new species of Trimerotropis grasshoppers from the western
United States are described. Four species are known only from California. T.
leucophaea is described from several localities east of the Sierra Nevada Moun¬
tains. The other three Californian species are restricted: T. bernardi to the San
Bernardino Mountains; T. infantilis to the Santa Cruz Mountains; and T. inyo to
the Whitney Portal area. T. whitei is known only from pink sandy substrate areas
of southwestern Utah.
In this paper we describe five new species of Trimerotropis Stal. Four species
are restricted to California and the fifth is found in southwestern Utah.
The genus presently includes 56 species (Rentz and Weissman, 1980, 1981;
Weissman, 1984), and is one of the most common orthopteran components of
insect communities in North America. Species occur, often sympatrically, in
habitats ranging from below sea level in desert areas, to coastal sand dunes, to
rocky slopes at or above timberline. White (1973) and Weissman and Rentz (1980)
divided the genus into two sections, A and B, based entirely on karyotypic pa¬
rameters. Section A included those species in which the entire chromosomal
complement was telocentric. Section B species possess metacentric chromosomes,
either fixed or polymorphic.
Species’ descriptions within the paper refer to the whole series. All specimens
examined except holotypes are paratypes. Holotypes and main series of the new
species are in the California Academy of Sciences (CAS), San Francisco. Paratypes
will be deposited in the Academy of Natural Sciences of Philadelphia (ANSP),
University of Michigan Museum of Zoology (UMMZ), and the Australian Na¬
tional Insect Collection, Canberra (ANIC).
Trimerotropis bernardi, New Species
(Figs. 1, 2)
Recognition characters.— Species with blue hind tibia, yellow wings, and meta¬
centric chromosomes. See Table 2 for comparison with other Trimerotropis species
with blue hind tibia and yellow wings. Most similar to T. fratercula McNeill
(which occurs only in the Rocky Mountains) but differs in having a broader
tegmen, generally less deeply sulcate frontal costa, and blue instead of yellow hind
tibia. The former resembles Circotettix splendidus Rehn and Hebard from which
it may be distinguished morphologically by its smaller size and reduced thickness
228
PAN-PACIFIC ENTOMOLOGIST
Table 1. Measurements (in mm) for new species of Trimerotropis.
Length
body
Length
pronotum
Length hind
femur
Length
tegmen
T. bernardi
Holotype 6
Paratypes
24.9
5.1
13.1
26.3
Males (6)
23.7-24.4
4.3-4.9
11.4-12.3
23.0-25.6
Females (8)
28.1-32.5
6.0-6.6
14.5-16.2
30.0-31.4
T. infantilis
Holotype <3
Paratypes
15.7
3.2
10.0
16.0
Males (17)
13.7-17.2
2.4-3.9
8.6-11.2
14.7-19.6
Females (7)
19.7-21.6
3.8-4.4
10.8-13.1
18.7-23.5
T. inyo
Holotype 3
Paratypes
23.0
4.5
11.7
23.3
Males (15)
19.8-21.0
3.8-4.3
10.9-12.3
21.8-23.0
Females (3)
27.7-31.2
5.7-6.0
14.3-15.0
28.1-28.8
T. leucophaea
Holotype 3
Paratypes
21.9
4.2
11.4
22.5
Males (51)
18.2-22.6
3.7-4.5
10.2-11.7
20.3-25.1
Females (36)
25.0-30.4
4.7-5.7
12.9-14.4
26.2-30.5
T. whitei
Holotype 3
Paratypes
17.3
3.7
10.4
20.9
Males (37)
12.1-13.6
3.4-3.9
9.6-10.9
19.3-22.2
Females (19)
22.0-23.7
4.6-4.7
12.3-12.5
24.5-25.7
of the anal veins of the hind wing, including the posterior axillary vein, and by
the typically low crepitational flights characteristic of Trimerotropis as opposed
to the very high flights accompanied by loud snapping characteristic of Circotettix.
Geographic distribution (Fig. 1). — Known only from the San Bernardino Moun¬
tains, California.
Holotype male. — “USA: California: San Bernardino Co., San Bernardino Mts.,
Mill Creek Camping area, 6000 ft. elev., #81-24, 25 July 1981. Cal. Acad. Sci.
Access. David Weissman Coll.” Holotype in California Academy of Sciences, No.
12787.
Description. — See Table 1 for measurements. Head with frontal carinae heavy,
parallel nearly their entire length, not extending to fronto-clypeal suture; fastigium
of vertex broad, shallow, lateral carinae moderately heavy, extending to about
middle of eye, median carina absent. Pronotum without lateral carinae, prozona
and mesozona deeply sulcate (Fig. 2), metazona smooth, not tuberculate; median
carina moderately high, prozona convex; lateral lobe without tooth (Fig. 2). Meso-
and metastemal interspace relatively broad. Tegmina wide, extending for a dis¬
tance of % its length beyond hind tibia. Ventral and lateral surfaces of thorax and
legs setose. Body and tegmina coloration speckled dark brown and greyish white.
Antenna annulate. Pronotum with median portion of disk dark brown, lateral
VOLUME 60, NUMBER 3
229
Table 2. Characters for distinguishing Trimerotropis species with blue hind tibia and yellow or
yellow-green wings. Most species are only distinguished by a combination of characters. See Weissman
and Rentz (1980) for crepitation terminology.
Species
Chromosomal
section 1
Pronotal tooth 2 :
present +,
absent —
Lateral
prozonal
carinae 2 :
present +,
absent -
Crepitation:
crackle (C),
buzz (B),
pulse buzz (PB)
Other
significant
characters
albescens McNeill
A
+
C, B
greyish white color
arenacea Rehn
A
+
B
probably not in
California
bernardi, n. sp.
B
C
San Bernardino Mts.
only
bifasciata Bruner
A
+
B
bilobata Rehn and
Hebard
A
~~ > +
B
distinctive median
carina
cincta (Thomas)
A
+
PB
not in California
fontana Thomas
A
+
+
B, C
tegminal bands
distinct
inconspicua Bruner
A
B
blue hind tibia
unusual
infantilis, n. sp.
A
+ , —
+
B
band through eye,
small size, northern
California
inyo, n. sp.
B
+
PB
blue tinge in
wing disk
koebelei (Bruner)
A
— > +
B
band through eye
occidentaloides Rentz
and Weissman
A
— , +
+
B
Santa Monica Mts.
suffusa Scudder
B
+
C
no tegminal bands
thalassica Bruner
B
+
+
B
topanga Rentz and
Weissman
A
+ , ~
+
B
southern California
1 Data from Weissman and Rentz (1980), Weissman (1984).
2 Usual condition listed first.
areas greyish white. Tegmina with 2 indistinct bands formed by coalescence of
speckles, absent in some specimens; wing disk opaque yellow at base becoming
semitranslucent near band; band narrow and smoky, apex clear with a few speckles.
Outer pagina of hind femur with 2 distinct, oblique dark brown bands; inner
pagina black, dorsal margin with 3 black bands separated by yellow areas; apex
with a yellow band; hind tibia blue.
Karyotype. — 2N $ = 23; Section B, subsection 2; 3 large, 6 medium, 2 small
autosomes. X metacentric, all 3 large autosomes homomorphic metacentrics,
floating inversion series in the two specimens examined involving 2 and 3 medium
autosomes respectively. Individuals examined: type locality, 2 6.
Specimens examined. — California, San Bernardino Co., San Bernardino Mts.,
Mill Creek Camping area, 1829 m, 25.vii. 1981, D. Weissman, 2 <3, 2 $, including
holotype (CAS); 5.x. 1947, P. Timberlake, 2 <3, 1 2 (CAS). San Bernardino Mts.,
Vivian Creek, 2190 m, heavy forest, 29.viii. 1919, Rehn and Hebard, 3 8, 6 2
(ANSP). San Bernardino Mts., High Creek, 2740 m, 29.viii.1919, Rehn and
Hebard, 2 <3 (ANSP).
230
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Known distribution of five new species of Trimerotropis.
Remarks. — T. bernardi is an inhabitant of the Ponderosa pine community from
1820 to 2743 meters elevation. The 1981 type series was found at the intersection
of Valley of the Falls and Quercus Roads on gravelly substrate with pine needle
cover. All 4 of these specimens flew 2-4 m; male crepitation was a crackle, similar
to that of T. p. pallidipennis (Burmeister) (see Otte, 1970:123). Both females
VOLUME 60, NUMBER 3
231
Figures 2-6. Lateral view, paratopotype males of Trimerotropis. 2. T. bernardi. 3. T. infantilis. 4.
T. inyo. 5. T. leucophaea. 6. T. whitei.
contained nearly mature eggs. The adjacent sandy, dry Mill Creek stream bed and
pine-oak woodland-grass banks were searched but only T. fontana Thomas was
found. The series shows variation primarily in the intensity of the color of the
hind wing; a single male has the entire apical portion of the wing smoky black.
This species is named with a dual purpose; we honor our friend Prof. Bernard
John and christen it after the San Bernardino Mountains.
Trimerotropis infantilis, New Species
(Figs. 1, 3)
Recognition characters. — Small species with blue hind tibia, yellow wings, a
band through the eye, and only telocentric chromosomes. See Table 2 for com-
232
PAN-PACIFIC ENTOMOLOGIST
parison with other Trimerotropis species with blue hind tibia and yellow wings.
Most similar to T. koebelei Bruner but differs in its smaller size, narrower and
more deeply sulcate frontal costa, higher pronotal crest, and less distinct lateral
pronotal carinae. This species can be confused with Conozoa species: in the latter
genus tegminal bands cover only half of the tegmen; in Trimerotropis the bands
are absent all together or continuous across the entire tegmen, although odd
individuals in the type series of T. infantilis resemble Conozoa in this character.
D. Otte (pers. comm.) is describing a small species from coastal California near
Lompoc which might be confused with T. infantilis. This former species is slightly
larger, more robust, and has a more poorly defined median pronotal carina. Its
color is pale, almost whitish, and the bands on the hind femur are more sharply
indicated. It lacks any trace of a band through the bottom of the eye.
Geographic distribution (Fig. 1). — Known only from the Santa Cruz Mountains,
California.
Holotype male. — “USA: California: Santa Cruz Co., Santa Cruz Mountains,
Felton, E. Zyante Rd., 1 mi E intersection of E. Zyante Rd. and Graham Hill
Road, 16 July 1977, leg. D. B. Weissman & D. C. Rentz, Calif. Acad. Sci. Coll.”
Holotype in California Academy of Sciences, No. 14100.
Description. — See Table 1 for measurements. Head with frontal carinae heavy,
constricted at median ocellus, carinae continuing almost to fronto-clypeal suture;
fastigium of vertex relatively narrow, lateral carinae heavy, extending beyond
middle of eye, convergent; occiput with a pair of impressions. Pronotum with
lateral carinae represented as tubercles; median carina heavy, prozona and me-
sozona raised; lateral lobe (Fig. 3) with a minute tooth; surface of disk minutely
rugose, lateral lobe smooth but not shining. Tegmina relatively short, extending
for about l A to V 5 its length beyond hind tibia. Legs and thorax very sparsely setose.
Body and tegmina coloration greyish to light brown; head with ventral half of eye
and adjacent intra-antennal area blackish, remainder of frons greyish white; an¬
tenna faintly annulate in basal half. Tegmina with 2 bands varying from indistinct
to distinct; wing light yellow, highly transparent, band narrow, relatively faint,
remainder of wing uniformly colorless except for cross veins. Outer pagina of
hind femur with 2 faint, oblique stripes; inner pagina dark brown in proximal
half, then with 2 yellow bands separated by a narrow dark brown band that
continues onto dorsal margin; hind tibia light blue grey.
Karyotype. — 2N <3 = 23; Section A; all telocentric; 3 large, 6 medium, 2 small
autosomes; of 8 <3 examined from the type locality, 5 had a supernumerary segment
heterozygous in one of the small autosomes, 1 had the segment homozygous, and
the remaining 2 had no segments.
Specimens examined. — California, Santa Cruz Co., Santa Cruz Mts., Felton, E.
Zyante Road, 1.6 km E intersection of E. Zyante and Graham Hill Roads,
16.vii.1977, D. B. Weissman, D. C. Rentz, 12 <3, 5 2 including holotype (CAS).
Santa Cruz Mts., no date, A. Koebele Collection, 1 6, 2 2 (CAS); 13.viii. 1938, J.
Russell, 1 <3 (ANSP); Alma, 16.viii. 1928, E. G. Linsley, 2 <3 (CAS); Felton,
13.viii. 1959, D. C. Rentz, C. Wemmer, 1 <3 (ANIC); Santa Cruz, 28.ix. 1941, K.
E. Frick, 1 <3 (CAS).
Remarks. — At the type locality T. infantilis was found on sandy substrate sparsely
covered with Lotus and grasses at the base of pines above a rock quarry (Lone
Star Industries, Olympia Plant #125). Individual flights were from 1 to 2 meters
VOLUME 60, NUMBER 3
233
with male crepitation a rapid buzz. Approximately 25% of the population was
fourth and fifth instar on 16 July 1977; females bore nearly mature eggs. No other
Trimerotropis was microsympatric although T. p. pallidipennis and T. thalassica
Bruner were caught nearby. This species is named with reference to its small size;
it is only slightly larger than T. pogonata Strohecker, the smallest known species
in the genus.
Trimerotropis inyo, New Species
(Figs. 1, 4)
Recognition characters. — Species with blue hind tibia, greenish yellow wing with
a blue tinge, and metacentric chromosomes. May be distinguished from other
Trimerotropis species with blue in the wing by the combination of the short lateral
pronotal carinae, distinct tegminal banding, greenish yellow (fades in some mu¬
seum specimens) wing with a blue tinge, narrow dark band, and high pronotal
crest. Morphologically most similar to T. pseudofasciata Scudder but differs in
lacking a lateral lobe tooth and always having the wing disk predominantly green,
in addition to the characters listed previously. Also similar to T. cyaneipennis
Bruner but differs in smaller size and narrower wing band. Also the flight in T.
inyo is shorter and the crepitation is a series of rapid buzzes (pulse buzz—see
Weissman and Rentz, 1980, Fig. 1), as opposed to the crackle of T. cyaneipennis
(Weissman, 1984).
Geographic distribution (Fig. 1). —Known from the eastern slopes of the Sierra
Nevada Mountains above the Owens Valley, California.
Holotype male. — “USA: California: Inyo County, 12.8 km W. Lone Pine on
Whitney Portal Road, 1920 m, 6.viii.l978, D. B. Weissman, D. Lightfoot Stop
120, Calif. Acad. Sci. Coll.” Holotype in California Academy of Sciences, No.
14101.
Description. — See Table 1 for measurements. Head with frontal carinae heavy,
nearly parallel two-thirds their entire length, persistent to fronto-clypeal suture;
fastigium of vertex broad, shallow, carinae heavy, continuing well past middle of
eye; occiput with a pair of inverted comma-like impressions. Pronotum with
minute lateral carinae, prozona and mesozona deeply sulcate, the former raised;
median carina well indicated; lateral lobe punctate, usually without a tooth, but
ventral margin (Fig. 4) sometimes with faint indication of tooth. Tegmina slender,
extending for a distance of about x h its length beyond hind tibia. Ventral surface
of legs and thorax sparsely setose. Body and tegmina coloration brown or grey
brown, speckled. Antenna intensely annulate. Tegmina with 2 distinct bands, a
third band indicated by a coalescence of speckles; wing mostly transparent, disk
greenish yellow with a blue tinge basally, band variable, from half as wide as
tegmina and broken, to almost as broad as tegmina and uninterrupted, dark brown,
apex colorless. Outer pagina of hind femur with 2 distinct, dark brown bands, a
fainter band present at base; inner pagina brown, with a single yellowish area and
2 dark brown or black bands continuous to dorsal margin; hind tibia blue.
Karyotype. — 2N $ = 23; Section B, subsection 1; 3 large, 6 medium, 2 small
autosomes. X metacentric, L l3 L 2 , L 3 , M 5 and M 6 homomorphic metacentric, rest
telocentric; no inversion polymorphism noted in the 4 males examined from the
type locality.
Specimens examined. —California, Inyo Co., 12.8 km W Lone Pine on Whitney
234
PAN-PACIFIC ENTOMOLOGIST
Table 3. Characters to distinguish T. leucophaea from T. cyaneipennis.
T. cyaneipennis
T. leucophaea
Chromosome
number
2N <3 = 21 (23 in central
New Mexico)
2N 3 = 23
Tegmina
usually with distinct
bands, slate grey-brown
without distinct bands,
brown
Wing disk
rich blue, translucent
(difficult to read label
through disk near center)
light blue, transparent
(easy to read label
through disk near
center)
Wing band
wide, complete
narrow, sometimes
interrupted
Behavior
loud crackle, difficult
to collect
soft crackle, easy
to collect
Distribution
Oregon, Arizona, Utah,
Nevada, Idaho, New Mexico,
? California
California, ? Nevada
Portal Road, 1920 m, 6.viii. 1978, D. B. Weissman, D. Lightfoot. 13 6, 3 9,
including holotype (CAS). 17.6 km W Lone Pine on Whitney Portal Road, 2530
m, 19.vii. 1961, D. C. Rentz, 2 6 (CAS). 9.6 km W Big Pine, l.ix. 1964, M. R.
Gardner, 1 <5 (U.C. Davis).
Remarks. — This species is found on bare, gravelly substrate in the Pinyon-Sage
community on the eastern slopes of the Sierra Nevada Mountains. It occurs
sympatrically with T. albescens McNeill with the latter being much more com¬
mon. Flights in T. inyo range from 3-7 m; male crepitation is a pulse-rapid buzz.
At the time of collection of the holotype, only two males crepitated of the 13
seen. Females at the time bore nearly mature eggs; one last instar female was
seen.
Trimerotropis leucophaea, New Species
(Figs. 1, 5)
Recognition characters. — Species with blue hind tibia and wings, and metacen-
tric chromosomes. Most similar to T. cyaneipennis but distinguishable by a com¬
bination of characters—see Table 3. Differs from T. pseudofasciata in having a
low metazonal carina and lacking a lateral lobe tooth. Also similar to T. sparsa
(Thomas) but differs in having blue rather than brown or yellowish-brown hind
tibia and having a distinctly blue wing disk. Differs from T. arizonensis Tinkham
(known only from holotype) in lacking green in the wing disk (always light blue
in leucophaea ), having uniform blue hind tibia, and less distinct tegminal bands.
The latter species was collected along the hot, barren Colorado River.
Geographic distribution (Fig. 1). —California; specimens at ANSP from Reno,
Nevada, may be this species.
Holotype male.— “California: Inyo County, Westgard Pass, 7200 feet elevation,
26 July 1962. D. C. Rentz, C. D. MacNeill, collectors.” Holotype in California
Academy of Sciences, No. 12774.
Description. — See Table 1 for measurements. Head with frontal carinae feeble,
VOLUME 60, NUMBER 3
235
weakly constricted and ending below median ocellus; fastigium of vertex (Fig. 5)
broad, shallow, lateral carinae weak, extending slightly beyond middle of eye,
median carina absent. Pronotum without lateral carinae; prozona and mesozona
relatively deeply sulcate; median carina low, absent in mesozona, lateral lobe (Fig.
5) without tooth; surface of disk relatively smooth, minutely, evenly punctate.
Tegmina elongate, extending for a distance of about Vi of its length beyond hind
tibia. Ventral surface of thorax and legs sparsely setose. Body and tegmina col¬
oration uniformly medium grey brown. Antenna faintly annulate in proximal half,
less so distad. Tegmina speckled without bands; wing light blue, transparent, cross¬
veins along anterior margin light brown, remainder blue, brown band faintly
indicated. Outer pagina of hind femur uniform color; internal pagina with 2 yellow
blotches, dorsal surface entirely yellow; hind tibia blue.
Karyotype. — 2N S — 23; Section B, subsection 2; 3 large, 6 medium, 2 small
autosomes. X metacentric, 2 largest autosomes fixed metacentrics; four other
autosomes from L 3 to S 10 were polymorphic for inversions. One Silver Canyon
male had 1 small B chromosome. Individuals examined: 1.6 km E type locality,
1 <3; Silver Canyon, 5 <3.
Specimens examined. — California, Inyo Co.: Westgard Pass, 2100 m,
26.vii.1962, D. C. Rentz, C. D. MacNeill, 10 6, 14 2 (CAS); 21.vii.1964, D. C.
and K. A. Rentz, J. D. Birchim, 4 6, 3 2 (CAS); 1.6 km E Westgard Pass summit,
2102 m, 6.viii.l978, D. B. Weissman, D. Lightfoot, 1 < 3 , 1 2 (CAS); 1.6 km W
summit, 24.vii.1965, D. C. Rentz, 1 6; White Mountains, Silver Canyon, 2010
m, 10.ix. 1919, Rehn and Hebard, 13 < 3 , 6 2 (ANSP); 2012 m, 19.viii. 1982, D. B.
Weissman, #82-73, 5 < 3 , 2 2 (CAS); Angus Range, 780 m, 9.ix.l922, Rehn and
Hebard, 1 <3 (ANSP); Amargosa Range, Boundry Canyon, Hole in the Rock Spring,
790 m, 17.viii.1919, Rehn and Hebard, 1 <3 (ANSP); Panamint Range, Wild Rose
Canyon Spring, 1102 m, 6 and 8.ix. 1922, Rehn and Hebard, 3 < 3 , 4 2 (ANSP).
Modoc Co., Warner Mts., Cedar Peak, 1920-2190 m, 22.viii. 1922, Rehn and
Hebard, 3 <3, 4 2 (ANSP); Cedar Peak, Stowe Reservoir Camp, 2130-2440 m,
18.vii. 1968, D. C. Rentz, 2 <3 (CAS); near Cedar Pass, 1830 m, 25.vii.1962, D.
C. Rentz, C. D. MacNeill, 1 <3 (CAS); Cedar Pass Camp, 28.viii. 1965, D. C. Rentz,
Stop 32, 2 <3, 1 2 (CAS); San Bernardino Co., Trona, 500-550 m, 5 and 9.ix.l922,
Rehn and Hebard, 4 < 3 , 1 2 (ANSP).
Remarks.— At Silver Canyon, T. leucophaea was found on the valley floor and
on gentle talus slopes covered with scattered vegetation. Individuals flew 2 to 10
m close to the ground and were easy to catch. Male crepitation was a crackle,
softer than that of T. cyaneipennis (DBW, personal observation, Zion National
Park, Utah; Hebard, 1935; Strohecker et al., 1968). Individuals from Westgard
Pass and Cedar Peak were found in dry arroyos and on rocky slopes. Two spec¬
imens collected on 6 August 1978 from the Westgard Pass area were both teneral.
In August of both 1978 and 1982, DBW was unable to find adults on the west
side of Westgard Pass, although the 1982 Silver Canyon adults were not teneral.
T. leucophaea occurs with Circotettix undulatus (Thomas) at both Westgard
Pass and Silver Canyon, although the latter is usually on steeper slopes. At Silver
Canyon, T. p. pallidipennis was intermingled with T. leucophaea.
T. leucophaea is presently known only from the mountains of eastern California.
Most California specimens considered previously (Strohecker et ah, 1968) to be
T. cyaneipennis are T. leucophaea. The occurrence of T. cyaneipennis in California
236
PAN-PACIFIC ENTOMOLOGIST
is now uncertain. Strohecker et al. (1968) listed the Westgard Pass records of this
species under T. sparsa.
Only material from the White Mountains has been examined cytologically—
they all have 2N 8 = 23 with inversion heterozygosity similar to that reported for
T. cyaneipennis, also a member of Section B. The latter species, though, has 2N
8 = 23 only in central New Mexico (White, 1951) whereas farther west it has 2N
8 = 21: western New Mexico (White, 1951), Arizona (King, 1923), Nevada (White,
1949—one population only some 100 km from Westgard Pass), and southeastern
Oregon (Weissman and Rentz, 1980).
The name is derived from leucophaeus, meaning grey or ash-colored.
Trimerotropis whitei, New Species
(Figs. 1, 6)
Recognition characters. — Species rusty brown in overall coloration with yellow
hind tibia and wings, and telocentric chromosomes. Similar to T. arenacea Rehn
but differs in having a brighter yellow disk and yellow instead of blue hind tibia.
Also similar to T. citrina Scudder and T. agrestis McNeill but smaller and without
red hind tibia.
Geographic distribution (Fig. 1). —Known only from pink sandy substrate areas
of Kane County, Utah.
Holotype male. — “Utah, Kane Co., Zion National Park, 2.6 km W. of Park’s
E boundary on road through Park. 1646 m, 21.viii.82. D. B. Weissman #82-82,
Cal. Acad. Sci. Coll.” Holotype in California Academy of Sciences, No. 14160.
Description. — See Table 1 for measurements. Head narrow, frontal carinae heavy,
extending to fronto-clypeal suture; sulcus deep; fastigium of vertex narrow, lateral
carinae strong, extending beyond middle of eye; median carina indicated solely
by color; occiput with a pair of shallow indentations. Pronotum with lateral carinae
very feebly indicated on prozona, distinct on metazona; transverse sulci contin¬
uous, deep; median carina distinct, higher on prozona than mesozona, low on
metazona; surface evenly punctate; ventral margin of lateral lobe without a tooth
(Fig. 6). Tegmina elongate, narrow, extending for a distance of about Vs its length
beyond apex of hind tibia. Ventral surface of thorax not setose, legs sparsely
setose. Thorax and tegmina rusty brown with whitish blotches on sides; head
speckled whitish brown; antenna reddish brown with distinct whitish annuli.
Tegmina lightly banded, the distal band not much more than a coalescence of
speckles; wing yellow, not highly opaque; black band narrow, continuous. Outer
pagina of hind femur with 2 very faint brown bands; internal pagina with 3
yellowish blotches separated by 2 narrow, brown bands; hind tibia uniformly pale
lemon yellow.
Karyotype. — 2N 8 = 23; Section A; all telocentric, 4 large, 5 medium, 2 small
autosomes. Individuals examined: type locality, 6 8.
Specimens examined.—Utah, Kane Co.: Zion National Park, 2.6 km W of
Park’s E boundary on road through Park, 1646 m, 21 .viii. 1982, D. B. Weissman,
12 8, 3 9, including holotype (CAS): 16 km NW Kanab on Hywy 89, 1737 m,
21.viii. 1982, D. B. Weissman, 1 2 (CAS); Sand Dunes 16 km NW Kanab,
2.viii. 1951, M. J. D. White, 7 8, 29 (ANSP); Kanab, 5.viii.l949, G. F. Knowlton,
1 8, 1 2 (ANSP); Pink Sandhills 40 km SE Mt. Carmel Junction, 1950 m,
VOLUME 60, NUMBER 3
237
22.viii. 1956, no collector, 1 9 (ANSP); Pink Sandhills, road between Virgin River
and Three Lakes, 1752 m, l.ix.1926, Rehn and Hebard, 18 6 , 11 9 (ANSP).
Remarks. —Vegetation at the type locality included Manzanita, pines, and other
shrubs. Although known only from pinkish sands, and despite having a pink hued
body, alighted specimens were conspicuous at the type locality. Flights by both
sexes were low, usually less than 2 m but occasionally to 5 m. Crepitation by both
sexes was a loud buzz made during most of flight. Individuals were easy to catch.
One individual at each 1982 locality was teneral. Sympatric oedipods included
T. cyaneipennis, Circotettix rabula Rehn and Hebard, and T. texana (Bruner).
This species is named in honor of Prof. M. J. D. White who has been interested
in the cytological characteristics of Trimerotropis for over 35 years, and who
recognized this as an undescribed species when he collected it.
Acknowledgments
V. F. Lee and C. Mullinex helped at the California Academy of Sciences; D.
C. Lightfoot and S. I. Weissman assisted in fieldwork; and T. J. Cohn and D.
Otte provided useful discussion. Financial assistance was provided to DBW by
American Philosophical Society, Explorers Club, and Sigma Xi. The California
Academy of Sciences is thanked for their usual generous use of facilities.
Literature Cited
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New Mexico. Proc. Acad. Nat. Sci. Philadelphia, 87:45-82.
John, B., D. C. Lightfoot, and D. B. Weissman. 1983. The meiotic behavior of natural F[ hybrids
between the grasshoppers Trimerotropis suffusa and T. cyaneipennis (Orthoptera: Oedipodinae).
Can. J. Genet. Cytol., 25:467-477.
-, and D. B. Weissman. 1977. Cytogenetic components of reproductive isolation in Trimer¬
otropis thalassica and T. occidentalis. Chromosoma (Berl.), 60:187-203.
King, R. L. 1923. Heteromorphic homologous chromosomes in three species of Pseudotrimerotropis
(Orthoptera: Acrididae). J. Morphol., 38:19-63.
Otte, D. 1970. A comparative study of communicative behavior in grasshoppers. Misc. Pubs. Mus.
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Rentz, D. C. F., and D. B. Weissman. 1980. An annotated checklist of the grasshopper species of
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Am. Entomol. Soc. (Philadelphia), 106:223-252.
-, and-. 1981. Faunal affinities, systematics, and bionomics of the Orthoptera of the
California Channel Islands. Univ. Calif. Pubs. Ent., 94:1-240.
Strohecker, H. F., W. W. Middlekauff, and D. C. Rentz. 1968. The grasshoppers of California. Bull.
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Weissman, D. B. 1984. Autecology, cytology, morphology, and crepitation notes on Trimerotropis
grasshoppers (Orthoptera: Oedipodinae). (Submitted)
-, and D. C. F. Rentz. 1980. Cytological, morphological, and crepitational characteristics of
the trimerotropine ( Aerochoreutes, Circotettix, and Trimerotropis) grasshoppers (Orthoptera:
Oedipodinae). Trans. Am. Entomol. Soc. (Philadelphia), 106:253-272.
White, M. J. D. 1949. A cytological survey of wild populations of Trimerotropis and Circotettix
(Orthoptera, Acrididae). I. The chromosomes of twelve species. Genetics, 34:537-563.
-. 1951. Cytogenetics of orthopteroid insects. Adv. Genet., 4:267-328.
-. 1973. Animal cytology and evolution. 3rd ed. Cambridge University Press, New York.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 238-243
Contributions to Nearctic Psychodidae
(Diptera, Nematocera)
Rudiger Wagner
Limnologische FluBstation des M.P.I. fur Limnologie, Schlitz (Federal Republic
of Germany).
Since Quate’s comprehensive work on Nearctic Psychodidae was published in
1955, only a few other publications have dealt with this interesting group of
Diptera (Quate, 1957, 1960a, 1960b, 1960c; Vaillant, 1959, 1963, 1968, 1973;
Wagner, 1980). Quate’s drawings simplify very often the complicated structure
of the male genitalia, which has lead to unnecessary synonymies by subsequent
authors.
In the present paper, I recognize a new synonymy, describe two new species
and one new subgenus of Nearctic Psychodidae.
Threticus bicolor (Banks)
(Figs. 1-4)
Psychoda bicolor Banks, 1894, Can. Ent., 26:333.
Psychoda nigra Dyar (nec Banks), 1928, Proc. Ent. Soc. Wash., 30:87. (Synonymy
after del Rosario, 1936:116.)
Threticus appalachicus Vaillant, 1973, Ann. Soc. Ent. Fr. (N.S.), 9(1):355. New
synonymy.
Material.— 1 6, U.S.N.M. type no. 4308, New York; 1 6, Erie County, Travis
Run, 4-VI-1980; 1 < 3 , Erie County, 4-mile Ck., 25-VIII-1980; 1 6, Erie County,
4-mile Ck., 4-IX-1980, and 8 further specimens from Erie County, 6-mile Ck.,
4-mile Ck., from June-August 1981, all leg. E. Masteller.
This species is easy to distinguish from other Nearctic Psychodidae by the
extremely long inner prolongations of the basistyles, and the strong laterally com¬
pressed basal part of the aedeagus. Quate (1955), and Vaillant (1973) both have
presented drawings of the male genitalia, but only the latter is adequate. Quate
simplified the very complicated build especially of the aedeagus, which is shown
very well by Vaillant (1973:353, 356).
This species has an asymmetric aedeagus, and as mentioned by Wagner (1979)
in species with asymmetric genitalia two sorts of males appear to be equally
abundant, whose genitalia are mirror symmetric. This happens also in Threticus
bicolor (Banks) (compare especially Figs. 3 and 4).
Quate (1957) placed Psychoda bicolor Banks in the genus Threticus Eaton, but
the genitalia show some specialized features, which are not found in the European
species. The basistyles have long flexible inner appendages. Their positions differ
according to a more or less extruded condition of the aedeagus (Figs. 1-4). The
two lateral parts (right one rastered) are laterally joined with the basistyles and
are also movable. Figure 1 shows the most everted, Figure 2 an intermedial, and
Figure 3 the most inverted position of the aedeagus.
VOLUME 60, NUMBER 3
239
Figures 1-4. Threticus bicolor (Banks), structure and function of genitalia. 1. Extruded position.
2. Intermedial position. 3. Inverted position of the aedeagus. 4. Mirror symmetric genitalia. AA =
aedeagus apodeme. BS = basistyle. DS = dististyle. LP = lateral parts of aedeagus. VB and DS omitted
in Figures 2-4.
Genus Stupkaiella Vaillant
Stupkaiella Vaillant, 1973, Ann. Soc. Ent. France (N.S.), 9(1):367.
Type-species.— S. furcata Vaillant, by original designation.
The generic description given by Vaillant (l.c.) lists the absence of sensory organs
on the thorax (patagiae, tegulae). I can confirm Quate’s observations (1955) that
at least S. kincaidi (Quate), S. bessophila (Quate), and S. mastelleri, n. sp. have
small patagiae.
Stupkaiella masterelli, New Species
(Figs. 5-8)
Material. — Holotype, < 3 , Erie County, 4-mile-Ck., Behrend Campus, 2-V-1980
(located in C.A.S.); paratype, 6, Erie County, 6-mile-Ck., 12-V-1980 (in my col¬
lection); further material, about 20 males, 4-mile-Ck. and 6-mile-Ck., May-June
1980, all leg. E. Masteller, to whom the species is dedicated.
Description. — Head: Eye bridge consists of 4 rows of facets, distance between
the eyes 3 facet diameters, interocular suture U-shaped; 6 postocular bristles
behind each eye. Antennae 16-segmented; scape elongate cylindric, 2.5 times
longer than wide and 2.5 times longer than the spherical pedicel; post-pedicel
240 PAN-PACIFIC ENTOMOLOGIST
Figures 5-8. Stupkaiella mastelleri, n. sp. 5. Patagium. 6. Ventral bridge, styles, and aedeagus
ventral view. 7. Antennal segments 1-4. 8. 9th tergite and cercopodia ventral view.
elongate, bearing 5-6 strong bristles; 16th segment with a thin eccentric tip;
segments 7-14 with a pair of short simple ascoids. Length ratio of the basal
antennal segments: 62-23-38-15. Length ratio of palpal segments: 40-62-64-82.
Pro thorax with a pair of small spherical patagia. Wing length 2.8-3.0 mm. M 2
with a basal peduncle. Medial angle: 145°.
Sternal bridge regularly curved, of equal width throughout. Basistyles with
straight medial and semicircular lateral margins. Dististyles forked. In ventral
view the dorsal branch is much shorter than the ventral one. Cercopodia strongly
bent with 15 slightly frazzeled tenacula. Ninth tergite 2 times wider than long.
Aedeagus with a long narrow spatula distally with two triangular spines, which
are homologous to the inner lamellae of Palaearctic Pericomini.
Relations.—S. mastelleri, n. sp. is a close relative of S. recurrens Vaillant, from
Great Smoky Mountains National Park. The shape of the aedeagus and of the
dististyles readily distinguish the two species.
Parapanimerus, New Subgenus
A number of Nearctic species presently included in Telmatoscopus Eaton are
in fact not members of that genus, but of Panimerus Eaton, in which species of
the group of Telmatoscopus basalis form a distinct subgenus, Parapanimerus new
subgenus.
Description.—Imago 6: Eyes separated by a distance of 1-2 facet diameters.
Eye bridge consists of 4 rows of facets. Antennae of 16 segments. Scape elongate
cylindric, 1.5-2 times longer than the spherical pedicel. Flagellar segments flask¬
shaped, the 16th segment is always the longest. About 35-40 simple short ascoids
on each flagellar segment. Head with a pair of flatspread sensory organs (cornicula),
VOLUME 60, NUMBER 3
241
Figures 9, 10. 9. Sagittal sections of genitalia of Panimerus (Parapanimerus) basalis. 10. Panimerus
(s. str.) goetghebueri. AA = aedeagus apodeme. GD = gonoduct. T = 9th tergite. ST = styles. C =
cercopodia. P = parameres. TG = tub-shaped tegmen. SF = slightly sclerotized flap.
which are normally carried between head and prothorax, and are everted only
during premating ceremonies.
Thorax: Front femora with a double row of 10-20 large bristles on the posterior
face. Wing of normal size and shape, radial fork closer to wing base than medial
fork. Head (without cornicula and antennae), thorax, and a more or less extended
area of the wing base are covered with black setae (androconia).
Genitalia have often been compared with those of Palaearctic Panimerus (sensu
stricto), (Figs. 9, 10), but their structure is different. The aedeagus apodem is
Y-shaped with two short distal parts, which surround the distal part of the gon¬
oduct. The bases of the parameres are close together and are medially linked with
the apodeme. They are placed very much towards the ventral side of the basistyles.
In contrast, in Palaearctic Panimerus they are laterally linked with the tips of the
apodeme, and they are placed near the dorsal side of the basistyles. The distal
part of the aedeagus (tegmen) is tubshaped, and covered by a thin slightly scle¬
rotized flap (possibly a distal closing mechanism of the gonoduct). Styles simple,
9th tergite rectangular, wider than long, cercopodia slightly curved or straight
with at least 10 tenacula, which often are distributed over the distal half of the
242
PAN-PACIFIC ENTOMOLOGIST
Figures 11-14. 11,12. Panimerus ( Parapanimerus) basalis (Banks). 11. Ventral bridge, styles, and
aedeagus, ventral view. 12. Tip of dististyle. 13, 14. P. (Parapanimerus) arnaudi, n. sp. 13. Genitalia,
ventral view. 14. Tip of dististyle.
cercopodia. In contrast, Palaearctic Panimerus do not have a tub-shaped tegmen,
and they have no slightly sclerotized flap.
Similar to the males. They do not have androconia, cornicula, and spines
on femore I. The antennal segments each bear a pair of ascoids, which are either
simple or have 2 or 3 short branches, differing in single specimens from segment
to segment.
No larvae and pupae of this genus have been described yet. The larva of “Pan¬
imerus” lucens Vaillant (1973:352), belongs to a species close to Telmatoscopus
quadripunctatus (Banks), and may possibly be closer related to Palaearctic Mormia
species (Vaillant, pers. comm.).
Type-species.—Psychoda basalis Banks, 1907, Proc. Ent. Soc. Wash., 8:150 (by
present designation). Aedeagus and styles in Figures 11, 12.
Further species included in Panimerus subgenus Parapanimerus : Panimerus
scalus (Haseman), New Combination; Panimerus dysmica (Quate), New Combi¬
nation; Panimerus cio (Quate), New Combination; Panimerus sierra (Quate), New
Combination; Panimerus arnaudi, New Species.
Panimerus (Parapanimerus) arnaudi New Species
(Figs. 13, 14)
Material. — Holotype, S, North Carolina, Cumberland County, Fort Bragg, 14-
V-1967, leg. Jim D. Birchim.
VOLUME 60, NUMBER 3
243
Etymology. — Dedicated to Dr. P. H. Arnaud, San Francisco, who made the
CAS collection available to me.
Description. — Head: Eye bridge consists of 4 rows of facets. Distance between
the eyes, 1.5 facet diameters. Scape twice as long as broad, and twice as long as
the spherical pedicel. Flagellar segments flask-shaped, each with about 40 simple
short ascoids. A pair of cornicula present. Length ratio of palpus segments: 28-
54-55-60.
Thorax: Femora I with double row of 5-7 spines. Wing length 1.5 mm, basally
with black anoconia. Medial angle 195°.
Genitalia: 9th tergite wider than long. Cercopodia straight, with 12 tenacula
distally, and 3 single tenacula in the middle of the cercopodia. Ventral bridge
slightly sclerotized with two pairs of tiny tips laterally, near the bases of the
parameres. Basistyles two times longer than wide. Dististyles slightly bent, with
two short tips (Fig. 14). Parameres strongly bent medially. Tegmen (tub-shaped
structure) slender with parallel sides and a distal incision. Aedeagus apodeme
small, short, and slender.
Relations.—P. arnaudi, n. sp. is the closest relative of P. basalis (Banks). How¬
ever, both species differ from the western Nearctic species with converging lateral
margins of the tegmen, in having a tegmen with parallel sides. P. arnaudi is
separated from P. basalis by the parameres, which are much more curved in P.
arnaudi, and by the dististyles, which have an entire tip in P. basalis.
Literature Cited
Banks, N. 1894. Some Psychodidae from Long Island. Can. Ent., 26:329-333.
Quate, L. W. 1955. A revision of the Psychodidae (Diptera) in America north of Mexico. Univ.
Calif. Publ. Entom., 10:103-273.
-. 1957. Classification of the Psychodini (Psychodidae, Diptera). Ann. Ent. Soc. Amer., 54:
441-451.
-. 1960a. New species and records of Nearctic Psychodidae (Diptera). Pan-Pac. Ent., 36:143-
149.
-. 1960b. Note on the synonymy of an American and Japanese species of Psychodidae (Diptera).
Pan-Pac. Ent., 36:156.
-. 1960c. Guide to the insects of Connecticut, VI. State Geol. Nat. Hist. Survey Connecticut,
92:1-54.
del Rosario, F. 1936. The American species of Psychoda (Diptera: Psychodidae). Philip. Jour. Sci.,
59:85-148.
Vaillant, F. 1959. The larvae of three Nearctic Diptera of the family Psychodidae. Joum. N.Y.
Entom. Soc., 67:39-50.
-. 1963. Les Maruina d’Amerique du Nord (Diptera, Psychodidae). Bull. Soc. Ent. Fr., 68:71—
91.
-. 1968. Un Diptere Psychodide nouveau d’Amerique du Nord et ses stades preimaginaux.
Bull. Soc. Zool. Fr., 93:423-436.
-. 1973. Somenew Psychodidae from the U nited States (Diptera, Psychodidae). Aquatic Insects,
1:55-63.
-. 1980. The Nearctic Trichomyiinae (Diptera,Psychodidae). Pan-Pac. Ent., 56:273-276.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, p. 244
Scientific Note
Wasps on Agave Flowers in Mexico
Although bats are the principal pollinators of Agave (Agavaceae) in the Amer¬
ican Southwest and Mexico (e.g., Howell and Roth, 1981, Ecology, 62:1-7), a
variety of insects, including bees and wasps also visit the flowers for pollen and
nectar (Schaffer and Schaffer, 1979, Ecology, 60:1051-1069; Schaffer et al., 1979,
Ecology, 60:976-987). Peak periods of visits to Agave flowers coincide with pe¬
riods of peak nectar and pollen availability (Schaffer et al., 1979, Ecology, 60:
976-987). In this note I report some observations on wasp associations with Agave
flowers at Mazatlan, State of Sinaloa, Mexico, 12-14 November 1979.
