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VOL. 91 JANUARY 1989 NO. 1
(ISSN 0013-8797)
PROCEEDINGS.
of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON /
_| (PUBLISHED
QUARTERLY
CONTENTS
BAKER, G. T. and M. M. ELLSBURY—Morphology of the mouth parts and antenna of the
larva of the clover stem borer, Languria mozardi Latreille (Coleoptera: Languriidae) .... 15
CLARK, W. E. and H. R. BURKE—Revision of the ornatus species group of the genus An-
thonomusiGermar (Coleoptera:\@urculionidae)) +. 3... 22... 2s2c.eccscene scene cee cesses 88
ELLSBURY, M. M. and G. T. BAKER—Biological and morphological aspects of the egg stage
and oviposition of Languria mozardi (Coleoptera: Languriidae) ....................... 9
FERNANDES, G. W., W. J. BOECKLEN, R. P. MARTINS and A. G. CASTRO—Ants as-
sociated with a coleopterous leaf-bud gall on Xylopia aromatica (Annonaceae) .......... 81
HANSSON, C.—New World species of Holcopelte and Jonympha (Hymenoptera: Eulophidae),
WAT HECeSCrID HONS) OlitwOme WiSDCCICSHStrr a eet eta ee My aios for Leer che ts Soe fides 59
HELLENTHAL, R. A. and R. D. PRICE— Geomydoecus (Mallophaga: Trichodectidae) from
the Texas and desert pocket gophers (Rodentia: Geomyidae) .......................... 1
HEYDON, S. L.—A review of the world species of Notoglyptus masi (Hymenoptera: Pterom-
LICL AC) Marcy eee ers oc ey nena ey Wo Rh MV a NR AK al cee a PS ea ee aad, ane 112
HURD, L. E. and R. M. EISENBERG—A mid-summer comparison of sizes and growth rates
among nymphs of three sympatric mantids (Mantodea: Mantidae) in two old-field habitats 51
HURD, L. E. and R. M. EISENBERG—Temporal distribution of hatching times in three sym-
patric mantids (Mantodea: Mantidae) with implications for niche separation and coexistence 55
MARSH, P. S.—Notes on the genus Hybrizon in North America (Hymenoptera: Paxylomma-
MILLER, S. E. and V.O. BECKER—North American moths described by L. A. G. Bosc D’Antic
(EepidonteranNoctuiddesbyralidae) p eates sok. ect ase wishes vik te lalate Beenie 22
PAOLETTI, M. G. and B. R. STINNER—Two new terrestrial Isopoda (Oniscidea) from coralline
CAYS Olay CHEZUC AIS: CariDDEAN COAST Monet. a tailed ale gradinis atoletesala dlaied hb wala thelle Rye ae ee 71
ROCK, E. A. and D. JACKSON —Naturally occurring host sites for xylophilic Cecidomyiidae
(DICT. 8 so ly be de es coe oe eee SRR EP ME cocks Ad ia ee 66
STARR, C. K.—The ins and outs of a tropical social wasp nest ............0.. ccc cueeeeee
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1989
F. CHRISTIAN THOMPSON President WARREN E. STEINER, JR. Program Chairman
JEFFREY R. ALDRICH President-Elect GEOFFREY B. WHITE, Membership Chairman
RICHARD G. Rossins, Recording Secretary ANNE M. WIEBER, Custodian
JOHN M. KINGSOLVER, Corresponding Secretary MANYA B. STOETZEL, Delegate, Wash. Acad. Sci.
NorRMAN E. WoobDLey, 7 reasurer
HirAM G. LAReEw, Editor
Publications Committee
REBECCA F. SURDICK GEORGE C. STEYSKAL
Book Review Editor
B. V. PETERSON
Honorary President |
Curtis W. SABROSKY
Honorary Members
LoulseE M. RUSSELL ALAN STONE THEODORE L. BISSELL
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of the Entomological Society of Washington for one year.
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in good standing receive the Proceedings of the Entomological Society of Washington. Nonmember subscriptions
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The Society does not exchange its publications for those of other societies.
PLEASE SEE P. 129 OF VOL. 90(1) FOR INFORMATION REGARDING |
PREPARATION OF MANUSCRIPTS.
STATEMENT OF OWNERSHIP
Title of Publication: Proceedings of the Entomological Society of Washington.
Frequency of Issue: Quarterly (January, April, July, October).
Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of Wash-
ington, c/o Department of Entomology, Smithsonian Institution, 10th and Constitution NW, Wash- |
ington, D.C. 20560.
Editor: Hiram G. Larew, B-470, ARS, USDA, Beltsville, MD 20705.
Managing Editor and Known Bondholders or other Security Holders: none.
This issue was mailed 13 December 1988
Second Class Postage Paid at Washington, D.C. and additional mailing office.
PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 1-8
GEOMYDOECUS (MALLOPHAGA: TRICHODECTIDAE) FROM THE
TEXAS AND DESERT POCKET GOPHERS
(RODENTIA: GEOMYIDAE)
RONALD A. HELLENTHAL AND ROGER D. PRICE
(RAH) Associate Professor, Department of Biological Sciences, University of Notre
Dame, Notre Dame, Indiana 46556; (RDP) Professor, Department of Entomology, Uni-
versity of Minnesota, St. Paul, Minnesota 55108.
Abstract.—Geomydoecus truncatus Werneck and G. quadridentatus Price.and-Emerson
are redescribed and illustrated. The new species G. neotruncatus 1S described, with the
type host being Geomys personatus streckeri Davis. Ae
Key Words:
Since the initial revision of the pocket
gopher lice by Price and Emerson (1971),
most of the taxa of the louse genus Geo-
mydoecus Ewing occurring on the host ge-
nus Geomys Rafinesque have been the sub-
ject of re-examination and further study and
analysis. The principal works dealing with
these lice are those by Price and Hellenthal
(1975) on the Geomydoecus texanus com-
plex, Price (1975) on the G. scleritus com-
plex, and Timm and Price (1980) on the G.
geomydis complex. This last work presents
keys to the males and females of all Geo-
mydoecus known to that time from Geomys
gophers. It is the purpose of the present pa-
per to complete the taxonomic study of lice
from Geomys by considering the Geomy-
doecus truncatus complex from the Texas
pocket gopher, Geomys personatus True, and
the Geomydoecus quadridentatus complex
from the desert pocket gopher, Geomys are-
narius Merriam.
Quantitative data for the lice studied in
this paper combined with host and locality
information form part of a computerized
pocket gopher-louse data base maintained
at the University of Notre Dame. Counted
or measured characters in the following de-
lice, Geomys arenarius, Geomys personatus
{ an Ar
JAN 77 7QaQ90
JINN EL Joy
scriptions are followed by the minimal and
maximal observed values, and, in paren-
theses, the sample size, mean, and standard
deviation. All measurements are in milli-
meters. In evaluating character usefulness
for specific discrimination, critical values
for each character were calculated at the
point where the likelihood of single char-
acter misidentification of the two compared
taxa was equal, given normality and equal
variance, and ignoring the probability of
collection. For characters offering moder-
ately good discriminating ability, these crit-
ical values and the corresponding probabil-
ities of misidentification are given. In an
abbreviated comparative description for a
species, quantitative data are given only for
those characters whose means differ at a sig-
nificance level of P < 0.01. The host dis-
tribution map was produced by a computer
from a pocket gopher/louse association data
base (Hellenthal and Price 1984). The map
projection is rectangular to simplify deter-
mination of the latitude and longitude for
individual collection sites. Original locality
data expressed in miles are followed par-
enthetically by the metric equivalent to 0.1
km; the English figure, rather than the met-
2 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ric, expresses the precision of the location
estimate. Abbreviations used for host acces-
sion numbers are KU (University of Kan-
sas), TAM (Texas A&M University), and
TT (Texas Tech University). Detailed de-
scriptions of the characters and quantitative
procedures used for Geomydoecus lice are
included in Hellenthal and Price (1980).
Geomydoecus truncatus Werneck
Figs. 1-6
Geomydoecus truncatus Werneck, 1950: 13.
Type host: Geomys personatus personatus
True.
Male.—As in Fig. 6. Temple width (TW)
0.435-0.470 (26: 0.450 + 0.0096); head
length (HL) 0.325-0.360 (26: 0.338 +
0.0089); submarginal and inner marginal
temple setae 0.030-0.045 (15: 0.036 +
0.0046) and 0.020-0.030 (24: 0.025 +
0.0015) long, respectively, with submargin-
al seta positioned near inner marginal seta
and both marginal setae blunt, spiniform
(Fig. 3). Antenna with scape length (SL)
0.180-0.200 (23: 0.191 + 0.0057), scape
medial width (SMW) 0.1 10—0.125 (23: 0.119
+ 0.0054), scape distal width (SDW) 0.1 10—
0.130 (23: 0.122 + 0.0054); without pro-
jection on posterior margin. Prothorax width
(PW) 0.320-0.345 (26: 0.330 + 0.0069).
Abdominal tergal setae: I, 2; II, 12-16 (26:
14.0 + 1.18); II, 16-24 (26: 19.6 + 1.68);
IV, 19-25 (26: 22.3 + 2.00); V, 16-25 (26:
19.9 + 2.13); VI, 13-20 (26: 15.8 + 1.83);
tergal and pleural setae on VII, 18-22 (26:
20.5 + 1.21). Abdominal sternal setae: II,
9-12 (25: 10.5 + 0.96); III, 9-15 (26: 11.5
+ 1.30); IV, 11-15 (26: 13.2 + 1.08); V, 8-
12y(253A10n ae 155) Vi, (6-9), (24: 7.7 =
0.85); VII, 6-9 (25: 6.9 + 0.91); VIII, 4-8
(25: 6.1 + 0.86). Total length (TL) 1.210-
1.385 (25: 1.285 + 0.0511). Genitalia as in
Fig. 5; spinose sac with 6 medium spines;
parameral arch flattened medioposteriorly,
width (PAW) 0.140-0.160 (23: 0.154 +
0.0057); endomeral plate broadly rounded,
with small medioposterior notch, width
(EPW) 0.075-0.090 (26: 0.085 + 0.0037),
length (EPL) 0.075-0.100 (24: 0.087 +
0.0054).