A single inflorescence, about 2 meters tall, bearing several panicles with flowers
in various stages of development, formed the basis of these observations. I noticed
that the vivid yellow flowers were visited heavily at various hours of the day by
at least three different kinds of large-bodied wasps. Very few other kinds of insects
were seen. I checked the flowers usually at 800, 1200, and 1600 hours on each of
three days, and made a small reference collection of wasps seen. For each of the
time periods, I saw from 0 to 7 wasps on the flowers, with the most commonly
observed species being Polistes ( Onerarius ) carnifex (F.) (Hymenoptera: Vespi-
dae). I saw from 1 to 4 individuals of this large wasp at various places in the
flower, including head-downward at the base. Another, less abundant visitor was
Polistes ( Aphanilopterus) instabilis de Saussere (also Vespidae). An unidentified
sphecid, probably subfamily Sphecinae, tribe Sceliphronini, was also seen at the
flowers. Typically these wasps positioned themselves in various ways on the
flower, but generally did not come into contact with the anthers. They were feeding
primarily on the viscous nectar at the base of each flower.
The frequent occurrence of wasps on flowers in various stages of development
suggests that these insects may be nectar thieves, contributing to a reduction in
the amount of potential reward for pollinators. Such an effect could be additive
if flowers are not pollinated the night before such thieving by wasps takes place.
Daytime thieving may also deprive other kinds of pollinators of nectar. Very likely
the wasps, being eusocial in the sense of Wilson (The Insect Societies, Harvard,
1971), make repeated visits to an Agave inflorescence once it is discovered. Such
behavior increases the impact of these insects, as nectar thieves, on the plant-
pollinator interaction.
I thank the Milwaukee Public Museum for support, and Dr. Robert L. Jeanne
for the determinations of the wasps.
Allen M. Young, Department of Invertebrate Zoology, Milwaukee Public Mu¬
seum, Milwaukee, Wisconsin 53233.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 245-247
A New Classification of the Subfamily Ephemerellinae
and the Description of a New Genus
Richard K. Allen
22021 Jonesport Lane, Huntington Beach, California 92646.
A single female nymph collected in central Idaho represents an undescribed
genus and species.
Caurinella Allen, New Genus
Nymph.— Head without tubercles, and without frontal or occipital projections;
head with long setae around lateral and ventral margins of head, above mouth-
parts; maxillary palpi two-segmented. Thoracic nota without tubercles or pro¬
tuberances. Abdominal terga without paired dorsal abdominal tubercles; postero¬
lateral projections on segment nine 50% longer than length of segment at midline,
and postero-lateral projections upturned at apices.
Type species. —Caurinella idahoensis Allen, New Species.
Remarks.— The nymph of this new taxon is not similar morphologically, to
any of the other known genera in the subfamily, but it does have some characters
in common with the genus Serratella. The nymphs of both are without tubercles
or projections on the head and thorax, they both possess well developed spines
on the femora, and they both have a reduced maxillary palpus and long spines
on the caudal filaments. The long postero-lateral projection on segment nine is
not unique as some or all species of Torleya, Acerella and Cincticostella have long
abdominal projections. The upturned apices of the postero-lateral projections of
segment nine and the row of long setae around the head are characters which are
unique to Caurinella.
Caurinella idahoensis Allen, New Species
Nymph. — Length: body 4.0 mm; caudal filaments 3.0 mm. General color light
brown. Head light brown; maxillary palpi 2-segmented. Thoracic nota light brown
with irregular brown markings; legs light brown; femora with dorsal spines and
ventral setae; dorsal spines set in elevated sockets; claws with 3-4 small denticles.
Abdomen light brown, median area terga 5-7 brown; imbricated gills terga 3-7
unusually small; postero-lateral projections segments 4-9, small on 4 and increas¬
ing in length posteriorly; postero-lateral projection segment 9 more than 50%
longer than segment length at midline and postero-lateral projections turned up¬
ward at apices. Caudal filaments light brown with spines and sparse setae.
Holotype. — Female nymph, Bushy Creek, trib. Lochsa River (between Lolo
Pass and Powell Ranger Sta.), Idaho Co., Idaho, 26-VI-80, Bill Platt et al., in
collection California Academy of Sciences, San Francisco.
Classification.—In 1980, Allen divided the subfamily Ephemerellinae into 2
tribes, 14 genera and 10 subgenera, and he placed the genus Vietnamella Tsher-
nova ,as a subgenus of Cincticostella Allen. This synonymy is incorrect. Vietna¬
mella is known only from the nymphal stage and the nymphs differ from those
246
PAN-PACIFIC ENTOMOLOGIST
of CincticoStella, and all other Ephemerellinae, in the number and placement of
the dorsal abdominal gills. The nymphs of Vietnamella have six pairs of lamellate
gills on segments 2-7, eleven other genera, including Cincticostella, have five pairs
of lamellate gills on segments 3-7, and four genera have four pairs of lamellate
gills on segments 4-7.
I apply the philosophy used in the separation of subgenera in the Ephemerel¬
linae, and herein erect subtribes in the tribe Ephemerellini which will eliminate
the necessity of referring to these nymphs as “those with gills on segments 3-7”
or “those with gills on segments 4-7”. I propose the name EPHEMERELLAE
for those genera with nymphal gills on abdominal segments 3-7, the name TIM-
PANOGAE for those genera with nymphal gills on segments 4-7, and the name
YIETNAMELLAE for the genus Vietnamella with nymphal gills on segments
2-7.
The subfamily Ephemerellinae is now divided into 2 tribes, 3 subtribes, 17
genera, and 9 subgenera as follows:
Tribe Ephemerellini Lameere, 1917
Subtribe EPHEMERELLAE Allen, New Subtribe
Type Genus. Ephemerella Walsh, 1862
Nymph. Abdominal terga with five pairs of lamellate gills on segments
3- 7.
Genera and Subgenera
1. Ephemerella {=Chitonophora Bengtsson, 1908)
2. Serratella Edmunds, 1959
3. Caurinella Allen, New Genus
4. Uracanthella Belov, 1979
5. Torleya Lestage, 1917
6. Teloganopsis Ulmer, 1939
7. Caudatella Edmunds, 1959
8. Drunella Needham, 1909
a. Drunella s.s.
b. Tribrochella Allen, 1980
c. Unirhachella Allen, 1980
d. Myllonella Allen, 1980
e. Eatonella Needham, 1927
9. Crinitella Allen and Edmunds, 1963
10. Acerella Allen, 1971
11. Cincticostella Allen, 1971
a. Cincticostella s.s.
b. Rhionella Allen, 1980
Subtribe TIMPANOGAE Allen, New Subtribe
Type Genus. Timpanoga Needham, 1927
Nymph. Abdominal terga with four pairs of lamellate gills on segments
4- 7.
Genera and subgenera
1. Attenella Edmunds 1971 {=Attenuatella Edmunds, 1959 nee Stehli,
1954)
2. Eurylophella Tiensuu, 1935
VOLUME 60, NUMBER 3
247
3. Dannella Edmunds, 1959
a. Dannella s.s.
b. Dentatella Allen, 1980
4. Timpanoga
Subtribe VIETNAMELLAE Allen, New Subtribe
Type Genus. Vietnamella Tshernova, 1972
Nymph. Abdominal terga with six pairs of lamellate gills on segments
2-7.
Vietnamella only known genus.
Tribe Hyrtanellini Allen, 1980
Hyrtanella Allen and Edmunds, 1976, only known genus.
Literature Cited
Allen, R. K. 1980. Geographic distribution and reclassification of the subfamily Ephemerellinae
(Ephemeroptera: Ephemerellidae). pp. 71-91 in J. F. Flannagan and K. E. Marshall (eds.),
Advances in Ephemeroptera Biology, Plenum Publ. Corp., N.Y., 552 pp.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 248-255
Bionomics of Nearctic Species of Aclypea Reitter:
Phytophagous “Carrion” Beetles
(Coleoptera: Silphidae)
Robert S. Anderson and Stewart B. Peck
Department of Biology, Carleton University, Ottawa K1S 5B6, Canada. Present
address: (RSA) Department of Entomology, University of Alberta, Edmonton
T6G 2E3, Canada.
Abstract.— Diagnoses, distributions, notes on bionomics, and a key for iden¬
tification are presented for the two Nearctic phytophagous “carrion” beetles Acly¬
pea opaca (L.) and A. bituberosa (LeC.). A brief discussion of the possible evo¬
lutionary history of the genus in the Nearctic region is given.
Recent studies on the biology, distribution and systematics of the Silphidae of
Canada and Alaska (Anderson, 1981) have revealed problems concerning the
identity and distribution of two Nearctic species of the genus Aclypea. The latest
reviews (Hatch, 1957; Miller and Peck, 1979) have indicated that two species live
in North America; the primarily Palearctic A. opaca (L.), 1758, and the Nearctic
A. bituberosa (LeConte), 1859. Adults of the two species are morphologically very
similar. We have found the characters in keys that attempt to separate the species
(Arnett, 1944; Hatch, 1927; Horn, 1880; Portevin, 1926) to be vague or imprecise.
Thus, there has been a problem in distinguishing them and in accurately docu¬
menting their distributions and habits in North America.
Adult Aclypea can be recognized by the following generic characters: labrum
deeply emarginate, with the inner faces of the emargination swollen; eyes relatively
small, and not bounded posteriorly by a row of erect setae. The genus is represented
by approximately 27 species, distributed throughout the Holarctic region, some
members of which have been previously assigned to the genera Blitophaga Reitter
and Silpha L. (Portevin, 1926; Hatch, 1927, 1928; and others). We use Aclypea
rather than Blitophaga following Seidlitz (1883:311) as first reviser in accordance
with article 24 (a) (i) in the International Code of Zoological Nomenclature.
The vast majority of the species are found in the Palearctic region, and no
attempt is made here to treat them. We have found that the two Nearctic forms
can be distinguished as follows:
1 a. Genitalia of both sexes thinly sclerotized (light brown in color); pronotum
often with impunctate area on anterior third behind eye; elytra with
punctures shallow, the distance behind punctures rarely less than the
width of a puncture (Fig. 3); isodiametric microsculpture of pronotum
and elytra distinct (Fig. 3); generally smaller in size (12-15 mm). Dis¬
tribution in Nearctic restricted to Alaska and extreme northwest N.W.T.
. A. opaca (Linn.)
VOLUME 60, NUMBER 3
249
lb. Genitalia of both sexes thickly sclerotized (dark brown in color); prono-
tum never with impunctate area behind eye, although other impunctate
areas may be present; elytra with punctures deep, the distance between
the punctures often less than the width of a puncture (Fig. 4); isodia-
metric microsculpture usually indistinct (Fig. 4); larger in size (14-17
mm). Widespread in western North America ... A. bituberosa (LeConte)
Aclypea opaca (Linnaeus), the Beet Carrion Beetle
(Figs. 1, 3, 5)
Silpha opaca Linnaeus, 1758:361.
Diagnosis. — Color blackish to brown, usually clothed with yellow hairs dorsally.
Pronotum with anterior margin usually narrow (Fig. 1), lateral margins not dis¬
tinctly raised; with occasional impunctate areas present behind eyes. Elytra with
punctures shallow, distance between them rarely less than their width (Fig. 3).
Isodiametric microsculpture of pronotum and elytra distinct. Genitalia in both
sexes thinly sclerotized, light brown in color. Smaller in size, 12-15 mm in length.
Synonymies are in Portevin (1926) and Hatch (1928).
Variation. — Variation in this species appears to occur primarily in the pattern
of the impunctate areas on the pronotum. Some specimens of A. opaca possess
impunctate areas on the pronotum in the anterior third directly behind the eyes
in addition to other impunctate areas which may be present near the midline.
These impunctate areas behind the eyes are not found in A. bituberosa, although
other impunctate areas may be present.
There is little structural variation in other characters in the material that we
have examined.
Distribution.—The species is known to us to occur in North America through
very few specimens from Alaska and the Northwest Territories (Fig. 5). It has
previously been recorded from various localities in the United States (Horn, 1880;
Hatch, 1927), but these appear (based on our re-examination of available speci¬
mens) to be misidentifications of A. bituberosa. Aclypea opaca also occurs through¬
out the northern areas of the Palearctic region (Hatch, 1928).
Material examined. — (24 specimens with the following data; abbreviations of
collections in Arnett and Samuelson, 1969).
CANADA. Northwest Territories. Aklavik, 10.VIII.31, O. Bryant, 1 (CASC).
UNITED STATES. Alaska. Circle, 3.VII.58, G. E. Ball, dry field, 1 (UASM);
Fairbanks, 26.VI-1.VII.79, B. Wright, 1 (NSMC); Fort Yukon, 17.VI.16, Har¬
rington, 2 under logs (USNM); 11.VI. 16, Evans, 1 (USNM); Kussiloff (Kasilof),
VII.98, Evans, 1 (USNM); Palmer, 15.VII.79, 1.VI.80, 25.VII.80, 1.VIII.80,
13.VIII.82, 3.IX.82, D. P. Bleicher, baited cone traps 8 (University of Alaska).
OTHER COUNTRIES. 8 specimens from Northern and Central Europe.
Bionomics. — Like other Aclypea, the species in Europe is phytophagous on
various Chenopodiaceae, and has also been recorded as a pest of sugar beets
(Martin, 1945; Heymons et al., 1929).
The natural history has been well documented by Heymons et al. (1929) and
Martin (1945) in Europe. According to these authors, adults overwinter and emerge
in the spring when the sugar beets are germinating. Larvae appear about two weeks
later and feed externally on the young shoots and leaves. When mature, the larvae
250
PAN-PACIFIC ENTOMOLOGIST
I-1
Figures 1—4. 1. Dorsal view of pronotum of Aclypea opaca (Northwest Territories: Aklavik); scale
line = 2.3 mm. 2. Dorsal view of pronotum of Aclypea bituberosa (Saskatchewan: Kenosee); scale
line = 2.7 mm. 3. Dorsal view of base of left elytron of Aclypea opaca (Northwest Territories: Aklavik);
scale line = 0.7 mm. 4. Dorsal view of base of left elytron of Aclypea bituberosa (Alberta: Calgary);
scale line = 0.9 mm.
move into the soil to pupate. Adults emerge 10-15 days later and also feed on
the plants, but not as extensively as the larvae.
Although recorded as a pest of sugar beets in North America (Cooley, 1906,
1917; Forbes and Hart, 1900), data presented here make it very likely that these
records were misidentihcations of A. bituberosa. There appears to be no indication
that A. opaca is a pest of any cultivated crops within the extent of its known
distribution in North America. In fact, we know of no reliable data on native
host plant relationships in North America.
Our Nearctic records indicate that adults are active from June through August.
Aclypea bituberosa (LeConte), the Spinach Carrion Beetle
(Figs. 2, 4, 5)
Silpha bituberosa LeConte, 1859:6.
Diagnosis. — Color blackish to brown, usually clothed with yellow hairs dorsally,
although these are often abraded. Pronotum with anterior margin thicker than in
VOLUME 60, NUMBER 3
251
Figure 5. Known distribution of Aclypea opaca (stars) and Aclypea bituberosa (dots) in North
America.
A. opaca (Fig. 2), lateral margins distinctly elevated; with occasional impunctate
areas, but not with any behind eyes. Elytra with punctures deep, often confluent,
especially towards base (Fig. 4). Microsculpture of pronotum and elytra not dis¬
tinct. Genitalia in both sexes thickly, darkly sclerotized. Larger in size, 14-17
mm in length.
Variation. — Although all prairie specimens of A. bituberosa are structurally
similar, examination of the few available specimens of A. bituberosa from montane
localities indicates a smaller overall size and a much less convex elytra than those
from the lowland prairie areas. In these montane forms, the genitalia tend to be
252
PAN-PACIFIC ENTOMOLOGIST
as heavily sclerotized as in all other A. bituberosa. However, the punctures of the
elytra tend to become smaller and shallower, thus approaching the state in A.
opaca. The punctures at the base of the elytra, although still large and deep, are
rarely confluent.
These patterns of morphological variation appear correlated with a change to
montane-alpine habitats and may have an ecogenotypic or ecophenotypic basis.
Possibly there is a continuous morphocline between the northern A. opaca and
the southern A. bituber os a through these intermediate montane populations. This
would indicate probable conspecificity but our collection data presently indicate
the existence of a geographic zone where neither species occurs. However, until
more representatives of this genus become available from this area (i.e., provided
the known distribution is a collecting artifact) we think it best to continue to
recognize the two forms as distinct species.
Distribution. — In Canada, the species is found throughout the provinces of
Alberta, Saskatchewan and Manitoba, extending northward into the southern
Northwest Territories (Fig. 5). It appears to be found primarily in open grassland
or prairie habitats, although some specimens have been collected in interior valleys
and alpine habitats in British Columbia. In the United States it occurs throughout
the northern midwest into the western coastal and Rocky Mountain states.
Material examined. — (278 specimens examined and other literature records.)
Only generalized locality and seasonality data (when available) are given here.
Full data are available from the authors. Specimen numbers are given only if the
record is based on more than one. Material is in the following collections: AMNH,
CASC, CDFA, CNCI, DZEC, FMNH, LACM, MCZC, SBPC, UASM, UBCZ,
UICM, USNM (abbreviations follow Arnett and Samuelson, 1969); the collections
of the Canadian Department of Agriculture at Lethbridge, Saskatoon, and Win¬
nipeg; the Saskatchewan Provincial Museum; the Universities of Manitoba and
Saskatchewan; and J. L. Carr collection, Calgary.
CANADA. Alberta. Brooks, August, 3. Calahoo, July. Calgary, May, 20; June,
2; July. Diamond City (Farstad, 1949). Drumheller, June. Gorge Creek (50 mi
W Calgary), June, 29; July, 23. Grand Prairie, July. High River, April. Holden,
May, 3. Hussar, May, 2. Leduc, June. Lethbridge, April, 3; May, 2; June, 2.
Medicine Hat, April; May, 6; June, 5. Millarville, May. Ralston, May, 2. Ray¬
mond, July, 2. Sherwood Park (suburb of Edmonton), April, 4; May, 5; September.
Tilley, June, 3. Viking (MacNay, 1954). British Columbia. Chilcotin, April, Oc¬
tober. Manning Prov. Park (Nicomen Ridge, 6600'), July. Vernon, May, 2. Man¬
itoba. Grandview, May, 3. Lauder, May. Northwest Territories. Fort Resolution,
June, 2; July. Fort Smith, May, 9; June, 11; July, 21; August, October. Fort
Simpson, June, July, August. Saskatchewan. Big River, July. Chaplin, July. Dun-
durn, April. Fort Walsh, July. Gravelbourg, July. Kenosee, June. Moosomin, July.
Norbery, May. Prince Albert (MacNay, 1954). Rockglen, July. Rosthern, June,
2. Saskatoon, March, April, May, July, October, November.
UNITED STATES. California. Alpine Co., Ebbetts Pass, July; Sonora Pass,
June, 2. Eldorado Co., Echo Lake, 7400'; July. Tallac, July. Mono Lake (1880
Horn record, in MCZ). Tuolumne Co., no other data; Yosemite National Park,
Mt. Lyell, August (see Miller and Peck, 1979, for full data). Colorado. Craig, May,
6200'. Custer Co., no other data. Idaho. Buhl, no other data. Downey, June.
Kansas. No locality, Forbes and Hart (1900); this could be an error resulting from
the type collection which was made in Wyoming, then in “Kansas Territory.”
VOLUME 60, NUMBER 3
253
Montana. Billings (Cooley, 1912, 1916); April, May. Bozeman, March, 2; April,
2; May, June. Butte, no date. Cartersville, July. Edgar, May, 2. Gallatin Co., April,
4800'; May, 4; June. Glacier Park, no date. Huntley, April, May. Jefferson Co.,
April, May, 18. Rapelje, October. Ravalli (Pepper, 1951). Teton Co., no date.
Yellowstone Park, June. Nebraska. Norfolk and West Point (Cooley, 1917). North
Dakota. Bottineau, June, 3; July. Maxbass, July. Oregon. Malheur Lake and Snake
River (Hatch, 1957); Alpine, June. Utah. Benson, March. Davis Co., May. Hoop¬
er, June. Logan, April. Logan Canyon, June. Moab, May. Orme, June. Ogden,
May. Petersboro, June. Provo, April, May. Salt Lake City, May. Trenton, June.
Vernon, May (most records from McComb and Knowlton, 1950). Washington.
Okanogan, Slate Peak, 6500', August. Wyoming. Fort Bridger (Holotype 8952 in
MCZ). Pinedale (20 mi NE), 10,400', August, 4. Togwotee Pass, 9600', August.
Bionomics.— The habits of this species have been well documented by Cooley
(1917). Adults and larvae are phytophagous, eating Chenopodium album (lamb’s
quarters, introduced), Monolepis nuttalliana (povertyweed) and other native
Chenopodiaceae, as well as Solanum triflorum (nightshade) (Solanaceae) (Hatch,
1957; Cooley, 1917). They have also been recorded from squash, pumpkin, spin¬
ach, beet, wheat, radish, rhubarb, potato, lettuce, cabbage, rapeseed, and turnip
(Cooley, 1906, 1916, 1917;MacNay, 1954, 1955; Forbes and Hart, 1900; Farstad,
1949, 1951) and are considered a pest of these crops in some areas. The species
is in some North American texts on agricultural entomology (sometimes under
the name Silpha opaca). We have found adults in montane meadows, where they
were crawling along the ground and not in direct association with any single plant
species. Adults could probably be collected in meadows and grassland areas by
using large pans as unbaited pit traps.
Cooley (1917) has reported on the life cycle of the species in Montana, sum¬
marized as follows: Adults emerge very early in the spring, and lay eggs in the
soil which hatch about 5 days later. Larvae eat leaves and young shoots of the
young plants during the day, hiding in the soil during the night. The first larval
instar lasts 5 days, the second also 5 days, and the third about 15 days. Pupation
takes place in the soil and the adults emerge about three weeks later. These new
adults feed, but do not lay eggs, returning to the soil to overwinter. Other rearing
(Forbes and Hart, 1900) found larvae in June and July, with adults emerging in
July. The above records show the adults to be active from March through No¬
vember, with later dates usually being more northerly or at higher elevations.
Immature Stages
Aclypea larvae are easily distinguished from all other silphine larvae by the
presence of a ventral as well as mesal serrate edge on the mandibles (Newton,
pers. comm.). The side margins of the tergites, noted as pale by Hatch (1927) and
used to distinguish Aclypea larvae from other silphine larvae, are black in some
of the material that we have seen and hence this character is unreliable.
Third instar larvae of Nearctic Aclypea will key to Heterosilpha ramosa in
Dorsey (1940) but can be distinguished from the latter by the aforementioned
mandibular structure and by the following key:
la. Maxillary apical palpomere approximately three times as long as wide;
labial palp with the basal segment 1.5 x the length of the apical segment
. H. ramosa
254
PAN-PACIFIC ENTOMOLOGIST
lb. Maxillary apical palpomere approximately 2 x as long as wide; labial palp
with the basal and apical segments equal in length. Aclypea
We are not presently able to distinguish the larvae of A. opaca from A. bitu-
berosa. Although larvae and other immature stages of these two have been treated
by Cooley (1917), Martin (1945), and Heymons et al. (1928), the descriptions in
these papers and the key in Hatch (1927) are insufficient to provide reliable species
identifications based on material that we have available.
Evolutionary Considerations
It has been long thought that A. opaca was introduced from Europe, probably
with shipments of plants (Hatch, 1927, 1957; Forbes and Hart, 1900) and hence
was not a regular member of our fauna. However, the apparent confinement of
the species to the extreme northwestern corner of North America, and its wide¬
spread distribution in the northern Palearctic region would suggest recent dispersal
via a Beringian land bridge connection. The direction of dispersal was probably
from the Palearctic to the Nearctic, as it was for many large mammals and some
other insects during the Quaternary, and as would be expected of a group whose
highest diversity is centered in the Palearctic.
The marked similarity of the two species suggests that they are sister species,
descended from a single common ancestor. However, we cannot now demonstrate
this by synapomorphies. We suggest that this ancestor originated in the Palearctic
and dispersed to the Nearctic across the Bering bridge in the Tertiary or Pleis¬
tocene. During a Pleistocene glacial some populations of the ancestor were isolated
in localities south of the ice margin, and differentiated into A. bituberosa. North
and west of the ice margin, in unglaciated refugia in Alaska and the Yukon, A.
opaca retained intermittent contact with Palearctic conspecihcs, or became extinct
and has reoccupied the Nearctic in the Recent. The lack of far northern specimens
of A. bituberosa indicates that it did not survive glaciation in the northern refugia,
but only in areas south of the ice sheets. Subsequent reinvasion of previously
glaciated land has therefore been from the south, resulting in the present distri¬
bution pattern.
Unfortunately, no fossil specimens of Aclypea have been recorded in North
America which can test these evolutionary hypotheses. 1
Acknowledgments
We thank the following curators for allowing study of material in collections
in their care: Lee Herman, AMNH; David Kavanaugh, CASC; Ales Smetana,
CNCI; Sharon Rose, DZEC; Henry Dybas, FMNH; A. F. Newton, MCZC; D.
Shepeley, UASM; S. G. Cannings, UBCZ; W. F. Barr, UICM: T. J. Spillman,
USNM; T. D. Galloway, Univ. of Manitoba, and J. L. Carr. The manuscript was
reviewed by A. F. Newton, Jr., S. E. Miller, J. V. Matthews, Jr., R. B. Madge,
and G. E. Ball.
Literature Cited
Anderson, R. S. 1981. The biology and distribution of the Silphidae and Agrytidae of Canada and
Alaska (Insecta: Coleoptera). M.Sc. thesis, Carleton University, Ottawa, 279 pp.
Arnett, R. H. 1944. A revision of the Nearctic Silphini and Nicrophorini based upon the female
genitalia. J. New York Entomol. Soc., 52:1-25.
VOLUME 60, NUMBER 3
255
-, and G. A. Samuelson. 1969. Directory of Coleoptera collections of North America (Canada
through Panama). Dept. Entomol., Purdue Univ., Lafayette, Indiana, 123 pp.
Cooley, R. A. 1906. Twelfth Annual Report, pp. 255-273. Montana Agric. Coll. Exp. Sta., Bozeman.
-. 1912. Tenth annual report of the state entomologist of Montana, pp. 51-67. Montana Agric.
Coll. Exp. Sta. Bull. 92.
-. 1916. Thirteenth annual report of the state entomologist of Montana, pp. 151-160. Montana
Agric. Coll. Exp. Sta. Bull. 109.
-. 1917. The spinach carrion beetle. J. Econ. Entomol., 10:94-102.
Farstad, C. W. 1949. Insects of the season in Alberta, 1949. Canadian Insect Pest Review, 28:
16-19.
-. 1950. Insects of the season in Alberta, 1950. Canadian Insect Pest Review, 29:16-22.
Forbes, S. A., and C. A. Hart. 1900. The economic entomology of the sugar beet. Univ. Illinois
Agric. Exp. Sta. Bull., 60:397-517.
Hatch, M. H. 1927. Studies on the Silphinae. J. New York Entomol. Soc., 35:3311-3371.
-. 1928. Coleopterorum Catalogus. Family Silphidae II. 95:1-244. Junk, The Hague.
-. 1957. The beetles of the Pacific Northwest. Part II. Staphyliniformia. University of Wash¬
ington, Publ. Biol. Vol. 16. Univ. Wash. Press, Seattle, 386 pp.
Heymons, R. H., H. von Lengerken, and M. Bayer. 1929. Studien fiber die Lebensercheinunger der
Silphini. IV. Blitophaga opaca L. Zeit. Morphol. Okol. Tiere, 14:234-260.
Horn, G. H. 1880. Synopsis of the Silphidae of the United States with reference to the genera of
other countries. Trans. Amer. Entomol. Soc., 8:219-322.
LeConte, J. L. 1859. The Coleoptera of Kansas and Eastern New Mexico. Smithsonian Contr.
Knowledge, 2:1-58.
Linnaeus, C. 1758. Systema naturae per regna tria naturae secundum classes, ordines, genera, species,
cum characteribus, differentis, synonymis, locis. Ed. 10, Vol. 1. Holmiae, 823 pp.
MacNay, C. G. 1954. Summary of important insect infestations, occurrences, and damage in Canada
in 1954. Canadian Insect Pest Review, 32:342-389.
-. 1955. Summary of important insect infestations, occurrences, and damage in Canada in
1955. Canadian Insect Pest Review, 33:311-380.
Martin, H. 1945. Contribution a l’etude des Silphes de la betterave en Suisse. Landwirtschaftliches
Jahrbuch der Schweiz, 59:757-819.
McComb, D., and G. F. Knowlton. 1950. Some Utah Coleoptera belonging to minor families. Utah
Agric. Exp. Sta., Utah Agric. Coll. Mimeo Ser. 369 (supp. 2, Coleoptera, of Mimeo Ser. 200,
1939).
Miller, S. E., and S. B. Peck. 1979. Fossil carrion beetles of Pleistocene California asphalt deposits,
with a synopsis of Holocene California Silphidae (Insecta: Coleoptera: Silphidae). Trans. San
Diego Soc. Natur. Hist., 19:85-106.
Pepper, J. H. 1951. Montana Insect Pests 1949-1950. Montana State Coll. Agric. Exp. Sta. Bull.
474, pp. 1-35.
Portevin, G. 1926. Les grands Necrophages du globe. Encyclopedie Entomologique 6. Lechevalier,
Paris, 270 pp.
Seidlitz, G. 1887-1891. Fauna Baltica. Die Kafer (Coleoptera) der Deutschen Ostseeprovinzen
Russlands. Zweite neu bearbeitete Auflage. Konigsberg: Haftungsche Verlagsdruckerei. (Sil¬
phidae issued in 1888.)
1 After this paper had gone to press, a single record of a fossil specimen of Aclypea was sent to us
by Dr. R. E. Nelson, Colby College, Maine. The specimen was collected from deposits of age 16,640
years BP at Seattle, Washington, and is assignable to A. bituberosa based on elytral sculpture and
punctation characteristic of that species. This discovery indicates that isolation and structural diver¬
gence of A. bituberosa did not occur in the glacial advance of the Late Wisconsinan.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 256-257
Scientific Note
The Occurrence of Trypoxylon clavicerum in
North America (Hymenoptera: Sphecidae)
Trypoxylon ( Trypoxylon ) clavicerum Lepeletier and Serville is a widespread
Palearctic wasp, occurring from Northern Europe to Japan (Bohart and Menke,
1976, Sphecid wasps of the world, University of California Press, 695 pp.). In the
course of a systematic study of Trypoxylon wasps in North America, I discovered
that T. clavicerum now occurs in the Great Lakes Region of the United States
and Canada. North American specimens were compared with examples of clav¬
icerum in the collections of the British Museum of Natural History (London),
Oxford University, and the Zoologisk Museum (Copenhagen). No differences
between the North American and European specimens were found.
Figures 1-4. 1. Head of female Trypoxylon clavicerum showing the width of the face (arrows)
below the transverse interantennal carina (t). 2. Head of female T. kolazyi showing width of face
(arrows). 3. Head of male T. clavicerum showing the two minute medial teeth (arrow) on the clypeal
apex. 4. Head of male T. kolazyi showing the single medial tooth (arrow) on the clypeal apex.
VOLUME 60, NUMBER 3
257
Trypoxylon clavicerum is probably a recent introduction into North America.
The oldest record is a female specimen from Van Buren Co., Michigan, taken on
11 June 1949. Specimens had been collected in Indiana, Ontario, Pennsylvania,
and several counties in Michigan by 1962.
In the most recent key to the Nearctic species in the subgenus Trypoxylon
(Sandhouse, 1940, Am. Midi. Nat., 24:133-174) T. clavicerum keys out to T.
backi Sandhouse, a junior synonym of T. kolazyi Kohl (see Krombein, 1979, in
Krombein et al., Cat. Hymen. Am. N. Mex., Vol. 2, Apocrita (Aculeata), p. 1644).
Trypoxylon clavicerum is easily confused with T. kolazyi. In females the face
below the transverse interantennal carina is much more narrow in T. clavicerum
(Fig. 1) than in T. kolazyi (Fig. 2). In males the clypeal apex of T. clavicerum has
two minute medial teeth (Fig. 3), whereas the clypeal apex of T. kolazyi has a
small medial tooth with an even smaller tooth or angulation on each side (Fig.
4).
Specimens ofT. clavicerum examined. — CANADA. Ontario. 1 6 London, VI-
1957, N. R. Couling (Canadian National Collection, Ottawa = CNC); 2 2 Guild
(?), 6 and 14-VII-1962 (CNC). UNITED STATES. Indiana. Hamilton Co.: 1 2
Cicero, 10-VII-1954, J. M. Kingsolver (Illinois State Natural History Survey).
Michigan. Branch Co.: 1 6 4-VII-1972, T. A. Bowling (Michigan State Univer¬
sity - MSU). Calhoun Co.: 12 6 Tekonsha, 2-VII-1972, T. A. Bowling (MSU).
Clare Co.: 1 2 23/28-VII-1959, R. R. Dreisbach (MSU). Gladwin Co.: 1 2 28/30-
VII-1959, R. R. Dreisbach (MSU). Ingham Co.: 1 6 East Lansing, 30-VIII-1960,
G. Eickwort (MSU). Iosco Co.: 24 6, 7 2 Tawas City, 8-VII-1973, I. J. Cantrall
(University of Michigan). Kalamazoo Co.: Gull Lake Biol. Sta., 1 2 2-VII-1956,
1 6 2-VII-1957, 6 5,52 1-VII/27-VIII-1958, 23 <5, 2 2 28-VI-1961, 16 1 2 26-
VI/11-VII-1965, 1 2 9-VII-1970, R. L. Fischer (MSU). Midland Co.: 1 2 17-IX-
1958, 1 2 12/16-VII-1960, R. R. Dreisbach (MSU). Saginaw Co.: 7 6, 16 2 St.
Charles, 28-V/16-VIII-1969, rotary trap at ground level-12 ft height, J. G. Truchan
(MSU). Van Buren Co.: 1 2 11-VI-1949, Royji Namda (MSU). Pennsylvania. Erie
Co.: 16 2 2 Presque Isle St. Pk., 15/20-VII-1961, F. E. Kurczewski (Museum of
Comparative Zoology, Harvard).
Rollin E. Coville, Division of Entomology and Parasitology, 201 Wellman Hall,
University of California, Berkeley, California 94720.
PAN-PACIFIC ENTOMOLOGIST
60(3), 1984, pp. 258-263
Neosminthurus bellingeri , a New Species from California
(Collembola: Sminthuridae ) 1
Suzanne F. Muzzio
111 South 12th St., San Jose, California 95112.
Abstract.—A new species, Neosminthurus bellingeri Muzzio, is described from
a survey of Collembola taken at Grover Gulch, near Bates Creek in Santa Cruz
County, California. This was the first time a member of this genus has been
collected on the West Coast. Some characteristics which separate it from other
members of the genus are the presence of ventral dental setae, absence of posterior
finger-like process on meso- and metafemora, weak subsegmentation on fourth
antennal segment and six cylindrical setae on the posterior margin of head.
A systematic and ecological study of Collembola was taken from April 1980
through April 1981. The study area was located approximately three miles north-
northeast from Soquel, in Grover Gulch near Bates Creek, in the Santa Cruz
Mountains, California. Sampled areas were fairly remote from human habitation.
During the survey a new and undescribed species of Neosminthurus was discov¬
ered. The purpose of this paper is to describe that new species.
1 mm
Figure 1. Neosminthurus bellingeri, n. sp. lateral view, habitus.
1 Partial fulfillment of Master’s Thesis at San Jose State University.
VOLUME 60, NUMBER 3
259
Figures 2-15. Neosminthurus bellingeri, n. sp. 2. Antenna, holotype. 3. Left eyepatch, paratype.
4. Cylindrical setae on posterior margin of head and interocular setae, holotype. 5. Metatrochanter
and femur, paratype. 6. Profemur, paratype. 7. Metatibia, paratype. 8. Fore foot, paratype. 9. Hind
foot (slightly damaged), paratype. 10. Abdominal setae, paratype. 11. Body setae, holotype. 12. Neo-
sminthuroid seta, paratype. 13. Furcula (=manubrium, dens, mucro), dorsal view, paratype. 14. Female
subanal appendage, lateral view, paratype. 15. Tenaculum, holotype.
260
PAN-PACIFIC ENTOMOLOGIST
VOLUME 60, NUMBER 3
261
14
262
PAN-PACIFIC ENTOMOLOGIST
Neosminthurus bellingeri. New Species
Antenna uniformly dark blue. Head mottled dark blue. Body dorsally mottled
dark blue to black, paler ventrally with fine intersegmental lines separating thoracic
segments. Legs, collophore and furcula speckled blue (Fig. 1).
Antenna IV with six to seven weak subsegments, without subapical bulb and
1.3-1.5 times as long as Ant. III. Ant. Ill with two subapical sense rods lying in
deep depressions (Fig. 2). Eyes 8 + 8 with ocellus A one-half the diameter of B;
D at least one-third diameter of B (Fig. 3). Head with six cylindrical setae located
on posterior margin of head; interocular setae short and blunt (Fig. 4). Metatro¬
chanter with posterior spine, five anterior setae and oval organ (Fig. 5). Profemora
with one appressed posterior spine (Fig. 6). Meso- and metafemora without pos¬
terior finger-like process. Metatibia with an oval organ located two-thirds distance
from apex on inner edge, five long setae on inner edge, four to five short setae on
outer edges and four setae on posterior surface (Fig. 7). Tarsus with anterior and
posterior setulae. Tenent hair acuminate. Unguis with inner tooth, tunica and
pseudonychium (=lateral serrations) present (Fig. 8). Unguiculus lamellate with
apical filament lanceolate (Fig. 9). Body with cylindrical scaled setae and smooth
setae (Figs. 10, 11). Neosminthuroid setae present and located between bothriotrix
C and base of furcula (Fig. 12). Manubrium with 14 dorsal setae. Dens with seven
inner dorsal setae, five external setae, two lateral internal setae, and one dorsal
seta; ventrally with one pair of subapical ventral setae, Ve 3 seta short, and located
one-third the distance from base of dens (Fig. 13). Mucro with inner lamella
serrate and outer lamella entire. Female subanal appendage curved and acuminate
with bilateral apical ciliations (Fig. 14). Sacs of ventral tube tuberculate. Tenacu¬
lum with four distal setae and 3 + 3 teeth (Fig. 15). Maximum length 1.2 mm.
Diagnosis
Neosminthurus bellingeri keys out closest to N. bakeri in Christiansen and
Bellinger (1980-81). It can be separated from other members of the group by the
following characteristics:
N. bellingeri
ventral setae present
on dens
dens with 5E, 7Id, 2L,
& ID setae
ant. IV weakly subseg¬
mented
N. bakeri
ventral setae absent
on dens
dens with 5E, 8Id, 3L,
& ID setae
ant. IV not subseg¬
mented
N. richardsi
ventral setae absent
on dens
dens with 4E, 7Id, &
3L setae
ant. IV not subseg¬
mented
N. clavatus
ventral setae absent
on dens
dens with 4E, 8Id, &
3L setae
ant. IV not subseg¬
mented
Features which commonly associate N. bellingeri with the Neosminthuroid
group are presence of the Neosminthuroid seta, the well-developed trochanteral
spine, the peculiar modification of head and body setae, and the slight bifid
appearance at the apex of the mucro.
It seems appropriate to extend the diagnosis for this genus to include the fol¬
lowing characteristics which may be present: Ant. IV weakly subsegmented, and
ventral setae on dens. In appreciation for his support and encouragement this
species was named after Dr. P. F. Bellinger, Professor of Biology, Northridge
University.
VOLUME 60, NUMBER 3
263
Ecological Notes
N. bellingeri was discovered during the wettest months of the year. Subadults
were collected in January, February, and April; adults were taken in March.
Specimens were extracted from Tanbark-oak Lithocarpus densiflora (H. & A.)
Rehd. and Pacific madrone Arbutus menziesii Pursh. leaf litter. Specimens were
found as far down as five centimeters in the soil.
The upper soil layer at the study site has a fine sandy clay loam composition.
The litter layer pH ranged from 4.45 to 4.80. Normal precipitation for the area
averages 1219 mm (48 inches), however, rainfall was below normal when the
survey was being conducted and amounted to only 482 mm (19 inches).
Species commonly found with N. bellingeri were Tomocerus reductus (Mills),
1949; Sinella sexoculata (Schott), 1896; Hypogastrura {Ceratophysella) spp.; Ta-
fallia robusta (Scott), 1961; Onychiurus Jlavescens Kinoshita, 1916; and Folsomia
Stella Christiansen-Tucker, 1977.
HOLOTYPE (female), ALLOTYPE (male) and PARATYPES from California,
Santa Cruz County, three miles north-northeast of Soquel, at Grover Gulch near
Bates Creek. Specimens were collected in Tanbark-oak and Pacific madrone leaf
litter in the following months: 11/Jan/81, 7/Feb/81, 7/Mar/81, and 11 /Apr/81
by Suzanne Muzzio. Holotype, allotype, two mounted paratypes, and fifteen spec¬
imens in alcohol deposited at California Academy of Sciences, San Francisco;
two at the Entomology Museum, San Jose State University; and one to Dr. P. F.
Bellinger, Northridge University.
Acknowledgments
The author wishes to thank Brian, Andrew and Michael Muzzio for their as¬
sistance with the collection of specimens, and Drs. P. F. Bellinger and R. J. Snider
for reviewing this manuscript.
Literature Cited
Christiansen, K., and P. F. Bellinger. 1980-81. The Collembola of North America, North of the Rio
Grande. Grinnell College, Grinnell, Iowa, 1322 pp.
264
PAN-PACIFIC ENTOMOLOGIST
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Vol. 60
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No. 4
THE
Pan-Pacific Entomologist
PHILIP, C. B .—Lanellus, a new subgenus of primitive, California “horse flies” (Diptera, Ta-
banidae). 265
EDMUNDS, G. F., JR. and D. N. ALSTAD—High summer mortality of black pineleaf scale
(Homoptera: Diaspididae). 267
WEISSMAN, D. B.—Notes on the autecology, cytology, morphology, and crepitation of Tri-
merotropis grasshoppers (Orthoptera: Oedipodinae) . 269
CHEMSAK, J. A. and E. G. LINSLEY—New genera and species of neotropical Cerambycidae
(Coleoptera). 279
LYON, R. J.—New cynipid wasps from California (Hymenoptera: Cynipidae). 289
DODSON, G. N. and D. T. GWYNNE—A digger wasp preying on a Jerusalem cricket. 297
CLEMENT, S. L. — Observations on the behavior of Triepeolus nr. eldredi Cockerell (Hyme¬
noptera: Anthophoridae). . 300
BLOM, P. E. and W. H. CLARK —Phobetus desertus, a new Melolonthine Scarabaeidae (Co¬
leoptera) from the central desert of Baja California, Mexico. 304
GARCIA-VIDAL, M.—The genus Phyllophaga Harris, 1826 (Coleoptera: Scarabaeidae) in
Cuba. II. Descriptions of eleven new species and illustrations of female genitalia of twelve
other Phyllophaga . 313
MANGAN, R. L. — Reproductive ecology of three cactophilic Diptera (Diptera: Drosophilidae,
Neridae, Syrphidae). 326
CAMPBELL, C. L. and K. S. PIKE—Descriptions of the life stages of Pyrausta orphisalis
(Lepidoptera: Pyralidae). 332
HEPPNER, J. B .—Fabiola quinqueferella : An obscure California moth formerly in Glyphip-
terigidae (Lepidoptera: Oecophoridae). 337
FREYTAG, P. H. and P. S. CWIKLA—Two new species of Idiocerine leafhoppers from Mal¬
agasy Republic (Homoptera: Cicadellidae)... 341
SCHWAN, T. G.—Nosopsyllus fasciatus parasitizing house mice on southeast Farallon Island,
California (Siphonaptera: Ceratophyllidae). 345
MURPHY, D. D. and R. R. WHITE—Rainfall, resources, and dispersal in southern populations
of Euphydryas editha (Lepidoptera: Nymphalidae). 350
DESCAMPS, M.—A new grasshopper of the genus Eumorsea from Baja California, Mexico
(Orthoptera: Eumastacidae). 355
SCIENTIFIC NOTES.357, 358, 360
PROCEEDINGS—420th to 427th meetings. 361
INDEX TO VOLUME 60 .,. 367
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Published by the PACIFIC COAST ENTOMOLOGICAL SOCIETY
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PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 265-266
Lanellus, a New Subgenus of Primitive, California
“Horse Flies” (Diptera, Tabanidae)
Cornelius B. Philip
California Academy of Sciences, San Francisco, California 94118.