Female.—As in Fig. 1. TW 0.475-0.500
(23: 0.483 + 0.0073); HL 0.310-0.345 (23:
0.327 + 0.0083); submarginal and inner
marginal temple setae 0.030-0.050 (17:
0.037 + 0.0055) and 0.035-0.045 (22: 0.040
+ 0.0031) long, respectively, with submar-
ginal seta positioned near inner marginal
seta (Fig. 2). PW 0.340-0.385 (23: 0.356 +
0.0119). Abdominal tergal setae: I, 2; I, 14—
18 (23: 16.3 + 1.05); TI, 20-25 (23: 22.4
s+) 1038); IV, 23-3032 2622--£ 1295) ave
24—28 (23: 25.6 + 1.08); VI, 21-25 (23: 23.4
+ 1.23); tergal and pleural setae on VII, 24—
34 (23: 28.4 + 2.31). Longest seta of medial
10 on tergite VI, 0.075—0.090 (22: 0.084 +
0.0040); on tergite VII, 0.090-0.120 (23:
0.102 + 0.0074), with 0-2 (23: 0.6 + 0.79)
of these longer than 0.100. Longer of medial
pair of setae on tergite VIII, 0.060-0.085
(22: 0.073 + 0.0070). Last tergite with 3
lateral setae close together on each side; out-
er, middle, and inner setae 0.070-0.095 (19:
0.082 + 0.0067), 0.080-0.105 (20: 0.092 +
0.0067), and 0.080-0.105 (19: 0.095 +
0.0057) long, respectively. Abdominal ster-
nal setae: II, 9-13 (23: 10.6 + 0.84); III, 9-
131@32 PS = 1:04)-1V, 12-1722 ara
+ 1.50); V, 10-14 (22: 12.0 + 1.21); VI, 8-
13(222 10:8 22 1760)? VII, 6=10) Q22 76
1.16). Subgenital plate with 18-23 (23: 21.2
+ 1.53) setae, with distribution and lengths
as in Fig. 1, with | seta on each side dis-
tinctly longer and thicker than others. TL
1.165-1.410 (21: 1.271 + 0.0525). Post-
vulval sclerite as in Fig. 1, with 2 subequal
short setae posterior to 1t on each side. Gen-
ital sac as in Fig. 4, width (GSW) 0.200-
0.280 (17: 0.248 + 0.0195), length (GSL)
0.100-0.180 (17: 0.132 + 0.0251); with
weak anterior papillose area and with 0-5
(17: 2.1 + 1.82) transverse anterior lines,
posteriormost line, when present, situated
0.020-0.060 (12: 0.042 + 0.0121) back from
anterior sac margin.
Discussion.—The male of G. truncatus 1s
VOLUME 91, NUMBER 1 3
VN
1 }-—_—
Figs. 1-6. Geomydoecus truncatus. 1, Female dorsal (left)— ventral (right) view. 2, Female dorsal left temple
margin. 3, Male dorsal left temple margin. 4, Female genital sac. 5, Male ventral genitalia. 6, Male dorsal (left)—
ventral (right) view. Measurements are in millimeters.
4 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
easily distinguished from all other Geomyd-
oecus by its uniquely shaped parameral arch;
no other described species of this genus has
the distinctive medioposterior flattening.
The female is not as readily differentiated,
but the combination of the genital sac struc-
ture, dimensions, and chaetotaxy features
should separate it.
Werneck (1950) described G. truncatus
from a series of six males taken off Geomys
personatus from Padre Island, Texas. This
locality would make the host G. p. persona-
tus, the only pocket gopher that Hall (1981)
lists from there. However, we have found
only Geomydoecus texanus texanus Ewing
on that host. The paucity of our records
cannot rule out the possibility that G. trun-
catus may also occur there, but, conversely,
we are unable to confirm that it does. Our
inability to do this becomes critical since
we have now determined that what has been
known as G. truncatus actually consists of
two species—one from Geomys p. streckeri
Davis and the other from G. p. fallax Mer-
riam. Price and Emerson (1971) had spec-
imens only from G. p. streckeri and named
them Geomydoecus truncatus. Numerous
subsequent collections from Geomys p. fal-
/ax and the determination that these were
different from the G. p. streckeri lice raised
the necessity of establishing which is the
true Geomydoecus truncatus. Fortunately,
we have been able to examine two of Wer-
neck’s paratypes and have determined that
they are conspecific with our series from
Geomys p. fallax.
Geomys p. fallax also has Geomydoecus
texanus texanus occurring on it. Of the six
gophers of this host taxon that yielded G. ¢.
texanus, only one also had G. truncatus. This
one gopher had 12 G. truncatus and only
one specimen of G. ¢. texanus, raising the
possibility that the latter might have been
a contaminant or straggler. It appears that
these two louse taxa, although found on the
same host subspecies, occur in exclusive
ranges.
Material examined.—2 ¢, Paratypes of
Geomydoecus truncatus, ex Geomys perso-
natus, Padre Island, Texas; 53 2, 57 4, ex G.
p. fallax, 9 gophers from 7 localities in San
Patricio Co., Nueces Co., and Live Oak Co.,
Texas.
Geomydoecus neotruncatus
Hellenthal and Price, NEw Species
Type host: Geomys personatus streckeri
Davis.
Male.— Much as for G. truncatus, except
as follows. TW 0.405-0.430 (20: 0.420 +
0.0053); HL 0.310-0.345 (20: 0.326 +
0.0077). Antennal SL 0.165-0.185 (19:
0.177 + 0.0051), SMW 0.100-0.120 (19:
0.109 + 0.0058), SDW 0.100-0.120 (19:
0.112 + 0.0047). PW 0.305-0.335 (20:
0.309 + 0.0078). Setae on sternite II, 7-11
(20: 9.1 + 1.02); VI, 8-12 (19: 9.5 + 0.90).
Genitalia PAW 0.140-0.155 (20: 0.145 +
0.0048).
Female.— Much as for G. truncatus, ex-
cept as follows. TW 0.440-0.465 (20: 0.448
+ 0.0077); HL 0.300-0.330 (20: 0.316 +
0.0075); inner marginal temple seta 0.035—
0.045 (20: 0.037 + 0.0030) long. PW 0.325-
0.340 (20: 0.329 + 0.0061). Tergal setae:
II, 13-17 (20: 15.0 + 1.23); III, 18-23 (20:
21.0 + 1.49); IV, 20-28 (20: 23.9 + 1.65);
V, 20-26 (20: 24.2 + 1.65). Longer seta of
medial pair on tergite VIII, 0.050-0.075 (19:
0.062 + 0.0068). Outer seta on last tergite
0.060-0.085 (20: 0.075 + 0.0057) long.
Sternal setae: II, 8-11 (20: 9.7 + 0.91); V,
10-15 (20: 13.0 + 1.10); VI, 9-14 (20: 12.3
+ 1.22); VII, 8-11 (20: 9.6 + 0.88).
Discussion.—Both sexes of G. neotrun-
catus are smaller than G. truncatus and tend
to have fewer abdominal tergal setae and
more sternal setae on the posterior seg-
ments. For males, the critical values for dis-
crimination and probabilities of misiden-
tification for the best discriminating
quantitative characters separating these two
taxa are the temple width 0.435 (0.034),
prothorax width 0.320 (0.085), and scape
length 0.184 (0.109). For females, the best
VOLUME 91, NUMBER 1
are temple width 0.466 (0.009), prothorax
width 0.343 (0.081), and setae on sternite
VII 8.69 (0.177).
The males of both species key to G. trun-
catus in the first half of couplet 6 in Timm
and Price (1980), where G. neotruncatus can
be separated by its temple width less than
0.435 and prothorax width less than 0.320.
The females of both species key either to G.
truncatus in couplet 2 or G. quadridentatus
Price and Emerson in couplet 9. Temple
width under 0.466 and prothorax width un-
der 0.343 will distinguish G. neotruncatus
from G. truncatus; both may be separated
from G. quadridentatus by their shorter se-
tae on pleurites III-IV (Fig. 1 vs. Fig. 7) and
differences in the genital sac configuration
(Fig. 4 vs. Fig. 8).
Material examined. — Holotype 8, ex Geo-
mys personatus streckeri, 14 mi (22.5 km)
W Crystal City, Zavala Co., Texas,
911.1953, KU-52238; in collection of the
University of Kansas. Paratypes, ex G. p.
streckeri: 2, 7 6, same as holotype; 22 9°, 17
3, same except KU-52239 or 10.11.1953,
KU-52245, 52246; 13 2, 11 6, E Carrizo
Springs, Dimmit Co., Texas, 4.1.1970, TT-
9665, 9666; 6 2, 4 6, 13 mi (20.9 km) N or
NE Carrizo Springs, Dimmit Co., Texas,
17.1.1970, TT-10126, 10131; 12, 1 mi (1.6
km) SW Carrizo Springs, Dimmit Co., Tex-
as, 23.V.1974, TAM-27613; 5 9, 4 4, Car-
rizo. Springs, Dimmit Co., Texas,
24.X1.1938, TAM-789: paratypes distrib-
uted among the United States National Mu-
seum of Natural History, Field Museum of
Natural History, University of Minnesota,
and Oklahoma State University.
Geomydoecus quadridentatus
Price and Emerson
Figs. 7-11
Geomydoecus quadridentatus Price and
Emerson, 1971: 240. Type host: Geomys
arenarius arenarius Merriam.
Male.—Grossly as in Fig. 6, except an-
tenna as in Fig. 10, and dorsal abdomen as
nn
in Fig. 11. TW 0.365-0.410 (80: 0.392 +
0.0100); HL 0.270-0.325 (79: 0.295 +
0.0126); submarginal and inner marginal
temple setae 0.040-0.065 (73: 0.052 +
0.0051) and 0.020-0.030 (79: 0.024 +
0.0023) long, respectively. Antenna with SL
0.145-0.175 (80: 0.164 + 0.0069), SMW
0.095-0.120 (80: 0.109 + 0.0062), SDW
0.115-0.150 (80: 0.135 + 0.0082); with
prominent process on posterior margin (Fig.
10). PW 0.265-0.315 (79: 0.289 + 0.0112).
Abdominal tergal setae: I, 2; II, 8-16 (80:
12.2 + 1.43); TI, 14-23 (78: 18.7 + 1.65);
IV, 17-27 (78: 21.1 + 2.03); V, 16-26 (78:
19.5 + 1.90); VI, 11-19 (78: 15.0 + 1.54);
tergal and pleural setae on VII, 15-24 (80:
20.2 + 1.69). Abdominal sternal setae: I,
9-15 (79: 11.7 + 1.49); III, 11-17 (79: 13.9
+ 1.39); IV, 11-19 (80: 14.3 + 1.62); V, 8—-
14 (80: 10.8 + 1.42); VI, 6-11 (79: 9.2 +
1.13); VII, 5-9 (77: 7.3 + 0.91); VIII, 5-7
(79: 5.9 + 0.51). TL 1.130-1.415 (79: 1.245
+ 0.0634). Genitalia as in Fig. 9; spinose
sac with 4 large central and 0-2 smaller lat-
erally displaced spines; parameral arch with
prominent medioposterior projection, PAW
0.130-0.155 (79: 0.144 + 0.0052); endo-
meral plate triangular with short apical di-
vision, EPW 0.065-0.080 (80: 0.072 +
0.0035), EPL 0.060-0.080 (80: 0.071 +
0.0049).
Female.—Grossly as in Fig. 1, except dor-
sal abdomen as in Fig. 7. TW 0.400-0.470
(80: 0.439 + 0.0122); HL 0.260-0.310 (80:
0.283 + 0.0098); submarginal and inner
marginal temple setae 0.040-0.070 (78:
0.054 + 0.0051) and 0.040-0.050 (78: 0.045
+ 0.0036) long, respectively. PW 0.280-
0.345 (80: 0.311 + 0.0120). Abdominal ter-
gal setae: I, 2; IJ, 13-19 (78: 15.2 + 1.40);
III, 19-27 (77: 21.8 + 1.94); IV, 20-30 (77:
24.6 + 2.40); V, 18-28 (78: 22.5 + 2.21);
VI, 16-26 (79: 20.9 + 2.38); tergal and pleu-
ral setae on VII, 25-39 (80: 32.4 + 2.95).
Longest seta of medial 10 on tergite VI,
0.070-0.100 (80: 0.087 + 0.0062); on ter-
gite VII, 0.085—-0.115 (80: 0.102 + 0.0069),
with 0-6 (80: 0.9 + 1.44) of these longer
6 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-11.
than 0.100. Longer of medial pair of setae
on tergite VIII, 0.050-0.085 (79: 0.067 +
0.0082). Last tergite with outer, middle, and
inner setae 0.045-0.075 (74: 0.058 +
0.0064), 0.060-0.090 (74: 0.074 + 0.0065),
and 0.060-0.090 (79: 0.076 + 0.0069) long,
respectively. Abdominal sternal setae: II, 8—
Kon 7/9 sel Ses 77): IM, WI—17 (772 14:3
+ 1.26); 1V, 11-19 (79: 15.0 + 1.75); V, 8-
LON S2 We8i se 1S). V1, 7=12 (78: 9.6: =
1.02); VII, 6-11 (78: 8.8 + 0.96). Subgenital
plate with 18-26 (80: 21.7 + 2.07) setae.
TL 1.090-1.335 (79: 1.198 + 0.0532). Gen-
ital sac as in Fig. 8, GSW 0.175-0.255 (79:
0.206 + 0.0144), GSL 0.155-0.200 (77:
0.181 + 0.0115), with O-4 (79: 2.1 + 0.82)
curved medioanterior loops, posteriormost
loop, when present, situated 0.040-0.105
(78: 0.071 + 0.0115) back from anterior sac
margin.
Discussion. — Both sexes of G. quadriden-
tatus are easily separated from G. truncatus
Geomydoecus quadridentatus. 7, Female dorsal abdomen. 8, Female genital sac. 9, Male ventral
genitalia. 10, Male ventral antenna. 11, Male dorsal abdomen. Measurements are in millimeters.
and G. neotruncatus. Males of G. quadri-
dentatus have conspicuously different gen-
italia (Fig. 9 vs. Fig. 5), the antennal scape
with a posterior process (Fig. 10), and dorsal
abdominal chaetotaxy (Fig. 11) with longer
setae on pleuron V, generally longer lateral
tergal setae, and the three short setae on
each side of the last tergite evenly spaced
and aligned with very short seta as shown.
Females of G. quadridentatus have a differ-
ent line configuration of the genital sac (Fig.
8 vs. Fig. 4) and longer pleural setae at least
on abdominal segments III-V (Fig. 7). These
three species of lice also are well separated
geographically, with G. quadridentatus dis-
tributed in north central Chihuahua, west-
ern Texas, and south central New Mexico,
and with G. truncatus and G. neotruncatus
in south central Texas (Fig. 12).
As originally described by Price and
Emerson (1971), males of G. quadridentatus
were said to have only four large genital sac
VOLUME 91, NUMBER |
spines, with no mention of one or two small-
er laterally displaced spines. However, re-
cent examination of much larger series of
lice than were available earlier has shown
94 of 179 (52.5%) males with only the four
large central spines, 46 (25.7%) with a single
smaller additional spine, and 39 (21.8%)
with two smaller spines as in Fig. 9. The
percentage of gophers with no, one, or two
smaller sac spines is essentially the same for
all gopher populations of G. quadridentatus
studied. The presence of these smaller spines
should not complicate proper identification,
if other characters and host association are
considered.
There is discussion among mammalogists
as to whether Geomys arenarius is a valid
species apart from G. bursarius (Shaw). Also
uncertain are the relationships among up to
five populations of Geomys possessing what
we here call Geomydoecus quadridentatus:
1) gophers around Gran Quivera, New
Mexico; 2) gophers around San Antonio,
New Mexico; 3) gophers considered to be
G. a. brevirostris Hall; 4) a “river” popu-
lation of gophers belonging to G. a. are-
narius; and 5) an “upland” population of
G. a. arenarius. We collected numerous lice
from all five of these groups, analyzed them
qualitatively and quantitatively, and could
find no meaningful differences. We could
demonstrate occasional quantatitive char-
acter differences at a relatively high prob-
ability of misidentification, but these showed
no consistent occurrence. We do not believe
these louse populations merit taxonomic
distinctions at this time. Speaking strictly
from the louse standpoint, the lice from all
five populations are sufficiently different
from lice from Geomys bursarius to support
G. arenarius as a separate taxon and sufh-
ciently similar to each other to group all five
gopher populations into G. arenarius.
Material examined.—244 9°, 267 6, ex
Geomys arenarius arenarius, 50 gophers
from 17 localities in New Mexico, Texas,
and Chihuahua; 67 2, 51 8, ex G. a. breviros-
tris, 15 gophers from 6 localities in New
Ww
Qa
=
E
=
S §
=I
107 102 97
LONGITUDE
Fig. 12. Geographic distribution of Geomydoecus
truncatus (closed circles), G. neotruncatus (open cir-
cles), and G. quadridentatus (triangles).
Mexico; 26 2, 22 6, ex Geomys, 7 gophers
from 2 localities near Gran Quivera, New
Mexico; 35 9, 28 6 ex Geomys, 4 gophers
near San Antonio, New Mexico.
ACKNOWLEDGMENTS
We wish to thank Sebastiao J. de Oliveira,
Curator, Fundacao Oswaldo Cruz, Rio de
Janeiro, Brasil, for the loan of the Werneck
type series. Partial support for this study
was supplied by a grant from the National
Science Foundation to the University of
Notre Dame (Grant No. BSR8&6-14456).
This is published as paper 15,754 of the
Scientific Journal Series of the Minnesota
Agricultural Experiment Station on re-
search conducted under Project No. Min-
17-015.
LITERATURE CITED
Hall, E.R. 1981. The Mammals of North America.
2nd Ed. Vol. I. John Wiley & Sons, New York,
NY. xv + 600 + 90 pp.
Hellenthal, R. A. and R. D. Price. 1980. A review of
the Geomydoecus subcalifornicus Complex (Mal-
lophaga: Trichodectidae) from Thomomys pocket
gophers (Rodentia: Geomyidace), with a discussion
of quantitative techniques and automated taxo-
nomic procedures. Ann. Entomol. Soc. Amer. 73:
495-503.
1984. Distributional associations among
Geomydoecus and Thomomydoecus lice (Mal-
lophaga: Trichodectidae) and pocket gopher hosts
of the Thomomys bottae group (Rodentia: Geo-
myidae). J. Med. Entomol. 21: 432-446.
Price, R. D. 1975. The Geomydoecus (Mallophaga:
Trichodectidae) of the southeastern USA pocket
gophers (Rodentia: Geomyidae). Proc. Entomol.
Soc. Wash. 77: 61-65.
Price, R. D. and K. C. Emerson. 1971.
the genus Geomydoecus (Mallophaga: Trichodec-
tidae) of the New World pocket gophers (Rodentia:
Geomyidae). J. Med. Entomol. 8: 228-257.
A revision of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Price, R. D. and R. A. Hellenthal. 1975. A review of
the Geomydoecus texanus complex (Mallophaga:
Trichodectidae) from Geomys and Pappogeomys
(Rodentia: Geomyidae). J. Med. Entomol. 12: 401-
408.
Timm, R. M. and R. D. Price. 1980. The taxonomy
of Geomydoecus (Mallophaga: Trichodectidae)
from the Geomys bursarius complex (Rodentia:
Geomyidae). J. Med. Entomol. 17: 126-145.
Werneck, F. L. 1950. Os Malofagos de Mamiferos.
Parte Il: Ischnocera (continuagao de Trichodec-
tidae) e Rhyncophthirina. Ed. do Inst. Osw. Cruz,
Rio de Janeiro. 207 pp.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 9-14
BIOLOGICAL AND MORPHOLOGICAL ASPECTS OF THE EGG
STAGE AND OVIPOSITION OF LANGURIA MOZARDI
(COLEOPTERA: LANGURIIDAE)
MICHAEL M. ELLSBURY AND GERALD T. BAKER
(MME) USDA-ARS, Crop Science Research Laboratory, Forage Research Unit, P.O.
Box 5367, Mississippi State, Mississippi 39762; (GTB) Department of Entomology, P.O.
Drawer EM, Mississippi State University, Mississippi State, Mississippi 39762.
Abstract. —Oviposition sites of Languria mozardi Latreille, the clover stem borer, were
determined on Trifolium incarnatum L., a previously unreported host. The 2nd and 3rd
internodes of primary flowering stems are the preferred sites on T. incarnatum. Oviposition
occurs in stems of 7. repens L., T. pratense L., T. hybridum L., T. resupinatum L., T.
striatum L., T. nigrescens Viv., and T. subterraneum L. but not on T. lappaceum L.. There
is usually one oviposition puncture and egg per stem, the egg being attached to the plant
tissue by an extension on the posterior end. The micropylar region is simple in form and
the chorionic surface has a granular appearance. The sensilla on the ovipositor differ in
the types, number, and distribution as compared to the sensilla on the ovipositor of other
beetle species. The ablation experiment indicates that these sensilla on the ovipositor are
possibly involved in selecting a suitable oviposition site.
Key Words:
The clover stem borer, Languria mozardi
Latreille, is endemic to North America on
a wide range of host plants including the
Leguminosae, Compositae, Umbelliferae,
and Gramineae. It is considered a pest of
alfalfa, Medicago sativa L., and red clover,
Trifolium pratense L. (Lintner 1881, Fol-
som 1909). Wildermuth and Gates (1920)
reported early studies on the biology, mor-
phology, and pest status of L. mozardi on
forage crops. Damage from larval feeding
inside stems weakens the plant and results
in lodging, loss of seed, and forage produc-
tion. Damage to stems also may predispose
plants to root rot pathogens. Increased fiber
content in damaged stems reduces forage
quality (Wildermuth and Gates 1920).
Except for a single report (Knight et al.
1976) from arrowleaf clover, 7. vesiculosum
Languria mozardi, sensilla, larva, mouth parts, antenna
Savi, L. mozardi is known as a forage pest
only on perennial legumes. This insect has
been collected annually in Mississippi from
1982-85 on T. vesiculosum, T. incarnatum
L. (crimson clover) and from 1983-85 on
T. alexandrinum L. (berseem clover). The
importance of seed production in annual
clovers to ensure stand persistence and the
ability of L. mozardi to reduce seed and
forage yield in other legumes makes this in-
sect a potential pest of annual clovers.
No information is available on the biol-
ogy of L. mozardi on annual clovers and
existing morphological studies are incom-
plete. In the present study, oviposition sites
and frequencies on crimson clover, a pre-
viously unreported host, are reported from
greenhouse studies. Host range for ovipo-
sition also are determined on several clover
10 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
species that are or have been of economic
importance. The morphology of the egg and
ovipositor sensilla are described.
MATERIALS AND METHODS
Adult L. mozardi were collected from
roadside plantings of crimson clover using
a motorcycle-mounted collection net (Ells-
bury and Davis 1982). Pairs of mating L.
mozardi were sorted from collections, pro-
vided with a water source, and held 24 h in
a 9-cm diam plastic Petri dish. To deter-
mine Oviposition sites on crimson clover, 3
pairs of insects were confined for 24 h on
each of 12 greenhouse-grown flowering
crimson clover plants in cylindrical screened
cages (14-cm diam x 90 cm). Host range
for oviposition by L. mozardi on other Tri-
folium species also was studied by similarly
caging 3 pairs of insects on each of 2 green-
house-grown flowering plants of 7. repens
L. (white clover), 7. pratense (red clover),
T. hybridum L. (alsike clover), 7. resupi-
natum L. (persian clover), 7. striatum L.
(pitts or striate clover), 7. nigrescens Viv.