Tabanid flies related to the generalized genus, Apatolestes, comprise a rather
primitive and sometimes pestiferous group in the Tribe Pangoniini. It consists of
13 described species from western North America, 11 of which are known to
occur in California (Middlekauff and Lane, 1980). Some deviation from the an¬
cestral stem has occurred, which permitted separation of the subgenus (now genus)
Brennania Philip. Information on the biologies of Apatolestes species has re¬
mained fragmentary until quite recently when studies conducted by Dr. Robert
S. Lane, University of California, Berkeley, and colleagues (Lane and Anderson,
1983; Lane et al., 1983) significantly added to our knowledge of Apatolestes actites
Philip and Steffan, an inhabitant of sandy ocean beaches in California. Morpho¬
logical and behavioral peculiarities of this species indicated the need to distinguish
it at least subgenerically from other members of the genus.
The new subgenus of Apatolestes, viz., Lanellus, n. subg., is proposed here with
actites (1962) designated as subgenotype species. Distinctive morphological char¬
acters of this subgenus are as follows: females with very wide fronts, strongly
convergent above, with subequal sides and basal widths and small but prominent
basal callosities; mandibular and maxillary stylets reduced and simple, the man¬
dibles lacking the usual serrated teeth characteristic of blood sucking tabanids.
Bodies of both sexes unusually pilose, obscuring the palpi and mouthparts. Palpi
of males more attenuate, less truncated than in most other Apatolestes species
(except rossi Philip and colei Philip). Eyes bare, unbanded in life.
Some unusual biological features of actites were observed initially by Dr. Lane,
for whom the subgenus is cordially named. Among tabanids, females of A. actites
are unique by laying their eggs subterraneously in amphipod or isopod burrows,
and by possessing the capability to produce two batches of eggs autogenously.
Neither flower visitation nor mating was noted in the vicinity of the breeding
sites. The psammophilous larvae are cylindrical, elongate, and non-tabaniform in
appearance; they live in the upper beach (supralittoral zone) in soils that are aridic
for at least six months annually.
Mackerras (1954) postulated that the primitive tabanid progenitors were gen¬
eralized flies that fed “on the juices of plants,” and that an “adaptation to blood
sucking” was an early trend in their evolution. Almost all modem tabanid species
females require a blood-meal to mature their developing eggs. By contrast, A.
actites is the first member of a predominantly blood-sucking brachycerous taxon
that can develop more than a single batch of eggs autogenously (Lane and An¬
derson, 1983), which suggests that A. actites never deviated from the progenitorial
non-biting line.
Apatolestes colei is the other member of the genus that may be subgenerically
266
PAN-PACIFIC ENTOMOLOGIST
related to A. actites. This rare horse fly was originally taken along the sandy banks
of the Santa Ana dry wash at La Quinta in southern California. The frons of the
female is likewise unusually wide, with an index of about 1/1. It also has a reduced
and bare basal callosity, the eyes are unbanded in both sexes, and males have
attenuated palpi. The mouthparts have not been examined because to do so would
require dissection, and only one more female has been reported (Middlekauff and
Lane, 1980) since description of the holotype in 1941. Consequently, non-he-
matophagy and autogenic egg development accompanied by reduced mouthparts,
though postulated, remain to be confirmed.
Literature Cited
Lane, R. S., and J. R. Anderson. 1983. Multiple autogeny and burrow oviposition by a marine horse
fly (Diptera: Tabanidae). J. Med. Entomol., 20:212-213.
-,-, and C. B. Philip. 1983. Biology of autogenus horse flies native to coastal California:
Apatolestes actites (Diptera: Tabanidae). Ann. Entomol. Soc. Am., 76:559-571.
Mackerras, I. M. 1954. The classification and distribution of Tabanidae (Diptera). I. General review.
Austral. J. Zool., 2:431-454 (III. Evolution and distribution, pp. 440-452).
Middlekauff, W. W., and R. S. Lane. 1980. Adult and immature Tabanidae (Diptera) of California.
Bull. Calif. Insect Surv., 22:1-99.
Philip, C. B., and W. A. Steffan. 1962. New North American Tabanidae. XIV. An undescribed
Apatolestes from the California coast. Pan-Pac. Entomol., 38:41-43.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 267-268
High Summer Mortality of Black Pineleaf Scale
(Homoptera: Diaspididae)
George F. Edmunds, Jr. and Donald N. Alstad
(GFE) University of Utah, Salt Lake City, Utah 84112; (DNA) University of
Minnesota, Minneapolis, Minnesota 55455.
Abstract .—Catastrophic summer mortality among first-instar larvae of the black
pineleaf scale occurred as a result of combined high temperatures and low relative
humidity.
Although density-independent factors are unlikely to regulate or stabilize animal
populations, infrequent catastrophes affect the density and abundance of insects.
Because long-term observation is required, documentation of such environmental
episodes is rare. We have studied populations of the black pineleaf scale ( Nucu-
laspis californica (Coleman)) in eastern Washington since 1955 and have quan¬
titative yearly data on reproductive success since 1973. The sedentary life history
of these insect herbivores allows direct observation of seasonal patterns of mor¬
tality and survivorship. Edmunds (1973) has shown that abrupt early winter or
spring episodes of freezing weather causes annual shifts in the lower elevation at
which black pineleaf scale survive, and excludes the species from large areas in
one eastern Washington valley. Fligher altitude kills are infrequent and unpre¬
dictable, occurring about one year in five.
Here we report a catastrophic episode of high summer mortality among first-
instar larvae of the black pineleaf scale during their dispersal period. Eggs hatch
in July as the new needle crop of ponderosa pines reaches its full elongation. The
tiny larvae move from beneath the mother’s scale, and travel 5 to 25 cm before
establishing positions on new pine needles. The entire population completes this
dispersal within a 10 day interval. During this period in July 1979, unusually
high daytime maxima ranged near 40.5°C, and midday relative humidity dropped
to 5%. The long-term means for this period in the Spokane area are 30°C and
28% relative humidity. We observed many moribund or dead larvae as a result
of desiccation, and more than two-thirds of the insects died before completing
their normal dispersal and establishment. The resulting colonization patterns were
highly variable and abnormal in distribution, and the number of established
colonists was substantially below that which we have observed in other years. No
similar summer catastrophe has occurred since our observations began in 1955.
We quantified the effect of this event on the 1979-1980 scale insect population
with random twig samples from 37 infested ponderosa pine in the spring of 1980,
and compared these observations with data collected in the spring of 1982. Scale
cover numbers on the pine needles were used to estimate insect population den¬
sities, which were scored in 11 density categories (0-10) representing infestations
ranging from 0 to 80 insects per needle. These data were compared with similar
268
PAN-PACIFIC ENTOMOLOGIST
density estimates from the 1978-1979 needles on the same twigs, and with 1981—
1982 needles from new twigs sampled from the same plot of pines.
On 4 of the 37 trees, the population density class was the same on 1979 needles
and 1978 needles (all carried low densities (class = 1)). Population density was
lower for the 1979 needles on 33 of 37 trees. Eighteen of the 37 trees showed a
reduction in density to one half or less of the 1978 level. One sample dropped
from class 10 to class 3. The comparison of the two years’ populations cannot be
precise because the 1978 needles also have some 1979 scale insects, and many of
the 1978 scale coverings were gone. Nevertheless, the work was done in long¬
term study plots where we have repeatedly sampled, and it was obvious that the
scale population fell to less than half its previous density. In recent years many
of the trees in the plot would have carried population class 8 to 10. The highest
population interval assigned to any of the twigs for 1979 needles was 5, although
previously a number of the trees would have had twigs of population intervals 6
to 10, and such densities were reestablished on 1981 needles.
Another evidence of the interference of the hot, dry period is the position of
the needles on which the larvae settled. Usually the scale population is distributed
rather evenly on the needles of a twig. The distribution of scale insects on 1979
needles was strongly biased toward basal needles on each twig, positions that
require the shortest movement. Twigs were randomly sampled from 44 infested
trees and the scale population estimated and assigned to population intervals on
the needles of the basal l A and of the apical l A of the twig. On 9 of 44 samples
the populations at the base and apex of the twig were assigned to the same
population class. The other 35 twigs all had lower population classes at the apex
with an average reduction of 2.0 population classes. Twigs longer than 6 inches
had a more striking reduction from base to apex than those shorter than 6 inches,
the average population class drop being 2.6 classes as contrasted with 1.77 on
those less than 6 inches. A count of scales on 10 twigs 6 or more inches long
showed an average of 81.8 scales on needles of the basal l A of the twig and 3.8
on the apical l A. We repeated such measurements in the same small plot in 1982.
Thirty-eight of 52 samples had the same population class at the base and apex.
Of the other 14, the apex on 4 had higher populations than the base and 10 had
lower populations than the base. There are no significant differences in populations
at the base and apex, and no difference in distribution on the 12 twigs longer than
6 inches (0.3 classes higher at apex vs. 0.1 higher at apex) as compared to the 40
twigs shorter than 6 inches.
The combination of direct observations on the moribund and dying scale larvae
in July 1979 and data from scale densities and within twig distribution in May
1980 and May 1982 indicate that this was a short-term climatic mortality episode
that directly affected the scales, although we cannot rule out that stress on the
host pine may have played a role. It is likely that high temperature-low humidity
episodes during the crawler stage affect the population dynamics of other scale
species.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 269-278
Notes on the Autecology, Cytology, Morphology, and
Crepitation of Trimerotropis Grasshoppers
(Orthoptera: Oedipodinae)
David B. Weissman
Department of Entomology, California Academy of Sciences, Golden Gate
Park, San Francisco, California 94118.
Abstract.— Numerous additions to the known cytological, morphological, and
crepitational characteristics of the Trimerotropis grasshoppers are presented. Aut-
ecological data are given for several infrequently collected species. T. santamonica
Rentz and Weissman is synonymized with T. thalassica Bruner.
Recent efforts have resulted in many additions and a few corrections making
a supplement to our 1980 paper (Weissman and Rentz, 1980) on the cytological,
morphological, and crepitational characteristics of the oedipod grasshopper genus
Trimerotropis Stal appropriate at this time. Specifically, characteristics of several
other species have been investigated; Baja California, Mexico, has been thoroughly
collected; large series at the Academy of Natural Sciences of Philadelphia (ANSP)
and University of Michigan Museum of Zoology (UMMZ) have been examined;
and overlooked literature references have been found. Several problem areas
concerning species’ ranges and species’ identifications have been uncovered.
The 62 species (Rentz and Weissman, 1980, 1981, 1984) presently constituting
the genus show chromosomal, morphological, and crepitational variation both
geographically and within populations. There are several reasons for supple¬
menting this catalogued variation, although the significance of such variation is
not always understood. First, chromosomal variation within species (e.g., White,
1951b) may be indicative of incipient speciation. Thus, the geographical vari¬
ability in karyotype number seen in the gracilis subspecies and in cyaneipennis
Bruner may actually represent different species. A similar situation exists in eastern
U.S. verruculatus (Kirby) (2N 3 = 21) and western U.S. sujfusa Scudder (2n 3 =
23). Both species are morphologically similar. Helwig (1955) was readily able to
hybridize the two forms yet unequivocally believed them not to be conspecific
based on ecology, behavior, and cytology. In any case, only detailed study of such
variable taxa, especially in areas of overlap, can resolve the question of conspe-
cificity, although there is also no agreement as to what degree of reproductive
isolation is necessary to constitute such speciation. Second, chromosomal vari¬
ation between species may serve as a species’ identification marker. For example,
until recently (Rentz and Weissman, 1981), any coastal California Trimerotropis
with blue legs and greenish wing disks was called occidentalis (Bruner) and Stro-
hecker et al. (1968) list its range as extending east to Idaho. Yet this species has
a unique karotype consisting of only 21 chromosomes, a telocentric X (sex) chro¬
mosome, and inversion polymorphism in several autosomes (see below). Such
270
PAN-PACIFIC ENTOMOLOGIST
information has enabled me to considerably restrict the range of this species (see
below). Also, chromosomal variation between species has permitted confirmation
of suspected hybridization (John and Weissman, 1977; John et al., 1983). Third,
knowledge of the range of morphological variation can facilitate discovery of
cryptic species; for example, the “abberant” blue legged specimens of californica
Bruner noted by Strohecker et al. (1968) were actually an undescribed species—
topanga Rentz and Weissman. Morphological variation in the genus is most
notable in wing disk and hind tibia color. Examination of large series at ANSP
and UMMZ shows greater color variation in hind tibia than previously noted
(Weissman and Rentz, 1980): species primarily with yellow hind tibia have some
individuals with brown ones; yellow tibia are also associated with orange ones,
and blue are associated with brown. When polymorphic, such variation is usually
present in each population. Some of this variation, when described from museum
specimens, may be related to changes after death. Those species variable for wing
disk color usually do not have all morphs represented in each population.
Species are discussed below in alphabetical order. Table 3 is arranged by karyo¬
typic characteristics and supplements information in Table 1 in Weissman and
Rentz (1980). Karyotypic Section A contains species whose chromosomes are all
rod shaped or telocentric. Section B species (but not all individuals) have some
autosomes metacentric with the X chromosome always (except in occidentalism
metacentric. The latter species has a telocentric X but is considered a Section B
species because of two characteristics only possessed by Trimerotropis taxa in that
group: inversion polymorphism and reduced chromosome number. Also it can
form viable hybrids with another Section B species, thalassica Bruner, from which
it differs by only one reciprocal translocation (John and Weissman, 1977). I believe
species within these respective chromosomal Sections are more closely related to
each other than to species in the other Sections (discussed in Weissman and Rentz,
1980). For ease of discussion the autosomes are arbitrarily divided into three size
classes: large (L), medium (M), and small (S), and are numbered from the largest
to the smallest. Crepitational terminology is after Weissman and Rentz (1980).
T. californica.—In California, this species was separated from strenua McNeill
by Rentz (pers. comm.) in Strohecker et al. (1968) on the following basis: in
californica the tegminal bands are light, narrow, and tend to be obscure or speckled;
in strenua the bands are heavy, dark, and not speckled. I feel these species may
still be distinct for the following reasons, although crepitationally, cytologically,
and ecologically inseparable. In California any one population does not contain
both above morphs. On the other hand several populations of strenua morphs
that I collected from San Luis Obispo and Santa Barbara Counties were variable
for tegminal band intensity: of 21 males and females from the Santa Ynez Moun¬
tains, 6 lacked the bands completely (making them the palest Trimerotropis spec¬
imens known), 5 had fully developed bands, and the rest had intermediate bands.
Similar intensity variation has never been observed in a California population
containing californica morphs (but see Hebard, 1906, for notes on specimens from
Utah) in which the tegminal bands are fairly uniform. Because of these differences
in population variation, further investigations on the relationship of these two
taxa seem warranted.
T. cyaneipennis.— Although recorded (Strohecker et al., 1968) from western
Texas to California (of questionable occurrence in the latter state—see Rentz and
VOLUME 60, NUMBER 4
271
Weissman, 1984 under leucophaea, new species) and north to southern Idaho,
several species may be involved. Specimens at ANSP and UMMZ from Utah,
Nevada, and Idaho consistently have blue wing disks. Some Oregon, Arizona
(Tinkham, 1948), and most New Mexico specimens (also see Hebard, 1935) have
disks that are green, green-blue, or yellow with a blue tinge at the base (some of
these may be hybrids between cyaneipennis and sujfusa— see John et al., 1983).
Crepitation is a crackle similar to fratercula McNeill in Otte (1970, p. 123) (in¬
cluding those from Harney Co., Oregon—listed in error in Weissman and Rentz,
1980) as noted in Rehn and Hebard (1908), Hebard (1935), Strohecker et al.
(1968) and confirmed in specimens from Kane Co., Utah (Weissman, unpubl.).
T. fontana Thomas.—This species is polymorphic for wing color and crepita¬
tion. Wing disk color is yellow-green, but specimens from near Laguna Hanson,
northern Baja California, have some blue in the wings. This case represents the
only Trimerotropis with blue in the wing that contains no metacentric chromo¬
somes ( arizonensis Tinkham is not yet known). Populations are variable for crep¬
itation pattern from northern Baja California to northern California. They vary
from a buzz to a pulse buzz to a crackle (Caudell, 1908; Fulton, 1930; Strohecker
et al., 1968; Weissman, unpubl.) frequently with two types heard in one popu¬
lation. One of three individuals from 9.6 km S Ruth, Trinity Co., California, was
heterozygous for a supernumerary segment in a small autosome. T. fontana from
Santa Rosa Island also had similar segments (Rentz and Weissman, 1981, Fig.
132). The X chromosome in the Ruth heterozygote also showed neometacentric
activity at metaphase I.
T. fratercula.—One male from Sandoval Co., New Mexico, was examined cy-
tologically: 2N <5 = 23, Section B, 3 large, 6 medium, and 2 small autosomes; LI,
L2, L3, and M5 were inversion homozygotes, X a metacentric. On the basis of
its yellow wings, yellow hind tibia (rarely individuals have a hint of green), definite
tegminal bands, crackle crepitation, and Section B chromosomal pattern, this
species is probably a member of the Pallidipennis Group (see Weissman and
Rentz, 1980).
T. helferi Strohecker.—This species was restudied 2 August 1980, at Cleone,
Mendocino Co., California, on a clear, sunny day. Individuals were extremely
difficult to collect, approaching Scirtetica Clementina Rentz and Weissman in
wariness (Rentz and Weissman, 1981, p. 75). Male crepitation was a pulse buzz
similar to cincta (Thomas) in Otte (1970), not a buzz as listed in Weissman and
Rentz (1980). Females made more of a flight noise than an actual crepitation.
Flights by both sexes were low and from 2 to 30 m in length. One last instar
female was seen; adult females contained nearly mature eggs. In the 14 captured
specimens, wing disk color was variable: clear (n = 3); greenish tinged (n = 7);
greenish yellow (n = 4). All these wing colors apparently fade out completely after
death as judged by most museum specimens.
Otte (pers. comm.) notes the morphological similarity in body shape and hir¬
suteness, among other characters, between this species and another California
oedipod Microtes occidentalis (Bruner) that likewise exclusively inhabits coastal
sand dunes. Such similarity is most probably the product of convergence from
inhabiting a similar habitat. Such convergence is widespread among the grass¬
hoppers, the genera Anconia Scudder, Xeracris Caudell, and Spaniacris Bruner
being a classic example. The three are found, often sympatrically, in sandy desert
272
PAN-PACIFIC ENTOMOLOGIST
Table 1. Karyotype frequency (2N <3 = 21) of 80 T. occidentalis from Woodside, San Mateo Co.,
California, during 1973 (habitat since destroyed for houses—see Murphy and Ehrlich, 1980).
Karyotype
(all autosomes telocentric
except those cited)
6 June
5 July
13 September
30 October
LI TM
0.41
0.33
0.35
0.45
LI TM 1 B
0.18
0.06
0.05
0.20
LI TM 2 B
0.00
0.06
0.00
0.00
LI MM
0.27
0.44
0.35
0.35
LI MM 1 B
0.05
0.00
0.10
0.00
LI MM M5 TM
0.05
0.06
0.00
0.00
Basic
0.05
0.06
0.15
0.00
Total specimens
22
18
20
20
TM = heterozygous bivalent.
MM = homozygous metacentric.
B = supernumerary chromosome.
Basic = no inversions present.
areas and were once considered by Hebard (1937) to form the Group Anconiae.
These three genera are now each considered to represent different subfamilies. T.
helferi has a metacentric X and inversion polymorphism (references in Weissman
and Rentz, 1980) placing it cytologically in Section B of T'rimerotropis. On the
other hand, M. occidentalis and M. nicola Rentz and Weissman have the usual
acridid karyotype of all telocentrics (Rentz and Weissman, 1981). So unusual
(White, 1973, p. 245; Hewitt, 1979, p. 12) are pericentric inversions in acridids
that their occurrence in a species would argue for phylogenetic relationship with
that genus that also possesses such inversions. I believe convergence in this
cytological character is far less likely than in the morphological characters cited
above.
T. munda (Scudder). — Seven males from 9.6 km S Ruth, Trinity Co., California,
were examined cytologically: 2N 6 = 23, Section A, 3 large, 6 medium, and 2
small autosomes. The X had occasional neometacentric activity; one male had a
supernumerary segment in the S10. Flights were low, usually 2 to 5 m, male
crepitation was a medium fast buzz similar to campestris Bruner in Otte (1970).
Some males and females were teneral on 3 August 1980. They were found in a
sandy, dry river bed bottom with fontana, 3 species of Dissosteira, and a Conozoa
species. The only munda female caught had small eggs (Stage I; see Weissman
and French, 1980). Strohecker et al. (1968) noted the similarity between munda
and pacifica Bruner. Except for the smaller size of the former, the taxa are indis¬
tinguishable in morphology, cytology, and crepitation.
T. occidentalis .—The inversion polymorphism (LI and M5) present in popu¬
lations from the San Francisco Bay Area was noted by Rentz and Weissman
(1981). Eighty individuals from one population had the karyotypes listed in Table
1. There is no apparent inversion or B chromosome frequency change through
the flight season. Specimens now have been examined cytologically (Table 2) from
Kern Co. in southern California to Klamath Co. in southern Oregon (these may
represent its south-north extremes). Populations in Oregon have an inversion in
a large autosome not found in California. When compared to the Woodside
VOLUME 60, NUMBER 4
273
Table 2. Karyotype frequency of T. occidentalis from California (first four columns) and Oregon
(last column). Abbreviations as in Table 1.
Karyotype
(all autosomes telocentric
except those cited)
Tehachapi,
Kem Co.
Zenia,
Trinity Co.
Bridgeville,
Humboldt Co.
Redding,
Shasta Co.
2 localities,
Klamath Co.
LI TM
0.33
0.20
0.58
0.40
0.38
LI MM
0.00
0.40
0.08
0.20
0.25
LI TM L2 TM
0.00
0.00
0.00
0.00
0.06
LI MM L2 TM
0.00
0.00
0.00
0.00
0.06
LI TM M5 TM
0.00
0.00
0.08
0.00
0.00
LI MM M5 TM
0.00
0.00
0.08
0.00
0.00
LI MM M5 TM 1 B
0.00
0.20
0.00
0.00
0.00
L2 TM
0.00
0.00
0.00
0.00
0.06
Basic
0.66
0.20
0.17
0.40
0.19
Total specimens
3
5
12
5
16
population (Table 1), these northern California and southern Oregon populations
are smaller in number of individuals and have less inversion heterozygosity,
consistent with the pattern found in pseudofasciata Scudder (Weissman, 1976;
and below) and helferi (Schroeter, 1968). The range of occidentalis probably ex¬
tends no farther east than western Nevada as morphologically similar individuals
from eastern Nevada identified in the ANSP and California Academy of Sciences
collections as occidentalis have 23 chromosomes and no inversion polymorphism
(Weissman, unpubl.) and probably represent fontana.
T. pacifica.— Ten males were examined cytologically from Bakersfield, Kern
Co., California: 2N S = 23, Section A, 3 large, 6 medium, and 2 small autosomes.
One male had a medium sized telocentric B chromosome. Males flew 3 to 10 m.
Crepitation was a medium fast buzz similar to campestris in Otte (1970) made
during the last % of flight. Only 1 of 5 females caught 6 August 1980, had developed
eggs. They were found on sandy substrate with short sparse grass associated with
p. pallidipennis (Burmeister).
T. p. pallidipennis. — Populations of this species from alkaline soil habitats (with
salt grass Distichlis) are morphologically very different from conspecifics on other
types of soil. The alkaline soil ecotypes are much lighter in body and tegminal
color, have reduced or no tegminal and wing bands, and a pale yellow or colorless
wing disk. These alkaline forms integrade into their darker conspecifics with
intermediates present over distances as short as 20 m (7.2 km N Beatty, Nevada)
to Vi km (Borax Lake, Alvord Hot Springs, Oregon (D. C. Lightfoot, pers. comm.)).
Crepitation and other flight characteristics were similar in both morphotypes. The
alkaline soil forms near Beatty may mature later. That these alkaline forms are
indeed distinctive in appearance is indicated by the population north of Beatty
being considered a distinct species by both White (1951a, p. 311; 1973, p. 331)
and Tinkham (pers. comm.). A similar population from Lake Abert, Oregon, was
even considered a second distinct species (Tinkham, pers. comm.). T. p. salina
McNeill may represent another ecotypic color phase on alkaline soils (see dis¬
cussion by Hebard, 1928, p. 254).
In North America hundreds of males of this widespread species have been
examined cytologically by White, Helwig, Coleman, and myself (see references in
Table 3. Supplement to cytological, morphological, and crepitational characteristics of Trimerotropis grasshoppers—see Table 1 of Weissman and Rentz (1980).
Only those species, and that specific category with additions herein, are listed below. Parentheses denote uncommon condition. Conditions without reference
numbers refer to museum specimens. Reference superscript numbers refer to citations at bottom of Table; reference superscript letters refer to specific localities
at bottom of Table. See Rentz and Weissman (1980) for species’ authors.
2N 3
Hind tibia color
Wing disk color
Crepitation
Comments
SECTION A
agrestis
crackle 3
a. barnumi
buzz 15
a. gracewileyae
crackle 10 ; buzz 15
a. hewitti
2313(C)
crackle 7
2 large, 7 medium, 2 small
autosomes 13(C)
albescens
supernumerary segments 13(B)
arenacea
(yellow)
(green)
crackle 7
B chromosomes, super-
numerary segments 13 ®
bilobata
B chromosomes 12
campestris
supernumerary segments 13 ® 1
cincta
(red)
buzz 5 ’ 8 20 ; crackle 13(E)
citrina
(yellow)
fontana
(blue tinge)
buzz to crackle 16
supernumerary segments 916
inconspicua
(blue)
yellow-green 1 ’ 7
buzz 7>13(A)
red tibia in Heifer 6 probably
another species
koebelei
(yellow)
latifasciata
crackle 13(H)
maritima
green-yellow,
(red) 2
munda
23 16
buzz 16
pacifica
buzz 16
B chromosomes 16
pistrinaria
(yellow)
pogonata
buzz 16
strenua
(yellow)
texana
(red)
crackle 13(D)
titusi
23 16
(red) 16
buzz 16
tolteca modesta
crackle 5 - 613 ®
female also crepitates 513 ®
274 PAN-PACIFIC ENTOMOLOGIST
Table 3. Continued.
2N <5
Hind tibia color
Wing disk color
Crepitation
Comments
SECTION B
cyaneipennis
(brown)
yellow, green-blue, blue
crackle only 5 - 619
fratercula
23 16
(green tinge)
crackle 20
gracilis sordida
crackle 311
helferi
clear to greenish yellow 16
pulse buzz 16
p. pallidipennis
colorless on alkaline soil;
B chromosomes 16
blue tinge at base
in typical 16
pseudofasciata
(yellow)
colorless, blue-green
buzz, pulse;
buzz, crackle 16(I)
saxatilis
green-yellow
sparsa
colorless, yellow, green,
buzz 5
?supemumerary segment 14
blue 417
suffusa
thalassica
yellow
yellow-green, blue-green
B chromosomes 16
Reference numbers for Table 3—complete citation in Literature Cited
1 Bamum, 1964
6 Heifer, 1953
11 Walker, 1902
16
Weissman, this report
2 Blatchley, 1920
7 Hewitt and Barr, 1967
12 Walters, 1968
17
Hagen, 1982
3 Brooks, 1958
8 Hubbell, 1922
13 Weissman, unpub.
18
Hebard, 1906
4 Hebard, 1928
9 Rentz and Weissman, 1981
14 White, 1951b
19
Rehn and Hebard, 1908
5 Hebard, 1935
10 Tinkham, 1960
15 Willey and Willey, 1971
20
Hebard, 1929
Reference letters for Table 3—locality of Cited Data
California
Mexico
Nevada
New Mexico
A Inyo Co.
B Baja California
C Humbolt Co.
D Dona Ana Cc
I Fresno Co.
Norte
E Sandoval Co.
Oregon
F Deschutes Co.
G Lake Co.
Arizona
H Coconino Co.
ho
C/1
VOLUME 60, NUMBER 4
276
PAN-PACIFIC ENTOMOLOGIST
Weissman and Rentz, 1980). The only reported populations with B chromosomes
were those from alkaline soils. Of 5 males I examined from near Beatty (also see
White, 1951a), one had a single B. One of 10 males from Summer Lake, Oregon,
had one B chromosome. Five normal colored males from nearby had no Bs.
Individuals of both morphs frequently had ditactic bivalents, a condition unusual
in the acrididae (Hewitt, 1979; Rentz and Weissman, 1981, p. 80 and Fig. 132).
T. pogonata Strohecker.—Individuals were locally very common at Los Osos
sand dunes, San Luis Obispo Co., California. Male flight was 1 to 2 m, crepitation
was a fast buzz similar to Encoptolophus costalis (Scudder) in Otte (1970), and
they were easy to catch. Females were more sluggish and had shorter flights. This
species was found microsympatric with Microtes occidentalis at Oso Flaco Lake,
San Luis Obispo Co., but the buzz crepitation in the latter species is slower.
Schroeter (1968, pers. comm.) found the karyotypes of 40 males to be uniform.
T. pseudofasciata. — Weissman (1976) noted that California Channel Island males
had a fixed L2 inversion homozygote, and had 5 autosomes polymorphic. Main¬
land populations from northern Baja California to northern Oregon and east to
Nevada had 4 autosomes fixed and 4 polymorphic. However, 3 males from Van-
denberg Air Force Base, coastal Santa Barbara Co., California, had only the L2
inversion fixed and only the M5 polymorphic. This mainland habitat resembled
that of San Nicolas Island, Ventura Co., as did the karyotype of the males ex¬
amined. Despite intensive searching at the former locality, only 5 (3 <3, 2 2)
specimens were collected. This pattern is consistent with that noted by Weissman
(1976) where ecologically marginal populations with low densities have little
inversion polymorphism.
California Central Valley populations have a reproductive dormancy (Weiss¬
man and French, 1980) and do not crepitate until late summer-early fall. Males
from Jacalitos Canyon, Fresno Co., in the Central Valley, collected on 19 No¬
vember 1982, represent the first Trimerotropis population to contain individuals
that buzz, pulse buzz, and crackle. Many males would buzz in the beginning of
flight and pulse buzz at the end.
T. strenua. — See under californica for discussion as to possible synonymy. Rare¬
ly individuals of the former have yellow hind tibia.
T. thalassica.— Under this species I here place santamonica Rentz and Weiss¬
man in synonymy. In our 1981 description of the latter, we noted its similarity
to thalassica but cited differences in wing disk color, tegminal bands, antennal
annulation, and ground color. Continued collecting at the type locality of san¬
tamonica south into Baja California demonstrated that we were dealing with a
cline for these characters. Baja California specimens were almost uniform for
blue-green wings, distinct tegminal bands, and definite antennal annulation.
Seven additional males from the Santa Monica Mountains, Los Angeles Co.,
have been examined cytologically: they agree in inversion diversity with those in
Tables 6 and 7 in Rentz and Weissman (1981). Specimens examined from near
Laguna Hanson and near Sierra San Pedro Martir National Park, both northern
Baja California, had only four autosomes heterozygous and 12% (2 of 17) had B
chromosomes.
This species normally occurs at low elevations in chaparral habitats. Flights
there are usually short and crepitation is a fast buzz heard only late in the flight
season (see Rentz and Weissman, 1981). These patterns are maintained in north-
VOLUME 60, NUMBER 4
277
em Baja California except for the populations bordering the Vallecitos meadow
area at 2350 m in the San Pedro Martir Park area. There, individuals morpho¬
logically and cytologically indistinguishable from those living in typical habitat
at lower elevations, inhabit open areas under and around pines. These populations
are dense, individuals fly long distances, and crepitation, which is made during
the entire flight season, is a pulse buzz similar to pallidipennis in Otte (1970).
Whether these differences reflect edaphic features or represent a cryptic species
need further investigation.
T. titusi Caudell.—Eight males were examined cytologically from Pinnacles
National Monument, San Benito Co., California: 2N 6 = 23, Section A, 2 large,
7 medium, and 2 small autosomes. Males flew 2 to 5 m, crepitation was a medium
fast buzz similar to campestris in Otte (1970) usually made in last x k of flight but
occasionally in last 3 4. Several females contained nearly mature eggs on 1 August,
1980. One female was teneral. Specimens were variable for tegminal band inten¬
sity; in one female the band was nearly absent. Four of 32 specimens had red
hind tibia rather than the usual yellow. At Pinnacles it was sympatric with oc-
cidentalis, p. pallidipennis, and Cibolacris parviceps (F. Walker) in a dry wash
with gravelly substrate and sparse grass cover.
Acknowledgments
The following contributed specimens, useful discussion, and unpublished in¬
formation: R. Bohart, T. J. Cohn, T. H. Hubbell, D. C. Lightfoot, A. S. Menke,
D. Otte, D. C. F. Rentz, G. L. Schroeter, E. R. Tinkham, and M. J. D. White.
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Blatchley, W. L. 1920. The Orthoptera of Northeastern America, with especial reference to the faunas
of Indiana and Florida. Nature Publishing Company, Indianapolis.
Brooks, A. R. 1958. Acridoidea of southern Alberta, Saskatchewan, and Manitoba (Orthoptera).
Can. Ent. Suppl., 9:1-92.
Caudell, A. N. 1908. Notes on some western Orthoptera: with the descriptions of one new species.
Proc. U.S. Nat. Mus., 34:71-81.
Fulton, B. B. 1930. Notes on Oregon Orthoptera with descriptions of new species and races. Ann.
Ent. Soc. Amer., 23:611-641.
Hagen, A. F. 1982. Variations in the wing band of Trimerotropis sparsa (Thomas) (Orthoptera:
Acrididae: Oedipodinae). J. Kansas Ent. Soc., 55:477-480.
Hebard, M. 1906. A contribution to the knowledge of the Orthoptera of Montana, Yellowstone Park,
Utah and Colorado. Proc. Acad. Nat. Sci., Philad., 58:358-418.
-. 1928. The Orthoptera of Montana. Proc. Acad. Nat. Sci., Philad., 80:211-306.
-. 1929. The Orthoptera of Colorado. Proc. Acad. Nat. Sci., Philad., 81:303-425.
-. 1935. Orthoptera of the upper Rio Grande Valley and the adjacent mountains in northern
New Mexico. Proc. Acad. Nat. Sci., Philad., 87:45-82.
-. 1937. Studies in Orthoptera which occur in North America north of the Mexican boundary.
IX. On Arphia conspersa, notes and a new race of Spharagemon, a new genus and its races of
the group Heliasti and a new generic name for the group Anconiae (Oedipodinae, Acrididae).
Trans. Amer. Ent. Soc., 63:361-379.
Heifer, J. R. 1953. How to know the grasshoppers, cockroaches and their allies. Wm. C. Brown
Company, Iowa.
Helwig, E. R. 1955. Spermatogenesis in hybrids between Circotettix verruculatus and Trimerotropis
sujfusa (Orthoptera: Oedipodinae). Univ. Colorado Stud. Ser. Biol., 3:47-64.
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PAN-PACIFIC ENTOMOLOGIST
Hewitt, G. B., and W. F. Barr. 1967. The banded-wing grasshoppers of Idaho. Univ. Idaho Agric.
Exp. Stat. Res. Bull., 72:1-64.
Hewitt, G. M. 1979. Animal cytogenetics. V. 3: Insecta I: Orthoptera. Bomtraeger, Berlin.
Hubbell, T. H. 1922. Notes on the Orthoptera of North Dakota. Occ. Papers Univ. Mich. Mus.
Zool., 113:1-56.
John, B., D. C. Lightfoot, and D. B. Weissman. 1983. The meiotic behaviour of natural F t hybrids
between the grasshoppers Trimerotropis suffusa Scudder and T. cyaneipennis Bruner (Orthop¬
tera: Oedipodinae). Canad. J. Gen. Cytol., 25:467-477.
-, and D. B. Weissman. 1977. Cytogenetic components of reproductive isolation in Trimerotro¬
pis thalassica and T. occidentalis. Chromosoma (Berl.), 60:187-203.
Murphy, D. D., and P. R. Ehrlich. 1980. Two California Checkerspot butterfly subspecies: one new,
one on the verge of extinction. J. Lep. Soc., 34:316-320.
Otte, D. 1970. A comparative study of communicative behavior in grasshoppers. Misc. Pubs. Mus.
Zool. Univ. Mich., 141:1-168.
Rehn, J. A. G., and M. Hebard. 1908. An orthopterological reconnoissance of the southwestern
United States. Part I: Arizona. Proc. Acad. Nat. Sci., Philad., 60:365-402.
Rentz, D. C. F., and D. B. Weissman. 1980. An annotated checklist of the grasshopper species of
Aerochoreutes, Circotettix, and Trimerotropis (Orthoptera: Acrididae: Oedipodinae). Trans.
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California Channel Islands. Univ. Calif. Pubs. Ent., 94:1-240.
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Schroeter, G. L. 1968. Pericentric inversion polymorphism in Trimerotropis helferi (Orthoptera:
Acrididae) and its effect on chiasma frequency. Ph.D. thesis, University of California, Davis,
University Microfilms, Ann Arbor, Michigan.
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Calif. Insect Surv., 10:1-177.
Tinkham, E. R. 1948. Faunistic and ecological studies on the Orthoptera of the Big Bend Region of
Trans-Pecos Texas, with especial reference to the orthopteran zones and faunae of midwestem
North America. Amer. Mid. Nat., 40:521-663.
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the genus Trimerotropis from the Utah dunes. Gr. Basin Nat., 20:49-58.
Walker, E. M. 1902. The Canadian species of Trimerotropis. Can. Ent., 34:1-11.
Walters, J. L. 1968. Chromosomal polymorphism in the grasshopper Trimerotropis bilobata, Rehn
and Hebard. Genetics, 60:234-235.
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Trimerotropispseudofasciata. Chromosoma (Berl.), 55:325-347.
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grasshopper with a reproductive dormancy. Acrida, 9:145-157.
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the trimerotropine ( Aerochoreutes, Circotettix, and Trimerotropis) grasshoppers (Orthoptera:
Oedipodinae). Trans. Amer. Ent. Soc., 106:253-272.
White, M. J. D. 1951a. Cytogenetics of orthopteroid insects. Adv. Genet., 4:267-328.
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PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 279-288
New Genera and Species of Neotropical
Cerambycidae (Coleoptera)
John A. Chemsak and E. G. Linsley
University of California, Berkeley, California 94720
Abstract. —Four new genera and five new species of Cerambycidae are described
from Mexico and Central America: Lycoplasma formosa from Panama (Hemi-
lophini); Noctileptura squamosa from Mexico and N. seriata from Guatemala
(Lepturinae); Oraphanes binotatum from Mexico (Hesperophanini); and Gies-
bertia rugosa from Mexico (Purpuricenini).
While identifying cerambycids from the faunas of Mexico and Central America,
a number of interesting new taxa were discovered. Because of their unique nature,
we are describing some of these here.
Lycoplasma, New Genus
Form moderate-sized, elytra very broadly explanate from behind humeri. Head
with front transverse, slightly convex, median line extending onto neck; mandibles
stout, gradually arcuate to apices; genae subparallel, as long as lower eye lobes;
eyes finely faceted, deeply emarginate, upper lobes small, widely separated; an¬
tennal tubercles flattened, widely divergent; antennae rather stout, shorter than
body, scape subequal to third segment, fourth slightly shorter than third, remaining
segments gradually decreasing in length, eleventh short, tapering, segments lightly
ciliate beneath, ciliae becoming less numerous toward apex. Pronotum broader
than long, sides obtusely tuberculate behind middle; apex and base broadly, shal¬
lowly impressed transversely; disk convex; prostemum narrow, intercoxal process
narrow, broady expanded at apex, coxal cavities closed behind; procoxae exserted,
cavities strongly angulate externally; mesostemum with intercoxal process rather
narrow, arcuate, lying well below tops of coxae; metastemum with a strongly
impressed median line, epistemum broad, gradually narrowing posteriorly. Elytra
very broadly expanding from behind numeri; sides briefly vertical behind humeri
above bases of lateral margins; disk rather strongly costate, costae uniting near
apex to form a somewhat reticulate pattern; lateral margins lightly fringed with
short hairs; apices rounded. Legs short, femora moderately broad, sublinear; tibiae
short, sulcations vague; tarsi broad, third segment cleft to base. Abdomen normally
segmented.
Type species. —Lycoplasma formosa, n. sp.
This genus is apparently very similar in appearance to Ites Waterhouse (1880).
Lane (1954) presented a lengthy discussion on the similarity of Ites with Lyco-
desmus Melzer (1927) and concluded that the two were identical. Since the second
and third antennal segments are of the same length, Melzer speculated that the
true second segment was hidden and listed Lycodesmus as having 12-segmented
280
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Lycoplasma formosa Chemsak and Linsley, 9.
antennae. Apparently the first two segments are equal in length and the antennae
contain only 11 segments. This unusual situation readily separates Lycoplasma
from Ites. In addition to the differences in the proportions of the antennal seg¬
ments, Lycoplasma has rather strong costae on the elytra.
The tribe Hemilophini contains a number of lycid-like genera in the Neotropical
region. These differ primarily in characters of the antennae and shape of the elytra.
Most can be separated from Lycoplasma on these bases. Assuming that Lane’s
characterization of Ites and Lycodesmus is accurate, Lycoplasma represents a
distinct genus.
Lycoplasma formosa, New Species
(Fig. 1)
Female.— Form moderate sized, elytra very broadly explanate behind humeri;
integument brownish, head yellowish except for broad dark bands behind eyes
and a median triangular dark band on vertex which narrows between eyes, prono-
tum with a broad, dark median, longitudinal band and a band on each side below
lateral tubercles, elytra with a broad post basal band which narrows toward lateral
margins, apical band broad, covering about apical x h, apical margin of apical band
notched medially, elytral bands with a strong violaceous caste, antennae dark,
legs dark except basal halves of femora, underside except prostemum dark. Head
with front moderately coarsely, rather densely punctate at middle, vertex more
VOLUME 60, NUMBER 4
281
coarsely punctate on subglabrous dark spot; pubescence dense, appressed, erect
ciliae sparse, more numerous on antennal tubercles; antennae extending to about
apical Vi of elytra, scape finely punctate, moderately densely clothed with short,
dark, appressed pubescence, outer segments densely clothed with very short, dark
pubescence, basal segments with a few erect setae beneath, these diminishing in
number toward apex. Pronotum broader than long, sides obtusely tuberculate
behind middle; median dark vitta subglabrous, sparsely, coarsely punctate; pu¬
bescence dense, appressed, pale yellowish, dark on dark vittae, long erect hairs
sparse at sides; prostemum densely pubescent; meso- and metastemum moder¬
ately densely clothed with appressed pale and dark pubescence, pubescence not
obscuring surface, punctation minute. Elytra about as long as greatest \ydth, base
slightly broader than pronotum; punctures on basal x h moderately coarse, contig¬
uous, becoming finer on middle l h and dense and confluent on apical Vi; each
elytron with 4 costae, two outside and two inside pairs joining near apex; pubes¬
cence dense, depressed, colored as background. Legs short, femora bicolored; front
tibiae arcuate. Abdomen minutely, densely punctate, densely clothed with fine,
appressed, pale and dark pubescence; last stemite longer than fourth, apex round¬
ed, lightly notched at middle. Length, 16 mm.
Holotype female (University of California, Davis) from Barro Colorado Island,
Panama, 14 May 1980 (H. Wolda). One female paratype with same data, 30 April
1980.
The only variation evident in the two available specimens is some reduction
of the pronotal dark vitta in the paratype. Although this species greatly resembles
some Mexican and Central species of Lycus, its model is probably a species of
Calopteron as evidenced by the violaceous caste and reticulated appearance of
the dark bands of the elytra.
Noctileptura , New Genus
Form small to moderate sized, subparallel; pubescence of body modified into
squamae. Head with front short, transversely impressed medially; tempora mod¬
erate, slightly convergent, abruptly constricted behind; palpi rather slender, un¬
equal; eyes rather large, coarsely faceted, deeply emarginate, upper lobes small,
widely separated; antennal tubercles prominent, widely divergent; antennae slen¬
der, 11 -segmented, longer than body, insertions almost in eye emarginations, scape
cylindrical, shorter than third segment, fourth shorter than third, fifth longer than
fourth, segments from third slightly expanded at apices, fifth more so, segments
from sixth carinate beneath, with sensory areas extending length of segments,
expansion of fifth segment containing sensory areas. Pronotum longer than broad,
sides angulate; apex much narrower than base, broadly impressed transversely;
base shallowly impressed; disk tumid behind apical impression; prostemum broadly
impressed, intercoxal process slender, arcuate, expanding at apex, coxal cavities
closed behind; mesostemal process almost plane, about even with tops of coxae;
metastemum with epistemum slender, gradually narrowing posteriorly. Elytra
subparallel, narrowing toward apex; apices subtruncate. Scutellum small. Legs
slender, femora linear, posterior tarsi short, first segment shorter than following
two together, third segment cleft to base. Abdomen normally segmented.
Type species.—Noctileptura squamosa, n. sp.
This genus is unlike any of the other New World Lepturinae. The shape of the
282
PAN-PACIFIC ENTOMOLOGIST
Figure 2. Noctileptura squamosa Chemsak and Linsley, 9.
head and pronotum, coarsely faceted eyes, and structure of the antennae make it
distinctive. The most unique characteristic is the presence of recurved, spatulate
scales on the body. These appear to be modified hairs and are not known to occur
on any other Western Hemisphere Cerambycidae.
Two undescribed species are presently known.
Noctileptura squamosa, New Species
(Fig. 2)
Female .—Form moderate sized, subparallel; integument reddish brown, opaque;
pubescence of body modified into whitish, flattened squamae, pubescence of ap¬
pendages normal. Head densely, confluently punctate, finely scabrous; surface
densely clothed with small, pale scales; antennae slender, slightly longer than
body, scape finely, densely punctate, opaque, outer segments slightly shining,
segments clothed with very fine, short, pale, appressed pubescence. Pronotum as
broad as long, sides angulate; disk tumid behind apical impression, tumosity
vaguely impressed at middle; punctures moderately coarse, very dense; surface
densely clothed with pale scales; prostemum densely punctate except on anterior
Vi, moderately densely clothed with scales; meso- and metastemum densely, shal¬
lowly punctate, sides densely clothed with scales. Elytra more than twice as long
VOLUME 60, NUMBER 4
283
as broad, subparallel to apical x h, then tapering; punctures coarse, contiguous;
scales dense, larger than those on pronotum; sides narrowly margined behind
middle; apices truncate. Legs finely, densely punctate, densely clothed with fine,
pale, appressed pubescence. Abdomen finely, shallowly punctate, finely scaled;
last stemite broadly rounded at apex. Length, 12 mm.
Holotype female (Canadian National Collection) from 12 mi N Tuxtla Gu¬
tierrez, Chiapas, Mexico, 7 June 1969 (J. M. Campbell).
This is one of the most distinctive and unique species of Lepturinae in this
hemisphere. The most unusual covering of scale-like hairs will readily identify it.
Noctileptura seriata, New Species
Female .—Form small; integument subopaque, reddish brown, head and prono¬
tum darker; pubescence rather sparse, finely scale-like. Head densely micro-punc¬
tate, densely clothed with appressed, narrow scales; antennae about as long as
body, segments densely clothed with fine, scale-like pubescence. Pronotum longer
than broad, sides vaguely angulate; apex narrower than base; base shallowly im¬
pressed transversely, apex broadly impressed; disk uneven, vaguely tumid behind
apical impression; punctures coarse, dense, subconfluent; scales rather sparse;
prostemum coarsely, shallowly punctate, scales sparse; meso- and metastemum
coarsely, shallowly punctate, scales sparse. Elytra slightly more than twice as long
as broad, sides subparallel, tapering near apex; punctures very coarse, deep, con¬
tiguous, arranged in longitudinal rows; scales narrow, recurved, serially arranged;
apices truncate. Legs moderately densely clothed with thin scales. Abdomen shal¬
lowly punctate, scales sparse; last stemite subtruncate at apex. Length, 9 mm.
Holotype female (National Museum of Natural History) from Livingston, Gua¬
temala, 6 May (Barber & Schwarz). One female paratype with same data, 8 May.
This species is readily separated from N. squamosa by the smaller size, much
sparser and serially arranged scales of the elytra, and by the more elongate, less
dorsally tumid pronotum.
Oraphanes, New Genus
Form moderately elongate, somewhat depressed, subparallel. Head with front
short, oblique; palpi unequal, apical segments rather small, triangular; mandibles
short, strongly arcuate at apical V 2 ; eyes large, moderately coarsely faceted, deeply
emarginate, upper lobes narrow, widely separated; genae very short; antennae
longer than body in male, eleven segmented, segments from third slightly produced
apically, obtusely carinate dorsally, scape conical, short, third segment about twice
as long as first, fourth shorter than third, fifth longer than fourth, eleventh longer
than tenth, basal segments with a few suberect setae beneath. Pronotum about as
long as broad, sides with small rounded tubercles slightly behind middle; disk
somewhat flattened, middle with a low glabrous callus; prostemum transversely
impressed, intercoxal process slender, not extending beyond coxae, coxal cavities
wide open behind, angulate externally; mesostemal process more than twice as
wide as prostemal, rather abruptly declivous anteriorly, coxal cavities open to
epimeron; metastemum with epistemum narrow, tapering posteriorly. Elytra about
three times as long as broad, sides subparallel; each elytron with two, fine, median
costae; apices narrowly rounded. Legs elongate; femora gradually enlarging toward
apices; tibiae slender, not carinate; posterior tarsi slender, first segment as long
284
PAN-PACIFIC ENTOMOLOGIST
as two following together, third segment cleft almost to base. Abdomen normally
segmented.
Type species. —Oraphanes binotatum, n. sp.
In the key to the genera of Mexican Hesperophanini (Chemsak and Linsley,
1963) this genus comes out to couplet 6 with Eucrossus. The lack of antennal and
elytral spines on Oraphanes will immediately separate the two. Oraphanes appears
unrelated to the other hesperophanine genera of Mexico.
Oraphanes binotatum, New Species
(Fig. 3)
Male .—Form moderately large, subparallel; integument reddish brown, elytra
pale brownish, with two, median, elongate, pale spots with small dark spots
anteriorly and larger, irregular dark spots posteriorly. Head finely, densely punc¬
tate, sparsely clothed with short appressed pubescence; antennae longer than body,
scape finely, densely punctate, remaining segments minutely, densely punctate,
densely clothed with short pubescence. Pronotum with disk very finely, densely
punctate, median callus and lateral tubercles glabrous; pubescence very fine, short,
pale, denser appearing at sides, sides with a few long erect hairs; prostemum
anteriorly plicate, glabrous, pubescence fine; meso- and metastemum minutely,
densely punctate, densely clothed with pale, subdepressed pubescence. Elytra
finely, densely punctate, punctures becoming finer and sparser toward apex; pu¬
bescence very short, recurved, sparse; pale median spots not ebumeous; costae
uniting a little before apex; apices narrowly dehiscent, narrowly rounded. Legs
minutely, densely punctate, femora finely pale pubescent, tibiae with pubescence
appearing golden in oblique light. Abdomen minutely, densely punctate, mod¬
erately densely pale pubescent; last stemite broady rounded at apex. Length, 24
mm.
Holotype male (California Academy of Sciences) from 20 mi SW Colima, Co¬
lima, Mexico, 24 June 1967 (A. Hardy).
The reddish integument and dark testaceous elytra with two pale spots bordered
anteriorly and posteriorly by dark patches make this species quite distinctive.
Giesbertia, New Genus
Form stout, subdepressed. Head with front subvertical, short, narrowly im¬
pressed transversely, with a deep pit below each antennal tubercle; palpi slightly
unequal, short, apical segments cylindrical; mandibles stout, broad, emarginate-
truncate at apices; genae narrow, subacute; eyes finely faceted, deeply emarginate,
upper lobes small, broadly separated; antennal tubercles slightly elevated, diver¬
gent; antennae slender, longer than body in males, shorter than body in females,
segments from third slightly expanded at apices, scape conical, shorter than third
segment in males, subequal in females, fourth subequal to third in males, shorter
in females, fifth longer than fourth. Pronotum broader than long, sides with very
large, blunt tubercles at middle; disk in males with a broad, distinctly delimited,
amphora-like median area, females with five, coarsely punctate calluses on disk;
prostemum narrow, transversely impressed, intercoxal process narrow, abmptly
declivous, lightly tuberculate, coxal cavities wide open behind, rounded externally;
mesostemum with intercoxal process gradually arcuate, lying below tops of coxae;
metastemum with epistemum rather narrow, subparallel, scent glands distinct.
VOLUME 60, NUMBER 4
285
Figure 3. Oraphanes binotatum Chemsak and Linsley, <?.
Elytra broad, slightly tapering, surface submetallic; apices slightly sinuately round¬
ed. Scutellum longer than broad, apex acute. Legs stout, femora gradually enlarging
toward apices, front pair subparallel; tibiae slender, not elongate; posterior tarsi
slender, first segment shorter than following two together, third segment cleft
almost to base. Abdomen normally segmented.
Type species. — Giesbertia rugosa, n. sp.
This genus appears to be related to the Crioprosopus and Deltaspis (sensu lato)
286
PAN-PACIFIC ENTOMOLOGIST
group of Purpuricenini. Giesbertia differs from Crioprosopus by the shorter genae,
more robust mandibles, sexual differences of the pronotum, much narrower pro-
sternal process, and less prominent mesostemal process. Additionally, the met-
epistemum is narrower and subparallel. Since the genus Deltaspis, as presently
constituted, contains a variety of forms, we can only compare Giesbertia with the
type species of Deltaspis, auromarginata Serville. That species has finer, apically
acute mandibles, smaller eyes, less prominently tuberculate sides of the pronotum,
thicker antennae (at least in females) and strongly metallic integument.
We cannot ascertain the exact nature of the male antennae of Giesbertia since
the only available specimen possesses only eight segments of one antenna.
It is a pleasure to dedicate this genus to E. Giesbert for his contributions to the
knowledge of Cerambycidae.
Giesbertia rugosa, New Species
(Fig. 4)
Female.— Form moderately large, subdepressed; integument black, head par¬
tially reddish, pronotum reddish along apical margin and on lateral tubercles,
elytra vaguely metallic, blueish, narrowly reddish basally and laterally, underside
medially reddish, femora reddish except apically; pubescence sparse dorsally, pale.
Head finely, rugosely punctate; pubescence fine, pale, suberect, dense on basal V 2
of mandibles; antennae extending to apical l h of elytra, scape densely punctate,
obtusely carinate, basal segments shining, outer segments opaque, segments three
and four with longer, subdepressed pubescence, outer segments densely clothed
with minute appressed pubescence, third segment subequal to scape, fourth shorter
than third, fifth subequal to fourth, eleventh short, slender. Pronotum much
broader than long, lateral tubercles prominent; disk coarsely, confluently punctate,
with five calluses, one on each side before middle, one median behind middle
and one on each side near base; pubescence long, pale, suberect; prostemum
deeply, confluently punctate, finely, densely pubescent; meso- and metastemum
densely, minutely punctate, densely clothed with pale, subdepressed pubescence.
Elytra about twice as long as broad; basal V 2 densely, coarsely, rugosely punctate,
punctures becoming finer toward apex; pubescence on basal ¥2 pale, long, suberect,
becoming short and appressed toward apex; apices slightly sinuately rounded.
Legs densely punctate, moderately densely pubescent. Abdomen minutely, densely
punctate, first four segments transversely glabrous at apices; pubescence fine, pale,
suberect and appressed; last stemite broadly subtruncate at apex. Length, 28 mm.
Male.— Form similar, sides tapering slightly. Antennae longer than body, third
segment longer than scape, fourth subequal to third, fifth longer than fourth.
Pronotum with lateral tubercles less robust; disk vaguely callused, middle with a
delimited amphora-like area enclosing dense, coarse, irregular punctures, outside
areas more deeply, closely punctate; prostemum with a delimited, deeply punctate
area. Abdomen with last stemite rather narrowly tmncate. Length, 26 mm.
Holotype female (California Academy of Sciences) from 5 mi N Huetamo,
Michoacan, Mexico, July 1976 (A. Lau). Paratypes as follows: 1 female, Canon de
Zopilote, 24 mi N Chilpancingo, Guerrero, Mexico, 11 July 1970 (E. Fisher, P.
Sullivan); 1 male, Cuernavaca, Morelos, Mexico, 7000 feet, 29 July 1961 (R. and
K. Dreisbach).
VOLUME 60, NUMBER 4
287
Figure 4. Giesbertia rugosa Chemsak and Linsley, 2.
Some color variation is expressed in the amount of red on the head, pronotum,
elytra, and underside.
Acknowledgements
Material was made available by the Canadian National Collection, Ottawa;
California Department of Food and Agriculture, Sacramento; University of Cal¬
ifornia, Davis; and E. Giesbert. Carolyn Tibbetts prepared the illustrations.
288
PAN-PACIFIC ENTOMOLOGIST
Literature Cited
Chemsak, John A., and E. G. Linsley. 1963. The genera of hesperophanine Cerambycidae presently
known from Mexico, with descriptions of several new species (Coleoptera). J. Kansas Entomol.
Soc., 36:207-230, 2 figs.
Lane, F. 1954. Notas sinonimicas 1. Lycodesmus Melzer, 1927 —Ites Waterhouse, 1880 (Coleoptera,
Lamiidae). Rev. Brasil. Entomol., 1:195-201, 2 figs.
Melzer, J. 1927. Longicomeos de Brasil, novos ou pouco conhecidos. Rev. Mus. Paul., 15:557-582.
Waterhouse, C. O. 1880. New South American Coleoptera, chiefly from Ecuador. Ann. Mag. Nat.
Hist., (5)5:297-298.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 289-296
New Cynipid Wasps from California
(Hymenoptera: Cynipidae)
Robert J. Lyon
Life Science Dept., Los Angeles City College, Los Angeles, California 90029.
Five new cynipid species from Southern California are described in this paper
plus a discussion of the biology of Andricus reticulatus Bassett. Andricus reticulatus
Bassett was previously thought to occur on species of white oaks ( Lepidobalanus )
in the Southwest United States only but is now known to occur on several white
oaks in California.
Gall-forming wasps are unique in the insect world in that the formations they
produce are usually distinctive for each species; thus, galls collected long after the
exit of the original makers give evidence of both described and undescribed
species. The galls produced by the cynipid wasps described in this paper, have
been known to other investigators; however, because of the difficulties involved
in rearing the insects, sufficient numbers of the adults have never been obtained.
In “Cynipid Galls of the Pacific Slope” (1957) Lewis Weld estimates that ap¬
proximately one-third of the oak, gall-forming wasps in this region are still un¬
described. A few new species have been described since this time (Dailey, 1973,
1977; Burnett, 1974) but there are many that are known from their galls only.
There is still great need for ecological studies in this group since many known
species represent the center of complex biological communities involving alter¬
nating generations and a host of parasites, hyperparasites and inquilines or guest
wasps. Eighteen of these life cycles have been demonstrated (Lyon et al.) and
circumstantial evidence indicates the existence of others. There are probably
alternating generations in some of the following new species but experimental
work will be necessary to demonstrate this. The terminology reviewed by Tulloch
(1929), Eady and Quinlan (1963), and Dailey (1973) is used in describing cynipid
thoracic and wing morphology in this paper.
Neuroterus c/trysolepis, New Species
Unisexual female.— Black except legs which are smoky yellow, proximal por¬
tions of coxae fuscous as are the femora and terminal tarsal segments. Front,
clypeus, mandibles and lower cheeks piceous. First three segments of the antennae
yellow, remaining segments fuscous. Ocelli brown. Head from above, transverse,
broader than the thorax, granulo-coriaceous; from in front, broader than high,
cheeks broadened behind the eyes; interocular area twice as broad as high, coarsely
granular; malar space one-fifth the length of the eye with striations; face and
clypeus beset with prominent, long, whitish bristles; antennae 14-segmented with
terminal segment sometimes indistinctly divided in smaller specimens giving the
appearance of 13 segments. Mesoscutum finely coriaceous with a few scattered
white hairs; faint traces of notaulices visible, also traces of anterior lines. Scutellum
joined to mesoscutum without a suture. Scutellum longer than broad with cori-
290
PAN-PACIFIC ENTOMOLOGIST
aceous disk, almost rugose in some lights. Disk margined on the sides with a
broad, deep, shining groove at the base. Mesopleuron granulorugose beneath
tegulae like the sides of the pronotum, rugose in lower portions. Mesostemum
prominently bulging below. Propodeum with several longitudinal wrinkles. Wing,
short, ciliate on front margin, longer cilia on posterior margin, pubescent, veins
heavy and smoky brown. First abscissa of radius arcuate, at right angles to the
second which is heavy and curves slightly upward; areolet well formed, reaching
one-fourth the distance to the basal (Rs+M). Mj reaching basal; distinct cloudy
areas around basal cross vein and also in the anal area; radial cell elongated, four
times as long as broad. Abdomen nearly as long as head and thorax combined,
longer than high, compressed laterally, collapsed in pinned specimens; all tergites
visible along the dorsal curvature. Ventral spine short, twice as long as broad,
ovipositor projecting straight out from behind. Tarsal claws simple. Range in
length of 59 specimens 1.7 mm. Average length 1.9 mm.
Types.— The holotype female and eight paratypes are in the collection of the
U.S. National Museum. Other paratypes are in the collections of the California
Academy of Sciences, Los Angeles County Museum, Weld Collection at Los
Angeles City College and the collection of Charles Dailey.
Host. — Quercus chrysolepis Liebm.
Gall (Fig. 1A).—An abrupt, blister-like, monothalamous swelling that arises on
the young stems of Quercus chrysolepis. The galls mature slowly and require at
least nine months to complete their development. Developing galls contained
large larvae in March, pupae in April and yielded adults in late April and early
May. At this time, adult females could be taken as they oviposited into the new
leaf buds and stem growth. Infestation was quite heavy on some trees and in
many instances, the new growth was severely damaged or completely killed. This
is the first Neuroterus to be described from Quercus chrysolepis or from related
oaks in this group. Insects of this genus had previously been known from galls
on white oaks only. This species shows distinct differences in sculpture from other
Pacific Slope members of the genus.
Locality. — Specimens in the type series were collected at Pasadena, California.
Andricus reticulatus Bassett
This species, originally described by Bassett, 1890, has been recorded from a
number of white oaks in the Southwest (Arizona, New Mexico and West Texas).
The cynipids of the Southwest have been considered to be sharply separated from
those of the Pacific Slope; therefore it is surprising to find this species producing
galls on Quercus engelmanni Greene in Southern California (Fig. IB). Specimens
were collected on a number of trees near Los Angeles and Pasadena with the
insects emerging in late February. Infestation was particularly heavy on some
trees, usually younger ones, and in several instances, the trees were nearly defo¬
liated. Female wasps were ovipositing in the unopened buds on 23 February in
La Canada, California. These sites were marked with colored thread and with the
opening of the new leaves, 23 March the young galls were immediately visible.
The young galls were covered with pinkish, white hairs and growth was rapid so
that galls were full sized by 23 April although the developing larvae were still
small. The appearance of the new galls only one month after emergence of the
adults indicates that an alternate bisexual generation does not exist in this species,
VOLUME 60, NUMBER 4
291
Figure 1. A. Blister galls of Neuroterus chrysolepis Lyon showing the typical arrangement of the
galls on a young twig from Quercus chrysolepis Liebm. B. Petiole gall of Andricus reticulatus Bassett
on Quercus engelmanni Greene with sagittal section showing arrangement of the larval cells. C.
Characteristic leaf cup spangles produced by Andricus bakkeri Lyon on Quercus dumosa Nutt. D.
Double chambered leaf gall of Heteroecus crescentus Lyon on Quercus chrysolepis Liebm. Larval
chamber is shown on left side. E. Leaf blister gall produced by Heteroecus devorus Lyon on Quercus
chrysolepis Liebm. F. Typical biconvex, lens-shaped galls produced by Callirhytis lentiformis Lyon.
Galls are spherical when fresh.
292
PAN-PACIFIC ENTOMOLOGIST
at least in this area. The adult females were compared with paratypes of Andricus
reticulatus Bass, from New Mexico and they appear to be the same, although the
notaulices are incomplete in some specimens from Quercus engelmanni. Lewis
Weld collected this California Andricus reticulatus many years ago and his notes
indicate that he could not separate it from Arizona specimens. Figure 172 in
“Cynipid Galls of the Pacific Slope,” Weld, shows the gall as an undescribed
species but notes similarity to Andricus reticulatus Bassett and an Eastern species,
Andricus petiolicola (O.S.). There are twelve specimens of the California reticulatus
in the Weld collection. Charles Dailey has sent in females of this species from
galls collected on Quercus turbinella Greene, ssp. turbinella Tucker growing at
El Condor, Baja California, Mexico and also from a hybrid oak, Quercus dumo-
sa x Quercus turbinella Greene, ssp. California Tucker growing at Gorman, Cal¬
ifornia. They are typical Andricus reticulatus Bass.
Andricus bakkeri, New Species
Unisexual female.— Uniform reddish brown except the posterior tergites and
ovipositor sheaths which are dark brown; eye, ocelli, tips of the antennae, sides
of the propodeum, metapleura and bases of the coxae, dark brown. Head trans¬
verse, coriaceous, as broad as the thorax; when viewed from the front, shows a
distinct flattened hump in the ocellar area; cheeks not broadened behind the eyes;
face beset with long, whitish hairs; interocular area more than twice as wide as
high; malar space striated, one-third the length of the eye. Antennae 14-segmented,
the third segment longer than the fourth and the terminal segment longer than
the preceding one. Mesoscutum as broad as long, microscopically coriaceous,
sparsely pubescent with scattered punctures. Notaulices percurrent and strongly
curved. A short median groove can be seen in some specimens. Lateral lines
smooth, slightly depressed. Scutellum rounded behind, rugose and pubescent; pits
deep, separated, with bottoms smooth and shining. Mesopleura smooth, polished
and bulging slightly below. Wing hyaline, pubescent, ciliate; veins brown, areolet
small and faint; Mj not reaching the basal (Rs+M); radial cell three times as long
as broad. Legs pubescent, tarsal claws toothed. Propodeum with arcuate carinae,
area between, smooth. Abdomen longer than high with tergites II and III only,
showing along the dorsal curvature. All tergites punctate along the posterior bor¬
ders. Ventral spine almost bare, six times as long as broad. Range in length of 26
females, 1.3-2.0 mm. Average length 1.6 mm.
Types. —The holotype female and six paratypes are in the collection of the U.S.
National Museum. Other paratypes are in the collections of the California Acad¬
emy of Sciences, the Los Angeles County Museum, the Weld collection at Los
Angeles City College and the collection of Charles Dailey.
Host. — Quercus dumosa Nutt.
Gall (Fig. 1C).—A monothalamous, cup-shaped leaf spangle, 3-3.5 mm in
height. The larval cell occupies the base of the cup whose margins are collapsed
at maturity. The galls appear on the leaves during the early summer months and
grow rapidly until they reach full size in August. At this time the galls are red
and contain small larvae. Pupation occurs in late October and early November.
Mature, fully winged, active adults can be cut out in early December; however,
normal emergence takes place in February and March, and later in some areas.
The gall resembles the gall of an Arizona species Andricus scutella Weld but is
VOLUME 60, NUMBER 4
293
somewhat smaller. The adult females of Andricus bakkeri can be distinguished
from A. scutella by the following characteristics: When viewed from the front,
the head shows a prominent, flattened hump in the ocellar area; the cheeks are
not broadened behind the eyes; the antennae are 14-segmented; wing more densely
pubescent; the abdomen is distinctly longer than high. A. scutella does not have
the flattened ocellar hump, the cheeks are slightly broadened behind the eyes, the
antennae are 13-segmented and the abdomen is almost as high as long. A similar
gall is frequently associated with the galls of A. bakkeri. This gall is also a mono-
thalamous, cup-shaped leaf spangle but the edges are inrolled like the gall of
Phylloteras cupella (Weld) from Arizona, although the insects do not resemble
the latter. This associated cup spangle is doubtfully distinct from A. bakkeri but
there are some minor morphological differences. Photographs of the galls are both
shown in Weld’s “Cynipid Galls of the Pacific Slope.”
Locality.— The type and paratypes were collected at La Crescenta, California.
Galls have also been taken on Quercus garryana Dougl., Quercus engelmanni
Greene and probably occur in many areas where these oaks are found.
Heteroecus crescent us, New Species
Bisexual Generation
Female.— Deep reddish brown, almost black. Parts of the head, mesoscutum
and abdomen yellowish. Antennae yellow. Head transverse, as wide as the thorax,
rugocoriaceous, only slightly broadened behind the eyes. Interocular space, twice
as wide as high. Malar space rugose, one-third the length of the eye. Ocellar area
appearing flattened when viewed from the front. Occiput concave. Antennae 13-
segmented, the third segment longer than the fourth, terminal segment twice as
long as the preceding one and appearing to be divided in some specimens. Meso¬
scutum broader than long, roughly coriaceous with scattered pubescence. No-
taulices incomplete, very deep posteriorly with ridged bottoms. Median groove
deep extending one-third the distance to the forward margin. Lateral lines rep¬
resented by smooth, raised lines. Scutellum rounded behind, coarsely rugose es¬
pecially on the posterior portion. Pits deep, separated, with ridged bottoms. Me-
sopleuron aciculate, smoother ventrally and slightly bulged. Wing hyaline,
pubescent and short ciliate. Veins very light, areolet present but faint. Mj not
reaching basal (Rs+M). Radial cell five times as long as broad. First abscissa of
radius angled. Legs pubescent, tarsal claws simple. Propodeum with arcuate ca-
rinae, area between ridged. Abdomen higher than long, tergite II only visible
tergite along the dorsal curvature, smooth and occupying nearly all the abdomen
in side view. Other tergites visible when abdomen distended with eggs. Ventral
spine bare, five times as long as broad. Range in length of 14 females, 1.5 mm
to 1.8 mm. Average length 1.65 mm.
Male. — Body black, except abdomen which is dark, reddish brown and legs
which are yellow. Head with very large eyes, malar space narrow, one-fifth the
length of the eye. Interocular space twice as wide as high. Ocelli protruding con¬
spicuously. Antennae fuscous with 15 segments. Mesoscutum uniformly rugose,
not as coarse as female. Pits deep and confluent. Wings pubescent with conspicuous
cilia along the posterior margins. Areolet small, absent in some specimens. M x
reaching the basal. Abdomen with tergites II and III visible along the dorsal
curvature. Range in length of 14 specimens, 1.4-1.8 mm. Average length 1.6 mm.
294
PAN-PACIFIC ENTOMOLOGIST
Types.— The holotype female, the allotype male and four paratypes are in the
collection of the U.S. National Museum. Other paratypes are in the collections
of the California Academy of Sciences, the Los Angeles County Museum, the
Weld collection at Los Angeles City College and collection of Charles Dailey.
Host. — Quercus chrysolepis Liebm.
Gall (Fig. ID).—A small, green blister two and one-half mm in length, showing
on both sides of the leaf. Galls are double chambered with the developing larvae
and pupae occupying the lower chamber. The adult insects emerge from the lower
side of the leaf.
Locality.— The type and paratypes were collected from galls on Quercus chry¬
solepis Liebm. in La Crescenta, California.
Heteroecus devorus, New Species
Bisexual Generation
Female.— Reddish brown, antennae and legs amber. Head transverse, as wide
as the thorax, roughly granular, not broadened behind the eyes. Interocular space
two-fifths as high as wide. Malar space slightly depressed, two-fifths the length of
the eye. Ocellar area somewhat flattened when viewed from in front; anterior
ocellus depressed. Occiput concave. Antennae 13-segmented, the third as long as
the fourth, terminal segment almost twice as long as the preceding one. Meso-
scutum broader than long, roughly coriaceous with scattered pubescence. No-
taulices incomplete, wide and deep posteriorly with ridged bottoms. Median groove
represented by a small notch. Lateral lines visible as smooth, raised ridges. Scu-
tellum rounded behind, coarsely rugose, especially posteriorly. Pits deep, sepa¬
rated, bottoms slightly ridged. Mesopleuron aciculate, slightly bulged. Wing hya¬
line, pubescent, not ciliate; veins light brown, areolet distinct, reaching one-fourth
the distance to the basal (Rs+M). M x not reaching basal; radial cell six times as
long as broad; first abscissa of radius arcuate. Legs pubescent, tarsal claws simple.
Propodeum with parallel carinae. Abdomen longer than high, all tergites visible
along the dorsal curvature. Tergite II the largest with posterior margin punctate.
Ventral spine long and slender, eight times as long as broad with a few scattered
bristles. Range in length of eight females, 1.4-2.2 mm. Average length 1.8 mm.
Male.— Body orange yellow, head and eyes black, ocelli amber. Eyes very large,
malar space narrow, only one-tenth the length of the eye. Interocular space three-
fifths as high as wide. Ocelli large and protruding with rugose surrounding area.
Antennae with 15 segments. Mesoscutum not as rough as female with a median
groove extending one-third the distance to the pronotum. Scutellar pits deep and
confluent. Wing pubescent with conspicuous cilia along the posterior margin.
Areolet small but well defined. M, reaching basal (Rs+M). Range in length of
four specimens 1.65 mm.
Type.— The holotype female and the allotype male are in the collection of the
U.S. National Museum. Paratypes are in the collection of the California Academy
of Science.
Host. — Quercus chrysolepis Liebm.
Gall (Fig. IE).—A small, monothalamous leaf blister 2.3 mm in length. The
galls are visible on both surfaces of the new leaves and mature rapidly. Adults
emerge from the upper surface of the gall 24 May-7 June.
VOLUME 60, NUMBER 4
295
Locality.—The types and paratypes were collected from galls growing on the
leaves of Quercus chrysolepis Liebm. in Pasadena, California.
Callirhytis lentiformis , New Species
Unisexual female.— Uniform reddish brown, ovipositor sheaths chocolate. Head
transverse, slightly narrower than the thorax, cheeks bulging prominently behind
the eyes. Malar space depressed, striated, less than one-half the length of the eye.
Interocular space three times as wide as high. Ocellar area coriaceous with a few
scattered punctures. Anterior ocellus depressed. Occiput strongly concave. An¬
tennae 14-segmented, the third segment slightly longer than the fourth, the ter¬
minal segment longer than the preceding one. Mesoscutum as long as broad,
coriaceous, sparsely pubescent with scattered, shallow punctures. Notaulices per-
current, very deep and strongly arcuate. Median represented by a small notch.
Lateral lines very short depressed. Scutellum coarsely rugose, margined, pits deep,
separated, with the bottoms smooth and shining. Mesopleuron shining, slightly
aciculate with a smooth area under the base of the wing. Wing hyaline, ciliate,
pubescent with brown veins. Areolet large, reaching one-fifth the distance to the
basal (Rs+M). Mj not reaching basal. Radial cell nearly four times as long as
broad. First abscissa of radius sharply angled and with a faint cloud. In some
specimens, this vein shows a distinct spur. Legs amber, claw simple. Propodeum
with parallel carinae, area between, smooth. Abdomen higher than long, smooth,
polished except posterior tergites which are slightly punctate. All tergites showing
along the dorsal curvature. Ventral spine long and slender, six times as long as
broad with a few bristles. Ovipositor sheaths protruding. Average length of 230
females, 2.8 mm.
Type.— The holotype female and ten paratypes are in the collection of the U.S.
National Museum. Other paratypes are in the collections of the Los Angeles
County Museum, the California Academy of Science and the collection of Charles
Dailey.
Host. —Quercus agrifolia Nee.
Gall (Fig. IF).—A spherical, deciduous, bud gall that drops to the ground in
October. The galls resemble the galls of Callirhytis agrifolia Bassett but are much
smaller and quickly shrivel to a biconvex, lens-shaped gall 3.5 mm in diameter.
The galls contain large larvae at the time they drop from the tree but adults do
not emerge until the following summer (July, August and September).
Locality.— The type and paratypes were collected from Quercus agrifolia Nee
growing in the San Gabriel Mountains north of Pasadena, California.
Acknowledgments
The writer wishes to express his appreciation to Charles Dailey of Sierra College
for his help in the completion of this paper; to Gerhard Bakker who has made
the illustrations for all of my papers; and to my wife, Devie, who has always
encouraged me in my work.
Literature Cited
Burnett, J. A. 1974. A new cynipid wasp from California. Pan-Pac. Ent., 50(3):298-302.
Dailey, D. C. 1973. A new species of Diplolepis from California. Pan-Pac. Ent., 49(2): 174-176.
296
PAN-PACIFIC ENTOMOLOGIST
-. 1977. Three new gall-inducing Callirhytis Foerster from Quercus cedrosensis Muller. Pan-
Pac. Ent., 53(1):43—46.
Eady, R. D., and J. Quinlan. 1963. Handbook for the identification of British insects, (Hymenoptera:
Cynipidae) key to families, subfamilies and Cynipinae. Including galls. Royal Ent. Soc. London,
7 pt. 1A:81 pp.
Tulloch, G. S. 1929. Proper use of terms “Parapsides” and “Parapsidal furrows.” Psyche, 36:376—
382.
Weld, L. H. 1957. Cynipid galls of the Pacific Slope. Printed privately, Ann Arbor, Michigan.
64 pp.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 297-299
A Digger Wasp Preying on a Jerusalem Cricket
G. N. Dodson and D. T. Gwynne
(GND) Biology Department, University of New Mexico, Albuquerque, New
Mexico 87131; (DTG) University of Toronto, Erindale Campus, Zoology De¬
partment, Mississauga, Ontario L5L 1C6, Canada.
Palmodes spp. (Hymenoptera: Sphecidae) are primarily predators on katydids
(Orthoptera: Tettigoniidae) (Bohart and Menke, 1976). The only known exception
to this is P. carbo Bohart and Menke whose single reported prey species is Cy-
phoderris strepitans Morris and Gwynne (Haglidae) (Evans, 1970). We observed
the nesting cycle of a P. carbo female preying on a Jerusalem cricket, Stenopel-
matus fuscus Haldeman (Stenopelmatidae) and are reporting it here both to add
to the paucity of prey records for P. carbo and to report this species as the first
known hymenopteran predator of Jerusalem crickets, a widespread group of Or¬
thoptera in the Southwestern U.S. (Tinkham and Rentz, 1969).
At approximately 0845 on 13 July 1981, 8 km N of Dutch John, Daggett Co.,
Utah, a female P. carbo was located by her intermittent buzzing as she excavated
her prey near the base of a sagebrush plant. With alternate movements of her
forelegs she excavated soil from around a Jerusalem cricket located 1-2 cm beneath
the soil surface. The prey was on its back and kicking vigorously when uncovered
by the wasp. Working rapidly, the wasp uncovered the cricket, grasped it with
her legs and stung it in the neck region. The paralyzed prey was then dragged
away by the wasp. At this point a plastic bag was placed over the pair and the
insects were observed in the bag. The wasp continued to grasp her prey and was
released from the bag at the point of capture a few minutes later. She straddled
the cricket and, holding it in her mandibles, dragged it headfirst over the ground
(Fig. 1). The prey was left three times while the wasp searched in the area. The
third time this occurred, the wasp located and opened the entrance to her nest.
She returned to her prey 2.3 min later and dragged it to the nest entrance. She
then proceeded to back into the burrow dragging the prey. At this point we
interrupted her by pulling the cricket a few cm from the entrance in order to take
photographs. The prey was then dragged into the nest headfirst and the wasp
reappeared at the nest entrance 3.1 min later. Nest closure took 26.5 min with
the female alternately throwing soil to form a mound at the nest entrance and
turning to push the soil into the burrow with her head and foretarsi. Occasionally,
she carried debris to the nest entrance and packed it into the burrow. At one point
she used a small stone, held in her mandibles, to push the soil. The entire nesting
cycle was estimated at ca. 50 min (subtracting the times that we disturbed the
wasp). The single prey, some 8 times heavier than the wasp (1.508-0.185 g wet
weight) was positioned on its right side, 5 cm deep and ca. 8 cm from the nest
entrance. The 3.5 mm egg was attached to the first abdominal segment and was
positioned transversely over the left metathoracic coxa (Fig. 2, arrow).
Thus, P. carbons only known prey are fossorial orthopterans in that it takes
both Jerusalem crickets and haglids (individuals of both groups are nocturnal and
298
PAN-PACIFIC ENTOMOLOGIST
Figures 1-2. 1. Palmodes crabro shown grasping her prey (a Jerusalem cricket) with her mandibles
while dragging it into her nest. 2. Arrow indicates position of wasp’s egg.
burrow into the soil during the day). This is in contrast to the known prey of the
six other Palmodes species, all of which take bush inhabiting Tettigoniidae (Bohart
and Menke, 1976).
VOLUME 60, NUMBER 4
299
Acknowledgments
We thank Dr. H. E. Evans for reading the manuscript, A. S. Menke for iden¬
tifying the wasp and Jill Jereb for held assistance.
Literature Cited
Bohart, R. M., and A. S. Menke. 1976. Sphecid wasps of the world. University of California Press,
Berkeley.
Evans, H. E. 1970. Ecological-behavioral studies of the wasps of Jackson Hole, Wyoming. Bull.
Mus. Comp. Zool., 140:451-511.
Tinkham, E. R., and D. C. Rentz. 1969. Notes on the bionomics and distribution of the genus
Stenopelmatus in central California with the description of a new species (Orthoptera: Gryl-
lacrididae). Pan-Pac. Ent., 45:4-14.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 300-303
Observations on the Behavior of Triepeolus nr. eldredi
Cockerell (Hymenoptera: Anthophoridae)
Stephen L. Clement 1
Department of Entomology, Ohio Agricultural Research and Development Cen¬
ter, Wooster, Ohio 44691.
Bees in the genus Triepeolus Robertson (Anthophoridae) are cleptoparasites
(nest parasites) of several species of the bee genus Melissodes Latreille (Antho¬
phoridae) (Bohart, 1970). Of the species of Melissodes whose nesting biologies
are known, it has been reported that the entrance tunnels in the nests of two
species (M. pallidisignata Cockerell, M. rustica (Say)) are continuously plugged
with loose sand from the time of initial nest construction to cell closure (Thorp,
1964; Clement, 1973). Thorp (1964) suggested that the absence of parasites from
the nests of M. pallidisignata was possibly due to the plugged tunnels. Clement
(1973) reported, however, that the plugged entrance tunnels of M. rustica nests
were no deterrent against parasitism by Triepeolus nr. eldredi Cockerell. In fact,
examination of 51 single celled nests of M. rustica revealed that 24 had been
parasitized by the cleptoparasite.
This paper makes available information about the behavior of T. nr. eldredi at
a nest site of M. rustica. Observations were carried out during the 1972 nesting
cycle and thereafter on one day during each of the nesting cycles of 1977 (July
30), 1978 (July 27), and 1979 (July 26). The study site, which is located in
Yellowstone National Park, was previously described during a study of M. rustica
(Clement, 1973).
On warm (>18°C) sunny days of observation numerous individuals of both
sexes of adult T. nr. eldredi were seen flying in a search pattern a few cm above
the ground of the M. rustica nest site. Additionally, both sexes of the parasitic
bee were seen visiting nearby flowers of Eriophyllum integrifolium (Hook.) Smiley
(Compositae). As reported earlier (Clement, 1973), M. rustica females relied ex¬
clusively on this floral source for pollen.
On several occasions, females of T. nr. eldredi alighted at the edge of an active
nest where an incoming host bee was in the process of searching for her nest
entrance (Fig. 1). While a host bee remained inside her nest (n = 12 obs.; 11-30
min), I observed a continuous turnover of parasitic bees waiting near the nest
entrance. In contrast to the observations of Rozen et al. (1978) on another parasitic
anthophorid bee, Protepeolus singularis Linsley and Michener, I did not observe
individual bees wait for extended periods near a nest entrance while a Melissodes
host was inside the nest. Although the significance of the behavior depicted in
Figure 1 is unclear, it may be used by this parasitic bee to learn the positions of
nest entrances, thereby allowing them to leave and then find the same nest again.
A discussion of nest-location learning behavior in other parasitic bees is provided
by Rozen et al. (1978).
1 Present address: USDA, American Embassy-Agric., APO New York 09794.
VOLUME 60, NUMBER 4
301
Figures 1-4. 1. Incoming female Melissodes rustica (arrow) searching for nest entrance with para¬
sitic females of Triepeolus nr. eldredi resting above her. 2. T. nr. eldredi female searching the ground
of the nest site. 3. T. nr. eldredi female entering the burrow of a M. rustica nest. 4. Same parasitic
bee penetrating loose sand of burrow plug.
On 31 occasions I observed parasitic bees enter and remain in nests for 7-20
min, leaving before the host bee returned. These bees appeared to locate nest
entrances by either randomly searching the ground of the nest site (Fig. 2), or by
observing host bees leave their nests. In the latter case, parasites alighted near
nest entrances as the host bees were in the process of digging out of plugged
burrows. In 5 instances, however, I observed a parasite enter a nest while the host
bee was in the nest, but these bees exited the nest within 1 min after disappearing
into the plugged burrow. Thus, it would appear that the avoidance behavior just
described serves to reduce disturbance of the host while she is provisioning her
nest.
Once a nest entrance was located it usually took several minutes of parasite
digging activity (n = 18 obs.; 2-10 min) before the head and most of the thorax
disappeared into the burrow plug (Fig. 3). After gaining this foothold in the burrow
plug, a bee usually all but disappeared (Fig. 4) in <1 min. During the initial
digging phase (Fig. 3 and before), female bees scraped loose sand particles out of
the tunnel entrance with their forelegs and probably their mandibles. The next
action involved backing away from the entrance for a short distance (<6 cm)
302
PAN-PACIFIC ENTOMOLOGIST
during which the kicking and scratching activity of the two posterior pairs of legs
propelled sand particles away from the nest entrance. It was apparent that T. nr.
eldredi experienced more difficulty in gaining entrance to a nest than did the host
bee. As previously reported (Clement, 1973), burrowing host bees quickly dis¬
appeared (12-28 sec) once they located their nest entrance.
I frequently observed parasitic females briefly grapple with each other on the
ground near nest entrances. Sometimes these encounters simultaneously involved
2-4 bees. Intraspecific aggressive behavior may have interfered with the ability
of individual bees to perch for extended lengths of time near nest entrances.
Interestingly, a specific form of antagonistic behavior was observed in one instance
where two parasites were seen attempting to pull a third from a nest entrance. I
did not observe any aggressive behavior between Melissodes and Triepeolus bees,
an observation that agrees with Bohart’s (1970) statement that host bees usually
ignore inquilines that approach or enter their nests. In contrast, aggressive inter¬
actions involving host bees and cleptoparasitic bees have been reported by Thorp
(1969), Rozen et al. (1978), and Eickwort and Abrams (1980).