(ball clover), 7. lappaceum L. (lappa clo-
ver), and 7. subterraneum L. (subterranean
clover). After 24 h exposure to test insects,
stems were split using a single-edge razor
blade and the location and number of eggs
were recorded for oviposition punctures on
primary and lateral stems.
An average internodal location of ovi-
position punctures was calculated for clo-
vers in the host range study from integer
values assigned to each internode beginning
with the lowermost internode (#1) and in-
creasing to the (last) internode below the
flower head of each stem.
Eggs were removed from the clover stems
and placed in 5% glutaraldehyde in 0.1 M
Na-cacodylate buffer at pH 7.2 for 12 h at
4°C. The specimens were washed 1n the same
buffer and then post-fixed in 4% OsO, in
cacodylate buffer for 8 h. After dehydration
the eggs were critical-point dried, affixed to
aluminum stubs with tape, sputter coated
with Au/Pd, and examined with a Hitachi
HHS-2R scanning electron microscope at
20 kV. The same preparation procedure was
used for the ovipositors. Measurements were
made with a light microscope with a cali-
brated ocular micrometer and are given as
a mean plus the range.
Mated females for the behavioral exper-
iments were anesthetized with CO, and the
ovipositor was gently pulled out and treated
with 16% HCl. They were placed in petri
dishes with a moist filter paper and allowed
to recover overnight. Six treated and 6 un-
treated females were placed in petri dishes
containing pieces of flowering clover stems
with the second and third internodes. The
specimens were examined every 10 minutes
for | h and the experiment was replicated
3 times.
RESULTS
Frequency distribution by internode of
Oviposition sites on crimson clover con-
taining | or 2 eggs is illustrated in Fig. 1.
Oviposition punctures were most frequent
(22/63 and 18/63, respectively) in the 2nd
and 3rd internodes of primary flowering
stems. No eggs were deposited in lateral
flowering stems. Of 78 stems examined, 11
had 2 oviposition punctures and 41 had one
puncture. Six of 8 sites where 2 eggs were
deposited were situated at the 3rd inter-
node. Mean number of eggs per stem was
0.86 + 0.11 (¢ + SE, N = 78).
Oviposition by L. mozardi in 11 species
of clovers during a no-choice test 1s sum-
marized in Table |. Lappa clover, 7. /ap-
paceum, was the only species in which eggs
were not deposited. The usual number of
eggs per site is | or 2 for all clovers studied,
although 3 eggs occurred in a single punc-
ture on 7. nigrescens and 4 eggs in a punc-
ture on a stem of 7. pratense. Eggs were
usually deposited in sites at the 3rd inter-
node or higher on all clovers studied. The
greatest number of eggs per stem (0.75 +
0.33) were deposited in red clover, 7. pra-
tense.
Behavior of the adult female during ovi-
VOLUME 91, NUMBER |
20
1 Egg per Puncture
n=55
2 Eggs per Puncture
INTERNODE NUMBER
|
10
NUMBER OF OVIPOSITION PUNCTURES
Fig. 1. Internodal frequency distribution of Lan-
guria mozardi egg punctures containing | or 2 eggs in
stems of crimson clover, Trifolium incarnatum.
15
20 1
position is similar on all Trifolium species
included in these studies. After chewing a
hole in the plant tissue the female uses her
Ovipositor to excavate a cavity in the pith
into which the egg is placed. Externally, the
opening is covered by shredded plant ma-
terial while internally the opening is sur-
rounded by a distinct rim (Fig. 2a, b). The
egg 1s attached to the plant tissue by an ex-
tension from the tapered end (Fig. 2c). The
egg is clyindrical, translucent yellowish, ta-
pered at one end and measures 1545 um
(1455-1800) in length and 495 wm (480-
Table 1.
during a no-choice test.
11
510) in width (Fig. 2d). The micropyle is
situated on the blunt end and consists of a
small, irregular shaped area from which sev-
eral ridges radiate (Fig. 2e). At low magni-
fication the surface of the chorion appears
smooth (Fig. 2d) but at higher magnifica-
tions the surface has a rough, granular ap-
pearance and minute openings are dispersed
randomly over the surface (Fig. 2f).
The ovipositor coxites are pointed dis-
tally and near the middle of the ventral sur-
face of each coxite there is an area with a
high concentration of long, slender spines
(Fig. 3a, b). Just posterior to the distal end
of each coxite a short style is present (Fig.
3a, b). It is 72 wm (69-74 um) long and 20
um (18—21.5 wm) wide and 10 sensilla trich-
odea are on the apex (Fig. 3b, d). These
sensilla vary in length from 23.5 wm (21-
25 um) to 98 wm (95-101 um) and they are
either slightly curved or straight (Fig. 3d).
Another group of 9 sensilla trichodea are
situated on the ectal surface of each coxite
near the base of the style (Fig. 3a, c). These
trichoid sensilla range in length from 19 wm
(17.5-21 wm) to 71.5 um (69-74 um). The
6 sensilla positioned between the straight
single posterior and 2 long straight anterior
sensilla are slightly curved (Fig. 3c). Sensilla
basiconica are situated on the dorsal and
Frequency and internodal sites of oviposition by Languria mozardi in eleven Trifolium species
Ave. Internodal Location
Trifolium Species! No. Stems Eggs/Stem* Mean No. Nodes/Stem of Oviposition Punctures
*T. subterraneum 19 47 + 16 6.42 + .55 4.67 + .69
T. lappaceum 22 0 9.14 + .60 -
*T_ nigrescens* 5 — 7200. + .32 _
*T. resupinatum 26 208; 5.69: 223) Ae 265
*T. repens 13 .38 + .14 7.96 + 1.04 3.40 = JAZ
*T. striatum 16 44+ 18 6.25 + .40 5.00% 82
T. pratense 12 ho: a: 633 5.25.37 5.00 + .50
T. vesiculosum 16 .69 + .24 10.50 + .61 RY PAT et Vf)
*T. alexandrinum 13 25 ee 12) 12.85 + .69 TOT = 145
*T. hybridum 9 222 218 8.22 + .78 4.00 + 1.00
*7T. nicarnatum 16 .69 + 1.8 6.50e=.35 4.22 + 0.43
' Trifolium species preceded by * are previously unreported hosts.
> All numbers are mean + standard error.
‘Only one 7. nigrescens plant used; 3 eggs laid in a single stem.
12 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
}
4
Fig. 2a-f. Egg of L. mozardi. a, External appearance of the oviposition site. b, Rim surrounding the oviposition
opening internally. c, Tapered end of the egg attached to the plant tissue. d, L. mozardi egg. e, Micropylar area.
f, Chorion surface. E = egg; Ex = egg extension; M = micropyle; R = nm.
ventral surfaces of the coxites (Fig. 3e). These
basiconica sensilla are 5.5 wm (4-6 um) in
length and are situated on a cuticular
depression (Fig. 3e). There are pores (ca. |
um wide) on both surfaces which are inter-
spersed among the basiconic sensilla (Fig.
3e). The sensilla trichodea and basiconica
are stained with the silver nitrate.
In the behavioral experiments, none of
the 18 treated females excavated any ovi-
position sites on the stems or laid any eggs
whereas the untreated females made ovi-
position excavations and deposited eggs.
Many of the treated females left the stems
after 20 to 30 minutes.
DISCUSSION
Oviposition behavior of L. mozardi in
crimson clover is consistent with that re-
ported by Folsom (1909) in red clover and
by Wildermuth and Gates (1920) in alfalfa.
Eggs are usually laid singly but occasionally
2 or more are deposited in a single puncture.
Normally one, and infrequently 2, punc-
ture(s) are found per stem.
The distribution of oviposition sites may
be an important consideration in sampling
for clover stem borer egg populations in
crimson clover. Most oviposition (54/63
sites) occurs in the stem interval between
the 2nd and 4th internodes. Sampling time
for egg-infested crimson clover stems could
be reduced by limiting samples to stem sec-
tions including only the 2nd through 4th
internodes.
Previously reported hosts of L. mozardi
in the genus 7rifolium include only T. pra-
tense (Lintner 1881, Folsom 1909) and 7.
vesiculosum (Knight et al. 1976). Since ovi-
position by L. mozardi occurs in 7 addi-
tional Trifolium species (Table 1) these clo-
vers also should be considered potential
hosts for this insect pending confirmation
of their suitability as hosts for larval de-
velopment.
The excavation for oviposition is similar
to that described for L. mozardi by Girault
(1907) but he does not mention the rim sur-
rounding the opening internally. The egg
shape of L. mozardi differs from Languria
VOLUME 91, NUMBER |
Fig. 3a-e. Ovipositor of L. mozardi. a, Dorsal surface. b, Ventral surface. c, Sensilla at the base of the stylus.
d, Sensilla on the apex of the stylus. e, Short sensilla basiconica on the dorsal surface.
erythrocephalus Blatchley and Acropteroxys
gracilis (Crotch) in that the end opposite the
micropyle is more tapered in L. mozardi as
compared to a blunt end in the other 2 lan-
guriid species (Piper 1978, Genung et al.
1980). The extension from the tapered end
of the egg is used to attach the egg to the
plant tissue and this structure was not ob-
served in other languriid species but is sim-
ilar in structure and function to the one
found on the egg of Lyctus brunneus (Ste-
phens) (Iwata and Nishimoto 1982).
Sensilla are situated on the apex of the
styli and near the base of the styli in other
beetles such as Acanthoscelides obtectus Say
(Bruchidae) (Szentesi 1976), Altica lythri
Aube (Chrysomelidae) (Phillips 1978), and
Thanasimus dubius (F.) (Cleridae) (Baker
and Nebeker 1986). There are.differences in
the types and numbers of sensilla on the
ovipositor of the three previously men-
tioned species as compared to those on L.
mozardi. Sensilla chaetica, trichoidea, and
basiconica are situated on the apex of the
styli and near the base of the styli in A.
obtectus (Szentesi 1976) and T. dubius (Bak-
er and Nebeker 1986) but L. mozardi has
no sensilla basiconica in this region. The
14 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
number of sensilla on the apex of the style
of L. mozardi (9-10) is similar to A. /ythri
(8-11) (Phillips 1978) but differs from A.
obtectus (22-24) and T. dubius (23-25)
(Szentesi 1976, Baker and Nebeker 1986).
The short basiconic sensilla which are sit-
uated on the dorsal and ventral surfaces of
the ovipositor of L. mozardi are lacking on
the ovipositors of the other three beetle
species.
The behavioral experiment indicates that
the sensilla on the ovipositor of L. mozardi
are needed for the deposition of an egg in
the plant substrate. Sensilla on the OVvipos-
itor of other insects are known to respond
to chemicals such as salts, water, and amino
acids (Rice 1976, 1977, Chadha and Roome
1980, Hood-Henderson 1982, Liscia et al.
1982). The sensilla trichoidea and basiconi-
ca on the ovipositor of L. mozardi may re-
spond to chemicals in the clover plant which
are released during the excavation of an ovi-
position site.
ACKNOWLEDGMENTS
The assistance of Mr. D. G. McMinn, Bi-
ological Technician, and Vincent McDaniel
in plant production and insect collection is
gratefully acknowledged. Contribution of
Mississippi Agricultural and Forestry Ex-
periment Station in cooperation with
USDA, ARS. Journal Article No. 64/3 of
the Mississippi Agricultural and Forestry
Experiment Station. Research contributes
to the Southern Regional Project No. S-201,
Role of Legume Cover Crops in Conser-
vation Tillage Production Systems. Men-
tion of trademark or proprietary products
does not constitute a guarantee or warranty
of the products by the U.S. Department of
Agriculture and does not imply their ap-
proval of the exclusion of other products.
LITERATURE CITED
Baker, G. T. and T. E. Nebeker. 1986. Sensory re-
ceptors on the ovipositor of Thanasimus dubius
(F.) (Coleoptera: Cleridae). Ann. Soc. Entomol. Fr.
22: 49-52.
Chadha, G. and R. Roome. 1980. Oviposition be-
havior and the sensilla of the ovipositor of Chilo
patellus and Spodoptera littoralis (Lepidoptera). J.
Zool., Lond. 192: 169-178.
Ellsbury, M. M. and F. M. Davis. 1982. Front-
mounted motorcycle net for mass collection of
clover insects., J. Econ. Entomol. 75: 251-253.
Folsom, J. W. 1909. Insect pests of clover and alfalfa.
Ill. Agric. Exp. Sta. Bull. No. 134. 197.
Genung, W. G., R. Woodruff, and E. Grissell. 1980.
Languria erythrocephalus: Host plants, immature
stages, parasites, and habits (Coleoptera: Langu-
riidae). Florida Entomol. 63: 206-210.
Girault, A. A. 1907. Oviposition of Languria moz-
ardi Latreille. Entomol. News 18: 366-367.
Hood-Henderson, D. E. 1982. Fine structure and
neurophysiology of a gustatory sensillum on the
ovipositors of Metasyrphus venablesi and Eu-
peodes volucris (Diptera: Syrphidae). Can. J. Zool.
60: 3187-3195.
Iwata, R. and K. Nishimoto. 1982. Observations on
the external morphology and the surface structure
of Lyctus brunneus (Stephens) (Coleoptera, Lyc-
tidae) by scanning electron microscopy. Kontyu
50: 10-22.
Knight, W. E., O. W. Barnett, L. L. Singleton, and C.
M. Smith. 1976. Potential disease and insect
problems in arrowleaf clover. Am. Soc. Agron.
Southern Branch (Abstract) 3: 7. Mobile, AL, Feb.
1-4, 1976.
Lintner, J. A. 1881. The insects of the clover plant.
Trans. N.Y. State Agric. Soc. 32: 187-207.
Liscia, A., R. Crnjar, A. Angioy, P. Pietra, and J. Stof-
folano. 1982. I chemosensilla dell ovopositore
in Tabanus nigrovittatus Macq.), Chrysops fuligt-
nosus (Wied.) e Rhagoletis pomonella (Walsh.).
Boll. Soc. Ital. Biol. Spmt. 58: 1325-1329.
Phillips, W. M. 1978. Sensilla types from the OvI-
positor of the flea beetle Altica /ythri (Coleoptera:
Chrysomelidae). Entomol. Exp. Appl. 24: 399-
400.
Piper, G. L. 1978. Life history of Acropteroxys gra-
cilis (Coleoptera: Languriidae) on common rag-
weed in northeastern Ohio. Ohio J. Sci. 78: 304—
309.
Rice, M. J. 1976. Contact chemoreceptors on the
ovipositor of Lucilia cuprina (Wied.), the Austra-
lian sheep blowfly. Aust. J. Zool. 24: 353-360.
1977. Blowfly ovipositor receptor neurone
sensitive to nonovalent cation concentration. Na-
ture 268: 747-749.
Szentesi, A. 1976. The effect of the amputation of
head appendages on the oviposition of the bean
weevil, Acanthoscelides obtectus Say (Coleoptera:
Bruchidae). Symp. Biol. Hung. 16: 275-281.
Wildermuth, V. L. and F. H. Gates. 1920. Clover
stem-borer as an alfalfa pest. USDA Bull. No. 889.
25.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 15-21
MORPHOLOGY OF THE MOUTH PARTS AND ANTENNA OF THE LARVA
OF THE CLOVER STEM BORER, LANGURIA MOZARDI LATREILLE
(COLEOPTERA: LANGURIIDAE)
GERALD T. BAKER AND MICHAEL M. ELLSBURY
(GTB) Department of Entomology, P.O. Drawer EM, Mississippi State, Mississippi
39762; (MME) USDA-ARS, Crop Science Research Laboratory, Forage Research Unit,
P.O. Box 5367, Mississippi State, Mississippi 39762.
Abstract.—The morphology of the mouth parts and antennae, including the sensilla on
these structures, is described. The mouth parts have several types of sensilla (basiconica,
digitiformia, chaetica, trichoidea, campaniformia, and coronal pegs). The apex of each
labial (10 sensilla) and maxillary (11 sensilla) palpus has the highest number of sensilla.
The striated region on each mandible is composed of several striae and each of these has
several rows of blunt, stout pegs. The antennae possess trichoid, basiconic, campaniform,
and placoid sensilla and an antennal sensory appendix which is situated on segment 2.
Key Words: lLanguriidae, Languria mozardi, larva, mouth parts, antennae, sensilla
The clover stem borer, Languria mozardi
Latreille, is nearly ubiquitous in North
America on a wide range of cultivated and
wild host plants (Wildermuth and Gates
1920). Adults occur frequently in the annual
clovers Trifolium incarnatum L. (crimson
clover), 7. vesiculosum Savi (arrowleaf clo-
ver), and 7. alexandrinum L. (berseem clo-
ver), grown for forage, soil stabilization, and
as cover crops in the Southeast.
The clover stem borer causes lodging, re-
duced seed set and poor forage quality as a
result of removal of stem tissue, loss of fo-
liage under stress, and increased fiber con-
tent (Wildermuth and Gates 1920). The im-
pact of L. mozardi on forage production has
probably been underestimated because of
the seclusive nature of the adults and the
cryptic habits of the larvae. The insect is
likely to be of economic importance where
clovers are grown for hay or seed produc-
tion.
Wildermuth and Gates (1920) recom-
mended cultural control through crop ro-
tation, early cutting for hay production, and
sanitation of field margins to reduce over-
wintering populations. Selection and breed-
ing of clovers for stem characteristics to re-
duce establishment and survival of the larva
would be an alternative control method. The
present studies were initiated to describe the
morphology, distribution, and number of
sensory receptors on larval mouth parts and
antennae of L. mozardi that may mediate
feeding or tunneling activity of the larva
through contact with the host plant.
MATERIALS AND METHODS
Specimens of final (Sth) instar larvae of
L. mozardi were reared on artificial diet
(Rose and McCabe 1973). Use of the diet
permitted verification of molting to ensure
that Sth instar larvae were examined. The
techniques for SEM and staining the spec-
imens are given in Baker et al. (1986). All
SEM observations are based on 8 last (Sth)
instar larvae.
16 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-5.
to the mandibles. 2.
several rows of stout, conical setae. C = campaniform sensillum; L = labrum; M = mandible; m = mola; P =
>
prostheca; Sc = sensillum chaeticum; St
RESULTS
The broad, rounded labrum has 24 sen-
silla chaetica that are mostly situated on the
posterior edge of the labrum where they
come in contact with the mandibles (Fig. 1).
Just above these sensilla, there are 2 cam-
paniform sensilla that are situated near the
mid-line of the labrum.
The mandibles have a tridentate apex, a
distinct mola, a small and pointed prosthe-
ca and a striated region near the mola (Figs.
Labrum and mandibles of L. mozardi. |. Distal row of sensilla chaetica on the labrum that extend
Mandible, apical view. 3. Mandible, ectal surface. 4. Molar region. 5. Stria composed of
= Stria; 1, 2, 3 = mandibular dentes.
2, 3, 4). The 2 apical teeth are similar in
size while the basal tooth is smaller. A pros-
theca is distad of the mola which is deeply
grooved and the grooves increase 1n density
from the basal to the distal region of the
mola. Laterad of the mola is a striated re-
gion (at low magnifications) but each stria
(at high magnifications) is composed of sev-
eral rows of blunt, stout spines (Figs. 3, 5).
Two sensilla chaetica and several campan-
iform sensilla are situated on the ectal sur-
face of the mandibles (Fig. 3).
VOLUME 91, NUMBER 1 17
Figs. 6-12. Labial and maxillary regions of the mouth parts. 6. Labium, maxillae, and mandibles. 7. Labial
palpus, apex. 8. Maxilla, inner surface. 9. Maxilla, outer surface. 10. Sensilla trichodea at base of maxillary
dentes. 11. Digitiform sensillum. 12. Sensilla on apex of maxillary palpus. Cp = coronal peg; DS = digitiform
sensillum; La = labium; M = mandible; Mx = maxilla; Sb = sensillum basiconicum (blunt); Sb, = sensillum
basiconicum (tapered); Ss = sensillum styloconicum; Str = sensillum trichodeum.
18 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
There is a total of 8 trichoid sensilla on
the ligula, mentum, and submentum of the
labium (Fig. 6). The labial palpi are 2-seg-
mented and the 2nd segment is longer. At
the apex of segment 2 there are 10 sensilla
(Fig. 7). The central sensory receptor is a
styloconic sensillum which has a short base
with a long peg (Fig. 7). There are 8 sensilla
basiconica, 7 that are blunt and one that has
a tapered tip (Fig. 7). These sensilla are lo-
cated on the periphery of the terminal palpal
segment. There is also one dome shaped
sensillum that is situated on a cuticular pro-
trusion (Fig. 7). This sensillum is known as
a coronal peg (Doane and Klingler 1978).
The maxilla is well developed with a
3-segmented palpus and a mala which is
long and curved with 2 large dentes at the
apex (Figs. 8, 9). Just below the dentes there
is a row of large sensilla chaetica on the
inner surface of the mala (Fig. 8), and on
the outer surface, 4 sensilla trichoidea are
located at the base of the dentes (Figs. 9,
10).
The maxillary palpi are 3-segmented and
the 3rd is the longest. There are 2 sensilla
chaetica on each palpal segment and | 1 sen-
silla are situated on the apex of segment 3
(Figs. 8, 9, 12). These sensilla are similar to
those on the apex of the labial palpus but
there are more tapered basiconic sensilla on
the maxillary palpus (Fig. 12). A single dig-
itiform sensillum 1s located on the inner sur-
face of the last palpal segment (Fig. 11). The
stipe and cardo are asperate (Fig. 8). There
are 2 pouch-like structures that are situated
between the submentum of the labium and
the cardo of each maxilla (Fig. 6). These
structures are also asperate.
There are 3 antennal segments and seg-
ment 2 is the longest (Fig. 13). Segment 1
is devoid of any hair-like sensory structures
but there are 4 campaniform sensilla (Figs.
13, 14). On the apical periphery of segment
2 there are 5 blunt trichoid sensilla (Figs.
14, 15). A single, long sensillum chaeticum
is situated on the outer, lateral surface of
subsegment 2 (Fig. 13). The antennal sen-
sory appendix is on the ventral side of seg-
ment 2 near the intersegmental membrane
between segments 2 and 3 (Fig. 15). Laterad
of this sensillum there are 2 sensilla basi-
conica (Fig. 15).