A unique feature of the behavior of T. nr. eldredi is its ability to dig through
the plugged tunnels of active nests of M. rustica. Moreover, a parasitic bee must
penetrate a plug that covers a linear distance of 6-20 cm in order to reach and
parasitize a cell (Clement, 1973 and unpubl. data). Other parasitic Anthophoridae,
namely melectine bees, are known to dig through the soil plug of completed host
nests (see Thorp, 1969). This is the first report that documents “digging behavior”
by a nomadine bee.
In their studies, Thorp (1964) and Clement (1973) noted that cell provisions
emitted a moderate or strong fermenting odor. Stephen et al. (1969) reported that
a similar type of odor has been detected in the abdominal venter of some adult
females in the Anthophorini and that this odor may serve as an attractant to the
nesting site. Is it possible that Melissodes and Triepeolus females are directed in
part by a chemical cue in nest plugs when they search (as shown in Fig. 2) for
nest entrances at the Yellowstone nesting site? In this context, Tengo and Berg¬
strom (1977) presented evidence suggesting that odor correspondence between
cleptoparasites {Nomada bees) and hosts ( Andrena bees) permitted female para¬
sites to locate and then gain access to a host nest more easily.
Acknowledgments
I dedicate this paper to Professor R. M. Bohart who first encouraged me to
study the biology and behavior of insects. I thank Dr. R. W. Rust for field assistance
and Dr. W. L. Rubink for reviewing the manuscript.
Literature Cited
Bohart, G. E. 1970. The evolution of parasitism among bees. Utah State Univ., Logan, 33 pp.
Clement, S. L. 1973. The nesting biology of Melissodes ( Eumelissodes ) rustica (Say), with a description
of the larva (Hymenoptera: Anthophoridae). J. Kansas Entomol. Soc., 46:516-525.
Eickwort, G. C., and J. Abrams. 1980. Parasitism of sweet bees in the genus Agapostemon by cuckoo
bees in the genus Nomada (Hymenoptera: Halictidae, Anthophoridae). Pan-Pacific Entomol.,
56:144-152.
Rozen, J. G., Jr., K. R. Eickwort, and G. C. Eickwort. 1978. The bionomics and immature stages
of the cleptoparasitic bee genus Protepeolus (Anthophoridae, Nomadinae). Amer. Mus. Nov-
itates, 2640:1-24.
VOLUME 60, NUMBER 4
303
Stephen, W. P., G. E. Bohart, and P. F. Torchio. 1969. The biology and external morphology of
bees. Ag. Exp. Sta., Oregon State Univ., Corvallis, 140 pp.
Tengo, J. and G. Bergstrom. 1977. Cleptoparasitism and odor mimetism in bees. Do Nomada males
imitate the odor of Andrena females? Science, 196:1117-1119.
Thorp, R. W. 1969. Ecology and behavior of Melecta separata callura (Hymenoptera: Anthophor-
idae). Am. Midi. Natur., 82:338-345.
-, and J. A. Chemsak. 1964. Biological observations on Melissodes ( Eumelissodes) pallidisig-
nata. Pan-Pacific Entomol., 40:75-83.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 304-312
Pho bet us desert us, a New Melolonthine Scarabaeidae
(Coleoptera) from the Central Desert of
Baja California, Mexico
Paul E. Blom and William H. Clark
Museum of Natural History, College of Idaho, Caldwell, Idaho 83605.
Phobetus was first reported from Baja California, Mexico, with the description
of Phobetus chearyi Hardy (1973). Since then Phobetus sleeperi Hardy was de¬
scribed from a single male specimen collected 11.7 km east of El Rosario. The
present paper describes a third endemic species to the peninsula, Phobetus desertus
n. sp. This species is distinguished from P. sleeperi by its larger size, black color,
white ventral thoracic pile and smooth elytra. Only limited biological information
has been reported for Phobetus comatus LeConte, Phobetus palpalis Saylor (Ca-
zier, 1937) and P. chearyi (Hardy, 1973). Shook (1979) reported an adult P.
comatus as a prey item for the tiger beetle Cicindela purpurea auduboni LeConte.
Males of P. desertus flew during a light rain shower in January 1976, on a clear
morning in January 1981 and following a night of showers in January 1982, being
attracted to and mating with stationary females. Little is known of the larvae and
host plants of the genus. Cazier (1937) reported larvae of P. comatus from roots
of Artemisia californica Less., and Ritcher (1949, 1966) provided a description
of the P. comatus larva. To our knowledge there is no published description of
the Phobetus egg. Some biological observations are reported for the new species,
as well as meteorological information and soil parameters for the type locality.
Resumen
El genero Phobetus se fue divulgado primero de Baja California, Mexico, con
la descripcion de Phobetus chearyi Hardy (1973). Desde entonces Phobetus sleeperi
Hardy se fue describido del unico ejemplar masculino colectado 11.7 km este de
El Rosario. Este papel describe la tercera especie endemica de la peninsula, Pho¬
betus desertus n. sp. Este especie se distingue de P. sleeperi por su tomano mas
grande, color negro, pelo bianco y ventral toracico, y elitros lisos. Sola la infor¬
mation biologica que es limitada ha divulgado de Phobetus comatus LeConte,
Phobetus palpalis Saylor (Cazier, 1937) y P. chearyi (Hardy, 1973). Shook (1979)
divulgo un P. comatus adulto como una cosa de rapina por Cicindela purpurea
auduboni LeConte. Los machos de P. desertus volaron durante una lluvia ligera
en el enero de 1976, en una manana clara en enero de 1981 y despues de un noche
de lluvia ligera en el enero de 1982, se fueron atraidos y se aparearon con las
hembras estacionarias. No mucho se sabe de las larvas y plantas de hostias del
genero. Cazier (1937) divulgo las larvas de P. comatus de las raices de Artemisia
californica Less., y Ritcher (1949, 1966) proveyo una descripcion de las larvas
de P. comatus. De nuestro conocimiento no hay una descripcion publicado del
huevo de Phobetus. Algunas observaciones biologicas se divulgan por las especies
nuevas, asi como la information meteorologica y parametros de la tierra por la
locilidad del tipo.
VOLUME 60, NUMBER 4
305
Phobetus desertus, New Species
Holotype male. — Length 13.6 mm, width 6.0 mm (elytra humeri). Head: Black,
clypeus and frons densely punctate, with scattered short brown setae; vertex smooth,
shining, with scattered punctures laterally. Antennae (Fig. 1 a) 9 segmented, with
6 segmented club, lamellae of segments four 0.52 and five 0.83 length of remaining
lamellae; ratio of length of scape to maximum lamellae 1:2.8, venter of scape
with long white setae, apex and lateral margins with erect brown setae. Palpi 3
segmented, first 0.47, second 0.31 mm, black, slender; terminal segment 0.62 mm
black to fuscous, apex white, slightly inflated. Venter covered with long, white to
yellow setae. Prothorax: Width 5.0 mm; black, shining, with fine punctures; mar¬
ginal bead complete; setae cream anteriorly, paler laterally, white posteriorly; disc
glabrous. Elytra: Black, shining, length 9.6 mm, expanded behind humeri, ap¬
pearing smooth to naked eye, but with faint striae, transverse rugulosities and
scattered micropunctures which are larger and more concentrated anteriorly; sur¬
face with long, white setae anteriorly, and a row of stout, brown to black setae
laterally, becoming shorter posteriorly, absent apically. Pygidium: Black, shining,
with scattered punctures, white setae. Legs: Profemora fuscous to black with long,
white setae ventrally, single median row of stout black setae dorsally; protibiae
black with brown spot dorsally, tridentate, apical spur subequal to distal tooth;
tarsal claws bifid; meso-, metatibiae with circle of apical teeth, two large posterior
apical spurs. Abdomen: Black, six free stemites, with scattered, fine punctures and
white setae. Genitalia: Figure 2. Brown, finely wrinkled. Parameres contiguous in
basal Vi, separate distal %; distal U swollen to twice medial paramere width; mesal
margins with two emarginations; pronounced lateral carina and declivity in basal
x h. Apex of parameres with median carina.
Allotype female. — Length 14.4 mm; width 6.5 mm (elytra humeri). Differs from
holotype as follows: Head: Antennae (Fig. lb) 9 segmented with 5 segmented
club; ratio of scape length to maximum lamellae length 1:1.5; setae brown. Pro¬
thorax: Width 5.3 mm, black with faint fuscous markings as stripe medially, two
spots laterally, band on posterior margin. Elytra: Length 9.2 mm; translucent,
yellow to brown, margins fuscous to black, punctulate, with internal features
resembling fracture lines. Legs: Profemora with dorsal row of brown setae.
Material examined.— Holotype male (CAS #14103), allotype female, 220 para-
types (206 male, 14 female): Mexico, Baja California Norte, 9 km NW Rancho
Santa Ines, Lat. 29°46'N, Long. 114°46'W, 550 m, 6 January 1982, William H.
Clark and Paul E. Blom. Twenty-four male paratypes, above locality, 22 January
1976, Dave Ward. Thirty-six paratypes (35 male, 1 female), above locality, 22
January 1976, William H. Clark. Two hundred male paratypes: Mexico, Baja
California Norte, 7.7 km N Catavinacito, 19 January 1981, E. L. Sleeper. Primary
types at California Academy of Sciences, San Francisco; paratypes deposited at
Orma J. Smith Museum of Natural History, College of Idaho, Caldwell; Univer-
sidad Nacional Autonoma de Mexico, Coyoacan; Instituto Nacional de Agricolas,
Chapingo; Los Angeles County Museum of Natural History, California; San Diego
Natural History Museum, California; California Department of Agriculture, Sac¬
ramento; Santa Barbara Museum of Natural History, California; California State
University, Long Beach; University of Idaho, Moscow; and collections of the
authors.
Variation in paratypes.— Males: Elytra length 8.1 to 11.0 mm. Pronotal width
306
PAN-PACIFIC ENTOMOLOGIST
Figures 1-3. 1. Antennae of Phobetus desertus primary types; la, holotype male; lb, allotype
female. 2. Genitalia of Phobetus desertus holotype male; 2a, frontal view; 2b, lateral view; MC =
median carina, LC = lateral carina, and LD = lateral declivity. 3. Phobetus desertus first instar larva
head and antenna, frontal view; L = labrum, PC = preclypeus, PSC = postclypeus.
4.0 to 5.7 mm. Head setae predominately brown, grading to black in a few
specimens. Palps brown to black. Elytra predominately black occasionally grading
to slightly fuscous. Legs black to fuscous. Females: Elytra length 7.5 to 10.3 mm.
Pronotal width 4.0 to 5.7 mm. Frons fuscous to black. Pronotum light brown
ULILLI o’L
VOLUME 60, NUMBER 4
307
with black markings to black. Elytra with or without internal features resembling
fracture lines.
Diagnosis.—Phobetus desertus males can be distinguished from all other species
of Phobetus except P. sleeperi by the six segmented club. Phobetus desertus keys
to P. sleeperi in Hardy’s key (1978), but differs from this species by the white
pile, black pronotum and elytra, larger size and significantly smoother elytra.
Specimens of P. desertus which approach fuscous in elytra color still appear black
to the naked eye, thus differing from the holotype of P. sleeperi. Male genitalia
of P. desertus have the lateral carina long and the declivity pronounced and deep,
while P. sleeperi has only a short lateral carina with a small shallow declivity.
Mesal margins of the parameres in P. desertus are usually with 1 or 2 emargi-
nations, whereas in P. sleeperi it is straight. An apical medial carina occurs in P.
desertus, but is lacking in P. sleeperi. Female P. desertus resemble males of Pho¬
betus saylori Cazier, but can be distinguished by their blunt, five segmented club.
Remarks.— There is a marked resemblance between P. desertus and P. sleeperi.
However it is impossible to assess the variation of P. sleeperi as the species is
known from only the single male holotype. The six segmented club of the male
and the dark coloring are the most evident similarities between the species. Close
examination of P. desertus reveals slight rugulosities in the elytra and a few of
the paratypes show a translucent character to the elytra and a shading to fuscous,
reinforcing the similarity. The genitalia of P. desertus are also similar to that of
P. sleeperi. However, specific distinction is proposed here for P. desertus based
on the following differences: Size of P. desertus is much larger than P. sleeperi.
Hardy (1978) measures the length of P. sleeperi as 11.7 mm; our measurements
(with an ocular micrometer from the dorsal aspect), which correspond to those
for male P. desertus of x = 13.4 mm ( n = 258), show the length of P. sleeperi at
10.5 mm. Hardy (1978) also reports a width for the thorax of P. sleeperi at 4.25
mm, comparing to a mean pronotal width of 5.0 mm for P. desertus (n = 265,
s = 0.58). The ratio of elytra length to pronotal width is 1.95 for male P. desertus
( n = 260, s = 0.061) and 2.5:1 for P. sleeperi, showing P. desertus to be much
more robust. While in P. desertus the ventral pile is consistently white grading to
cream only on the anterior margins of the pronotum, the thoracic setae are uni¬
formly brown in P. sleeperi. Head setae of P. desertus like those of P. sleeperi are
brown. The consistently black appearance of P. desertus differs obviously from
the fuscous color of P. sleeperi, though with microscopic examination a few of
the P. desertus shade to fuscous. Transverse rugosities and striations are obvious
in P. sleeperi, but obscure in P. desertus.
At present the genitalia of Phobetus provide minimal information for specific
determination. Strong genital similarities between P. sleeperi and P. desertus are
also shared by P. saylori. Such interspecific similarities coupled with intraspecific
variation in P. comatus and P. desertus accord this character little value. Cazier
(1937) reports genitalic similarity between the externally distinct Phobetus mo-
javus Barrett and P. palpalis and considers the character of limited value. See
illustrations in Cazier (1937) and Hardy (1978).
Locality information on the holotype of P. sleeperi and subsequently reported
in Hardy (1978) is “5 mi E El Rosario.” However, according to the collector, E.
L. Sleeper (pers. comm.), this is an error for the collection was actually made 11.7
km (7.25 mi) east of El Rosario, Baja California, Mexico, placing the collection
308
PAN-PACIFIC ENTOMOLOGIST
Table 1. Soils data for six areas at our Phobetus desertus collection site 9 km NW Rancho Santa
Ines, Baja California, Mexico (— parameter not analysed).
Parameter
Elec¬
trical
conduc¬
tivity
Sam¬
ple
Depth
(cm)
of satu¬
ration
extract
(Aimhos)
Sodium-
adsorp¬
tion
ratio
pH
Texture
Calcium
carbonate
Phos¬
phorus
(ppm)
Potas¬
sium
(ppm)
Nitrate
nitrogen
(ppm)
1 a
305
0.7
5.25
7.8
Coarse sand
Slight
2.6
73
<10
b
915
0.6
3.44
8.6
Coarse sand
Moderate
4.3
56
<10
c
1220
0.20
0.80
8.5
Coarse sand
Slight
8.7
1230
20
2 a
0-50
0.35
0.22
6.7
Coarse sand
0
18.4
128.1
2
b
150-200
0.35
0.78
7.1
Coarse sand
0
9.3
54.7
2
3 a
50-150
0.5
0.40
7.4
Coarse sand
0
17.2
195
b
305
0.25
0.64
7.9
Moderately
coarse sand
0
2
132
<10
c
458
0.35
1.57
8.4
Coarse sand
Moderate
11.8
1220
20
4 a
50-150
0.30
0.37
7.5
Coarse sand
0
5.8
215
10
b
305
0.45
1.43
7.9
Coarse sand
0
9.7
381
<10
5 a
3
0.7
0
7.1
Coarse sand
0
6.7
25
b
20-25
0.35
0.894
7.8
Coarse sand
0
5.9
25
6 a
3
0.6
0.555
6.9
Coarse sand
0
7.0
25
b
20-25
0.2
1.15
7.4
Coarse sand
0
7.0
25
on the northern edge of the Central Desert, approximately 180 m (590 ft) elevation.
While these two locations are in the Central Desert they differ ecologically: El
Rosario is still subject to coastal climatic influence, whereas the Catavina Region
is a granitic boulder field equidistant from the Gulf of California and the Pacific
Ocean. This difference is exemplified by the various definitions and subdivisions
given to regions of the Central Desert by various authors (Nelson, 1922; Tinkham,
1957; Shreve and Wiggins, 1964; Aschmann, 1967; and Wiggins, 1969, to list a
few).
Type locality.— The Catavina area is characterized by the shrubs Larrea tri-
dentata (Sesse and Moc. ex DC.) Coville, Ambrosia dumosa (A. Gray) Payne and
Ambrosia chenopodifolia (Benth.) Payne as well as several cactus species: Opuntia
echinocarpa Engelm. and Bigel., Opuntia cholla Weber, Opuntia molesta Bran-
degee, Pachycereus pringlei (S. Wats.) Britt, and Rose, Lophocereus schottii (En¬
gelm.) Britt, and Rose and Ferocactus gracilis Gates. For a more detailed botanical
and zoological survey of this site see Bratz (1976).
Soils of the Catavina site are characterized in Table 1 by eight parameters.
Fourteen soil samples from six selected soil pits near the Phobetus collection
locality are listed in the table. Each soil profile is characterized by a surface or
near surface sample and one or two deeper samples depending on depth possible.
The entire study site is dominated by coarse grus produced by the weathering of
Cretaceous granite (tonalite) of the Jaraguay block (Gastil et al., 1975). The table
shows that all samples are considered to be coarse sandy soil (one is moderately
coarse indicating a sandy loam situation). The electrical conductivity was low,
0.2-0.7 /umhos at the site. Sodium values were generally low, ranging 0-5.25
VOLUME 60, NUMBER 4
309
(sodium adsorption ratio). The soil was usually neutral to slightly basic (alkaline),
pH 6.7-8.6. The pH values were always lower on the surface. In 70% of the
samples calcium carbonate was not detected, while in four samples slight to
moderate amounts were found. Phosphorus concentrations varied greatly, from
<2 to 18.4 ppm. Half of the areas showed the highest phosphorus concentrations
at or near the ground surface. Potassium levels were low in about 50% of the
samples (<25-73 ppm) and higher in the rest (128.1-1230 ppm). Nitrate as
nitrogen was not examined for each sample, but was low when detected (2-20
ppm).
The mean annual precipitation for the Catavina area is about 96 mm (3.8 in.)
(Hastings, 1964; Hastings and Humphrey, 1969). The data was collected at several
weather stations in the area operated by the Secretaria de Recursos Hidraulicos.
According to a rain gauge established at the type locality on 9 July 1981, the
cumulative precipitation recorded on 4 January 1982 was 46 mm (1.81 in.).
Biology .—Collections of P. desertus were made on 6 January 1982, 22 January
1976 and on 19 January 1981 (by E. L. Sleeper) at the Catavina type locality.
Sleeper’s 1981 collection of 7.7 km N Catavinacito is within 1 km of our 1976
and 1982 collections, and considered the same locality. The 1976 collections were
made between 0700 and 0900 h PST following a brief rain. A temperature of 15-
19°C and barometric pressure of 1028 mb (30.04 in.) was recorded during this
time by a Weathermeasure Meteorograph. In 1982a large emergence of P. desertus
was encountered beginning at 0545 h, just prior to daylight (ca. 0620h), and was
continuing at 0830 h. Flight activity was initially sporadic but became extensive
by dawn. Barometric pressure was estimated to be 999 mb (29.5 in.) by a vehicle
altimeter and a temperature range of 9-10.5°C was recorded with a Reotemp dial
thermometer. This last emergence was preceded by an evening of rain that soaked
the substrate to a depth of 2.5 cm by the following morning, and took place during
heavy fog conditions. Major activity was in the sandy areas between the large
granitic outcrops. Males actively flew within 1 m of the soil surface and congregated
around emerging females. The females were not observed to fly. Females would
frequently be found in open spaces between vegetation and at or near the base of
the dominant shrub, Larrea tridentata. Approaching males attempted to copulate
(Fig. 4a) and would often displace one another. In one instance, an estimated 20
males were observed flying about a L. tridentata with a single female at its base.
During copulation the female would enter her burrow, usually followed by the
remaining unattached males. As many as six males were removed from the bur¬
rows of individual females, to a depth of 10 cm. Some burrows were open, others
were closed with loose soil but easy to recognize by the tumulus. Initially it
appeared that males were attracted to the female. In some cases, however, a female
emerged from her burrow and the males continued their downward digging rather
than following the female (Fig. 4b). Dissipation of the fog around 0830 h did not
seem to influence the flight activity. The time of flight termination was not ob¬
served in 1982 as we drove north along the highway to try and determine how
widespread the emergence was.
The 1981 collections by E. L. Sleeper (pers. comm.) were made on a “bright
and sunny” morning. The most recent rain had been eight days previous, with
the last noticeable dew four days after. The overnight low had been 5.0°C, and
when the flight activity began at 0830 h the temperature was 5.8°C. Activity was
310
PAN-PACIFIC ENTOMOLOGIST
Figure 4. 4a, copulating Phobetus desertus on rim of burrow, 6 January 1982; 4b, copulating pair
of Phobetus desertus on edge of burrow with one of several males continuing downward movement
into burrow, 6 January 1982. Photos by William H. Clark.
heavy by 0900 h (6.2°C). Little air movement was noted and none registered by
his meteorological equipment. For many of the males activity lasted for only an
hour, when they began to drop to the ground in a “stupor.” A number of these
individuals were attacked by ants. Acromyrmex versicolor (Pergande) and Pheidole
yaqui Creighton and Gregg were identified from parts still grasping tarsi of pre¬
served P. desertus.
Figures 5-7. 5. Lateral view of antennal segments 4 and 5. 6. Right maxilla of first instar Phobetus
desertus larva, dorsal view. 7. Mandible of Phobetus desertus first instar larva; 7a, dorsal; 7b, ventral;
stippling indicates relative sclerotization.
Several copulating pairs of the 1982 P. desertus were placed in quart jars with
0.5-0.8 cm of loose soil. Inspection of the soil after two months revealed up to
18 eggs and larvae per jar. Most of the eggs were laid around the periphery of the
jar bottom; one egg was located only four centimeters below the soil surface in a
small (ca. 2 cm diameter) earthen cell. Embryological progression is evident by
the development of segmentation and progressive degrees of mandibular sclero¬
tization. Larvae worked their way to the surface after hatching. Females were
recovered from a variety of depths in the soil; all but one were dead.
312
PAN-PACIFIC ENTOMOLOGIST
Immature forms.— The egg is white. The chorion is translucent, without surface
sculpturing. Length 3.9-4.2 mm ( x = 4.05, s = 0.138, n = 6), width 2.5-3.2 mm
(x = 2.81, s = 0.23, n = 6). First instar larvae of P. desertus generally agree with
Ritcher’s (1949) description of third instar P. comatus. Larvae of P. desertus do
appear to have long setae on the anterior margin of the post-clypeus, which are
lacking in Ritcher’s (1949) illustration of P. comatus. Figures 3, 5-7 illustrate the
front, antenna, palp and mandible of P. desertus larva. Total length x = 10.2 mm
( n = 12). Head capsule width x = 2.1 mm ( n = 12). Pigmentation of the head
capsule takes place after hatching.
Acknowledgments
The authors were partially funded by grants from EARTHWATCH and the
Center for Field Research and Sigma Xi. Dave Ward, Jr., along with the College
of Idaho and EARTHWATCH team members provided field assistance. We are
grateful to the Senora Josefina Antonia Zuniga and Francisco Espinoza Quintero
of Rancho Santa Ines for providing accommodations over the years. Peter L.
Comanor provided the soils analysis. David H. Kavanaugh of the California
Academy of Sciences graciously loaned the holotype of Phobetus sleeperi. James
B. Johnson, Donald R. Frohlich, William F. Barr, Alan R. Hardy and Frank T.
Hovore contributed guidance and manuscript comments. Jane C. Luther provided
the Spanish translation of the resumen. We thank Marc J. Klowden, Elbert L.
Sleeper, Irving and Minni Belle Imholf, Mary Clark and Ann Blom for their
suggestions, support and encouragement.
Literature Cited
Aschmann, H. 1967. The Central Desert of Baja California: Demography and ecology. Manessier
Publ. Co., Riverside, CA. 315 pp.
Bratz, R. D. 1976. The Central Desert of Baja California, Mexico. Jour. Id. Acad. Sci., 12:58-72.
Cazier, M. A. 1937. A revision of the Pachydemini of North America. Jour. Zool. Entomol., 29:
73-87.
Gastil, R. G., R. P. Phillips, and E. C. Allison. 1975. Reconnaissance geology of the state of Baja
California. Geol. Soc. Amer. Mem. 140. 170 pp. + maps.
Hardy, A. R. 1973. A new species of Phobetus (Coleoptera: Scarabaeidae). Pan-Pac. Entomol., 49:
127-131.
-. 1978. Three new Pachydemini and a key to the species of the genus Phobetus (Coleoptera:
Scarabaeidae). Coleopt. Bull., 32:47-52.
Hastings, J. R. 1964. Climatological data for Baja California. Univ. Az. Inst. Atmos. Physics Tech.
Rep. 14. 132 pp.
-, and R. R. Humphrey. 1969. Climatological data and statistics for Baja California. Univ.
Ariz. Inst. Atmos. Physics Tech. Rep. 18. 96 pp.
Nelson, E. W. 1922. Lower California and its natural resources. Natl. Acad. Sci. Mem. 16. 194 pp.
Ritcher, P. O. 1949. Larvae of Melolonthinae with keys to tribes, genera and species (Coleoptera:
Scarabaeidae). Ky. Agr. Exp. Sta. Bull. 537. 36 pp.
-. 1966. White grubs and their allies. Or. State Univ. Monogr., Stud. Entomol. 4. 219 pp.
Shook, G. 1979. A note on a prey and a predator of Cincindela purpurea auduboni. Cincindela, 11:
12 .
Shreve, F., and I. L. Wiggins. 1964. Vegetation and flora of the Sonoran Desert. 2 vols. Stanford
Univ. Press, Stanford, Calif. 1740 pp.
Tinkham, E. R. 1957. The Califomia-Sonoran-Sinaloan vegetation province. Proc. 8th Pac. Sci.
Cong., 4:139-147.
Wiggins, I. L. 1969. Observations on the Vizcaino desert and its biota. Proc. Ca. Acad. Sci., 4th
Ser., 36:317-346.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 313-325
The Genus Phyllophaga Harris, 1826
(Coleoptera: Scarabaeidae) in Cuba. II. Descriptions of
Eleven New Species and Illustrations of Female
Genitalia of Twelve Other Phyllophaga
Miguel Garcia-Vidal
586 Hickory St. San Francisco, California 94102
When I first started the study of the family Scarabaeidae in Cuba, I concentrated
my attention on the genus Phyllophaga, because it was the most economically
important. All species of Phyllophaga are polyphagous and are pests of numerous
agricultural crops, among them sugar-cane, which is the principal Cuban crop.
Moreover, some species of this genus have been reported as hosts of some parasites
(Garcia, 1975).
During my preliminary organization of the material available in the Cuban
collections, I found some specimens misidentified and many without any iden¬
tification. After a long period of work I separated the species already described
from those that did not fit the previous descriptions. This was possible only with
male Phyllophaga. Females are very difficult, almost impossible to identify, if
they are separated from males of the same species.
The descriptions of the first five new species were published in 1978 (Garcia,
1978), as a first part of a series about the genus.
The majority of the species have been established based upon male descriptions,
with illustrations of genitalia, but the genitalia of the female of these species, in
most cases, are not available. Even when the species has been established from
female material, the illustrations of the genitalia have not been published, which
makes identification more difficult.
In the present paper I present illustrations of genitalia of the females of twelve
species already described (P. explanicollis Chapin, P. expansa Chapin, P. oncho-
phora Chapin, P. tuberculifrons Chevrolat, P. bimammifrons Saylor, P. puberula
Du Val, P. patruelis Chevrolat, P. crenaticollis Blanchard, P. baracoana Chapin,
P. signaticollis Burmeister, P.jaronua Chapin and P. cubana Chapin. In the Cuban
collections I could not find identified females of the other Cuban Phyllophaga.
There are many females in these collections that could not be determined. I hope
the plates of the female genitalia will help in identification.
In this paper also, I include the descriptions of eleven new species, including
the illustrations of male and female genitalia.
The plates of the female genitalia were made by the author of this paper. I wish
sincerely to thank Dr. Alan Hardy, from the CDFA in Sacramento for the pictures
of the male genitalia and for reviewing this paper. I also want to express my
gratitude to the late Dr. Fernando de Zayas for the loan of the specimens in his
collection for my studies and for his advice and support.
Phyllophaga bimaculata, New Species
Male. — Submedium, pruinose. Elytra, legs, underparts rufocastaneous; sutural
margin, disc of pronotum dark brown; margin of pronotum lighter with an almost
314
PAN-PACIFIC ENTOMOLOGIST
Figures 1-12. 1. Phyllophaga onchophora Chapin. Female genitalia. 2. Phyllophaga cubana Chap¬
in. Female genitalia. 3. Phyllophaga baracoana Chapin. Female genitalia. 4. Phyllophaga puberula
Du Yal. Female genitalia. 5. Phyllophaga expansa Chapin. Female genitalia. 6. Phyllophaga signa-
ticollis Burmeister genitalia. 7. Phyllophaga crenaticollis Blanchard genitalia. 8. Phyllophaga explan-
icollis Chapin. Female genitalia. 9. Phyllophaga jaronua Chapin. Female genitalia. 10. Phyllophaga
tuberculifrons Chevrolat genitalia. 11. Phyllophaga patruelis Chevrolat. Female genitalia. 12. Phyl¬
lophaga bimammifrons Saylor. Female genitalia.
VOLUME 60, NUMBER 4
315
circular spot medio-laterally; head piceous. Clypeus densely, coarsely punctured;
punctures with different shapes; margins abruptly reflexed; median indentation
deep, angulate; clypeo-frontal suture almost obliterated, absent on median line.
Frons densely, coarsely punctured, but less than clypeus; punctures deep, of dif¬
ferent sizes; basal margin and median line impunctate. Pronotum less densely
punctured than head; punctures umbilicate, more orderly distributed; margins
entire; anterior angles acute, posterior obtuse, rounded; sides broadly dilated
medially. Scutellum more sparsely punctured; small punctures forming rows, disc
impunctate. Elytra slightly more densely punctured than pronotum, but punctures
smaller; with short, thick, straight hairs; sutural margin tumid, darker, not ele¬
vated, wider posteriorly. Pygidium more densely punctured than elytra but punc¬
tures smaller, shallower, with short, thick, straight hairs; dark spot covering %
from anterior margin; x h light castaneous; apex broadly rounded; margins bearing
hairs longer than those on disc and reflexed. Antennal club 3-segmented, elongate,
about twice the stem. Protibiae tridentate; upper tooth distant; median stouter
than distal. Stemites densely, finely punctured; hairs fine. Coxal plates with few
thick, straight hairs. Urostemites somewhat less densely punctured than sternites;
fifth segment with a tuft of long, fine, straight hairs laterally; last segment narrowly,
transversely grooved, with fine, long, straight hairs. Longer calcar of metatibia
acuminate, about as long as first tarsal segment with its spines. Claws slightly
curved; median tooth fine. Length: 12 mm. Figures 13, 14.
Holotype.— Male, collected by J. Valdes, from Puerto Boniato, Oriente, June
1963. Deposited in CAS Collection No. 14281.
This species is very characteristic by its coloration and antennal club, which is
one of the longest I have observed in a Cuban species.
Phyllophaga caneyensis, New Species
Male. — Submedium, pruinose, slightly ovoid. Elytra, pronotum, head piceous;
legs, underparts castaneous-ferrugineous. Frons densely, coarsely punctured;
punctures irregular; base impunctate; irregular depression toward median line;
clypeo-frontal suture almost obliterated. Clypeus punctured as frons; punctures
smaller toward clypeo-frontal suture; irregular depression toward median line,
but smaller than on frons; margins broadly reflexed, moderately elevated; median
indentation not very deep, subangulate; punctures umbilicate, very irregularly
distributed. Pronotum scarcely punctured; punctures big; margins entire; sides
broadly dilated medially; long, thick, reflexed hairs on lateral margins; anterior
angles slightly acute, almost straight; basal curved; median line conspicuous.
Scutellum scarcely punctured on sides; disc impunctate. Elytra more densely
punctured than pronotum, but less than frons; punctures smaller than on prono¬
tum but more regularly distributed; sutural margins not elevated. Pygidium cas¬
taneous-ferrugineous, punctured as elytra but punctures smaller, more regularly
distributed; apex broadly curved, with fine, straight, long hairs on margins. An¬
tennal club very well developed, 5-segmented, segments about same size, longer
than stem. Protibia tridentate; upper tooth smaller, distant from others; median
as stout as distal. Coxal plates scarcely punctured; punctures different, with thick,
straight hairs. Stemites densely, finely, regularly punctured, pubescent; hairs fine,
long. Urostemites punctured as sternites; fifth segment with tuft of long, fine,
straight hairs laterally; last segment transversely grooved. Longer calcar of meta-
316
PAN-PACIFIC ENTOMOLOGIST
VOLUME 60, NUMBER 4
317
tibia stout, elongate, acuminate. Claws slightly curved; distal tooth elongate, slen¬
der. Length: 11.5-12 mm. Figures 17, 39.
Holotype. — Male, collected by F. de Zayas, June 1967, from Gran Piedra, Caney
Oriente. In CAS, No. 14282.
Paratype. — Same data as holotype, male. In USNM.
This species is close to P. cardini Chapin, but is smaller. The antennal club is
very unusual because of its size and well developed fifth segment.
Phyllophaga complexipennis, New Species
Male.— Medium, piceous, pruinose. Frons sparsely punctured with moderate
irregular punctures; base impunctate. Clypeus slightly more densely punctured
than frons with bigger punctures; almost flat; margins slightly reflexed; median
indentation deep, angulate. Pronotum moderately punctured; punctures small;
margins entire; sides moderately dilated medially; anterior angles acute, posterior
obtuse, curved. Scutellum punctured as pronotum, but punctures more regularly
distributed. Elytra slightly more densely punctured than pronotum but punctures
similar; sutural margin tumid, slightly elevated. Pygidium punctured as pronotum;
apex subtruncate; margins bearing short, thick, straight hairs. Antennal club 3-
segmented, slightly smaller than four preceding segments as whole. Protibia tri-
dentate; teeth almost equidistant; upper slightly smaller than median; distal elon¬
gate, blunt. Stemites densely punctured; punctures fine, regularly distributed;
pubescent, hairs straight, short. Coxal plates with some thick, straight hairs. Uro-
stemites less densely punctured; hairs similar; fifth segment with tuft of long fine,
straight hairs laterally; last segment shallowly grooved. Longer calcar of metatibia
elongate, fine, acuminate, slightly longer than first tarsal segment with its spines.
Claws strongly curved; apical tooth finer than median, equal in length. Length:
17 mm. Figures 15, 16, 36.
Holotype.— Male, collected by F. de Zayas, from Vinales, Pinar del Rio, April
1964. In CAS, No. 14283.
The antennal club is atypically short. The aedeagus is very complex. It possesses
two long spines, curved downward, arising from the internal sac. From the back¬
sides of each arises another spine, one of which is smaller that its precedent and
the other projects out of the aedeagus.
Phyllophaga chalumeaui, New Species
Male.— Large, shining, elongate, yellowish-castaneous. Frons densely, coarsely
punctured; punctures of different sizes and shapes, irregularly distributed; base
impunctate; clypeo-frontal suture conspicuous. Clypeus more densely, coarsely
Figures 13-24. 13. Phyllophaga bimaculata, n. sp. Aedeagus, frontal view. 14. Phyllophaga bi-
maculata, n. sp. Aedeagus, lateral view. 15. Phyllophaga complexipennis, n. sp. Aedeagus, lateral view.
16. Phyllophaga complexipennis, n. sp. Aedeagus, frontal view. 17. Phyllophaga caneyensis, n. sp.
Aedeagus, lateral view. 18. Phyllophaga chalumeaui, n. sp. Aedeagus, lateral view. 19. Phyllophaga
franciscana, n. sp. Aedeagus, lateral view. 20. Phyllophaga hernandezi, n. sp. Aedeagus, lateral view.
21. Phyllophaga hernandezi, n. sp. Aedeagus, frontal view. 22. Phyllophaga independentista, n. sp.
Aedeagus, lateral view. 23. Phyllophaga hardyi, n. sp. Aedeagus, lateral view. 24. Phyllophaga maes-
trensis, n. sp. Aedeagus, lateral view.
318
PAN-PACIFIC ENTOMOLOGIST
Figures 25-33. 25. Phyllophaga migratoria, n. sp. Aedeagus, lateral view. 26. Phyllophaga migrato-
ria, n. sp. Aedeagus, frontal view. 27. Phyllophaga maestrensis, n. sp. Aedeagus, dorsal view. 28.
Phyllophaga maestrensis, n. sp. Aedeagus, dorso-frontal view. 29. Phyllophaga independentista, n. sp.
Aedeagus, frontal view. 30. Phyllophaga independentista, n. sp. Aedeagus, dorsal view. 31. Phyllophaga
spinicola, n. sp. Aedeagus, dorsal view. 32. Phyllophaga hernandezi, n. sp. Aedeagus, frontal view.
33. Phyllophaga hardyi, n. sp. Aedeagus, frontal view.
VOLUME 60, NUMBER 4
319
punctured, but punctures more regularly distributed; margins broadly reflexed,
more elevated on both sides of median indentation, which is deep, angulate;
margins bearing long, thick, reflexed hairs. Pronotum punctured as frons; punc¬
tures bigger, deeper; margins entire; sides broadly dilated medially; anterior angles
slightly acute, posterior broadly curved; lateral margins with long, thick, reflexed
hairs. Scutellum punctured as pronotum; disc impunctate. Elytra somewhat more
densely punctured than pronotum, but punctures slightly smaller; sutural margin
tumid; very sparsely punctured, punctures shallower, irregularly distributed; apex
moderately curved; short, straight hairs on margins. Antennal club 3-segmented,
about as long as four preceding segments as a whole. Protibia tridentate; distal
tooth elongate, acuminate. Stemites densely pubescent; hairs long, fine, straight,
arising from small, shallow punctures. Coxal plates sparsely pubescent; hairs thick,
long, straight. Urostemites less pubescent; hairs shorter, finer; last segment widely,
transversely, posteriorly grooved; sparsely pubescent. Longer calcar of metatibia
elongate; sides almost parallel, very slender, blunt, about as long as first tarsal
segment with its spines. Claws slightly curved; median tooth stouter than distal.
Length 24 mm. Figures 18, 38.
Holotype.— Male, collected by J. Acuna, from Loma de Cala to P. Mocha, Sierra
Maestra, Cuba, 16 May 1948 at 3600-3900 ft altitude. In CAS, No. 14284.
This species is dedicated to Dr. F. Chalumeau, from the Institut de Recherches
Entomologiques de la Caraibe.
Phyllophaga franciscana, New Species
Male. — Submedium, pruinose. Elytra dark castaneous; legs, underparts yellow¬
ish-brown; head piceous; pronotum dark castaneous with light ferrugineous band
on each side of median anterior region. Clypeus densely punctured; punctures
moderate; margins abruptly reflexed; median indentation moderately deep, sub-
angulate. Frons densely punctured, but less than clypeus; base impunctate, with
shallow depressions along median line. Pronotum less densely punctured than
head; punctures similar but more regularly distributed; band brighter laterally;
anterior, posterior margins entire; lateral subcrenulate; sides broadly dilated pos¬
teriorly, their widest part almost on posterior angles, which are obtuse; anterior
slightly acute. Scutellum moderately, irregularly punctured; punctures similar to
those on pronotum but deeper; disc impunctate. Elytra densely, coarsely punc¬
tured; punctures irregular, deeper than on pronotum; sutural margin tumid, wider
posteriorly. Pygidium irregularly punctured; punctures fine, shallow, more nu¬
merous on disc; apex truncate. Protibia tridentate; upper tooth distant from others;
median same length as distal. Stemites densely punctured; punctures fine, shallow.
Coxal plates with large punctures; hairs thick, short. Urostemites densely punc¬
tured, but less sparsely than stemites; punctures similar; last segment almost
completely, transversely grooved. Longer calcar of metatibia acuminate. Tarsi,
antenna missing. Length: 12.5 mm. Figures 19, 35, 37.
Holotype.— Male, collected by Noel Gonzalez, from Sierra del Grillo, Havana,
Cuba, 14 December 1974, on the ground. In CAS, No. 14286.
Phyllophaga hardyi, New Species
Male. — Small, shining. Body castaneous to dark brown; head piceous. Frons
densely, coarsely punctured; punctures of different sizes, irregularly distributed;
320
PAN-PACIFIC ENTOMOLOGIST
Figures 34-41. 34. Phyllophaga hardyi, n. sp. Aedeagus, dorsal view. 35. Phyllophaga franciscana,
n. sp. Aedeagus, dorsal view. 36. Phyllophaga complexipennis, n. sp. Aedeagus, dorso-frontal view.
37. Phyllophaga franciscana, n. sp. Aedeagus, frontal view. 38. Phyllophaga chalumeaui, n. sp. Ae¬
deagus, dorso-frontal view. 39. Phyllophaga caneyensis, n. sp. Aedeagus, frontal view. 40. Phyllophaga
spinicola, n. sp. Aedeagus, frontal view. 41. Phyllophaga spinicola, n. sp. Aedeagus, lateral view.
VOLUME 60, NUMBER 4
321
base impunctate. Clypeus punctured as frons; margins broadly reflexed, elevated;
median indentation inconspicuous; margins bearing long, fine, reflexed hairs.
Pronotum moderately punctured; punctures more regularly distributed than on
frons; anterior and posterior margins entire; lateral subcrenulate, bearing long,
fine, reflexed hairs; sides broadly, medially dilated; anterior angles acute; posterior
slightly obtuse, curved. Scutellum with small punctures toward margins; disc
impunctate. Elytra more densely punctured than pronotum; punctures slightly
deeper; sutural margin tumid, slightly elevated, wider posteriorly, with some
punctures. Pygidium more densely punctured than elytra with smaller punctures;
apex broadly curved; margins with long, fine, straight hairs. Mesosternite densely
punctured; punctures small, regularly distributed; with fine, long hairs. Coxal
plates with thick straight hairs. Urostemites less densely punctured than stemites;
long, fine hairs arising from punctures; last segment transversely, posteriorly
grooved. Pro tibia tridentate; teeth equidistant. Antennal club 3-segmented, longer
than funicle. Longer calcar of metatibia slender, acuminate, longer than first tarsal
segment. Claws very slightly curved; distal tooth long and fine, median small.
Length: 11 mm. Figures 23, 33, 34.
Holotype. — Male, collected by F. de Zayas, from Tortuguilla, Provincia Oriente,
Cuba, June 1965. In CAS, No. 14287.
Paratypes .—One male, collected by F. de Zayas from Tortuguilla, Prov. Oriente,
Cuba, June 1965, in CAS. One male, col. F. de Zayas, from Tortuguilla, Prov.
Ote. Cuba, June 1964, in USNM.
This species is dedicated to Dr. Alan Hardy from the CDFA in Sacramento.
Phyllophaga hernandezi, New Species
Male.— Large. Elytra yellowish-castaneous; pronotum, legs, underparts rufo-
testaceous; head dark brown. Clypeus densely, coarsely punctured; margins broad¬
ly reflexed, slightly elevated; median indentation moderately deep, subangulate;
clypeo-frontal suture conspicuous. Frons densely punctured, but less than clypeus;
punctures small, more numerous toward impunctate base. Pronotum densely
punctured, but less than frons; punctures bigger, umbilicate, more regularly dis¬
tributed; margins entire; sides broadly, medially, posteriorly dilated; anterior an¬
gles almost straight, posterior curved; lateral margins with thick, long, reflexed
hairs; one small depression on side of medio-posterior region aligned with eyes.