Segment 3 has a raised area at the apex
on which are situated 3 sensilla, a campan-
iform sensillum, and a trichoid and basi-
conic peg (Fig. 16). Just proximal of these
receptors are 3 trichoid sensilla (Fig. 16). A
placoid sensillum is situated on the ventral
surface of segment 3 near the antennal sen-
sory appendix (Fig. 15). Only the sensilla
chaetica and campaniformia that are situ-
ated on the mouth parts and antennae do
not stain with crystal violet or reduced sil-
ver, thus indicating their nonporosity. All
other sensilla that are mentioned pick up
these stains indicating that these sensilla
have a porous cuticular peg.
DISCUSSION
The close proximity of the labral sensilla
chaetica to the underlying mandibles indi-
cates that these sensilla probably act in
monitoring the position of the labrum and
mandibles during feeding. Campaniform
sensilla are known to act as stress receptors
and the 2 sensilla situated on the middle
portion of the labrum may function as cu-
ticular stress receptors.
In previous descriptions of larval langu-
riid mandibles (Comstock and Comstock
1916, Peterson 1951) there is no mention
of the structure of the molar surface and the
striae that occur laterad the mola. The deep-
ly grooved molar surface aids in grinding
plant material that is to be ingested. Several
rows of blunt spines comprise each of the
striae that are present on the mandibles.
They are also present on the same region of
Tribolium larvae (Kvenborg 1977). The
function of the striae is unknown, but they
could be involved in stridulation or groom-
ing. The 2 nonporous sensilla chaetica and
several campaniform sensilla on the ectal
surface of each mandible probably function
as those located on the labrum, to monitor
VOLUME 91, NUMBER |
Figs. 13-16.
cuticular stress and to relay information on
the position of the mouth parts in relation
to one another.
The general structure of the labium is
similar to the descriptions for other langu-
riid species (Wildermuth and Gates 1920,
Piper 1978). L. mozardi differs from Acro-
pteroxys gracilis (Newman) in the number
and types of sensilla situated on the apex of
the labial palpi. A. gracilis has 11-12 basi-
conic sensilla on the apex of the palpus (Pip-
er 1978), while L. mozardi has a total of 9-
10 sensilla but there are 2 types of basiconic
Larval antenna of L. mozardi. 13. Antenna, dorsal surface. 14. Antenna, ventral surface. 15.
Intersegmental region between antennal segments 2 and 3. 16. Antenna, apex. Asa = antennal sensory appendix;
C = campaniform sensillum; Ps = placoid sensillum; Sb = sensillum basiconicum; Str = sensillum trichodeum.
sensilla, a sensillum styloconicum and a co-
ronal peg.
A similar situation exists when one com-
pares the number and types of sensilla on
the maxillary palpal apex of L. mozardi and
A. gracilis. L. mozardi has 10-11 sensilla,
1 styloconic and coronal sensillum and 2
types of basiconic sensilla while A. gracilis
has 11-12 basiconic sensilla. The total
number of apical sensilla on the maxillary
(11) and labial (10) palpi of L. mozardi is
similar to what 1s found on 7ribolium lar-
vae (13 & 12) (Ryan and Behan 1973) and
20 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Hypera larvae (12 & 11) (Bland, 1983). But
the various types of apical sensilla on these
structures of L. mozardi resemble the apical
sensilla on wireworm larvae (Doane and
Klingler 1978).
The maxillary and labial apical basiconic
and styloconic sensilla stain with crystal vi-
olet and reduced silver indicating that these
sensilla have a porous cuticle. Also, behav-
ioral and electrophysiological studies on
other coleopterous larvae have shown that
similar receptors respond to CO, and other
chemicals and therefore may be considered
to be chemoreceptors (Klingler 1966, White
et al. 1974, Doane and Klingler 1978,
Mitchell 1978).
The external structure of the digitiform
sensillum is similar to what is found on oth-
er coleopterous larvae (Zacharuk et al. 1977,
Doane and Klingler 1978, Guse and Hono-
michl 1980, Honomichl 1980). Ultrastruc-
tural and electrophysiological evidence in-
dicate that this sensillum is a type of
mechanoreceptor that is involved in mon-
itoring the larva’s tunneling activity (Za-
charuck et al. 1977).
The total number of antennal sensilla on
L. mozardi (22) is similar to Tenebrio mol-
itor L. (24 sensilla) (Pierantoni 1969) and
Tribolium larvae (21 sensilla) (Behan and
Ryan 1978). The morphology of the basi-
conic and porous trichoid sensilla resembles
that found on other coleopterous larvae, and
ultrastructural studies indicate that these
sensilla are chemoreceptors (Behan and
Ryan 1978, Bloom et al. 1982a, b). The
antennal sensory appendix of L. mozardi is
similar in morphology and position on the
antenna as that on elaterid larvae (Zacharuk
1962). The fine structure of the sensory ap-
pendix suggests that it functions as an ol-
factory receptor (Scott and Zacharuk 1971).
The ultrastructure of the placoid sensillum
on tenebrionid larvae indicates a chemo-
sensory function (Behan and Ryan 1978,
Bloom et al. 1982), but electrophysiological
data is lacking.
ACKNOWLEDGMENTS
Assistance of Mr. D. G. McMinn, Bio-
logical Technician, and Vincent O. Mc-
Daniel during these studies is gratefully ac-
knowledged. Contribution of Mississippi
Agricultural and Forestry Experiment Sta-
tion in operation with USDA, ARS. Journal
Article No. 64/4 of the Mississippi Agri-
cultural and Forestry Experiment Station.
Research contributes to Southern Regional
Project No. S-201, Role of Legume Cover
Crops in Conservation Tillage Production
Systems. Mention of trademark or propri-
etary products does not constitute a guar-
antee or warranty of the products by the
U.S. Department of Agriculture and does
not imply approval to the exclusion of other
products.
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Behan, M. and M. F. Ryan. 1978. Ultrastructure of
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Bland, R. G. 1983. Sensilla on the antennae, mouth
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212:
Bloom, J. W., R. Y. Zacharuk, and A. E. Holodniuk.
1982a. Ultrastructure of the larval antenna of Te-
nebrio molitor L.(Coleoptera: Tenebrionidae):
Structure of the trichoid and uniporous peg sen-
silla. Can. J. Zool. 60: 1528-1544.
1982b. Ultrastructure of the larval antenna
of Tenebrio molitor L. (Coleoptera: Tenebrioni-
dae): Structure of the blunt-tipped peg and paillate
sensilla. Can. J. Zool. 60: 1545-1556.
Comstock, J. H. and A. B. Comstock. 1916. Manual
for the Study of Insects. Comstock Publ. Co., Ith-
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Doane, J. F. and J. Klingler. 1978. Location of CO,-
receptive sensilla on larvae of the wireworms
Agriotes lineatus-obscurus and Limonius califor-
nicus. Ann. Entomol. Soc. Am. 71: 357-363.
Guse, G. W. and K. Honomichl. 1980. Die digiti-
VOLUME 91, NUMBER 1
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(L.) and Hydrobius fuscipes (L.). Protoplasma 103:
55-68.
Honomichl, K. 1980. Die digitiformen Sensillen auf
dem Maxillarpalpus von Coleoptera I. Verglei-
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Klingler, J. 1966. Uber den Sitz der CO,-Receptoren
bei der Larve von Otiorhynchus sulcatus. Ento-
mol. Exp. Appl. 9: 271-277.
Kvenborg, J. E. 1977. Scanning electron microscopic
study of adult stored product beetle mandibles. J.
Assoc. Off. Analyt. Chem. 60: 1185-1209.
Mitchell, B. K. 1978. Some aspects of gustation in
the larval red turnip beetle, Entomoscelis ameri-
cana, related to feeding and host plant selection.
Entomol. Exp. Appl. 24: 340-349.
Peterson, A. 1951. Larvae of Insects, Part II. Edwards
Bros. Inc., Ann Arbor, MI, 416 p.
Pierantoni, R. L. 1969. Mechano- and chemo-recep-
tors in the antenna of Tenebrio molitor in the larval
stage. Proc. Elect. Micros. Soc. Am. 2: 246-247.
Piper, G. L. 1978. Life history of Acropteroxys gra-
culis (Coleoptera: Languriidae) on common rag-
weed in northeastern Ohio. Ohio J. Sci. 78: 304—
309.
21
Rose, R. I. and J. M. McCabe. 1973. Laboratory
rearing techniques for the southern corn root-
worm. J. Econ. Entomol. 66: 398-400.
Ryan, M. F. and M. Behan. 1973. The sensory re-
ceptors of Tribolium larvae. Physiol. Zool. 46: 238-
244.
Scott, D. A.and R. Y. Zacharuk. 1971. Fine structure
of the antennal sensory appendix in the larvae of
Ctenicera destructor (Brown) (Elateridae: Coleop-
tera). Can. J. Zool. 49: 199-210.
White, R. A., U. Paim, and W. D. Seabrook. 1974.
Maxillary and labial sites of CO,-sensitive recep-
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leoptera: Cerambycidae). J. Comp. Physiol. 88:
235-246.
Wildermuth, H. L. and F. H. Gates. 1920. Clover
stem borer as an alfalfa pest. USDA Bull. No. 889.
25 pp.
Zacharuk, R. Y. 1962. Sense organs of the head of
larvae of some Elateridae (Coleoptera): their dis-
tribution, structure and innervation. J. Morphol.
111: 1-34.
Zacharuk, R. Y., P. J. Albert, and F. W. Bellamy. 1977.
Ultrastructure and function of digitiform sensilla
on the labial palp ofa larval elaterid (Coleoptera).
Can. J. Zool. 55: 569-578.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 22-25
NORTH AMERICAN MOTHS DESCRIBED BY L. A. G. BOSC D’ANTIC
(LEPIDOPTERA: NOCTUIDAE, PYRALIDAE)
Scott E. MILLER AND V1TOR O. BECKER
(SEM) Bishop Museum, Box 19000-A, Honolulu, Hawaii 96817; (VOB) Centro de
Pesquisa Agropecuaria dos Cerrados, Caixa postal 70-0023, 73300-Planaltina, DF, Brasil.
Abstract.—Three species of moths were described from ‘Caroline’ by Bosc [1800].
Crambus adspergillus Bosc was recognized by Franclemont and Todd (1983) as Renia
adspergillus (Noctuidae), but the other two names have remained unplaced. We recognize
Alucita cereella Bosc as conspecific with Nola sorghiella Riley, n. syn. (Noctuidae) and
Pyralis sacculana Bosc as conspecific with C/ydonopteron tecomae Riley, n. syn. (Pyral-
idae).
Key Words:
The frenchman Louis Auguste Guillaume
Bosc d’Antic (1759-1828) lived in the
United States during the French Revolution
and collected insects, especially in South
Carolina (Blake 1952; Zimsen 1964: 16-17).
His collection, which contained many type
specimens including several of those de-
scribed by J. C. Fabricius, went to the Mu-
s¢um National d’Histoire Naturelle, Paris
(MNHN). At least two of the three species
described by Bosc [1 800] from North Amer-
ica were present in the MNHN around the
turn of the 18th century, because they were
illustrated by Coquebert (1801). However,
no Lepidopteran types from the Bosc col-
lection are now extant in the Paris museum
(Zimsen 1964, P. Viette pers. comm.).