Scutellum semicircular, completely punctured; punctures as on pronotum. Elytra
more densely punctured than pronotum but with finer punctures; hairs long, thick,
straight. Sutural margins tumid. Pygidium densely punctured, slightly more than
elytra, but punctures shallower; apex narrowly rounded. Antennal club 3-segment¬
ed, about as long as funicle. Protibia tridentate; upper tooth distant from others;
median wider than distal at base. Coxal plates with thick, long, straight hairs,
which arise from fine punctures. Stemites pubescent; hairs long, fine, straight.
Urostemites less densely punctured, with similar punctures, sparsely pubescent;
last segment medially, transversely grooved. Longer calcar of metatibia slender,
acuminate. Claws somewhat curved; median tooth stouter than distal. Asym¬
metric aedeagus with a long spine arising backward from internal sac. Length: 24
mm. Figures 20, 21, 32.
Female .—Median line conspicuous on frons; antennal club as long as three
preceding segments as whole; apex of pygidium truncate; last urostemite wide,
322
PAN-PACIFIC ENTOMOLOGIST
Figures 42-43. 42. Phyllophaga maestrensis, n. sp. Female genitalia. 43. Phyllophaga hernandezi,
n. sp. Female genitalia.
convex; longer calcar of metatibia slightly longer than first tarsal segment, slender,
blunt, not typical for females. Length: 24.5 mm. Figures 20, 21, 32.
Holotype.— Male, col. Zayas and Valdes, from Sierra Maestra, June 1965. In
CAS, No. 14288.
Allotype. — Same data as holotype. In CAS.
Paratype.— Female, same data as types. In USNM.
This species is dedicated to the Cuban Entomologist Luis R. Hernandez.
Phyllophaga independentista, New Species
Male.— Medium. Elytra, pronotum, legs light yellowish-castaneous; head pi-
ceous; underparts yellowish-brown; close to Ph. alayoi Garcia. Frons densely,
VOLUME 60, NUMBER 4
323
coarsely punctured; punctures fine, irregular, more scarce on base. Clypeus densely
punctured also, but punctures more regular; margins abruptly reflexed, not very
elevated, lateral almost flat; median indentation deep, angulate; long, thick, re¬
flexed hairs on margins. Pronotum densely punctured, but less than head; punc¬
tures large, umbilicate; anterior, posterior margins entire; lateral crenulate, with
long, thick, reflexed hairs; sides broadly, medially, posteriorly dilated; anterior
angles slightly acute; basal obtuse. Scutellum moderately punctured; punctures as
on pronotum, almost in rows; disc impunctate. Elytra more densely, coarsely
punctured than pronotum; punctures finer, shallower; sutural margins tumid,
wider posteriorly. Pygidium densely punctured with fine, shallow punctures; thick,
straight hairs of different lengths, longer posteriorly; apex truncate. Antennal club
3-segmented, about as long as four preceding segments together. Protibia triden-
tate; upper tooth small, median stouter than distal. Stemites densely pubescent,
with fine, long, straight hairs arising from small, shallow punctures. Coxal plates
with thick, long, straight hairs. Urostemite densely pubescent, but less so than
stemites; hairs shorter, thicker; last segment completely grooved, with some hairs.
Longer calcar of metatibia elongate, slender, acuminate, longer than first tarsal
segment with its spines. Claws slightly curved; median tooth stouter than distal.
Length: 18-19 mm. Figures 22, 29, 30.
Holotype.— Male, col. M. R. from Santiago de las Vegas, 10 October 1971. In
CAS No. 14289.
Paratype. —Male, col. from Los Jazmines, 9 June 1971, col. unknown. In USNM.
Phyllophaga maestrensis, New Species
Male.— Medium, shining. Upper, underparts, legs yellowish-castaneous to cas-
taneous-ferrugineous; sides almost parallel. Frons densely punctured; punctures
fine, irregularly distributed; base impunctate; median line conspicuous. Clypeus
densely punctured; punctures as on frons but more regularly distributed; margins
broadly reflexed, elevated; median indentation moderately deep, subangulate.
Pronotum less densely, coarsely punctured than frons, punctures bigger; margins
entire; sides broadly, medially dilated; anterior angles almost straight, basal ob¬
tuse; lateral margins bearing long, reflexed hairs anteriorly. Scutellum punctured
as pronotum; punctures regularly distributed. Elytra more densely, coarsely punc¬
tured than pronotum but less than frons; hairs short, straight; sutural margins
tumid; lateral, posterior margins with fine hairs. Pygidium punctured as elytra;
hairs fine, straight, mainly posterior; apex broadly curved. Antennal club with
three very long segments, an incomplete fourth and a fifth shorter; longer than
all preceding segments together. Protibia tridentate; upper tooth tiny, distant from
others. Stemites densely punctured; punctures fine with fine, long hairs. Coxal
plates sparsely punctured; punctures with short, thick hairs. Urostemites less
densely punctured with shorter hairs; last segment transversely grooved. Longer
calcar of metatibia blunt, slender, sides almost parallel, longer than first tarsal
segment. Claws slightly curved, teeth rather slender. Length: 18-19 mm. Figures
24, 27, 28.
Female.— Apex of pygidium more narrowly curved; antennal club shorter but
5-segmented also, about as long as three preceding segments together; last uro¬
stemite convex; longer calcar of metatibia compressed dorsoventrally, about one
and a half as long as first tarsal segment. Length: 18-19 mm. Figure 42.
324
PAN-PACIFIC ENTOMOLOGIST
Holotype.— Male, col. J. Ferra from “Capitolio,” Rio Yara, Cuba, 15-19 May
1948, at about 1150 feet altitude. In CAS, No. 14290.
Allotype.—C ol. J. Acuna, F. Valdes and C. Fortum from Rio Yara, Oriente,
15-20 May 1948, at about 1000 feet altitude. In CAS.
Paratypes .—One male, same data as holotype; one female, from Loma de Cala
to P. Mocha, Sierra Maestra, Cuba, 16 May 1948, at 3600-3900 feet altitude,
collector unknown. Both in USNM.
Phyllophaga migratoria, New Species
Male. — Submedium, pruinose. Elytra, underparts, legs rufocastaneous to cas-
taneous; pronotum dark castaneous; clypeus dark brown. Clypeus densely punc¬
tured; punctures rather large, irregular; margins abruptly reflexed; median inden¬
tation deep, angulate. Frons less densely punctured; punctures as on clypeus but
more irregularly distributed; depression on disc and on median line; base im-
punctate. Pronotum less densely, coarsely punctured, punctures similar but shal¬
lower, regularly distributed; margins entire; borders rufocastaneous, disc dark
castaneous; sides broadly, medially dilated; anterior angles slightly acute, posterior
obtuse. Elytra more densely punctured than pronotum, but with finer punctures,
more regularly distributed; sutural margin wider posteriorly, moderately elevated.
Scutellum triangular, sparsely punctured; disc impunctate. Pygidium punctured
as elytra, but more densely; short, straight hairs arising from punctures; apex
subtruncate. Antennal club 3-segmented, longer than funicle. Protibia tridentate;
upper tooth distant, median stouter than apical. Coxal plates sparsely pubescent;
hairs thick, straight. Stemites finely densely punctured; hairs fine, long, straight.
Urosternites less densely punctured than stemites with similar punctures; last
segment transversely, posteriorly grooved. Longer calcar of metatibia elongate,
slender. Claws slightly curved; median slightly stouter than distal. Length: 11.5-
12 mm. Figures 25, 26.
Holotype.— Male, col. C. and P. Vaurie, from San Vicente, Pinar del Rio, 25-
28 July 1956. In CAS, No. 14291.
Paratype.— Male, collected from Vinales, Pinar del Rio, June 1964, at light,
collector unknown.
The paratype has lost it pruinosity, probably during the extraction of the gen¬
italia.
Phyllophaga spinicola, New Species
Male.— Large, shining. Head, protibiae castaneous to piceous, body rufotes¬
taceous. Clypeus densely, coarsely punctured, especially toward median line; mar¬
gins broadly, slightly reflexed, almost flat; median indentation deep, angulate;
clypeo-frontal suture conspicuous. Frons punctured as clypeus, punctures small,
base almost impunctate. Pronotum slightly less densely, coarsely punctured than
head, but with larger punctures; anterior, posterior margins entire, lateral cren-
ulate; anterior angles acute, posterior obtuse; sides medially, broadly dilated, with
long, thick, reflexed hairs. Scutellum punctured as pronotum, but punctures small¬
er; disc impunctate. Elytra sparsely punctured; punctures smaller than on head;
sutural margins conspicuous, not tumid. Pygidium punctured as elytra but slightly
more coarsely; apex narrowly rounded. Antennal club 3-segmented, about as long
as the four preceding segments. Protibia tridentate, upper tooth apart but well
VOLUME 60, NUMBER 4
325
developed. Coxae very sparsely punctured, with thick, straight hairs. Stemites
densely pubescent with long, straight hairs. Urostemites very sparsely punctured,
punctures fine with few hairs; last segment shallowly, transversely grooved. Longer
calcar of metatibia longer than first tarsal segment with its spines, blunt. Claws
curved; distal tooth longer than median, which is fine. Length: 24-26 mm. Figures
31,40,41.
Female .—Antennal club about as long as three preceding segments; apex of
pygidium narrower; longer calcar of metatibia about twice as long as first tarsal
segment, compressed, wider than in male, acuminate; last urostemite not grooved,
convex. Length: 22-26 mm.
Holotype. — Male, from Gran Piedra, Oriente, July 1953, col. Zayas and Alayo.
Allotype.— Col. Zayas-Alayo, from Gran Piedra, Oriente, June 1954. Both in
CAS, No. 14285.
Paratypes .—Three males, col. Zayas-Alayo, from Gran Piedra, Oriente, July
1953; one in CAS, one in USNM, one in Illinois State Natural History Survey
Division. Three females, col. Zayas-Alayo, from Gran Piedra, Oriente, June 1954;
in CAS, USNM and ISNHSD. Two females from Gran Piedra, nr. Santiago, Prov.
Oriente, 30-31 May 1959, Col. M. W. Sanderson; in ISNHSD.
This species is very characteristic by the punctures on pronotum.
Literature Cited
Garcia-Vidal, M. 1975. Compilation sobre los principals aspectos relativos al genero Phyllophaga
Harris. Revista Agrotec. Cuba, 7(1-2): 17-22.
-. 1978. El genero Phyllophaga Harris, 1826 (Coleoptera: Scarabaeidae) en Cuba. I. Description
de cinco nuevas especies. Poeyana, 182:1-14.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 326-331
Reproductive Ecology of Three Cactophilic Diptera 1
(Diptera: Drosophilidae, Neridae, Syrphidae)
Robert L. Mangan
Department of Entomology, The Pennsylvania State University, and U.S. Re¬
gional Pasture Research Laboratory, University Park, Pennsylvania 16802.
If habitat is really the template for ecological strategies (Southwood, 1977), one
should be able to predict reproductive and developmental characteristics of a
given species from a knowledge of its habitat. Means for translating habitat char¬
acteristics into life history strategies have been proposed and are reviewed in
Stearns (1976). Of numerous proposals, two general groups of theories are useful
in explaining variations in life history characteristics. The first, derived from
arguments of MacArthur and Wilson (1967), proposes that selection may favor
either rapid population growth at the expense of competitive efficiency (r selection)
or competitive ability and efficiency at the expense of population growth rate (K
selection). A second proposal, the so-called “trade-off’ theory, was developed
largely by Schaffer (1974) from more general treatments of Murphy (1968). This
theory proposes that habitat factors, which determine survival predictability at
different stages in life history, in turn select levels of parental effort expended in
immediate versus future reproduction. A number of theoretical studies have at¬
tempted to show how predictability and magnitude of environmental fluctuations
can result in varying degrees of r or K selection or levels of reproductive effort
(Schaffer and Rosenzweig, 1977; Felsenstein, 1979; Boyce, 1979; Livdahl, 1979).
Any attempt to correlate theory and observation requires habitat interpretation.
Here I examine life history characteristics of dominant dipterans using the same
tissues of a single plant species, the major purpose of this study being to examine
habitat characteristics in order to explain certain life history strategies. Secondly,
I examine levels of oviposition effort with respect to predictions of trade-off models
relating immediate reproductive effort to adult self-maintenance for future re¬
productive effort.
Ecology
In this paper three nonrelated dipterans which feed largely or exclusively on
decaying stem tissues of saguaro ( Carnegea gigantia ) cacti in the Sonoran desert
of Arizona and Sonora, Mexico, are compared. The species studied are Drosophila
nigrospiracula Patterson and Wheeler (Drosophilidae), Odontoloxozus longicornis
Coquillett (Neriidae), and Volucella isabellina Williston (Syrphidae). Drosophila
nigrospiracula is restricted to the Sonoran desert and Baja, California; larvae in
the upper Sonoran desert use stem tissues of saguaro cactus with rare incursions
into similar tissues of other columnar cactus species. Larvae of O. longicornis use
several columnar catci in the Sonoran desert; the species is distributed from
1 Contribution No. 8109 of the U.S. Regional Pasture Research Laboratory, University Park, PA
16802; and authorized as a Paper in the Journal Series of the Pennsylvania Agricultural Experiment
Station.
VOLUME 60, NUMBER 4
327
Arizona and California south to Costa Rica. Larvae probably inhabit prickly pear
(Opuntia ) cacti extensively at higher elevations. Volucella isabellina is apparently
restricted to Sonoran desert areas where saguaro cacti are distributed. I have not
reared this species from Opuntia, but it uses several other columnar cactus species
in Mexico. Generally, the larval habitat of these species over most of their Arizona
distribution is limited to saguaro cacti. In studies of cactophilic Drosophila at the
University of Arizona, hundreds of individuals of these three species have been
reared from naturally occurring saguaro necroses.
Adults of these species appear to be attracted by similar olfactory and perhaps
visual factors. Cacti with tissues just beginning to decay and having juice running
down the surface of the stem are most likely to attract adults. This suggests that
adult substrates are equally distributed for the three species.
It should be noted that while these dipterans are all classified in the suborder
Cyclorapha, V. isabellina is in the division Aschiza. Drosophila nigrospiracula
and O. longicornis are placed in the same division, Schizophora, and section,
Acalyptratae, but are placed in widely separated superfamilies. Similarities and
differences also appear in the adaptive potentials of the genera of these species.
The genus Odontoloxozus is restricted to cacti and at present only one species is
described. The genus Volucella includes at least 39 species in North and Central
America where larval habitats include decaying succulent plant material and nests
of social hymenoptera (Spradbery, 1973). The genus Drosophila may be one of
the most adaptable animal taxa. Worldwide, substrates include decaying tissues
of nearly all angiosperm families (Throckmorton, 1975).
Ecological and phylogenetic evidence suggests that genetic potential of the taxa
and rigorous selection regime experienced by these species should have interacted
to produce reproductive strategies relevant to larval and adult substrates and
ambient communities.
Methods
In order to compare these species on an r-K continuum, two main factors are
evaluated: egg production potential is measured by ovariole number; and mat¬
uration rate is measured by egg to adult development time. Estimation of im¬
mediate versus future reproductive efforts for comparisons among these species
is mainly by examination of a subjective degree of oviposition effort. General
levels of adult fitness are estimated by thorax size which is assumed to be related
to body volume and migratory ability. While these comparisons do not lend
themselves to rigorous quantification, they do allow comparisons which suggest
the more suitable hypotheses to explain differences in reproductive behaviors
among these species.
Observations discussed here were made on naturally occurring saguaro necroses
in Pima Co., Arizona and the region of Sonora, Mexico north and west of Her-
mosillo. Field data were collected in 1977 and 1978 for O. longicornis and 1975—
1978 for D. nigrospiracula and V. isabellina.
Data included here for D. nigrospiracula were collected for a study of compet¬
itive interactions and host plant specificity (Mangan, 1978; Mangan and Heed,
1980). Dissection of reproductive systems employed methods described by Kam-
bysellis and Heed (1971). Observations and dissections of O. longicornis were
collected for study of mating behavior (Mangan, 1979). Observations of V. isa¬
bellina were taken from notes and dissections performed to gain comparative
328
PAN-PACIFIC ENTOMOLOGIST
Table 1. Characteristics of three cactophilic Diptera.
Characteristic
D. nigrospiracula
O. longicornis
V. isabellina
Ovariole number
30-60
100-190
?
Egg number/adult
200
33-156
200-500
Egg number/ovariole
1-4
1
?
Oviposition sites
single inserted
grouped inserted
grouped on
and pattern
fresh tissue
old tissue
protrusions
outside cactus
Mating behavior
crepuscular on
substrate
diurnal on sub¬
strate
mainly crepus¬
cular in air
Adult number/cactus
10 2 -10 4
10-20
1-3
Larval medium
freshly rotted
tissue
old, dark firm
tissue
old, anaerobic
liquid tissue
Pupation
grouped on sub¬
strate surface
single, burrow in
old substrate
migratory, bur¬
row in sand
Development time
(22°C; egg to adult)
16 days
30-50 days
>50 days
Adult thorax length
(mm ± SD)
1.24 ± 0.32
3.68 ± 0.49
6.35 ± 0.38
ecological insight into species utilizing saguaro cacti in possible competition with
D. nigrospiracula.
Observations of larval feeding and development were made under both labo¬
ratory and field conditions during 1976, 1977, and 1978. Laboratory rearings
were conducted at room temperatures (19-22°C) at the University of Arizona in
1 gallon widemouth jars, Vi pint milk bottles, or 5 gallon aquaria. Development
times were determined from naturally occurring eggs in saguaro necroses; both
eggs and substrate were placed in the containers, with sand or newspaper to absorb
excess exudate.
Results
Observations and data are summarized in Table 1. The species show about as
much interspecific variation in all aspects of life history strategies as can be found
among any saprophagous diptera.
Female D. nigrospiracula are quite exacting in their oviposition requirements.
While a typical saguaro rot pocket contains thousands of larvae, laboratory stocks
can seldom be maintained without cactus tissue as an oviposition medium. Larvae
develop well on laboratory media once eggs are elicited. Females may accumulate
up to four eggs per ovariole in the laboratory if suitable media are not provided.
Eggs are always inserted singly into the medium with space between consecutive
eggs. In crowded population cages, however, thousands of eggs may be deposited
on a single gram of cactus. Larvae are only weakly motile and pupation occurs
in groups on open surfaces, usually inside the cactus or on the sides of rearing
bottles.
Oviposition characteristics of O. longicornis differ from D. nigrospiracula in
that while eggs are inserted into the medium, masses of eggs from one female are
usually inserted through a single opening in the cactus epidermis or a single spot
of laboratory medium. As described in Mangan (1979), males of this species hold
VOLUME 60, NUMBER 4
329
territories at oviposition sites and restrict the movements of females until eggs
are deposited. This behavior may increase the tendency of females to group eggs.
Females, however, will oviposit on nearly any medium of soft, wet texture, though
larval dietary requirements are quite restrictive and adult emergence from pupae
reared in Drosophila medium is only about 20%. Larvae are quite motile, bur¬
rowing rapidly into the tissue when disturbed. Pupation occurs in the tissue mass;
pupae have not been found exposed on the medium surface.
Female V. isabellina spend little time around rotting cacti except for oviposition.
Eggs are commonly found attached to cactus spines or suspended in spider webs
on the outside of decaying cacti. Female V. isabellina were difficult to capture or
rear to reproductive maturity so fecundity estimates were made from numbers
of eggs per egg mass. These masses are made up of groups of eggs which adhere
to each other and surfaces to which they are attached. Eclosing larvae apparently
drop to the ground then crawl up the cactus and burrow into the decaying tissues.
Larvae feed in pockets of decaying, mucoid necrotic tissue which emit strong odor
of anaerobic sulfur decay products. Larvae leave the cactus at the end of the third
stadium and pupate in the soil.
Discussion
These data and observations indicate that habitat characteristics described in
terms of host plant tissue, decaying saguaro stem cortex, and host plant com¬
munity, upper Sonoran desert, provide little predictive information relative to
reproductive ecology and behavior. Mating behavior, oviposition behavior, larval
development time, and pupation behavior are all shown to vary considerably
among these species.
In attempting to explain such differences in traits among these species, two
general classes of adaptations may be distinguished. Adaptations may be plesio-
morphic traits which were retained during desert adaptation or they may be in
situ apomorphic responses. Migratory ability, generation time, and perhaps pre¬
ferred larval medium are probably ancestral traits. Other characteristics such as
oviposition site preference, pupation site, and mating behavior are more likely
adaptations to the saguaro substrate and evolved in the desert.
The greatest differences in habitats which would affect reproductive behavior
are larval microhabitats. The short developmental period for D. nigrospiracula
coincides with the microflora-rich larval medium (Starmer et al., 1976) and rel¬
atively small adult size. The larger species, V. isabellina and O. longicornis, use
nutrient depleted, older tissue which, along with their larger adult size, incurs a
two to three times longer developmental period. The adults, however, are probably
more resistant to environmental fluctuations than D. nigrospiracula owing to their
large size.
The use of sparsely distributed columnar cactus tissues over the large area of
the upper Sonoran desert, by all three species, suggests that if substrate distribution
is a selective factor in reproductive adaptation, differing migratory abilities of
these species result in different predictabilities of adult migratory success. In terms
of future versus immediate reproduction, the smaller D. nigrospiracula adults are
more susceptible to environmental fluctuations in such factors as necrosis density
than the larger species; this may affect future reproductive success.
The daily probability of larval substrate degeneration, mainly through des¬
iccation, is identical for each species. Probability of habitat degeneration during
330
PAN-PACIFIC ENTOMOLOGIST
the larval development period increases with larval development time. Since
larvae cannot migrate to new cacti, the slower developing species face greater
uncertainty in maturation than D. nigrospiracula. Assuming that female flies are
unable to predict substrate degeneration, large investments in finding suitable
larval substrates are not as likely to increase fitness for the larger species as for
D. nigrospiracula.
Interaction of migratory success and larval development success probabilities
are hypothesized to explain differing oviposition efforts among these species.
Quick developing Drosophila with environmentally susceptible adults invest greater
effort in oviposition site choice than the slower developing, but environmentally
buffered and larger O. longicornis and V. isabellina. This situation approximates
theoretical conditions suggested by Schaffer (1974) and Stearns (1976) for estab¬
lishment of differing optimal reproductive strategies through immediate versus
future reproduction trade-offs.
Acknowledgments
I thank Chris M. van Dyck, William C. Templeton, Jr., Charles W. Pitts, and
Jeremy R. Montague for reviewing earlier versions of this manuscript. Theoretical
treatments presented are my interpretations of ideas offered by William B. Heed,
William Schaffer, and others in informal seminars and discussions.
Literature Cited
Boyce, M. S. 1979. Seasonality and patterns of neutral selection for life histories. Am. Nat., 114:
569-583.
Felsenstein, J. 1979. r- and K-selection in a completely chaotic population model. Am. Nat., 113:
499-510.
Kambysellis, M. P., and W. B. Heed. 1971. Studies of oogenesis in natural populations of Droso-
philidae. I. Relation of ovarian development and ecological habitats of the Hawaiian species.
Am. Nat., 185:265-278.
Livdahl, T. P. 1979. Environmental uncertainty and selection for life cycle delays in opportunistic
species. Am. Nat., 113:835-842.
MacArthur, R. H., and E. O. Wilson. 1967. The theory of island biogeography. Princeton Univ.
Press, Princeton, N.J.
Mangan, R. L. 1978. Competitive interactions among host plant specific Drosophila species. Ph.D.
thesis, Univ. of Arizona, 204 pp.
-. 1979. Reproductive behavior of the cactus fly, Odontoloxozus longicornis, male territoriality
and female guarding as adaptive strategies. Behav. Ecol. Sociobiol., 4:265-278.
-, and W. B. Heed. 1980. Community ecology of Sonoran Desert Drosophila. In M. Ashbumer,
H. L. Carson, and J. N. Thompson (eds.), Genetics and biology of Drosophila. Vol. 3. Academic
Press, London (in press).
Murphy, G. I. 1968. Pattern in life history and environment. Am. Nat., 102:390-404.
Schaffer, W. 1974. Optimal reproductive effort in fluctuating environments. Am. Nat., 108:783—
790.
-, and M. L. Rosenzweig. 1977. Selection for optimal life histories. II. Multiple equilibria and
the evolution of alternate reproductive strategies. Ecology, 58:60-72.
Southwood, T. R. E. 1977. Habitat, the templet for ecological strategies. J. Anim. Ecol., 40:337-
365.
Spradbery, J. P. 1973. Wasps. Univ. of Wash. Press, Seattle, 408 pp.
Starmer, W. T., W. B. Heed, M. Miranda, M. W. Miller, and H. J. Phaff. 1976. The ecology of yeast
flora associated with cactophilic Drosophila and their host plants in the Sonoran desert. Microb.
Ecol., 3:11-30.
Steams, S. 1976. Life history strategies: A review of ideas. Q. Rev. Biol., 51:3-47.
VOLUME 60, NUMBER 4
331
Throckmorton, L. H. 1975. The phytogeny, ecology, and geography of Drosophila. In R. C. King
(ed.), Handbook of genetics. 3:421-469. Plenum, New York, 874 pp.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 332-336
Descriptions of the Life Stages of Pyrausta orphisalis
(Lepidoptera: Pyralidae) 1
C. L. Campbell and K. S. Pike
Department of Entomology, Washington State University, Irrigated Agriculture
Research and Extension Center, Prosser, Washington 99350.
Abstract. — The life stages of Pyrausta orphisalis Walker (Lepidoptera: Pyralidae)
are described, and the immature stages illustrated.
Pyrausta orphisalis Walker (Lepidoptera: Pyralidae) is a small, orange and brown,
diurnal moth, which according to Munroe (1976), occurs across Canada’s southern
tier and along both coasts of the United States. In Washington, this species feeds
on all commercial types of mint {Mentha spp.). The adult is compared briefly to
two closely related species by Munroe (1976), but aside from this and the very
short original description (Walker, 1859), no descriptions exist for any of the life
stages. This paper describes all life stages of P. orphisalis, utilizing specimens
collected (and also deposited) at the Washington State University, Irrig. Agric.
Res. and Ext. Center, Prosser, Washington.
Egg (Fig. la).—The egg is creamy white, ca. 0.4 mm wide and 0.65 mm long,
oval-shaped with a flattened venter, appearing as an elongated dome when viewed
laterally. A distinct irregular sculpting is visible under scanning electron micros¬
copy, which gives the egg an appearance of being encased by a net.
Larvae. — Chaetotaxy utilizes the terminology of Fracker (1915).
First instar (Fig. lb). —Body length 1.2 to 2.5 mm. Head capsule width 0.21 to
0.23 mm. Head colored uniformly brown. Six stemmata arranged in a near semi¬
circle, lenses usually appearing to surround a central collection of pigment cells.
Antennae pale. Clypeus pale, labrum slightly reddish brown. Apices of mandibles
reddish brown. Maxillae pale to lightly pigmented. Labium pale except for dark¬
ened anterior margin of mentum. Anterior corner of hypostomal lobe black. Head
capsule with a small notch located at posterior-lateral margin and marked by a
black patch; posterior margin of head capsule trimmed in black from black patch
to sub-vertex. Prothoracic shield somewhat rectangular, lighter brown than head
capsule, and bearing setae. Remaining thorax and abdomen pale, except for spot
pigmentation at base of setae. Setae simple, primary, and prominent. Spiracles
circular, pale, and brown rimmed, present on prothorax and abdominal segments
1 to 8. Suranal plate concolorous with prothoracic shield and bearing setae. Tho¬
racic legs normally developed, bearing one pretarsal claw. Prolegs of abdominal
segments 3 to 6 subequal and normally developed; pygopods appearing slightly
larger. Crochets uniserial, triordinal, and arranged in a mesal penellipse, remaining
this way throughout all instars.
1 Scientific Paper No. SP 6298, Washington State University, College of Agriculture Research Center,
Pullman. Project 4412. This research was made possible in part by funds provided by the Washington
Mint Commission and the Mint Industry Research Council.
VOLUME 60, NUMBER 4
333
Figure 1. Immature stages of Pyrausta orphisalis Walker, a. egg; b. first instar; c. second instar; d.
third instar; e. fourth instar; f. fifth instar; g. pupa. (Scale line with reference to egg = 0.2 mm; larvae
and pupa = 1.0 mm.)
Second instar (Fig. lc). —Body length 2.0 to 3.5 mm. Head capsule width 0.32.
to 0.38 mm. Head beige to light brown and mottled. Six stemmata arranged in
a near semi-circle, lenses of stemmata 1 to 5 overlaying obvious pigment cells.
Antennae pale. Clypeus and labrum beige to light brown. Apices of mandibles
reddish brown. Maxillae pale to light beige. Labium pale, except for darkened
anterior margin of mentum. Anterior corner of hypostomal lobe black. Head
capsule with a small, inconspicuous notch located at posterior-lateral margin and
usually marked by a black patch, posterior margin of head capsule trimmed in
black for a varying distance between black patch and vertex; these markings
remaining unchanged through remaining instars. Prothoracic shield slightly lighter
than head capsule, mottled and bearing setae. Remaining thorax and abdomen
pale, except for spot pigmentation at base of setae. Setae simple, primary and
subprimary, and prominent; setae remaining this way through subsequent instars.
Spiracles same as first instar. Suranal plate pale to lightly pigmented and bearing
setae. Thoracic legs and prolegs normally developed.
Third instar (Fig. Id).—Body length 4.4 to 6.8 mm. Head capsule width 0.48
334
PAN-PACIFIC ENTOMOLOGIST
to 0.57 mm. Head beige to light brown and mottled. Stemmata unchanged. An¬
tennae beige to light brown. Clypeus and labrum concolorous with head capsule.
Apices of mandibles reddish brown. Maxillae pale to beige. Labium pale to beige
except for darkened anterior margin of mentum. Anterior corner of hypostomal
lobe black. Head capsule, appendages, and markings remaining fairly constant
through remaining instars. Thorax and abdomen beige to light yellow-green. Pro-
thoracic shield pale to beige and bearing setae. A dorsal stripe present, appearing
beige to light green if ingesta present. An irregular faint subdorsal stripe, reaching
nearly to alpha seta, may be present but is never prominent. Pinnacula bearing
setal group rho of meso- and metathorax large, black and prominent. Remaining
setae usually with dark ringed papillae. A small black spot located anterior to
alpha setae of abdominal segments 2 to 8. Spiracles circular, pale, and brown or
darker ringed; location same as earlier instars. Suranal plate pale to lightly pig¬
mented and bearing setae.
Fourth instar (Fig. le). —Body length 5.6 to 10.7 mm. Head capsule width 0.75
to 0.83 mm. Head beige to light brown and mottled. Stemmata, pigmentation of
appendages, spiracles, and various cephalic markings remaining the same as in
third instar. Thorax and abdomen beige to green. Prothoracic shield pale to beige
and bearing setae. A green dorsal stripe present, flanked on each side by a faint
whitish subdorsal stripe nearly reaching thoracic beta and abdominal alpha setae.
A second subdorsal stripe, whitish but fainter, located ventrolaterally to alpha
and beta setae; extending from the mesothorax to abdominal segment 9. A faint
whitish subspiracular stripe extends from the prothoracic kappa group to abdom¬
inal segment 9, encompassing the kappa group of the meso- and metathorax, and
reaching to the abdominal kappa groups. Setal group rho of meso- and metathorax
borne on a large, black pinnaculum. Alpha and beta setae from mesothorax to
abdominal segment 8 borne on slightly smaller black pinnacula. Remaining setae
with papillae dark ringed. A small black spot located anterior to each alpha seta
of abdominal segments 2 to 8. Suranal plate pale to beige, may be slightly mottled,
bearing setae.
Fifth instar (Fig. If).—Body length 8.8 to 15 mm. Head capsule width 0.98 to
1.2 mm. Head beige to light brown and mottled. Stemmata, pigmentation of
appendages, and various cephalic markings as in previous instar. Thorax and
abdomen beige to green. Prothoracic shield pale to beige and bearing setae. A
prominent green dorsal stripe visible in the living specimen; an irregular dark
dorsal stripe, inconspicuous during life, visible in preserved specimens. Dorsal
stripe flanked on each side by two separate whitish subdorsal stripes as in previous
instar. A distinct greenish black supraspiracular stripe extends from the meso¬
thorax to abdominal segment 9, encompassing the pinnacula bearing rho setae.
A faint whitish subspiracular stripe, as seen in previous instar, is also present.
Intersegmental grooves often yellow. Setae and setal groups borne upon promi¬
nent, black pinnacula as in previous instar. Remaining setae with darkly ringed
papillae. Alpha setae with a dark spot located anteriorally. Spiracles circular,
largely pale, and black rimmed. Suranal plate pale to beige, may be slightly mot¬
tled; bearing setae. Near the end of the ambulatory stage of this instar, after feeding
has ceased, the trunk loses its green color and becomes dark beige to light orangish
brown except for striped areas. Dorsal stripe becomes light to medium brown,
subdorsal stripes become more beige, supraspiracular stripe appears pinkish to
VOLUME 60, NUMBER 4
335
orangish brown with areas of darker brown interspersed, and subspiracular stripe
appears light beige. Venter remains a beige color as does prothorax. Pigmentation
tends to lighten to a cream color prior to pupation with only the dorsal and
supraspiracular stripes being prominently visible. Overwintering, full grown larvae
show similar pigmentation.
Larvae which pass through six larval instars instead of the typical five have
their first three instars no different than the “typical.” The fourth and fifth instars
appear essentially the same as a typical fourth instar, except that their head
capsules and body lengths range differently from the typical. Head capsule width
for the fourth instar ranges from 0.65 to 0.71 mm; body length, 4.4 to 7.6 mm.
Head capsule width for the fifth instar range from 0.85 to 1.0 mm; body length,
7.2 to 10 mm. The sixth instar has a head capsule, body length, and distinctive
supraspiracular stripe comparable to the typical fifth instar.
Pupa (Fig. lg).—Obtect, length ca. 7.0 to 8.5 mm. Impunctate, shiny, golden
brown color, darkening prior to eclosion. Vertex has assumed a dorsal position,
causing frons to be mostly cephalic; this arrangement contributes to the “shoul¬
dered” appearance reported for Pyraustinae pupae (Mosher, 1916). Epistomal
suture and anterior tentorial pits well defined. Labrum broadly triangular, bor¬
dered caudo-laterally by large pilifers which are separated medially by narrow,
chisel-shaped labial palpi. Eye-pieces large, covering easily the visible compound
eye. Galea of maxillae large and prominent, extending slightly past apex of fore¬
wing. Galea bordered laterally and subproximally by spindle-shaped sclerite rep¬
resenting prothoracic femur; this in turn is bordered laterally by prothoracic tibia
and tarsi. Tibia and tarsi extend slightly past midpoint of galea. Prothoracic leg
bordered laterally by mesothoracic leg which extends nearly to distal end of galea.
Antenna borders mesothoracic leg and extends nearly to apex of forewing. At the
junction of antenna, eye-piece, and bases of tibiae is a small subrectangular sclerite
which represents the maxillary palpus.
Pronotum distinct, subrectangular, bounded laterally by antennae. Mesonotum
large; sclerite continuous, covers forewings. Metanotum smaller, hindwings largely
covered by forewings. Abdomen with 8 demarcated segments. Functional spiracles
present on segments 2 to 7, segment 8 with a non-functional spiracle. Male with
visible phallomeres on venter of ninth segment, female with visible vulva on
venter of eighth segment. Cremaster dark brown, distinct, possessing 8 hooked
setae.
Adult.— Adult male and female are quite similar in appearance. [See Munroe
(1976) for colored photographs of male and females.] Wing expanse of male
approximately 14 to 15 mm, female approximately 15 to 16 mm. The following
descriptions give liknesses and differences between sexes, except for genitalia,
which are outlined by Munroe (1976).
Male. — Labial palpi porrect and prominent, cream colored scaling basally, a
mixture of rust-orange and fuscous scaling distally. Proboscis well developed with
cream colored scaling basally. Maxillary palpi filiform, a mixture of rust-orange
and fuscous scaling. Frons rounded, anterior and lateral margins rust-orange,
center fuscous; scaling in front of and between antennae primarily long, thin, rust-
orange scales and some fuscous. Antennae long and filiform, scape and pedicel
moderately scaled rust-orange and fuscous, flagellum with small brownish scales
dorsally. Compound eye amber, sometimes appearing speckled. Ocelli dark amber
336
PAN-PACIFIC ENTOMOLOGIST
and fairly prominent. Vertex with a mixture of long, thin rust-orange and fuscous
scaling; rim of occiput mostly with erect, rust-orange scales.
Pronotum with a mixture of rust-orange and fuscous scales. Mesonotum and
metanotum primarily fuscous with some rust-orange scaling laterally. Partagia
covered with a combination of long, thin, rust-orange scales, long paddle-shaped
rust-orange scales, and long, paddle-shaped fuscous scales. Forewings subtrian-
gular, dorsal surface primarily a combination of rust and fuscous scales, and
usually with at least two distinct patches of orange—one antemedially and the
other postmedially, from the costal margin extending posteriorally about one third
the wing’s width. Termen heavily fringed, inner margin less so. Basal two thirds
of ventral surface mostly orange with two fuscous patches, except for anal region
which is cream colored. Distal third mostly fuscous. Hindwings subtriangular,
but more rounded. Dorsal surface primarily fuscous, but with a median orange
patch in the basal third and a wide, orange postmedial band which extends across
wing, parallel to termen. Few rust colored scales present. Termen and inner margin
fringed with fuscous scales. Basal two thirds of ventral surface orange, but with
a poorly defined, fuscous antemedial band present. Distal third of wing fuscous.
Frenulum present as a strong, orange colored spine. Venter of thorax and all legs
covered by cream to light buff colored scales. Legs normally developed.
Dorsum of abdomen primarily fuscous, but mixed with rust and orange scales.
Venter mostly cream to light buff, with increasing fuscous scaling posteriorally.
First abdominal sternum with a pair of tympana. Terminal visible segment com¬
pressed and appearing somewhat oblique in profile.
Female .—Female is nearly identical to the male except dorsal surface of fore¬
wing is slightly more variegated with more orange patches which may nearly form
a postmedial band; ventral aspect may have more rust-colored scales along the
termen. Dorsal surface of hindwing appears to have slightly more rust scales.
Ventral aspect as in male, but may have fuscous antemedial band more defined
and may also have more rusty scales along termen and inner margins. Frenulum,
a single spine in male, is a bifurcate spine in female. Visible terminal segment of
abdomen somewhat truncated and slightly compressed. Tip of antovipositor, with
its fine setae, often visible.
Literature Cited
Fracker, S. B. 1915 (revised in 1930). The classification of Lepidopterous larvae. Univ. of Ill., Biol.
Mono., Vol. 2, No. 1.
Mosher, E. 1916. A classification of Lepidoptera based on characters of the pupa. Illinois Natur.
Hist. Survey Bull., 12:17-85.
Munroe E. 1976. The moths of America north of Mexico. Pp. 119-120, pi. 5 in R. B. Dominick et
al., Fasc. 13-2B, Pyraloidea (in part). The Curwen Press, London.
Walker, F. 1859. List of specimens of Lepidopterous insects in the collection of the British Museum.
Part 17:310, London.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 337-340
Fabiola quinqueferella : An Obscure California Moth Formerly in
Glyphipterigidae (Lepidoptera: Oecophoridae)
John B. Heppner
Department of Entomology, Smithsonian Institution, Washington, D.C. 20560.
Abstract.—Fabiola quinqueferella (Walsingham) is illustrated to demonstrate
its affinity to other Oecophoridae and to Fabiola species.
Walsingham (1881) described a small colorful moth from northern California
that he collected during his expedition of 1871. Superficially it resembles other
moths of the genus Glyphipterix from California, now properly in Glyphipteri¬
gidae, so he described the new species as Glyphipteryx [sic] quinqueferella Wal¬
singham. While studying the North American fauna of Glyphipterigidae this
species was quickly noted to belong to the Oecophoridae (Heppner, 1978), most
noticeably by the scaled haustellum (Glyphipterigidae all have a naked haustellum)
and by male genitalia resembling other small oecophorids.
The moth appears best placed in the genus Fabiola (Hodges, pers. comm.) and
has been transferred there in the new North American checklist (Hodges et ah,
1983). This paper has been prepared to illustrate the species, especially the gen¬
italia, and describe characters not noted by Walsingham.
Fabiola quinqueferella (Walsingham)
Glyphipteryx [sic] quinqueferella Walsingham, 1881:322.
Size. — 6.0—7.5 mm base to apex forewing length.
Head.— Frons and vertex buff; labial palpus buff, 2nd segment nearly twice
length of apical segment; antenna without pecten, x h forewing length, buff, with
ventral sensory setae in males twice length of antennal segments.
Thorax. — Buff; patagia buff; venter shining light buff; legs buff with some white
on apices of each tarsal segment.
Fore wing. — Elongated with rounded apex; R2 from end of cell; buff-gray ground
color, with white crosswing fascia angled at basal l h, another to midwing at Vi
wing length, angled toward basal fascia; wing base with yellow line along cubital
vein, reaching first white fascia, with orange along radius and anal veins; orange
between each white fascia and between midwing fascia and apical wing patch;
apical x h of wing with pale gray to whitish area with black and intense white scales
mixed in, with white prominent between 4 black spots along tornal margin; each
black spot having a small distal silvery spot; fringe around apical x h of wing very
long, dark gray.
Hindwing. —Uniformly gray, with darker gray fringe.
Abdomen. —Buff
Male genitalia.— Uncus pointed, narrow; gnathos unspined, subtending tuba
analis, hooklike, pointed; tegumen as wide as valva at base, narrowing to width
338
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Fabiola quinqueferella (Walsingham), male, California.
of uncus; valva with rounded apex; sacculus coming to an upcurved free point,
with broadened base; vinculum rounded, no saccus; juxta with a pair of dorsal
elongated appendages and a pair of small pointed basal lobes; aedeagus curved,
with small phallobase; vesica without cornuti but with small spines.
Female genitalia.— Unknown.
Types. — Lectotype S (designated by Heppner, 1978). California: Hatchet Creek,
Shasta Co., 14-17 Jul 1871, Walsingham (92020), (BMNH). Paralectotypes (7 <$).
California: Hatchet Creek, Shasta Co., 14-17 Jul 1871 (3 8), Walsingham (92021-
92023), (BMNH); Burney Falls, Shasta Co., 18-20 Jul 1871 (2 8), Walsingham
(92024, 92025), (BMNH); Bear Creek, Shasta Co., 27-28 Jul 1871 (1 8), Wal¬
singham (92026), (BMNH); “Cal,” (1 8), Walsingham, [no date], (USNM). Another
putative syntype is without locality data but apparently collected by Walsingham
(MCZ, ex Chambers Coll. [Labelled paralectotype in 1978]). Walsingham (1881)
noted only 8 syntypes in his original description.
Additional specimens.— California: Colusa Co.: “24” [Cache Cr., 24-25 Jun
1871] (1 8), Stainton Coll. 401445 (BMNH). Mariposa Co.: Yosemite Natl. Park,
30 Jun 1941 (1 8), R. M. Bohart (LACM), and 28 Jul 1915 (2 8), (ANSP). Tulare
Co.: Monache Meadows, 8-14 Jul (1 8), (USNM). Oregon: Klamath Co.: “62”
[near Summit Lk., 26 Sep 1871] (1 <$), Walsingham (BMNH). [Walsingham field
numbered specimens dated with Essig (1941) paper.]