Bosc [1800] described three species of
North American moths, but the paper was
overlooked for over a century. The part of
the journal in which Bosc’s paper appears
does not beara date, but is headed **Prairial,
an 8 de la République,” meaning it was pub-
lished in 1800, the eighth year of the French
Revolution (an earlier part bears the head-
ing “Germinel, an 6 de la République (Mars
1798)”). It appears that the first use of any
nomenclature, Nearctic, moths, Noctuidae, Pyralidae
of Bosc’s names is that of Kaye and Lamont
(1927), who applied one to a Trinidad
species of Chrysauginae (Pyralidae). Of the
three species described by Bosc [1800], only
one, Crambus adspergillus, has been in-
cluded in the North American literature
(Franclemont and Todd 1983).
Although the types are not available for
study, the descriptions and the figures pre-
sented by Bosc allow identification of the
species. We have also checked the relevant
bibliographies for homonymy and found
that the three names are nomenclatorally
available.
Two of the species, Crambus adspergillus
and Pyralis sacculana, were illustrated in
color by Coquebert de Montbret (1801), but
no mention of A/ucita cereella was included.
NOcTUIDAE
Renia adspergillus (Bosc)
Figs. 1, 4
Crambus adspergillus Bosc [1800]: 114, fig.
2 [Type locality: United States, “Caro-
line”’].—Coquebert 1801: 72, pl. XVII, fig.
1OA-B.
VOLUME 91, NUMBER |
Figs. 1-3.
Alucita cereella, Fig. 3: Pyralis sacculana.
Figs. 4-6.
cereella (Bosc); Fig. 6: Clydonopteron sacculana (Bosc). Figs. 4-6 approximately twice natural size.
Renia larvalis Grote 1872: 26 [Type local-
ity: United States, Pennsylvania and Tex-
as] [Synonymy by Franclemont and Todd
1983]
Renia restrictalis Grote 1872: 26 [Type lo-
cality: United States, Pennsylvania] [Syn-
onymy by Franclemont and Todd 1983]
Renia adspergillus (Bosc).—Franclemont
and Todd 1983.
Of the thirteen species of Renia recog-
nized in North America, the figure resem-
bles two species, previously described as fla-
vipunctalis (Geyer) and /arvalis (Grote). We
follow Franclemont and Todd (1983: 121),
Reproductions of illustrations from Bosc ({1800]: pl. VII), Fig. 1: Crambus adspergillus, Fig. 2:
Figures of specimens matching Bosc’s illustrations: Fig. 4: Renia adspergillus (Bosc); Fig. 5: Nola
who know the group well and were the first
revisers, in recognizing adspergillus as con-
specific with /arvalis.
Nola cereella (Bosc),
New ComMBINATION
Figs. 2, 5
Alucita cereella Bosc [1800]: 115, fig. 4 [Type
locality: United States, “Caroline’’]
Nola sorghiella Riley 1882: 187, pl. XI, fig.
1 [Type locality: United States, Alabama
and Florida], n. syn.
Nola portoricensis Moeschler 1890:
[Type locality: Puerto Rico], n. syn.
118
24 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Because of the small size of the figure and
the quality of printing of the plate, this was
the most difficult of the three Bosc species
to identify. The pattern shown in the plate
resembles species belonging to various fam-
ilies, and the plate shows two protruding
appendages in front of the head that could
be interpreted as either palpi or front legs.
The shape and pattern resemble those of
some species of Donacaula Meyrick (Pyr-
alidae), especially the species belonging to
the melinella group. The figure also resem-
bles the pattern of some species of Ethmia
Huebner (Oecophoridae), especially F. tri-
furcella (Chambers).
We have two main reasons for consid-
ering cereella as conspecific with sorghiella.
First, Bosc stated that it is a serious pest of
grain. Secondly, although his description of
the damage seems to be confused with that
caused by the Hessian fly (Mayetiola de-
structor (Say), Diptera: Cecidomyiidae), and
Sitotroga cerealella (Olivier) (Lepidoptera:
Gelechiidae), no other known grain pest re-
sembles the figure of cereella. Bosc’s original
description follows:
“Elle est cendrée. Les bords des ailes sont
brans et paroissent renflés. On voit sur le
disque de chaque aile supérieure deux points
bruns. Les ailes posterieures sont d’un gris
brillant.
“Cette Alucite est la teigne qui, sous le
nom d’Hessian fly, a fait, il ya environ douze
ans, de grandes ravages dans les bleds
d’Ameérique, et a menace de les étendre sur
ceux d’Europe, ce qui a oblige le parlement
d’Angleterre de proscrire les bleds qui veno-
ient de ce pays. Sa larve a seize pattes; elle
est d’un blanc verdatre. Elle mange d’abord
le germe du grain, puis la farine, ne laissant
que l’écorce. I] n’y a qu’une larve dans cha-
que grain: elle est trés-feconde et les gén-
érations se succédent rapidement. Au défaut
de bled, elle attaque le mais, et se multiplie
considérablement dans les greniers ot l’on
conserve ce grain.
“Le meilleur moyen pour détruire la larva
de cette Alucite, c’est de faire passer les grains
dans une étuve trés-chaude.”
PYRALIDAE
Clydonopteron sacculana (Bosc),
New ComMBINATION
Figs. 3, 6
Pyralis sacculana Bosc [1800]: 115, fig. 3A,
3B [Type locality: United States, ““Caro-
line”’].—Coquebert 1801: 71, pl. X VII, fig.
TA-C.
Clydonopteron tecomae Riley 1880: 286,
figs. 152-153 [Type locality: United
States, probably Missouri], n. syn.
There is no doubt that sacculana is the
same species described by Riley (1880) as
tecomae. The peculiar shape of the wings
and palpi, combined with the male struc-
tures on the costa of the forewing makes
this species readily recognized. Kaye and
Lamont (1927) and Amsel (1956) identified
specimens of Chrysauginae from Trinidad
and Venezuela as sacculana and placed it
in the genus Salobrena Walker. Cashatt
(1969) and Munroe (1983: 79), however,
regard Salobrena and Clydonopteron Riley
as distinct genera. Following their arrange-
ment, we decided to place sacculana in Cly-
donopteron. Specimens that are similar to
the North American sacculana have been
collected throughout the neotropics to
southern Brazil, and have been identified as
tecomae (Hampson 1897: 649, Lima 1950:
20). However, a full revision of the group
is needed to establish the number of species
involved, their distribution, synonymy, and
generic assignments. Also, we are not the
first to recognize the synonymy of sacculana
with tecomae. In the USNM collection there
isa label, added by Carl Heinrich, proposing
this synonymy, based on Coquebert (1801),
but Heinrich never published his discovery.
ACKNOWLEDGMENTS
We thank the Museum of Comparative
Zoology Library, Harvard University, for
providing copies of Bosc’s illustrations, and
the lepidopterists of the National Museum
of Natural History and Systematic Ento-
mology Laboratory, U.S. Department of
VOLUME 91, NUMBER 1
Agriculture (especially R. W. Hodges and
D. C. Ferguson), for their assistance in ver-
ifying the identities of Bosc’s species. R.
Stanger translated Bosc’s paper for us. The
photographs of specimens were taken by V.
Krantz of the Smithsonian Institution. M.
J. Scoble helped verify the date of Bosc
[1800]. Research for this paper was con-
ducted while both authors had Smithsonian
Institution fellowships. N. L. Evenhuis, D.
C. Ferguson, and R. W. Hodges reviewed
the manuscript.
LITERATURE CITED
Amsel, H. G. 1956. Microlepidoptera Venezolana.
Bol. Entomol. Venez. 10: 1-336, pl. I-CX.
Blake, D. H. 1952. American Chrysomelidae in the
Bosc collection (Coleoptera). Proc. Entomol. Soc.
Wash. 54: 57-68.
Bosc d’Antic, L. A. G. [1800]. Description de trois
espécies de Lépidoptéres de la Caroline. Bulletin
des Sciences, par la Societe Philomathique, Paris
[series 1] 2(39): 114-115, pl. VII. [German trans-
lation in Archiv fiir zoologie und zootomie 3(1):
188-190, 1802]
Cashatt, E. D. 1969. Revision of the Chrysauginae
of North America (Lepidoptera: Pyralidae) [ab-
stract only]. Diss. Abstr. (B) 29: 4696.
Coquebert de Montbert, A. J. 1801. Illustratio icono-
graphica insectorum quae in Musaeis Parisinis ob-
servavit et in lucem edidit Joh. Christ. Fabricius,
praemissis ejusdem descriptionibus; accedunt
25
species plurimae, vel minus aut nondum cognitae.
Volume II, pp. [45]-90, pl. XI-XX.
Franclemont, J. G. and E. L. Todd. 1983. Noctuidae,
pp. 120-159. In Hodges, R. W., ed., Check List
of the Lepidoptera of America north of Mexico.
London, E. W. Classey Ltd. xxiv + 284 pp.
Grote, A. R. 1872. Descriptions of North American
Noctuidae.—No. |. Trans. Amer. Entomol. Soc.
4: 20-28.
Hampson, G. F. 1897. On the classification of the
Chrysauginae, a subfamily of moths of the family
Pyralidae. Proc. Zool. Soc. London 1897: 633-
692, 74 figs.
Kaye, W. J. and N. Lamont. 1927. A catalogue of
the Trinidad Lepidoptera Heterocera (moths).
Mem. Dept. Agr., Trinidad and Tobago 3: vii +
144 + xv pp.
Lima, A. M. da Costa. 1950. Insetos do Brasil. 6.°
tomo. Lepidopteros. 2a parte. Rio de Janeiro, Es-
cola Nacional de Agronomia. 420 pp., 331 figs.
Moeschler, H. B. 1890. Die Lepidopteren-Fauna der
Insel Portorico. Abh. Senkenb. Naturforsch. Ges.
16: 77-360, 1 pl.
Munroe, E.G. 1983. Pyralidae, pp. 67-85. Jn Hodges,
R. W., ed., Check List of the Lepidoptera of Amer-
ica North of Mexico. London, E. W. Classey Ltd.
xxiv + 284 pp.
Riley, C. V. 1880. Ona new pyralid infesting the seed
pods of the trumpet vine. C/ydonopteron tecomae,
nov. gen., n. sp. Amer. Entomol. 3: 286-288.
1882. Report of the entomologist, in Report
of the Commissioner of Agriculture for the years
1881 and 1882. Washington, U.S. Government
Printing Office.
Zimsen, E. 1964. The type material of I.C. Fabricius.
Copenhagen, Munksgaard. 656 pp.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 26-28
THE INS AND OUTS OF A TROPICAL SOCIAL WASP NEST
CHRISTOPHER K. STARR
Department of Entomology, National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C. 20560 (Present address: Department of Horticulture, University
of Georgia, Athens, Georgia 30602).
Abstract. —An extraordinarily large nest of the social wasp Synoeca septentrionalis is
recorded from Costa Rica. The wasps had formed two holes through which they could
enter or leave the nest, unlike the usual single-hole condition in the genus. A statistical
test shows that one nest-hole was preferentially used for entering and the other for leaving,
which presumably increased the colony’s traffic efficiency. It is postulated that this ten-
dency was a statistical result of small individual behavior differences, so that it need not
require any organizing mechanism at the colony level.
Key Words:
Vespine and swarm-founding polistine
wasps (Hymenoptera: Vespidae), except for
the small genus Apoica, characteristically
nest either in secure cavities or, More com-
monly, construct an envelope around the
brood-combs (Jeanne 1975, Wenzel, in
press). This allows them to restrict access
to the interior of the nest. There have been
occasional observations of nests with two
or more entrance holes (Chopra 1925, Maidl
1934, Richards and Richards 1951, R. S.