Distribution.— Western Oregon to the southern Sierra Nevada of California.
Host. —Unknown.
Remarks.—Fabiola quinqueferella appears to be a primitive member of the
Fabiola group of species on the basis of characters such as the relatively simple
VOLUME 60, NUMBER 4
339
Figures 2-4. Fabiola quinqueferella (Walsingham). 2. Male genitalia (lectotype, California; BMNH
slide 20230). 3. Aedeagus (lectotype). 4. Male genitalia (paralectotype, California; USNM slide 77105)
[reduced].
340
PAN-PACIFIC ENTOMOLOGIST
genitalia and the length of setae on the antennae (similar to Fabiola shaleriella
Chambers). The species may require a new genus but it is preferable at this time
to defer any such change until the female is discovered.
The key to species of Fabiola in Hodges (1974) may be amended as follows to
include F. quinqueferella :
1. Apical l h of forewing white with various markings . quinqueferella
Apical x h of forewing mostly black and orange, plus various markings, with
some white, but not with most of area white . 2
2. [remainder of key as in Hodges (1974) but with couplet numbers advanced
by one].
Acknowledgments
Specimens were made available from the following collections: Academy of
Natural Sciences, Philadelphia, Pennsylvania (ANSP); British Museum (Natural
History), London, England (BMNH); Los Angeles County Museum of Natural
History, Los Angeles, California (LACM); Museum of Comparative Zoology,
Harvard University, Cambridge, Massachusetts (MCZ); and Smithsonian Insti¬
tution, Washington, D.C. (USNM).
Literature Cited
Essig, E. O. 1941. Itinerary of Lord Walsingham in California and Oregon, 1871-1872. Pan-Pac.
Ent., 17:97-113; 1 pi.
Heppner, J. B. 1978. Transfers of some Nearctic genera and species of Glyphipterigidae (auctorum)
to Oecophoridae, Copromorphidae, Plutellidae, and Tortricidae. Pan-Pac. Ent., 54:48-55.
Hodges, R. W. 1974. Gelechioidea Oecophoridae. In R. B. Dominick et al., The moths of America
north of Mexico. Fasc. 6.2. Classey, London. 142 pp.; 7 col. pi., 1 monoch. pi.
-, et al. 1983. Checklist of the Lepidoptera of America north of Mexico. Classey, London. 284
pp.
Walsingham, T. de Grey. 1881. On some North American Tineidae. Proc. Zool. Soc. Lond., 1881:
301-325; pi. 35-36.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 341-344
Two New Species of Idiocerine Leafhoppers from
Malagasy Republic (Homoptera: Cicadellidae ) 1
Paul H. Freytag and Paul S. Cwikla
Department of Entomology, University of Kentucky, Lexington, 40546-0091.
Present address: (PSC) Department of Entomology, Ohio State University, Co¬
lumbus, Ohio 43210.
Abstract. — Two new species of idiocerine leafhoppers are described from Mal¬
agasy Republic (Madagascar). These are Idiocerus rossi and Nesocerus trimacu-
latus.
Two additional species can now be added to the idiocerine leafhoppers known
from Malagasy Republic (Madagascar). Freytag and Knight (1966) reviewed the
subfamily Idiocerinae and included 17 species. All 19 species, including the two
described at this time, appear to be endemic to this island.
We thank Curtis Dunn, Academy of Natural Sciences, Philadelphia for the loan
of these specimens.
Idiocerus rossi, New Species
(Figs. 1-4)
Length. — Male 3.5 mm (approximately); female unknown.
Color.— Head yellow; ocelli dark brown. Pronotum yellow; lateral margins be¬
hind eyes with brown patches; scutellum yellow. Forewings hyline; veins brown
except costal veins yellow.
Male genitalia. — Pygofer short, posterior margin broadly concave, with a thin
pair of internal bifurcate processes inserted on anterior margin in lateral view.
Aedeagus simple, similar to sakarahensis but with shaft longer and a sharp lip¬
like projection under gonopore (best seen in lateral view). Connective as in sak¬
arahensis. Style simple, apical third, in dorsal view, curved lateral.
Holotype. — Male—Madagascar, Ankarafantsika (Forest Reserve), near Maro-
voay, XII-1-1959, E. S. Ross, Collector. Deposited in the Academy of Natural
Sciences, Philadelphia.
Etymology. — This species is named in honor of the collector, E. S. Ross, curator
at the California Academy of Science.
Diagnostic features.— This species is related to sakarahensis, but can be sepa¬
rated from it by the aedeagus having a linear shaft without projections on the
dorsal margin and the lip-like projection under the gonopore.
Notes.— This species is known only from the holotype. The thorax of the hol¬
otype has been damaged making it necessary to give an approximate measurement
of the length of this species.
1 The investigation reported in this paper (No. 82-7-199) is in connection with a project of the
Kentucky Agricultural Experiment Station and is published with approval of the Director.
342 PAN-PACIFIC ENTOMOLOGIST
Figures 1-4. Idiocerus rossi, n. sp. 1. Lateral view of pygofer. 2. Ventral view of aedeagus. 3.
Lateral view of aedeagus. 4. Latero-ventral view of style. Figures 2 and 3, to the same scale.
VOLUME 60, NUMBER 4
343
Figures 5-9. Nesocerus trimaculatus, n. sp. 5. Dorsal view of head, pronotum and scutellum. 6.
Ventral view of aedeagus. 7. Lateral view of aedeagus. 8. Lateral view of pygofer. 9. Latero-ventral
view of style. Figures 6, 7 and 9 drawn to same scale.
344
PAN-PACIFIC ENTOMOLOGIST
Nesocerus trimaculatus, New Species
(Figs. 5-9)
Structural features. — Head wider than pronotum. Forewing normal except me¬
dian vein evanescent near base.
Color. — Head light yellow with 3 brown patches along anterior margin in dorsal
view; face light yellow with 2 brown patches near anterior margin of postclypeus
in ventral view; postclypeus with a median longitudinal brown band; brown
patches circling ocelli; second antennal segment brown; postclypeus with 2 small
brown spots on disc, longitudinal medial brown stripe and 2 small brown spots
on lateral posterior margin. Scutellum light yellow with 4 longitudinal bands
starting at anterior margin and ending near disc. Lateral edges of thorax light
yellow, legs light yellow, except hindleg with brown spot on anterior-distal margin.
Forewing transparent, veins brown.
Male genitalia. — Pygofer produced posterior margin along with a few setae
scattered on apical half. Plates spatulate; apex bluntly rounded, macrosetae only
on lateral margin. Aedeagus stout, curved dorsad in lateral view with 2 pairs of
processes at apex in caudal view, processes directed dorsad. Connective broadly
Y-shaped. Style linear, apex curved dorsad in lateral view.
Holotype.— Male—Madagascar, Ankarafantsika (Forest Reserve), near Maro-
voay, XII-1-1959, E. S. Ross, Collector. Deposited in the Academy of Natural
Sciences, Philadelphia.
Etymology.— Latin, tri (three) and maculat (spotted) refers to the 3 dark brown
patches on the head in dorsal view.
Diagnostic features.— This species will key to N. duospinus in Freytag and
Knight’s key to the Madagascar Nesocerus (1966:82). It can be separated from it
by the aedeagus having 2 pairs of apical processes and the head with 3 brown
patches in dorsal view.
Literature Cited
Freytag, Paul H., and W. J. Knight. 1966. The Idiocerinae of Madagascar (Horn. Cicadellidae) Ann.
Soc. Ent. Fr. (N.S.), 2(1):75-103, 120 figs.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 345-349
Nosopsyllus fasciatus Parasitizing House Mice on
Southeast Farallon Island, California
(Siphonaptera: Ceratophyllidae)
Tom G. Schwan
Division of Entomology and Parasitology, University of California, Berkeley,
California 94720. (Present address: Yale Arbovirus Research Unit, 60 College
St., New Haven, CT 06510)
Southeast Farallon Island is a small granitic island located near the edge of the
continental shelf approximately 48 km WSW from the Golden Gate Bridge, San
Francisco, California. Presently the island is under the jurisdiction of the San
Francisco Bay National Wildlife Refuge and the 12th U.S. Coast Guard District.
The Point Reyes Bird Observatory (PRBO) maintains a research station at the
island with logistic support provided by the Coast Guard and the Oceanic Society.
The island serves as an important sanctuary and breeding ground for 12 species
of marine birds and 5 species of pinnipeds (Ainley and Lewis, 1974). Besides
humans the only other land mammals presently residing on the island are house
mice, Mus musculus Linnaeus (DeSante and Ainley, 1980). European rabbits,
Oryctolagus cuniculus Lilljebord, exterminated from the island in 1974, had been
resident beginning sometime between 1855-1876 (Pinney, 1965). Various do¬
mestic mammals including dogs, cats and mules have resided there during the
past but are no longer present (DeSante and Ainley, 1980) and hoary bats, Lasiurus
cinereus (Palisot de Beauvois), and red bats, L. borealis (Muller), occur in small
numbers for brief periods during their fall migration (Tenaza, 1966; PRBO, un-
publ.). The absence of rats, Rattus norvegicus (Berkenhout) and R. rattus (Lin¬
naeus), (DeSante and Ainley, 1980) is an interesting aspect of the island’s ecology
considering the rather intense and continuous human activity since 1849 (Dough¬
ty, 1971; Shanks and Shanks, 1978).
During 23 August to 6 September 1980 I visited the island to continue studies
on the marine bird flea, Ceratophyllus pelecani Auguston, parasitizing western
gulls, Larus occidentalis Audubon. During this period the house mouse population
was at its annual peak (R. Boekelheide, pers. comm.) and the PRBO staff biologists
were snap-trapping mice inside their house in an attempt to control the rather
high indoor mouse infestation. House mice were present on both the main island
and West End, a smaller island to the west separated by the narrow Jordan
Channel, and mice were quite active on the southern marine terrace where large
numbers of western gulls had recently nested. Because of the high mouse popu¬
lation, the dispersed gull population (gulls were no longer territorial about their
nests and most hatching year gulls were capable of flight), and mouse activity in
the gull nesting area, it appeared possible that the mice were serving as fall
alternative hosts for the bird fleas. There is one record of C. pelecani collected
from Peromyscus maniculatus elusus Nelson and Goldman, Santa Barbara Island,
Channel Islands, California (Hubbard, 1947) and this flea will readily engorge on
white mice in the laboratory (Schwan, unpubl.). Therefore, during my stay on the
346
PAN-PACIFIC ENTOMOLOGIST
island I examined house mice snap-trapped in the PRBO house and the nearby
Coast Guard (CG) house, as well as a few I captured by hand outdoors. The results
of these examinations are presented herein.
Results
During 26 August-6 September 1980 I examined 48 house mice for fleas; 20
mice from the PRBO house, 18 from the CG house, and 10 from outdoors. While
adult C. pelecani were still active in abandoned gull nests located near these houses,
I found none on these mice. However, C. pelecani is a nest flea, spending little
time on its avian host and one might not find many on mice either even though
these fleas might be feeding on them. However, 26 of the 48 mice (54.2%) were
infested with the northern rat flea, Nosopsyllus fasciatus (Bose), a flea not pre¬
viously known to occur on the island. I collected 39 N. fasciatus (27 males, 12
females) from the mice: flea index for all mice = 0.81, flea index for only infested
mice = 1.50. One male and one female were also found in a mouse nest located
under a board on the marine terrace.
The percentages of mice infested from the CG house, PRBO house and outdoors
were 72.2%, 60.0% and 10.0%, respectively, while the N. fasciatus indices were,
in the same order, 1.33, 0.70 and 0.10 fleas per mouse. These data suggest that
N. fasciatus is more abundant on indoor mice compared to outdoor mice, although
Eskey (1938) found the opposite for N. fasciatus infesting rats in San Francisco.
The CG house, which has only occasional and short term human occupancy and
no mouse control, contained mice which were more frequently infested and had
a N. fasciatus index nearly double that found on the mice from the PRBO house.
The PRBO house has continued human occupancy and mouse control and I
suggest that the lower infestation rate and flea indices for PRBO house mice are
probably due to a continual influx of relatively clean mice from outside.
Male mice outnumbered females 33 to 15. Forty-three of the 48 mice, weighed
to the nearest 0.1 g, had a mean weight of 15.1 g (range 8.1-21.1 g) and 33 of
these 43 mice (76.7%) weighed over 14 g and were therefore adult (DeFong, 1967).
The mice were in good condition: they had clean pelage and no scars, most were
molting and many had large fat deposits associated with their reproductive tracts,
and none were infested with any mites, lice or ticks.
Discussion
Prior to this study N. fasciatus was unknown on SE Farallon Island and the
presence of an established N. fasciatus population on house mice in the absence
of Rattus is interesting. For example, during November 1970 to December 1972
I trapped 253 house mice from a feral population in an annual grassland com¬
munity near Hayward, Alameda Co., California (Schwan, unpubl.). No Rattus
were present and from the house mice I collected 85 Leptopsylla segnis (Schonherr)
(32 males and 53 females), 9 Malaraeus telchinus (Rothschild), (1 male and 8
females) and 1 male Hystrichopsylla occidentalis Holland. No A. fasciatus were
found. The M. telchinus and H. occidentalis were primarily associated with sym-
patric voles, Microtus californicus (Peale) (Schwan, 1975). Finsdale and Davis
(1956) examined 77 house mice in an area free of rats in Monterey Co., California
and found no N. fasciatus. Other studies in the San Francisco Bay area have shown
occasional light infestations of N. fasciatus on house mice in areas where mice
VOLUME 60, NUMBER 4
347
and rats occur together. Miles et al. (1957) reported 9 N. fasciatus from 227 house
mice and Stark and Miles (1962) found an unreported low number on 27 mice.
In Hawaii, Haas et al. (1972) commonly found N. fasciatus on house mice living
with R. rattus and R. exulans (Peale) in a Eucalyptus plantation.
N. fasciatus is a rat flea endemic to Europe and the British Isles and has a
world-wide distribution due to its association with commensal rodents (Buxton,
1941; Jordan, 1948; Lewis, 1975) and it is an important plague vector among
rats (Pollitzer, 1954). The species is well established throughout the United States
(Benton, 1980; Prince, 1943) and was first reported from San Francisco by Fox
(1908) where it occurs on R. norvegicus during all months of the year (Eads and
Barnes, 1976). It is found throughout much of California (Prince, 1943) and has
recently been collected on San Miguel Island, Channel Islands, off the southern
California coast (2 females ex: Peromyscus maniculatus streatori Nelson and Gold¬
man, collected 25 May 1978; B. C. Nelson, pers. comm.), an island also inhabited
by roof rats. N. fasciatus occurs on the Hawaiian Islands (Haas et al., 1972) and
has been recorded from Macquarie Island in the Subantarctic (Dunnet, 1961).
Yet the species is not known to occur on R. rattus or M. musculus on the Galapagos
Islands (Smit, 1970).
The time of introduction of both house mice and N. fasciatus to SE Farallon
Island is not known. Pinney (1965) and Marshall and Nelson (1967) were the first
to report house mice even occurring on the island, but mice probably reached the
island many years before the 1960s. An early report of burrowing owls, Athene
cunicularia (Bonaparte) nesting on the island (Dawson, 1911) and reported sight¬
ings of this owl back to 1887 (Bryant, 1888) may suggest the presence of mice
before 1900. Burrowing owl pellets collected on the island during March 1980
contained skeletal remains of house mice (PRBO, unpubl. journal, 17 March
1980) but burrowing owls are known to include many prey items, including insects,
in their diet (Bent, 1938). Therefore, past nesting of these owls does not necessarily
prove the presence of mice at those times.
Regarding the introduction of N. fasciatus, this flea may have arrived on house
mice, most likely from San Francisco, where Fox (1908) reported this flea pri¬
marily from rats but also small numbers from mice. Lewis (1967) stated that “TV.
fasciatus appears to be an unspecialized species of flea capable of colonizing new,
unpopulated areas with relative ease” and “being an adaptable species, an ex¬
pansion of the population in one locality may involve the transfer to a different,
though related, host.” However, house mice generally carry few fleas and Worth
(1950), although experimenting with Xenopsylla cheopis (Rothschild), concluded
that house mice would unlikely participate significantly to the dispersal of rat
fleas. Another possible route of introduction, although remote, could have in¬
volved an infested avian predator, flying from the mainland, as discussed by
Jellison (1939) regarding the role of predatory birds in the dispersal of fleas and
plague. However, I believe the presence of N. fasciatus on the island indicates
past unsuccessful introductions of R. norvegicus or R. rattus. As mentioned earlier,
the absence of rats is interesting in spite of much human disturbance to the island,
such disturbance even being reflected by the high percentage of exotic plant species
(Coulter, 1971). One possible explanation is that rats were exterminated many
years ago during the unsuccessful poisoning campaigns to control rabbits.
In conclusion, the flea Nosopsyllus fasciatus is established as a house mouse
348
PAN-PACIFIC ENTOMOLOGIST
flea on SE Farallon Island and its presence suggests previous unsuccessful intro¬
ductions of rats to the island. N. fasciatus is the fourth species of flea now known
to occur on the island, the others being C. pelecani, a nest flea primarily of gulls
and cormorants, Actenopsylla suavis Jordan & Rothschild, a nest flea of Cassin’s
auklets, and Dasypsyllus gallinulae perpinnatus (Baker), a nest flea of passerine
birds.
Acknowledgments
I thank the San Francisco Bay National Wildlife Refuge and the Pt. Reyes Bird
Observatory for permission to work on SE Farallon Island and the Oceanic Society
for providing transportation. R. Boekelheide, T. Harvey and P. Henderson assisted
me in various aspects while working on the fleas of SE Farallon Island. This
manuscript was reviewed and improved by the following: R. Boekelheide, PRBO;
Dr. D. P. Furman, Division of Entomology and Parasitology, University of Cal¬
ifornia, Berkeley; Dr. R. E. Lewis, Department of Entomology, Iowa State Uni¬
versity, Ames; Dr. B. C. Nelson, Vector Biology and Control Branch, California
Department of Health Services, Berkeley. To all of these individuals and orga¬
nizations, I express my gratitude. This paper is Contribution No. 234 of Pt. Reyes
Bird Observatory for the Farallon Islands.
Literature Cited
Ainley, D. G., and T. J. Lewis. 1974. The history of the Farallon Island bird populations, 1854—
1974. Condor, 76:432-446.
Bent, A. C. 1938. Life histories of North American birds of prey. U.S. Natl. Bull. 170, 482 pp.
Benton, A. H. 1980. An atlas of the fleas of the eastern United States. Marginal Media, Freonia,
N.Y., 177 pp.
Bryant, W. E. 1888. Birds and eggs from the Farallon Islands. Proc. Calif. Acad. Sci., 2nd Series, 1:
25-50.
Buxton, P. A. 1941. The recorded distribution of certain fleas. Bull. Entomol. Res., 32:119-122.
Coulter, M. 1971. A flora of the Farallon Islands, California. Madrono, 21:131-137.
Dawson, W. L. 1911. Another fortnight on the Farallones. Condor, 13:171-183.
DeLong, K. T. 1967. Population ecology of feral house mice. Ecology, 48:611-634.
DeSante, D. F., and D. G. Ainley. 1980. The avifauna of the South Farallon Islands, California.
Studies in Avian Biology No. 4, 104 pp.
Doughty, R. W. 1971. San Francisco’s nineteenth century egg basket: The Farallones. Geog. Review,
61:554-572.
Dunnet, G. M. 1961. Fleas from Macquarie Island, with a description of a new species of Parapsyllus
Enderlein. Proc. Roy. Ent. Soc. London (B), 30:43-49.
Eads, R. B., and A. M. Barnes. 1976. Status of Nosopsyllus londiniensis (Rothschild, 1903) in the
United States (Siphonaptera: Ceratophyllidae). Calif. Vector Views, 23:1-3.
Eskey, C. R. 1938. Flea infestation of domestic rats in San Francisco, Calif. Pub. Health Reports,
53:948-951.
Fox, C. 1908. Identification of fleas at San Francisco, Calif. Pub. Health Reports, 23:1371.
Haas, G. E., N. Wilson, and P. Q. Tomich. 1972. Ectoparasites of the Hawaiian Islands. I. Siphon¬
aptera. Contr. Amer. Entomol. Inst., 8:1-76.
Hubbard, C. A. 1947. Fleas of Western North America. Ames, Iowa, Iowa State Coll. Press, 533
pp.
Jellison, W. L. 1939. Sylvatic plague: Studies of predatory and scavenger birds in relation to its
epidemiology. Pub. Health Reports, 54:792-798.
Jordan, K. 1948. Suctoria. Pp. 211-245 in J. Smart, A handbook for the identification of insects of
medical importance. British Museum (Natural History), London, 303 pp.
Lewis, R. E. 1967. Contributions to a taxonomic revision of the genus Nosopsyllus Jordan, 1933
(Siphonaptera: Ceratophyllidae). I. African species. J. Med. Entomol., 4:123-142.
VOLUME 60, NUMBER 4
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-. 1975. Notes on the geographic distribution and host preferences in the Order Siphonaptera.
Part 6. Ceratophyllidae. J. Med. Entomol., 11:658-676.
Linsdale, J. M., and B. S. Davis. 1956. Taxonomic appraisal and occurrence of fleas at the Hastings
Reservation in central California. Univ. Calif. Publ. Zool., 54:293-370.
Marshall, A. G., and B. C. Nelson. 1967. Bird ectoparasites from South Farallon Island, California.
J. Med. Entomol., 4:335-338.
Miles, V. I., A. R. Kinney, and H. E. Stark. 1957. Flea-host relationships of associated Rattus and
native wild rodents in the San Francisco Bay area of California, with special reference to plague.
Amer. J. Trop. Med. Hyg., 6:752-760.
Pinney, T. C. 1965. The biology of the Farallon rabbit. Unpubl. Ph.D., Stanford University, 107
pp.
Pollitzer, R. 1954. Plague. WHO Monogr. Ser. No. 22, 698 pp.
Prince, F. M. 1943. Species of fleas on rats collected in states west of the 102d meridian and their
relation to the dissemination of plague. Pub. Health Reports, 58:700-708.
Schwan, T. G. 1975. Flea reinfestation on the California meadow vole (Microtus calif or nicus). J.
Med. Entomol., 11:760.
Shanks, R. C., Jr., and J. T. Shanks. 1978. Lighthouses and lightboats on the redwood coast. Costana
Books, San Anselmo, Calif., 255 pp.
Stark, H. E., and V. I. Miles. 1962. Ecological studies of wild rodent plague in the San Francisco
Bay area of California. VI. The relative abundance of certain flea species and their host rela¬
tionships on co-existing wild and domestic rodents. Amer. J. Trop. Med. Hyg., 11:525-534.
Smit, F. G. A. M. 1970. A new species of flea from the Galapagos Islands. Entomol. Berichten,
Deel, 30:244-247.
Tenaza, R. R. 1966. Migration of hoary bats on South Farallon Island, California. J. Mamm., 47:
533-535.
Worth, C. B. 1950. House mice and commensal rats in relation to the dissemination of rat fleas.
Amer. J. Trop. Med., 30:917-920.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 350-354
Rainfall, Resources, and Dispersal in Southern
Populations of Euphydryas edit ha
(Lepidoptera: Nymphalidae)
Dennis D. Murphy and Raymond R. White
Department of Biological Sciences, Stanford University, Stanford, California
94305.
While the dispersal capacity of an insect species probably changes little from
one generation to another, its actual dispersal or vagility may vary considerably.
This is well documented for insects which have migratory generations, such as
locusts, aphids, and butterflies, but remains virtually undocumented in non-mi-
gratory species. One exception is the butterfly, Euphydryas editha. Individuals of
this species were originally shown to be extremely sedentary (Ehrlich, 1965), but
in some populations the mean dispersal distance of individuals was recently
observed to change dramatically from one generation to the next. These changes
in vagility appeared to be influenced by the availability of oviposition plants and
adult nectar resources; significantly greater vagility occurred in drier years when
those resources were sparse (White and Levin, 1981).
However, subsequent field observations of four E. editha populations in South¬
ern California indicate that the determinants of vagility are more complicated.
In 1977 these populations exploded in size and mass dispersal was observed. Here
we discuss how weather and butterfly population size interact to determine host
plant availability and butterfly population dynamics. Additionally, we consider
the biogeographic and genetic consequences of dramatic population fluctuations
in E. editha, and the mechanisms by which both the butterflies and their host
plants respond to environmental stress.
Life History
Euphydryas editha populations in San Diego County usually fly in a four to
eight week period, starting as early as mid-January and ending as late as the end
of April, depending on the timing of winter rainfall. Eggs hatch about ten days
after oviposition and about two weeks later larvae enter an obligatory fourth instar
diapause. Diapause terminates after winter rainfall when larval host plants are
again available. Postdiapause larvae then feed through another four instars, pu¬
pate, and emerge as adults to complete the life cycle. The temporal proximity of
the postdiapause larval feeding period to the oviposition and prediapause larval
feeding periods is key to understanding year-to-year changes in host plant densities
and in butterfly dispersal.
Observations
During the spring of 1977 rainfall preceding the flight of E. editha in the mesa
grasslands of San Diego County and northern Baja California was just above the
long-term average (11.48 cm versus average 11.15 cm, November-January; NOAA
VOLUME 60, NUMBER 4
351
Climatological Data). Based on the hypothesis that normal rainfall would provide
adequate plant resources, we predicted limited adult butterfly movements. Instead,
in an early and extended flight period, February through April, colonies exhibited
local population explosions and very high rates of dispersal. Several San Diego
area populations studied by our group for almost a decade (Ehrlich et al., 1975;
Singer and Ehrlich, 1979) increased nearly two orders of magnitude, to hundreds
of thousands of individuals at each of four population centers: Lower Otay (LO),
Upper Otay (UO), Brown’s Field (BF) on Otay Mesa, and La Pressa Rodriquez
(LPR) near Tijuana.
Although butterflies were extremely abundant, there was no apparent scarcity
of nectar. Flowers most commonly visited ( Eriogonum fasciculatum Bentham,
Vigueria laciniata Gray, Linanthus dianthiflorus (Bentham) Greene, and Diche-
lostemma pulchellum Heller) were available in large numbers. Thus, adult move¬
ments induced by a lack of proximate nectar sources, such as those observed by
Gilbert and Singer (1973), seemed unlikely. However, the larval host plants,
Plantago insularis Eastwood and P. hookeriana F. and M., were scarce or absent.
Nearly eight hours spent searching for these normally abundant plants resulted
in only a few dozen being found at UO and BF, and none at either LPR or LO.
On the basis of this search we estimate that the portion of the plant crop consumed
by postdiapause larvae was more than 99%, considerably higher than the maxi¬
mum of 80% previously reported (White, 1974). Defoliation of this degree results
in significant larval starvation and competition for host plant resources. None¬
theless, enough food was apparently available to allow emergence of very large
numbers of adults. The very nearly total defoliation of larval host plants produced
a scarcity of oviposition sites, and this elicited movements far in excess of any
previously recorded for either sex (White and Levin, 1981). Egg-laden females
were captured several kilometers north of LO and UO in unsuitable chaparral
habitat and similar distances west, over barley fields and other inappropriate
habitats lacking host plants. At LO the flight observed of both sexes was direc¬
tional, away from the population centers described by White and Levin (1981).
Determinants of Vagility
Given these latest observations, we can now attempt a more comprehensive
model to explain generation-to-generation changes in vagility in this species.
Figure 1 illustrates how factors acting on butterfly host plants determine adult
population dynamics. Dry winters result in adult Euphydryas editha that exhibit
increased vagility, because: 1) nectar plant quantity and quality are locally reduced,
2) fewer suitable oviposition plants are available and these senesce earlier, so
females must fly farther to oviposit, and 3) postdiapause larvae defoliate many
of these plants before adult females emerge to oviposit. When a wet year is followed
by a dry year dispersal is intensified; and, the greater the difference in rainfall
from one year to the next, the greater the change in dispersal. This is because the
larger the difference in rainfall, the greater will be the imbalance in the densities
of the emerging adult population and the available oviposition plants. When
defoliation by postdiapause larvae is particularly extreme, competition for limited
oviposition plants may become intense for those individuals who become adults.
In the more usual course of events, however, postdiapause larval starvation is
352
PAN-PACIFIC ENTOMOLOGIST
Figure 1. Row chart of the influence of rainfall on host plant availability, larval survival, fecundity
of adults, and gene flow. Discussion in text.
minimal, adult resources and oviposition plants are abundant, and adults are
sedentary (this is what White and Levin observed in 1973).
Even without a decline in rainfall, consecutive years of average or better rainfall
apparently result in such large population build-ups that postdiapause larvae
almost totally defoliate the Plantago crop regardless of its abundance. This is
what seems to have occurred in 1977. Whether larval food-stress predisposes
adults to migrate is not known, but the presence of nectar appears not to inhibit
emigration under these circumstances. These populations of butterflies are thus
regulated by rainfall (which determines plant crop density) in a largely density-
independent manner most years. But, intermittent generations are subject to severe
competition, hence are regulated in a density-dependent manner.
Adaptations to Stress in Plants and Butterflies
The long-term status of larger local populations of Plantago and Euphydryas
in Southern California appears to be little affected by host plant defoliation, larval
starvation, and adult dispersal. Though we observed essentially total defoliation
of Plantago in 1977, so that no seed was set, the autumn rains that year produced
large standing crops from the seed bank remaining in the soil from previous years.
In February of 1978, an average of 19 plants per square decimeter were counted
in 15 randomly selected plots (range: 0-67) at LO. We have observed similar
phenomena in Northern California where Euphydryas use annual larval host
plants, which nearly disappeared due to drought rather than to defoliation, forcing
small E. editha populations to extinction (Ehrlich et al., 1980).
Three characteristics of Euphydryas editha, particularly well developed in the
wrighti subspecies, seem to be key to its survival in this relatively unpredictable
environment. First, adult size shows a positive correlation with rainfall and is
VOLUME 60, NUMBER 4
353
greatest when rainfall-dependent host plant density and quality are high (White
and Levin, 1981; Murphy et al., in prep.). In seasons of poor host plant conditions,
larvae pupate small, rather than feeding to some “optimum” size. Though smaller
adult females produce fewer eggs in such years, offspring are provided sufficient
time to reach diapause before host plants senesce. Second, postdiapause larvae
experiencing food-stress have been observed to re-enter diapause (Singer and
Ehrlich, 1979). This may explain why in 1978 the adult population was of mod¬
erate size: larvae from 1976 re-entered diapause in 1977 to emerge successfully
in 1978. Third, emergence from diapause may be controlled more by rainfall and
host plant availability than by time of year, and appears to be less obligatory than
in other subspecies such as Northern California’s E. editha bayensis. Abundant
host plant crops are used whenever available, even when they occur out of season
due to unusual rainfall. The sizes of fall-flying generations, such as the one reported
for November 1936, may resemble those of normal February-March generations
(Fred Thorne, pers. comm.). The pool of diapausing larvae is thus replenished
whenever rainfall is substantial, rather than only in years with good winter rainfall.
Biogeographic and Genetic Consequences of Mass Dispersal
Clearly important in the immigration-extinction dynamics of Southern Cali¬
fornia Euphydryas editha is the role of migrants moving into unoccupied sites
during these infrequent population explosions. Populations of E. e. wrighti have
long been known to vanish for years, then to suddenly reappear (Orsak, 1977; J.
W. Johnson, pers. comm.). We suspect that in many years rainfall, and therefore
host plants, are insufficient to support populations in many small, topographically
homogeneous or otherwise marginal habitats and that local population extinction
is likely common there. Between population outbreaks, populations may survive
only in the largest, most diverse habitats. During consecutive years of adequate
rainfall these populations increase in size and exhaust host plant availability
which leads to mass dispersal. Larger populations then act as sources of migrants
to uninhabited areas, which due to adequate rainfall in these years support suf¬
ficient plant resources to make establishment or re-establishment of populations
possible. The largest, most persistent populations thus support the long-term
existence of many other populations.
This extinction/recolonization dynamic has implications for conservation ef¬
forts aimed at this species (Murphy and Ehrlich, 1980). (The wrighti subspecies
has been placed on several review lists for potential endangered status.) The
extirpation of a single, large reservoir population of wrighti may effectively deny
other habitats necessary migrants, creating a ripple effect of irreversible long¬
term extinctions. We suspect that just such a circumstance has eliminated Eu¬
phydryas editha wrighti from Orange County and much of coastal San Diego
County, and now threatens populations in Riverside and inland San Diego Coun¬
ties in California.
The role of infrequent episodes of mass dispersal in affecting gene flow between
Euphydryas editha populations is not so clear. There is a lack of any significant
genetic difference among populations at UO, LO, and BF (McKechnie et al., 1975;
C. E. Floldren, pers. comm.). In most years the migration rates among these
populations seem to be very low, probably less than that necessary to keep the
frequencies of selectively neutral alleles the same (Lewontin, 1974). It is tempting,
354
PAN-PACIFIC ENTOMOLOGIST
therefore, to ascribe the observed genetic similarities to population explosions of
the sort seen in 1977. However, the environmental conditions that result in
increased dispersal make reproduction nearly impossible for migrants arriving at
occupied population sites. Hence, while adult movements are necessary for gene
flow to occur, they are not equivalent to gene flow in the case of established
populations.
Conclusions
After extensive study of several populations of Euphydryas, movement patterns
appeared to be characteristic of local populations and determined by local resource
distribution (Gilbert and Singer, 1973). Further research showed that for one
population, year-to-year changes in rainfall produce changes in resource avail¬
ability, which in turn result in changes in vagility (White and Levin, 1981). Our
current understanding contains an additional element: prior build-up of insect
population numbers may cause resource depletion resulting in host plant scarcity
even in years of favorable weather. Thus the dynamic history of a population can
have a significant effect on dispersal patterns of its individuals, and on the overall
distribution of the organism.
Acknowledgments
We gratefully acknowledge support by a series of grants from the National
Science Foundation to Paul R. Ehrlich, including DEB78-22413. D. D. Murphy
is presently supported by a grant from the Koret Foundation of San Francisco.
Bruce A. Wilcox, Paul R. Ehrlich, M. Deane Bowers, and two reviewers com¬
mented on drafts of this manuscript. Secretarial support was provided by Old
Dominion University, Norfolk, Virginia.
Literature Cited
Ehrlich, P. R. 1965. The population biology of the butterfly, Euphydryas editha. II. The structure
of the Jasper Ridge colony. Evolution, 19:327-336.
-, D. D. Murphy, M. C. Singer, C. B. Sherwood, R. R. White, and I. L. Brown. 1980. Extinction,
reduction, stability, and increase: The responses of checkerspot butterfly populations to the
California drought. Oecologia, 46:101-105.
-, R. R. White, M. C. Singer, S. W. McKechnie, andL. E. Gilbert. 1975. Checkerspot butterflies:
An historical perspective. Science, 188:221-228.
Gilbert, L. E., and M. C. Singer. 1973. Dispersal and gene flow in a butterfly species. American
Naturalist, 10:58-73.
Lewontin, R. C. 1974. The genetic basis of evolutionary change. Columbia University Press, New
York.
McKechnie, S. W., P. R. Ehrlich, and R. R. White. 1975. Population genetics of Euphydryas but¬
terflies. I. Genetic variation and the neutrality hypothesis. Genetics, 81:571-594.
Murphy, D. D., I. L. Brown, and M. D. Bowers. In prep. Annual variation in butterfly size in
populations of Euphydryas editha.
-, and P. R. Ehrlich. 1980. Two California checkerspot butterfly subspecies: One new, one on
the verge of extinction. J. Lepid. Soc., 34:316-320.
NOAA Climatological Data. 1976-1977. California.
Orsak, L. J. 1977. The butterflies of Orange County, California. University of California, Irvine.
Singer, M. C., and P. R. Ehrlich. 1979. Population dynamics of the checkerspot butterfly, Euphydryas
editha. Fortschr. Zool., 25:53-60.
White, R. R. 1974. Food plant defoliation and larval starvation of Euphydryas editha. Oecologia,
14:307-315.
-, and M. P. Levin. 1981. Temporal variation in insect vagility: Implications for evolutionary
studies. Am. Midi. Nat., 105:348-357.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 355-356
A New Grasshopper of the Genus Eumorsea from Baja
California, Mexico (Orthoptera: Eumastacidae)
Marius Descamps
Museum national d’Histoire naturelle, 45 rue Buffon, 75005, Paris.
The genus Eumorsea Hebard is currently known only from the state of Arizona
from which two species have been described: E. balli Hebard, 1935 (Huachuca
Mts., Glen Oaks) and E. pinaleno Rehn and Grant, 1959 (Pinaleno Mts.). Among
the Eumastacoidea sent to me for study from Baja California, Mexico, was a third
species described below.
At present, the three known species are easily differentiated by the form of the
sub-genital plate of the female.
1. Fastigium of vertex with latero-apical angles very broadly rounded,
anterior margin curved outward (Fig. 6) . 2
Fastigium of vertex with latero-apical angles less broadly rounded, an¬
terior margin incurved or sub-rectangular (Fig. 4); female sub-genital
plate apically broadly rounded (Fig. 3) . E. truncaticeps, n. sp.
2(1). Apex of female sub-genital plate triangular with two strong shoulders
laterally (Fig. 1) . E. balli Hebard
Apex of female sub-genital plate with tongue-shaped process (Fig. 2). .
E. pinaleno Rehn and Grant
Eumorsea truncaticeps, New Species
Holotype.— Female: Mexico, Baja California Norte, Sierra San Pedro Martir
National Park, 4.8 km E of park entrance on road from Hwy. 1 (2316 m), 28-
VII-1978, D. B. Weissman and D. C. Lightfoot. Holotype (in alcohol) in California
Academy of Sciences, No. 14190.
Size relatively small. Fastigium of vertex (Fig. 4) short, slightly convex, anterior
edge sinuate, gently incurved, and forming a narrowly rounded 90° angle with
lateral edges which are parallel and straight, with a fine median carinule. Length
of the head from end of fastigium to occipital suture distinctly shorter than length
of pronotum on median line.
Pronotal disc bounded by distinct fine carinules, sinuate, broken, divergent
caudad in anterior two-thirds, then convergent to caudal margin; metanotum with
lateral carinae strong and sharp, convergent caudad. Median carina narrow and
distinct.
Posterior femur with dorsal carina ending in a rather strong spine.
Subgenital plate well developed posteriorly, edges evenly convex, apex broadly
rounded. Ventral valves of ovipositor small, external margin with two well-marked
teeth, internal margin with one postbasal tooth.
Color in preserved specimens generally dark brown mottled with black, in life
somewhat lighter and greyer, matching granite substrate. Lateral lobes of prono¬
tum with irregular black band extending across meso- and metanotum, ventro-
356
PAN-PACIFIC ENTOMOLOGIST
Figures 1-6. 1-3. Female subgenital plate of Eumorsea. 1. E. balli. 2. E. pinaleno. 3. E. trunca-
ticeps. 4-5. E. truncaticeps. 4. Fastigium of vertex, female, dorsal view. 5. Male cercus in profile. 6.
E. balli, fastigium of vertex, female, dorsal view.
posterior angle marked by a light triangle; first and second abdominal tergites
without black marks; other tergites with lateral black marks forming a broken
band; femora, tibiae and tarsi with dark and light rings, more or less distinct in
individual specimens.
Allotype. — Male: Mexico, Baja California Norte, Sierra San Pedro Martir Na¬
tional Park, 4.8 km E of park entrance on road from Hwy. 1 (2316 m), 28-VII-
1978, D. B. Weissman and D. C. Lightfoot. Holotype (in alcohol) in California
Academy of Sciences, No. 14190.
Cercus thick, slightly incurved, length about 2.5 times basal width (Fig. 5).
Pronotum Posterior femur
1 <3 1.8 mm 7.5 mm
2 2 2.0-2.4 mm 10.0-10.2 mm
Records.— MEXICO: Baja California Norte, Sierra San Pedro Martir National
Park, 4.8 km E of park entrance on road from Hwy. 1 (2316 m), 28-VII-1978,
D. B. Weissman and D. C. Lightfoot, 1 $ allotype, 1 2 holotype (both in alcohol),
5 2 paratypes (pinned); 0.8 km W (outside) of entrance to Sierra San Pedro Martir
National Park, 20-VII-1977, D. Weissman, 1 2 paratype (pinned). Types deposited
in California Academy of Sciences, San Francisco; University of Michigan Mu¬
seum of Zoology, Ann Arbor; and Academy of Natural Sciences, Philadelphia.
Literature Cited
Descamps, M. 1973. Revision des Eumastacoidea (Orthoptera) aux echelons des families et des
sous-familles (genitalia, repartition, phylogenie). Acrida, 2:161-298.
Hebard, M. 1935. Studies in the Orthoptera of Arizona. I. New genera, species and geographic races.
Trans. Amer. Ent. Soc., 61:111-154.
Rehn, J. A. G. and H. J. Grant. 1959. A review of the genera Psychomastax and Eumorsea (Or¬
thoptera, Acridoidea, Eumastacidae). Trans. Amer. Ent. Soc., 84:273-302.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, p. 357
Scientific Note
Association of Mydas xanthopterus (Loew) (Diptera: Mydidae)
and Pepsis formosa Say (Hymenoptera: Pompilidae)
in the Chiricahua Mountains of Southeastern Arizona
During mid-July of 1980, the adults of Mydas xanthopterus (Loew) were col¬
lected in association with Pepsis formosa Say in Pinery Canyon 25 km E of Portal,
Cochise County, Arizona, USA. Geographically, M. xanthopterus is sympatric
with Pepsis species distributed from southeastern Arizona to the trans-Pecos
region of western Texas (Cole, 1969, The flies of western North America, p. 181).
Males and females of M. xanthopterus, which resemble red-winged Pepsis species
and are considered possible Batesian mimics by Cole (Ibid.), were collected sym-
patrically with P. formosa from the yellowish-white blossoms of squaw bush ( Rhus
sp.) (plants 0.5-2.0 m high) along Pinery Creek in the Chiricahua Mountains. In
flight, M. xanthopterus was mistaken for P. formosa. Both species fly along a
direct flight path with the fore legs extended forward beyond the head, the middle
and hind legs directed rearward beside the abdomen, and with all tarsi curved
upward. Prior to alighting, P. formosa would characteristically fly from side to
side in a horizontal plane as individuals approached within ca. 0.5 m of the
blossoms. Adults of M. xanthopterus did not exhibit any of these lateral move¬
ments before landing. Either males or females would normally approach a group
of blossoms at the top of bushes, descend slowly, alight and begin nectar feeding.
The largest of the M. xanthopterus collected was equivalent in size to a medium
sized female or large male Pepsis. Males of M. xanthopterus were typically smaller
than females.
Though we did not quantify our collections, both M. xanthopterus and P.
formosa (including perhaps other Pepsis species not collected) appeared to be
equally abundant. In three hours of intensive collecting along Pinery Creek, the
authors took 9 specimens (5 6, 4 2) of the mydaid, including one female of M.
abdominalis Adams, and several P. formosa. Specimens were identified by com¬
paring our material to that deposited in the insect repository at the University of
California, Davis (courtesy of R. O. Schuster). A small series of M. xanthopterus
has been placed in the museum at U. C. Davis.
The authors wish to thank Dr. D. Clark of Arizona State University for iden¬
tifying the Rhus collected from Pinery Canyon and Mr. R. O. Schuster of the
Department of Entomology, University of California, Davis, who gave us per¬
mission to use the identified specimens deposited in the U. C. Davis museum as
an aid towards the identification of M. xanthopterus and P. formosa.
R. P. Meyer, 1 T. L. McKenzie and F. G. Zalom, Department of Entomology,
University of California-Davis, Davis, California 95616.
1 Present address: Arbovirus Field Station, University of California, Berkeley, P.O. Box 1564,
Bakersfield, California 93302.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 358-359
Scientific Note
Tibicen Cicada (Homoptera: Cicadidae) Associated With A
Potted Ornamental Palm Tree
The purpose of this note is to call attention to what appears to be a feeding
association between a nymph of the cicada Tibicen canicularis (Harris) (Homop¬
tera: Cicadidae) and the roots of a potted ornamental palm tree, Chamaedorea
elegans Martius (Dwarf Date Palm), in a Milwaukee, Wisconsin household. Al¬
though actual feeding by the nymph was not observed, other observations suggest
a feeding relationship with the potted plant. This tentative conclusion is of par¬
ticular interest since there are not many records of nymphal feeding associations
for North American cicadas such as the widespread T. canicularis (Beamer, 1928,
Univ. Kans. Sci. Bull., 18:155-263; Moore, 1966, Pap. Mich. Acad. Sci. Arts and
Lett., 51:75-96).
On 6 April 1981 a male Tibicen emerged from a hole in the soil of a potted C.
elegans kept throughout the winter in the living room of a Milwaukee residence.
This palm tree was about 0.75 m tall and kept in a large plastic pot outfitted with
a series of 3 /4-inch diameter holes in the bottom. The previous summer the palm
was kept out-of-doors for about 2 months, on a stone patio, and the remaining
time on a gravel bed along one side of the house and adjacent to several small
rosebushes. While out of doors, the basic pot was enclosed inside a large redwood
decorator pot, outfitted with a single 3 A-inch hole in the center of the bottom. This
arrangement blocked all but one of the several holes in the basic pot during this
period. The plant was purchased in the spring of 1979 from a local ornamental
plant dealer who related that the palms were grown from seeds in Del Ray Beach,
near Miami, Florida. The soil of the potted plant was never completely replaced,
and for the small amounts that were replaced from time to time, “sterile” potting
soil purchased from a local vendor was always used. At the farm site in Florida,
it takes about a year for the palm to grow to a meter, and the plants are then
trucked north in paper containers with the soil.
The nymphal cast and adult cicada were sent to Dr. Thomas E. Moore who
determined them tentatively as T. canicularis. As the cicada was still teneral when
it was preserved, conclusive species identification could not be confirmed. The
emergence hole in the pot was very close to the base of the palm (Fig. 1) but
because the soil had been disturbed, it was not possible to make a plaster-of-paris
cast of the nymph’s burrow to confirm feeding on the plant’s roots. The palm was
left undisturbed for several weeks to see if other cicadas emerged, and none did;
then the plant was dug up and a thorough check made of the soil and roots. No
other nymphs were found.
Suspecting that the nymph might have accidentally or deliberately crawled up
into the pot through the single unobstructed bottom hole while the plant was out-
of-doors in the previous summer, I reasoned that the warmth of the house in the
winter accelerated the development of the nymph, thereby causing the adult to
emerge much earlier than it normally would outside. In this region, T. canicularis
usually begins emerging in early July (T. E. Moore, pers. comm.). One piece of
valuable data, therefore, would be the discovery of other nymphal casts of this
VOLUME 60, NUMBER 4
359
cicada species in the yard during the summer months of 1981. If found, they
would add credence to the association between the cicada and potted palm being
from a nymph present in the yard the previous summer. But none were found
when checks were made in late August. The yard contains one Seedless Mountain
Ash tree about 10 years old, a few lilac bushes and rosebushes. Since T. canicularis
females commonly lay eggs in the dead, pithy stems of shrubs such as rosebushes
and lilacs within three feet of the soil (T. E. Moore, pers. comm.), and the potted
plant was placed near these bushes the previous summer, I suspected that if any
nymphs were present, they would turn up near these bushes, some of which are
between 7 and 13 years old. T. E. Moore (pers. comm.) suspects a 10-year life
cycle for this cicada in this region. Beamer (op. cit.) reared T. canicularis auriferei
in Kansas to the beginning of the fourth instar in three years, and determined
that fifth instar nymphs require an undetermined period lasting more than one
year to complete development.
Based upon the above observations, I conclude that this cicada was probably
introduced into the potted palm in Florida where these ornamentals are grown
out-of-doors. The nymph, in moving through the soil, either entered the pot or
else became associated with the palm if it was grown in free soil (both methods
are used). During the approximately two-year period in the Milwaukee household,
the nymph probably fed upon the roots of the palm, only to then have an accel¬
erated emergence prompted by the warmth of the house and “long-day” cues
from artificial lights. In my experience in the Milwaukee area, T. canicularis tends
to emerge in clusters in that several nymphal casts are usually found together
under a tree or bush in the course of a summer. Scattered, single individuals are
encountered far less frequently. But T. E. Moore has noted at two sites in Michigan
(Ann Arbor area) that the same study sites produce no nymphal casts in some
years while many in others, even though adults are heard singing in the area every
year. Tentatively, the complete absence of nymphal casts from the yard in the
summer following the one in which the plant was kept out-of-doors is circum¬
stantial evidence against the numph being from a local population there. Tibicen
canicularis occurs in Florida (T. E. Moore, pers. comm.).
Acknowledgments
I thank Dr. Thomas E. Moore, Museum of Zoology, The University of Mich¬
igan, for determining the cicada, discussion of the observations, and reviewing
the manuscript. Dr. Natalie W. Uhl of the L. H. Bailey Hortorium, Cornell
University, acting in the stead of the late Dr. Harold C. Moore, determined the
palm tree. Mr. Mark Schafer of Flower City in Milwaukee provided information
on the source of the palm. Finally, very special thanks to the Pousha family for
alerting me of the cicada emergence in their living room, and for allowing nu¬
merous trips to their home for observations. The cicada and its nymphal cast are
deposited in the collections of the Milwaukee Public Museum.
Allen M. Young, Department of Invertebrate Zoology, Milwaukee Public Mu¬
seum, Milwaukee, Wisconsin 53233.
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, p. 360
Scientific Note
Pachodynerus californicus (Saussure) New to the U.S.
(Hymenoptera: Eumenidae)
On April 1, 1978 at Deep Canyon, Boyd Desert Research Institute, Palm Desert,
Riverside Co., California, I collected Pachodynerus californicus (Saussure) visiting
Encelia farinosa Gray. This species has previously been collected from Baja Cal¬
ifornia Norte but this is the first United States record. Deep Canyon is more than
80 miles north of the nearest Baja site. It is unlikely that it would migrate that
distance and a small population probably exists in Deep Canyon.
Pachodynerus californicus is a robust wasp approximately 13 mm long, and 4
mm wide, mostly black with 2 pale yellow stripes on the gaster and pale yellow
markings on the clypeus, propodeum and upper mesopleuron. It probably nests
in pre-existing cavities in stems and twigs and in abandoned mud-dauber nests.
Caterpillars are probably used as prey (Krombein, K. et al., 1979., Cat. Hymen.
Amer. North of Mexico, 2:1499).
Pachodynerus, a member of the Vespoid family Eumenidae, is a small genus
with only 3 species known previously from America North of Mexico. P. erynnis
(Lepeletier) occurs in the southeast; P. astraeus (Cameron) is found from south¬
western Texas to southern California and P. nasidens (Latreille) is found in south¬
ern Arizona, Texas, Florida and Hawaii.
Norman J. Smith, Staff Entomologist, Fresno Co. Department of Agriculture,
1730 S. Maple, Fresno, California 93702.
VOLUME 60, NUMBER 4
361
Proceedings
FOUR HUNDRED AND TWENTIETH MEETING
The 420th meeting was held Friday, 21 January 1983, at 8:15 p.m., in the Morrison Auditorium,
California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L.
Smith presiding.
The minutes of the meeting held 17 December 1982 were read and accepted. Three persons were
elected as members of the Society: Mr. Randall J. Mitchell as student member, and Dr. Peter R. Atsatt
and Mr. Charles D. Hunter as regular members.
Dr. Smith reported on the meeting of the executive board which decided that the 1983 editorial
board would consist of an executive editor, several assistant editors, and a secretary.
Dr. Smith also pointed out that the sixtieth volume of The Pan-Pacific Entomologist is due in 1984
and suggested that a special commemorative volume be published.
Mr. Gary A. Freed, a graphic artist and member of the Society, offered for sale t-shirts which bear
the Society’s logo. Dr. Vincent D. Roth announced the recent publication of his Handbook for Spider
Identification and offered it for sale.
Dr. David H. Kavanaugh introduced Dr. Roth, resident director of the Southwestern Research
Station, Portal, Arizona. Mr. John A. Skinner introduced four persons from the University of Cali¬
fornia, Davis.
Dr. Smith reported that researchers, in a recent article in Science, demonstrated that the stabili-
mentum of spiders acts as a signal for birds and insects to avoid the web. He also reported on two
groups of fossil myriapods, the Euthycarcinida which were aquatic and the Arthropleurida.
The featured speaker Dr. Richard Karban, University of California, Davis, presented “Ecology of
the Periodical Cicadas (Homoptera).” His slide-illustrated lecture reported on the natural history and
the effect of density on reproductive success of some periodical cicadas of eastern North America.
The social hour was held in the entomology conference room following adjournment of the meeting.
A total of 39 persons was present, of which 28 signed as members and 11 as guests.—Vincent F.
Lee, Secretary.
FOUR HUNDRED AND TWENTY-FIRST MEETING
The 421st meeting was held Friday, 18 February 1983, at 8:05 p.m., in the Morrison Auditorium,
California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L.
Smith presiding.
The minutes of the meeting held 21 January 1983 were read and accepted. Two persons were elected
as members of the Society: Mr. Larry D. Corpus as student member, and Dr. Jacqueline L. Robertson
as regular member.
Dr. Smith reported that the Society will have to face some serious decisions regarding financing a
paid secretary to the editor since the present officers of the Society who are employed by the California
Academy of Sciences cannot afford to carry more responsibilities for the Society.
Several guests from faraway places such as Germany and New York were introduced or introduced
themselves.
Mr. Vincent F. Lee gave a note on the Biological Institute of Tropical America (or BIOTA) and
solicited donations for this organization. He also announced that Dr. David B. Weissman will be
conducting a research cruise this summer to several islands in the Gulf of California and along the
Pacific Coast of Baja California. Dr. Smith noted the great proliferation of an eriophyid mite, introduced
from South America, on fuchsias in San Francisco and offered the formula of a mixture to control
the infestation.
The featured speaker Dr. Edward S. Ross, Department of Entomology, California Academy of
Sciences, presented “Insect Diversity in an Amazonian Forest.” His well-illustrated slide lecture
reported primarily on a recent trip to Explorer’s Inn along the Tambopata River of Peru, with additional
slides from his previous trips to other Amazonian areas.
The social hour was held in the Goethe Room following adjournment of the meeting.
A total of 98 persons was present, of which 30 signed as members and 44 as guests.—Vincent F.
Lee, Secretary.
362
PAN-PACIFIC ENTOMOLOGIST
FOUR HUNDRED AND TWENTY-SECOND MEETING
The 422nd meeting was held Friday, 18 March 1983, at 8:10 p.m., in the Goethe Room, California
Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L. Smith presiding.
The minutes of the meeting held 18 February 1983 were read, corrected, and accepted. Dr. Philippe
de Wailly was elected as a regular member of the Society.
Dr. Smith announced that Dr. Paul H. Amaud Jr. had volunteered to be the interim editor of The
Pan-Pacific Entomologist. Mr. Steve Engle introduced himself as a new guest of the Society. Mr.
Vincent F. Lee announced that the Department of Entomology of the California Academy of Sciences
has used hard-bottomed insect trays for sale. Dr. David H. Kavanaugh showed slides of his trip to
the Steens Mountain, Oregon last year.
The featured speaker Dr. Rollin E. Coville, Division of Entomology and Parasitology, University
of California, Berkeley, presented “Nesting Biology and Behavior of Trypoxylon Spider Wasps (Hy-
menoptera: Sphecidae).” His slide-illustrated lecture reported primarily on his research in Costa Rica.
Special mention was made of the nesting biology and enemies of the wasps.
The social hour was held in the entomology conference room following adjournment of the meeting.
A total of 40 persons was present, of which 25 signed as members and 13 as guests.—Vincent F.
Lee, Secretary.
FOUR HUNDRED AND TWENTY-THIRD MEETING
The 423rd meeting was held Friday, 15 April 1983, at 8:05 p.m., in the Morrison Auditorium,
California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L.
Smith presiding.
The minutes of the meeting held 18 March 1983 were read and accepted. Mr. David A. Rider was
elected as a student member of the Society.
Dr. David H. Kavanaugh introduced Dr. Larry E. Watrous and Dr. Craig Dietrick. Dr. Smith gave
an update on the status of The Pan-Pacific Entomologist. Mr. Vincent F. Lee announced again that
the Department of Entomology of the California Academy of Sciences has used hard-bottomed insect
trays for sale. Dr. William E. Ferguson noted the migration of painted lady butterflies in the Borrego
Desert of Southern California. Dr. Smith summarized recent fossil finds of Dr. Jarmilla Kukalova-
Peck.
The featured speaker Mr. Charles P. Hansen, San Mateo County Mosquito Abatement District,
presented “Canine Heartworm in the San Francisco Bay Area.” His slide presentation reported on
the occurrence, detection, and spread of canine heartworm by mosquitoes in dogs and coyotes.
The social hour was held in the entomology conference room following adjournment of the meeting.
A total of 20 persons was present, of which 13 signed as members and 7 as guests.—Vincent F. Lee,
Secretary.
FOUR HUNDRED AND TWENTY-FOURTH MEETING
The 424th meeting was held Friday, 20 May 1983, at 8:05 p.m., in the Goethe Room, California
Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L. Smith presiding.
The minutes of the meeting held 15 April 1983 were read and accepted. Mr. Roy R. Snelling and
Dr. David B. Weissman were elected as regular members of the Society.
Dr. Smith announced that Dr. Robert S. Lane had kindly volunteered to replace Dr. Paul H. Amaud
Jr., now interim editor of The Pan-Pacific Entomologist, on the Publications Committee. He will
complete Dr. Amaud’s term through 1985. Dr. Smith commented on some changes in editorial policy,
and Dr. Amaud summarized the current status of the journal.
Dr. David H. Kavanaugh introduced Dr. Rob E. Roughley, and Mr. Gary W. Ulrich introduced
Mr. Michael Pitcairn. Dr. Kavanaugh announced the existence of the National Science Foundation’s
Long Term Ecological Network (LTEN) sites which systematists might be interested in sampling. Dr.
Smith mentioned that the American Association for the Advancement of Science will be meeting in
San Francisco in 1984. He suggested that the Society participate in a joint meeting with the Academy
at these meetings. He also gave further details on Monura, a fossil machilid of phylogenetic significance.
The featured speaker Dr. Hugh Dingle, University of California, Davis, presented “Ecology and
Evolution of Insect Migration.” He demonstrated how migration and diapause are interrelated in
some insects of the temperate regions. Examples of migration in the monarch, Oncopeltus, Spodoptera,
and others were discussed.
VOLUME 60, NUMBER 4
363
The social hour was held in the entomology conference room following adjournment of the meeting.
A total of 37 persons was present, of which 26 signed as members and 11 as guests.—Vincent F.
Lee, Secretary.
FOUR HUNDRED AND TWENTY-FIFTH MEETING
The 425th meeting was held Friday, 21 October 1983, at 8:10 p.m., in the Morrison Auditorium,
California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L.
Smith presiding.
The minutes of the meeting held 20 May 1983 were read and accepted. Twenty-four persons were
proposed and elected as new members: Mr. Mark A. Bamby, Mr. Ryan Church, Mr. Anthony J.
Galluccio, and Ms. Leslie B. Weeks as student members; and Dr. Richard L. Brown, Mr. Jim Cope,
Mr. David K. Faulkner, Dr. George R. Ferguson, Dr. Paul H. Freytag, Dr. Dan Gerling, Mr. Robert
R. Hansen, Mr. Lawrence R. Hummer, Dr. Urless N. Lanham, Dr. William B. Muchmore, Dr. John
L. Neff, Dr. Mark F. O’Brien, Mr. Carl A. Olson, Dr. George R. Roemhild, Ms. Adrienne E. Roth,
Dr. Tom G. Schwan, Dr. Todd E. Shelly, Dr. Alexander Sokoloff, Dr. Gary M. Stonedahl, and Dr.
Donald B. Thomas Jr. as regular members.
Dr. Smith announced that a new membership category, sponsoring member, was recently created
by the Publications Committee, and Mr. Vincent F. Lee read the names of the members who kindly
elected this category for 1984: Dr. Phillip A. Adams, Mr. Robert P. Allen, Dr. Richard M. Bohart,
Mr. and Mrs. Robert Buickerood, Dr. George R. Ferguson, Mr. Wayne C. Fields Jr., Dr. John G.
Franclemont, Dr. Cornelius B. Philip, Mr. David B. Scott, and Dr. David B. Weissman. He also
mentioned a donation from Dr. David M. Gordon. Dr. Smith announced that the estate of Dr. Charles
P. Alexander bequeathed a large sum of money for the publication of The Pan-Pacific Entomologist.
He also gave an account of a dinner honoring Dr. Richard M. Bohart held on 28 September at Hs.
Lordship’s Restaurant in Berkeley where two leather-bound copies of the special volume of our journal
(volume 59) in his honor were presented to Dr. Bohart and his wife Margaret on the occasion of his
seventieth birthday. Dr. Amaud summarized the current status of the journal. Dr. Smith also an¬
nounced that the XVII International Congress of Entomology will be held 20-26 August 1984 in
Hamburg, Federal Republic of Germany. Dr. Paul H. Amaud Jr. announced a special price of Memoir
4 of the Society publications by the late Dr. Robert L. Usinger, Autobiography of an Entomologist.
Dr. Ronald E. Stecker showed two excellent slides of Dr. Bohart taken several years apart. Dr. Kirby
W. Brown showed slides of lepidopteran eggs taken with a microscope and camera. Dr. Thomas S.
Briggs passed around a live ricinuleid, a rare arachnid, collected when he attended the IX International
Congress of Arachnology in Panama this past August. Dr. Smith exhibited recent publications on
Carboniferous arthropods from the Burgess Shale of Canada and summarized the unique and unusual
features about them.
The featured speaker Mr. Martin Galindo-Ramirez, University of California, Santa Cruz, presented
“Population Genetics and Spider Biogeography.” He showed results of his research using gel electro¬
phoresis on Bothriocyrtum tarantulas and how these tie in with field studies of their natural distri¬
butions. He also reported on his on-going research with this technique on Lutica spiders of sand-
dunes of southern California and Baja California and speculated on the biogeographical distributions
of them in California.
The social hour was held in the entomology conference following adjournment of the meeting.
A total of 42 persons was present, of which 27 signed as members and 15 as guests.—Vincent F.
Lee, Secretary.
FOUR HUNDRED AND TWENTY-SIXTH MEETING
The 426th meeting was held Friday, 18 November 1983, at 8:05 p.m., in the Morrison Auditorium,
California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L.
Smith presiding.
The minutes of the meeting held 21 October 1983 were read and accepted. Six persons were proposed
and elected as new members: Mr. Michael F. Day, Mr. Lawrence E. Serpa, and Ms. Sally M. Swales
as student members; and Mr. Peter Ghormley, Dr. Glenn E. Haas, and Dr. Frank W. Merickel as
regular members.
Dr. Smith called on the audience for volunteers to serve on the program, membership, and pub¬
lication committees of the Society for 1984. He also announced that the Society should chair a section
of the meetings of the Pacific Branch of the American Association for the Advancement of Science
in June 1984 in San Francisco and asked for volunteers. Mr. Vincent F. Lee read these additional
364
PAN-PACIFIC ENTOMOLOGIST
names of members who elected the sponsoring member category for 1984: Dr. Richard K. Allen, Dr.
William F. Barr, Dr. Donald J. Burdick, Dr. Leopoldo E. Caltagirone, Dr. Kenneth W. Cooper, Dr.
J. Gordon and Alice Edwards, Dr. Theodore W. Fisher, Dr. E. Eric Grissell, Dr. Kenneth S. Hagen,
Dr. Alice S. Hunter, Mr. Johannes L. Joos, Mr. Robert L. Langston, Dr. Robert J. Lyon, Mr. Gordon
A. Marsh, Mr. David G. Marqua, Dr. Woodrow W. Middlekauff, Mr. Robert B. Miller, Mr. Calvert
E. Norland, Mr. Harry W. Oswald, Dr. Robert W. L. Potts, Dr. Jacqueline L. Robertson, Ms. Leslie
S. Saul, Dr. Harvey I. Scudder, Dr. Terry N. Seeno, Mr. Frank E. Skinner, and Mr. Baldomero Villegas.
He also mentioned the following persons who gave generous donations to the Society: Mr. John C.
Downey, Dr. Norville M. Downie, Dr. Edward S. Ross, and Mr. Stephen R. Scott.
Mr. Alan I. Kaplan announced that the Northern California Spider Society has for sale t-shirts
bearing his society’s name and a beautifully executed silk screen of a spider. Dr. Smith commented
on the recent findings of gigantic fossil arthropods, notably eurypterids, scorpions, trilobites, millipedes,
and spiders and posed questions for speculation on how these organisms are able to function, given
their sizes.
The featured speaker Dr. Edward S. Ross, California Academy of Sciences, presented “Entomological
Encounters in China, Malaya, and Sumatra.” He showed slides and lectured on his trip to these areas
in the summer this year with a tour group in China and on his own in the other countries. He talked
about the structure and insect associates of Rafflesia, Earth’s largest flower.
The social hour was held in the entomology conference room following adjournment of the meeting.
A total of 92 persons was present, of which 44 signed as members and 35 as guests.—Vincent F.
Lee, Secretary.
FOUR HUNDRED AND TWENTY-SEVENTH MEETING
The 427th meeting was held Friday, 16 December 1983, at 8:15 p.m., in the Morrison Auditorium,
California Academy of Sciences, Golden Gate Park, San Francisco, with President Dr. Edward L.
Smith presiding.
The minutes of the meeting held 18 November 1983 were read and accepted. Ms. Carol Evkhanian
was proposed and elected as a regular member.
Dr. Smith called for reports from the standing committees. Dr. Wojciech J. Pulawski, Treasurer,
announced that the Society’s finances are in excellent condition, especially after receiving the Charles
P. Alexander bequest. Mr. Vincent F. Lee, a member of the membership committee, gave a breakdown
of the 1983 membership. Dr. John E. Hafemik Jr., chairperson of the nominating committee, read
the slate of officers proposed for 1984: Dr. Harvey I. Scudder as president, Dr. J. Gordon Edwards
as president-elect, Dr. Wojciech J. Pulawski as treasurer, and Mr. Vincent F. Lee as secretary. Dr.
Smith called for and received no additional nominees from the audience. The nominees were unan¬
imously elected to office. Dr. Paul H. Amaud Jr. summarized the report of the auditing committee
and reported that the accountant Mr. H. Vannoy Davis found the accounts in good order.
Dr. Smith gave further details on the Society’s participation in the meetings of the Pacific Branch
of the American Association for the Advancement of Science in June 1984. He issued a call for papers
and suggested that abstracts be published in the October 1984 issue. He also suggested that articles
in The Pan-Pacific Entomologist should have an author’s abstract and keywords before each article.
He then handed the gavel over to Dr. Scudder, the new president for 1984. Mr. Lee announced that
this meeting has approval for the State of California Agricultural Pest Control Adviser continuing
education accreditation and asked those persons who wish to receive credit to sign the sign-up sheet.
He then read additional names of members who elected to be sponsoring members for 1984: Mr. Eric
M. Fisher, Dr. John E. Hafemik Jr., Mr. Charles E. Kennett, Mr. Dennis M. Kubly, Dr. Robert L.
Mangan, Dr. Richard L. Penrose, Dr. Evert I. Schlinger, and Drs. Marius S. Wasbauer and Joanne
Slansky. On behalf of the Society, he thanked Dr. James W. Tilden and William M. Upholt, both
Life Members, who gave generous donations to the Society. Dr. Woodrow W. Middlekauff introduced
Dr. David C. F. Rentz, a member of the Society, who was visiting from the CSIRO, Canberra City,
Australia. Dr. Scudder announced titles of the talks for the January and Febuary meetings.
The featured speaker Dr. Edward L. Smith, California Academy of Sciences, gave the presidential
address, “Sawfly Galls: How Hymenoptera-Symphyta Provision Their Larvae with Plant Tumors.”
He showed slides and lectured on gall formation by these insects, with excellent photographs of
microtomic sections under different lighting techniques.
The social hour was held in the entomology conference room following adjournment of the meeting.
A total of 24 persons was present, of which 20 signed as members and 4 as guests.—Vincent F. Lee,
Secretary.
VOLUME 60, NUMBER 4
365
PACIFIC COAST ENTOMOLOGICAL SOCIETY
STATEMENT OF INCOME, EXPENDITURES AND
CHANGES IN FUND BALANCES
Years Ended September 30, 1983 and 1982
1983 1982
Income
Dues and subscriptions. $10,037 $10,078
Reprints and miscellaneous. 6,504 6,046
Sales of Memoirs. 226 256
Interest. 2,509 5,727
Dividends, American Telephone & Telegraph Company. 468 432
Increase (Decrease) in value of capital stock
of American Telephone & Telegraph Company. 590 (150 )
$20,334 $22,389
C. P. Alexander bequest. 29,732 _
$50,066 $22,389
Expenditures
Publication costs—Pan-Pacific Entomologist. $10,262 $ 6,418
Reprints, postage and miscellaneous. 694 5,039
$10,956 $11,457
Increase in fund balances. $39,110 $10,932
Fund balances October 1, 1982 and 1981 . 45,885 34,953
Fund balances September 30, 1983 and 1982 . $84,995 $45,885
STATEMENT OF ASSETS, September 30, 1983 and 1982
1983 1982
Cash in bank
Commercial account. $ 3,385 $12,023
Savings accounts & certificates of deposit
General fund. 54,995 9,909
Memoir fund . 21,415 19,229
Total cash in bank. $79,795 $41,161
Undeposited receipts. 0 114
Investment in 80 shares of American Telephone
& Telegraph Co. common stock (Life Member¬
ship and Fall Funds), at market value. 5,200 4,610
$84,995 $45,885
See accompanying notes to the financial statements.
PACIFIC COAST ENTOMOLOGICAL SOCIETY
NOTES TO THE FINANCIAL STATEMENTS
Year Ended September 30, 1983
Summary of significant accounting policies.
Accounting Method: Income and expenses are recorded by using the cash basis of accounting. Mar¬
ketable Securities: American Telephone & Telegraph Co. common stock is carried at market value.
Increases and decreases in value are reflected in income. Income Tax: The Society is exempt from
Federal income and California franchise tax. Accounts Receivable: As of September 30, 1983, accounts
receivable aggregated $3,617. Accounts Payable: As of September 30, 1983, there was no material
amount of unpaid bills.
366
PAN-PACIFIC ENTOMOLOGIST
As Chairman of the Auditing Committee, and in accordance with its bylaws, I have reviewed the
financial records of the Society. During the course of this review nothing was noted which indicated
any material inaccuracy in the foregoing statements.
H. Vannoy Davis
Chairman of the Auditing Committee
PAN-PACIFIC ENTOMOLOGIST
60(4), 1984, pp. 367-368
The Pan-Pacific Entomologist
Index to Volume 60
(New taxa in boldface)
Acanthodoryctes tomentosus, 99
Aclypea opaca, 249; bituberosa, 250
Ambracyptus, 163; minutissimus, 163
Andricus reticulatus, 290; bakkeri, 292
Anthicidae, 69
Anthophoridae, 119, 155, 188, 300
Aradidae, 76
Asilidae, 53
Bembidion, 165
Braconidae, 97
Callirhytis lentiformis, 295
Camelopsocus, 193;bactrianus, 197;hiemalis, 199;
monticolus, 201; similis, 202; tucsonensis, 204
Campyloneura virgula, 4
Capnia hornigi, 30
Capniidae, 30
Carabidae, 165
Caurinella, 245; idahoensis, 245
Cerambycidae, 63, 67, 73, 94, 114, 279
Ceratophyllidae, 345
Chemetidae, 20
Chinessa arnaudi, 76
Chironomidae, 88
Cicadellidae, 341
Cicadidae, 358
Coleoptera, 23, 63, 65, 69, 73, 94, 114, 122, 163,
165, 248, 279, 304, 313
Collembola, 258
Corydalidae, 1
Cynipidae, 289
Dasymutilla, 219
Dendrocoris maculosus, 8
Diaspididae, 267
Dichaetocoris anasazi, 33
Diptera, 37, 53, 88, 213, 238, 265, 326
Drosophila pseudoobscura, 16
Elateridae, 65
Elmidae, 23
Epeolus minimus, 119
Ephemerellae, 246
Ephemerellinae, 245
Ephydridae, 37
Eremapis parvula, 155
Eumastacidae, 355
Eumenidae, 12, 360
Eumorsea truncaticeps, 355
Euphydryas editha, 350
Euvelia advena, 56; lata, 59; concava, 60; discala,
61
Exomalopsis crenulata, 188
Fabiola quinqueferella, 337
Filacus, 102; doanei, 106; pluricinctus, 108; plu-
ricinctellus, 110; provancheri. 111
Formica haemorrhoidalis, 79
Formicidae, 79, 151
Gabrius, 122; unzenensis, 125; subdepressus, 125;
kobayashii, 128; egens, 130; yamanei, 130;
io, 133; damon, 135; abas, 135; demades, 138;
philo, 140; kuanshanensis, 142; sharpianus,
142; ophion, 146; nepos, 146
Gastrodynerus, 12; searsi, 13; stangei, 15; van-
duzeei, 15
Giesbertia, 284; rugosa, 286
Heleniella nr. ornaticollis, 90
Hemiptera, 4, 8, 33, 47, 55, 76
Heteroecus crescentus, 293; devorus, 294
Homoptera, 65, 267, 341, 358
Hymenoptera, 12, 79, 97, 101, 119, 151, 155,
188, 219, 289, 300
Idiocerus rossi, 341
Lacon profusa, 65
Lanellus, 265
Lepidoptera, 332, 337, 350
Leptelmis, 23; philomina, 24
Linsleyella, 114; ricei, 115; virens, 115; michel-
bacheri, 117
Lycoplasma, 279; formosa, 280
Megaloptera, 1
Mesocricotopus thienemanni, 90
Miridae, 4, 33, 47
Mucrochernes hirsutus, 20
Mutillidae, 219
My das xanthopterus, 357
Mydidae, 357
Neosminthurus bellingeri, 262
Nesocerus trimaculatus, 344
Neuroterus chrysolepis, 289
Noctileptura, 281; squamosa, 282; seriata, 283
Nosopsyllus fasciatus, 345
Nymphalidae, 350
367
368
PAN-PACIFIC ENTOMOLOGIST
Obrium californicum, 67
Oecophoridae, 337
Oedipodinae, 269
Oraphanes, 283; binotatum, 284
Orohermes, 1
Orthoptera, 227, 269, 355
Pachodynerus californicus, 360
Paracantha cultaris, 213
Parapanimerus, 240; arnaudi, 242
Pentatomidae, 8
Pepsis formosa, 357
Pheidole hyatti, 152; pilifera pacifica, 153
Phobetus desertus, 305
Phyllophaga bimaculata, 313; caneyensis, 315;
complexipennis, 317; chalumeaui, 317; fran-
ciscana, 319; hardyi, 319; hernandezi, 321;
independentista, 322; maestrensis, 323; mi-
gratoria, 324; spinicola, 324
Phy mat odes shareeae, 63; oregonensis, 67; tysoni,
73
Phytocoris nobilis, 47; occidentalis, 50
Plagithmysus bilineatus, 94
Plecoptera, 30
Pompilidae, 357
Protochauliodes cascadius, 1
Pseudoscorpionida, 20
Pseudosmittia gracilis, 88
Psocidae, 193
Psocoptera, 193
Psychodidae, 238
Pyralidae, 332
Pyrausta orphisalis, 332
Scarabaeidae, 304, 313
Silphidae, 248
Siphonaptera, 345
Sminthuridae, 258
Solenopsis maniosa, 151
Sphecidae, 256, 297
Staphylinidae, 122, 163
Stenopogon setosus, 53
Stupkaiella, 239; masterelli, 239
Tabanidae, 265
Tanarthrus andrewsi, 69; salinus, 71; alutaceus,
71; brevipennis, 71; coruscus, 71; inhabilis,
71; inyo, 71; iselini, 71; tartarus, 71
Tenthredinidae, 101
Tephritidae, 213
Threticus bicolor, 238
Tibicen canicularis, 358
Timpanogae, 246
Triepeolus heterurus, 120
Triepeolus nr. eldredi, 300
Trimerotropis, 269
Trimerotropis bernardi, 227; infantilis, 231; inyo,
233; leucophaea, 234; whitei, 236
Trypoxylon clavicerum, 256
Veliidae, 55
Vietnamellae, 247
Published by the
Pacific Coast Entomological Society
in cooperation with
The California Academy of Sciences
VOLUME SIXTY
1984
EDITORIAL BOARD
J. A. CHEMSAK, Editor
R. S. LANE, Associate Editor
R. M. BOHART
J. T. DOYEN
J. A. POWELL
J. E. HAFERNICK, JR.
W. J. PULAWSKI, Treasurer
PUBLICATION COMMITTEE
1984
1985
R. M. Bohart P. H. Amaud, Jr.
J. G. Edwards M. S. Wasbauer
San Francisco, California
1984
The Pan-Pacific Entomologist
Contents for Volume 60
Allen, R. K.—A new classification of the subfamily Ephemerellinae and the description of a
new genus. 245
Alstad, D. N.—See Edmunds, G. F., Jr.
Anderson, R. S., and S. B. Peck—B ionomics of Nearctic species of Aclypea Reitter: Phytoph¬
agous “Carrion” beetles (Coleoptera: Silphidae) . 248
Barnby, M. A., and V. H. Resh—D istribution and seasonal abundance of brine flies (Diptera:
Ephydridae) in a San Francisco Bay salt marsh. 37
Baumann, R. W., and A. L. Sheldon —Capnia hornigi, a new winter stonefly from the western
Great Basin (Plecoptera: Capniidae). 30
Bezark, L. G. — Description of the first Central American species of Stenopogon Loew (Diptera:
Asilidae) . 53
Blom, P. E., and W. H. Clark —Phobetus desertus, a new Melolonthine Scarabaeidae (Co¬
leoptera) from the central desert of Baja California, Mexico. 304
Bohart, R. M . — Gastrodynerus, a new genus of Eumenidae from western North America (Hy-
menoptera: Eumenidae). 12
Boussy, I. A. —See Coyne, J. A.
Brown, H. P., and M. P. Thobias—W orld synopsis of the riffle beetle genus Leptelmis Sharp,
1888, with a key to Asiatic species and description of a new species from India (Coleoptera,
Dryopoidea, Elmidae) . 23
Bryant, — See Coyne, J. A.
Campbell, C. L., and K. S. Pike—D escriptions of the life stages of Pyrausta orphisalis (Lepi-
doptera: Pyralidae) . 332
Cavender, G. L., and R. D. Goeden—T he life history of Paracantha cultaris (Coquillett) on
wild sunflower, Helianthus annuus L. ssp. lenticularis (Douglas) Cockerell, in Southern
California (Diptera: Tephritidae). 213
Chandler, D. S.—A new species and records of Tanarthrus (Coleoptera: Anthicidae) . 69
Chemsak, J. A.—Description of a new purpuricenine genus, Linsleyella (Coleoptera: Ceram-
bycidae). 114
Chemsak, J. A., and E. G. Linsley—A new California species of Phymatodes Mulsant (Co¬
leoptera: Cerambycidae) . 73
Chemsak, J. A., and E. G. Linsley—N ew genera and species of neotropical Cerambycidae
(Coleoptera) . 279
Clark, W. H.—See Blom, P. E.
Clement, S. L.—Observations on the behavior of Triepeolus nr. eldredi Cockerell (Hymenoptera:
Anthophoridae). 300
Cope, J. S.—A new species of Phymatodes from New Mexico (Coleoptera: Cerambycidae) ... 63
Coville, R. E.—The occurrence of Trypoxylon clavicerum in North America (Hymenoptera:
Sphecidae) . 256
Coyne, J. A., I. A. Boussy, and S. Bryant—I s Drosophila pseudoobscura a garbage
species? . 16
Cwikla, P. S. —See Freytag, P. H.
Descamps, M.—A new grasshopper of the genus Eumorsea from Baja California, Mexico (Or-
thoptera: Eumastacidae) . 355
Dodson, G. N., and D. T. Gwynne—A digger wasp preying on a Jerusalem cricket. 297
Edmunds, G. F., Jr., and D. N. Alstad—H igh summer mortality of black pineleaf scale
(Homoptera: Diaspididae). 267
Erwin, T. L.—Studies of the Tribe Bembidiini (Coleoptera: Carabidae): Lectotype designations
and species group assignments for Bembidion species described by Thomas L. Casey and
others. 165
Evans, E. D.—A new genus and a new species of Dobsonfly from the far Western United States
(Megaloptera: Corydalidae). 1
Freytag, P. H., and P. S. Cwikla—Two new species of Idiocerine leafhoppers from Malagasy
Republic (Homoptera: Cicadellidae). 341
ii
Garcia-Vidal, M.—The genus Phyllophaga Harris, 1826 (Coleoptera: Scarabaeidae) in Cuba.
II. Descriptions of eleven new species and illustrations of female genitalia of twelve other
Phyllophaga . 313
Gibson, G. A. P.—See Smith, D. R.
Goeden, R. D.—See Cavander, G. L.
Gwynne, D. T. —See Dodson, G. N.
Haraguchi, J. E. —See Stein, J. D.
Heppner, J. B .—Fabiola quinqueferella : An obscure California moth formerly in Glyphipte-
rigidae (Lepidoptera: Oecophoridae). 337
Kormilev, N. A.—A new species of the genus Chinessa Usinger and Matsuda, 1959, from
Papua New Guinea (Hemiptera: Aradidae). 76
Lattin, J. D., and G. M. Stonedahl— Campyloneura virgula, a predacious Miridae not pre¬
viously recorded from the United States (Hemiptera) . 4
Linsley, E. G.—See Chemsak, J. A.
Lundgren, R. W .—Ambracyptus, a new name for Paracyptus Seevers (Coleoptera: Staphylin-
idae). 163
Lyon, R. J.—New cynipid wasps from California (Hymenoptera: Cynipidae) . 289
MacKay, E. E., and W. P. MacKay—B iology of the thatching ant Formica haemorrhoidalis
Emery (Hymenoptera: Formicidae) . 79
MacKay, W. P. —See MacKay, E. E.
Mangan, R. L.—Reproductive ecology of three cactophilic Diptera (Diptera: Drosophilidae,
Neridae, Syrphidae) . 326
Manley, D. G.—Predation upon Velvet Ants of the genus Dasymutilla in California (Hyme¬
noptera: Mutillidae) . 219
McCluskey, D. K. —See McCluskey, E. S.
McCluskey, E. S., and D. K. McCluskey—H our of mating flight in three species of ants
(Hymenoptera: Formicidae) . 151
McKenzie, T. L.—See Meyer, R. P.
Meyer, R. P., T. L. McKenzie, and F. G. Zalom—A ssociation of Mydas xanthopterus (Loew)
(Diptera: Mydidae) and Pepsis formosa Say (Hymenoptera: Pompilidae) in the Chiricahua
Mountains of Southeastern Arizona . 357
Mockford, E. L.—A systematic study of the genus Camelopsocus with descriptions of three
new species (Psocoptera: Psocidae) . 193
Muchmore, W. B.—Further data on Mucrochernes hirsutus (Banks) Pseudoscorpionda, Cher-
netidae) . 20
Murphy, D. D., and R. F. White—R ainfall, resources, and dispersal in southern populations
of Euphydryas editha (Lepidoptera: Nymphalidae). 350
Muzzio, S. F .—Neosminthurus bellingeri, a new species from California (Collembola: Smin-
thuridae) . 258
Neff, J. L.—Observations on the biology of Eremapis parvula Ogloblin an anthophorid bee
with a metasomal scopa (Hymenoptera: Anthophoridae) . 155
Parker, F. D.—Biological notes on the bee Exomalopsis crenulata Timberlake (Hymenoptera:
Anthophoridae). 188
Peck, S. B. —See Anderson, R. S.
Philip, C. B .—Lanellus, a new subgenus of primitive, California “horse flies” (Diptera, Taban-
idae). 265
Pike, K. S. —See Campbell, C. L.
Polhemus, D. A.—See Polhemus, J. T.
Polhemus, D. A.—A new species of Dichaetocoris Knight from the western United States, with
notes on other species (Hemiptera: Miridae) . 33
Polhemus, J. T., and D. A. Polhemus—N otes on Neotropical Veliidae (Hemiptera) VI. Re¬
vision of the genus Euvelia Drake. 55
Quicke, D. L. J.—The genus Acanthodoryctes Turner (Hymenoptera: Braconidae): Redescription
and new generic synonymy. 97
Rentz, D. C. F., and D. B. Weissman—F ive new species of the band-winged grasshopper genus
Trimerotropis StM (Orthoptera: Oedipodinae) . 227
Resh, V. H.—See Barnby, M. A.
iii
Rust, R. W.—Synonymy in California Channel Island Epeolini bees (Hymenoptera: Antho-
phoridae) . 119
Schwan, T. G.—Nosopsyllus fasciatus parasitizing house mice on southeast Farallon Island,
California (Siphonaptera: Ceratophyllidae) . 345
Sheldon, A. L. —See Baumann, R. W.
Smetana, A. — Review of the Japanese species of the genus Gabrius Stephens (Coleoptera:
Staphylinidae) . 122
Smith, N. J .—Pachodynerus californicus (Saussure) new to the U.S. (Hymenoptera: Eumeni-
dae). 360
Smith, D. R., and G. A. P. Gibson— Filacus, a new genus for four species of sawflies previously
placed in Macrophya or Zaschizonyx (Hymenoptera: Tenthredinidae) . 101
Stein, J. D., and J. E. Haraguchi— Meridic diet for rearing of the host specific tropical wood-
borer Plagithmysus bilineatus (Coleoptera: Cerambycidae) . 94
Steiner, J. W.—Descriptions of the Nearctic larvae of Pseudosmittia gracilis, Mesocricotopus
thienemanni and Heleniella nr. ornaticollis (Diptera: Chironomidae). 88
Stonedahl, G. M.— Two new conifer-inhabiting Phytocoris from western North America (He-
miptera: Miridae) . 47
Stonedahl, G. M. —See Lattin, J. D.
Thobias, M. P.—See Brown, H. P.
Thomas, D. B., Jr.— A new species of Dendrocoris Bergroth from Mexico (Hemiptera: Penta-
tomidae) . 8
Tilles, D. A. —Feeding behavior of Lacon profusa (Candeze) (Coleoptera: Elateridae) in Car¬
penter Ant attended colonies of Cinara spp. (Homoptera: Aphididae). 65
Turnbow, R. H., Jr.— New records for two cerambycids from the Pacific Northwest . 67
Wagner, R.—Contributions to Nearctic Psychodidae (Diptera, Nematocera) . 238
Weissman, D. B.—Notes on the autecology, cytology, morphology, and crepitation of Trimero-
tropis grasshoppers (Orthoptera: Oedipodinae). 269
Weissman, D. B. —See Rentz, D. C. F.
White, R. F.—See Murphy, D. D.
Young, A. M.—Wasps on Agave flowers in Mexico . 244
Young, A. M . — Tibicen Cicada (Homoptera: Cicadidae) associated with a potted ornamental
palm tree . 358
Zalom, F. G. —See Meyer, R. P.
iv
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