Jacobson, pers. comm.; pers. obs.), and
Réaumur (1722) intimated that it is usual
for Vespula sp. (probably V. germanica and/
or vulgaris) nests to have two holes. It now
seems certain, though, that all social wasps
and bees with regular envelopes character-
istically have one entrance hole. This is ap-
parently taken for granted, although the nests
of ants and termites often have many such
holes.
Synoeca_ septentrionalis Richards is a
widespread, conspicuous, swarm-founding
polistine of Central and South America
(Richards 1978). Its nest begins as a single
comb flat against a tree trunk or limb or
nest, Synoeca, Vespidae, wasp
other substantial surface, surrounded by a
domed envelope with prominent ridges run-
ning across it and typically with a round
hole at the high end. As the colony grows,
it may add a new lobe to the nest at the high
end, obscuring the old hole and replacing it
with a new one on the new lobe. I have seen
occasional nests with two or three lobes in
Costa Rica; Rau (1933) mentioned a five-
lobed nest in Panama, and Buysson (1906)
figured one from Mexico with six lobes, ap-
parently the largest nest reported for the ge-
nus.
In early 1979, on a large fig tree (Ficus
sp.) near the headquarters of the Santa Rosa
National Park of Costa Rica, I found an
active S. septentrionalis nest with two re-
markable features. First, it consisted of nine
lobes with a total length of about 3 meters
(Fig. 1). Second, it had paired holes, each
of the usual form and in the usual position
(Fig. 2), with a combined perimeter of about
10 to 12 cm.
Réaumur (1722) reported that Vespula
workers consistently enter through one of
the two nest-holes and leave through the
VOLUME 91, NUMBER |
Fig. 1.
Nine-lobed nest of Synoeca septentrionalis
in Santa Rosa National Park, Costa Rica. It is on the
underside of a large branch at an angle of about 60°
from horizontal.
Fig. 2. The uppermost lobes of the nest, showing
the two entrance holes. Wasps on the envelope are
about 20 mm long.
27
other. Although the quantitative study of
behavior was nearly unknown in his time,
we know that Réaumur made original ob-
servations on Vespula colonies, and it is rea-
sonable to suppose that he found at least
one nest with two holes and watched it long
enough to gain an impression of directed
traffic. Indeed, if a colony has considerable
trafhe through a bottleneck, it makes bio-
logical sense that any separation of the in-
ward and outward streams will increase the
efficiency of passage. Accordingly, I pre-
dicted that the very large Santa Rosa Sy-
noeca colony would preferentially treat one
of its two nest-holes as an entrance and the
other as an exit.
Traffic at the nest was usually so heavy
that I could not simultaneously monitor the
direction of movement through both holes.
In each of eight observations periods, I se-
quentially recorded a) the hole-choice of 50
wasps without regard to whether they en-
tered or left the nest, b) direction of passage
of 25 wasps at one hole, and c) direction of
25 wasps at the other hole. The observation
periods were at various times when the
wasps were active over the course of 10 days
and totaled about two hours.
The hole-choice results (a) show greater
use of the right-hand hole, which was the
site of 280 (70%) of the 400 movements.
The direction-of-passage data (b and c) show
apparently more wasps leaving than enter-
ing the nest (Table 1). If the hole-choice bias
is entered as a correction factor (e.g. by mul-
tiplying the left-hole figures by ¥ or the right-
hole figures by % in Table 1), though, the
overall inward and outward traffic during
the observation time is found to be almost
Table 1. Cumulative numbers of wasps leaving and
entering the nest out of 25 moving through each hole
during each of eight observations periods.
Left Hole Right Hole Total
Entering 64 114 178
Leaving 136 86 222
Total 200 200
28 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Percentage breakdown of total traffic dur-
ing the observation time. Based on Table | and the
hole-choice results, as explained in text.
Left Hole Right Hole Total
Entering 9.6 39.9 49.5
Leaving 20.4 30.1 50.5
Total 30.0 70.0 100.0
identical. Table 2 thus gives the percentage
breakdown of the traffic which would pre-
sumably have been recorded if I had been
able to monitor all passages at once.
The bias-corrected results show a signif-
icant difference between the two holes in the
directional distribution of traffic (Chi-
square, P < 0.01). As seen in Table 2, it is
equivalent to what would be recorded if 20%
of the wasps consistently leave the nest by
the left hole and return by the right hole,
while the other 80% each tend to leave and
return by the same hole.
There is some indication in Synoeca spp.
ofan unusual flexibility in nest construction
(Vecht 1967, Overal 1982), which may ac-
count for this nest’s two-holed condition. It
makes little difference here whether this
originated and was maintained as an adap-
tive response to extraordinarily large col-
ony-size or through ordinary building errors.
On the other hand, the origin of the ten-
dency toward a plausibly more efficient flow
of traffic is pertinent. There is no evidence
for a higher organizing mechanism in Sy-
noeca which could account for this, and we
need not postulate any. The most parsi-
monious hypothesis is that the small be-
havioral differences which unavoidably arise
between individuals have been summed to
produce a meaningful phenomenon at the
colony level.
ACKNOWLEDGMENTS
My stay in Costa Rica was supported by
a graduate assistantship from the Depart-
ment of Entomology, University of Geor-
gia. A grant-in-aid from Sigma Xi provided
research supplies. I am grateful to Jorge
Morales and other staffof the National Parks
Service of Costa Rica for making it con-
venient and pleasant to work in Santa Rosa.
Criticism from Bob Jeanne, John Wenzel
and the journal’s reviewers led to substan-
tial improvements in the paper.
LITERATURE CITED
Buysson, R. Du. 1906. Monographie des vespides
appartenant aux genres Apoica et Synoeca. Ann.
Soc. Entomol. France 75: 333-362.
Chopra, B. 1925. Notes on a nest of the common
Indian hornet, Vespa cincta Fabr. J. Bombay Nat.
Hist. Soc. 30: 858-860.
Jeanne, R. L. 1975. The adaptiveness of social wasp
nest architecture. Quart. Rev. Biol. 50: 267-287.
Maidl, F. 1934. Die Lebensgewohnheiten und In-
stinkte der staatenbildenden Insekten. Vienna: Fritz
Wagner 823 pp.
Overal, W. L. 1982. Acoustical behavior and variable
nest architecture in Synoeca virginea (Hymenop-
tera, Vespidae). J. Georgia Entomol. Soc. 17: 1-4.
Rau, P. 1933. Jungle Bees and Wasps of Barro Col-
orado Island. Kirkwood, Missouri, Publ. by au-
thor 324 pp.
Reaumur, R. A. F. de. 1722. Histoire des guépes.
Mem. Acad. R. Sci. Paris 21: 302-364.
Richards, O. W. 1978. The Social Wasps of the
Americas, Excluding the Vespinae. London, Brit-
ish Museum (Natural History) 580 pp.
Richards, O. W. and M. J. Richards. 1951. Obser-
vations of the social wasps of South America (Hy-
menoptera, Vespidae). Trans. Entomol. Soc. Lon-
don 102: 1-170.
Vecht, J. van der. 1967. Bouwproblemen van sociale
wespen. Verh. K. Ned. Akad. Wetensch. (Afd. Na-
tuurkunde) 76: 59-68.
Wenzel, J. W. In press. Evolution of nest architecture
in social vespids. /n K. G. Ross, and R. W. Mat-
thews, eds., The Social Biology of Wasps. Ithaca,
Cornell Univ. Press.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 29-34
NOTES ON THE GENUS HYBRIZON IN NORTH AMERICA
(HYMENOPTERA: PAXYLOMMATIDAE)
PAUL M. MARSH
Systematic Entomology Laboratory, U.S. Department of Agriculture, Agricultural Re-
search Service, % U.S. National Museum of Natural History, NHB 168, Washington,
D.C. 20560.
Abstract.—The two North American species of the unusual genus Hybrizon Fallén are
redescribed from a large collection of specimens made in Virginia. Brief comments are
made on the taxonomic placement of the genus and on the observed sex ratio of the
collected material.
Key Words:
The genus Hybhrizon Fallen is one of the
most peculiar and taxonomically confusing
groups in the Ichneumonoidae. It contains
seven species (five Palearctic, two Nearctic)
and is the only member in the family Paxy-
lommatidae except for one undescribed ge-
nus from Japan. Because it lacks a second
recurrent vein in the fore wing, the genus
has often been classified as a subfamily of
the Braconidae (Wesmael 1835, Curtis 1837,
Haliday 1840, Muesebeck and Walkley
1951, Marsh 1963, Shenefelt 1969, van
Achterberg 1976, Watanabe 1984). How-
ever, it also has been classified as a subfam-
ily of the Ichneumonidae (Rasnitsyn 1980,
Gauld 1984), or in a distinct family (Wa-
tanabe 1946, Tobias 1968, Marsh 1971,
1979, Mason 1981, van Achterberg 1984,
Marsh et al. 1987). Mason (1981) argued
convincingly that Hybrizon should be ex-
cluded from the Braconidae because it lacks
a critical synapomorphy of the family,
namely, the fusion of abdominal terga 2 and
3. Furthermore, van Achterberg (1984) gave
two synapomorphies of wing venation that
show the Paxylommatidae are more closely
related to the Ichneumonidae than to the
Braconidae. The same conclusion was
taxonomy, Ichneumonoidea, ant-parasites
reached by Sharkey and Wahl (1987), who
suggested that Hybrizon might be placed
within the Ichneumonidae. This action had
already been proposed by Rasnitsyn (1980)
who classified Hybrizon as a subfamily of
the Ichneumonidae. However, Mason
(1981) argued against this in favor of a sep-
arate family classification, the Paxylom-
matidae, and I have followed his classifi-
cation in this paper.
During the summers of 1986 and 1987,
my colleague, David R. Smith, operated
several Malaise traps in two locations in
Virginia, at his home in Annandale (a sub-
urb of Washington, D.C.) and near Cuckoo
in Louisa County. Approximately 200 spec-
imens of Hybrizon were collected during
these two years representing two species.
Prior to this the U.S. National Museum
contained only about 50 specimens of the
genus. Approximately 7 of the specimens
collected by Smith are ri/eyvi (Ashmead); the
other '/ are a distinct species which I thought
was undescribed but now have identified as
the previously unknown female of /lavo-
cinctus (Ashmead). I have provided descrip-
tions and a key to separate the species be-
low. Additional specimens were borrowed
30 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
from the Canadian National Collection, Ot-
tawa, Canada (M. Sharkey), the American
Entomological Institute, Gainesville, Flor-
ida (H. Townes), the Museum of Compar-
ative Zoology, Harvard University, Cam-
bridge, Massachusetts (S. Shaw), and the
Riksmuseum van Natuurlike Histoire,
Leiden, The Netherlands (C. van Achter-
berg).
The biology of these unusual wasps has
not been satisfactorily established. They are
associated with ant nests and are likely to
be endoparasitoids of ant larvae. Donis-
thorpe and Wilkinson (1930) give the most
extensive review of the biology.
Of interest is the high ratio of females to
males of the North American species in the
National Collection, 241:14 in ri/eyi and
137:5 in flavocinctus. Female biased sex ra-
tios are predicted by the local mate com-
petition (LMC) model of Hamilton (1967),
. where females place offspring in dis-
crete patches of the resource (in this case,
ant colonies), and those offspring mate ran-
domly in their patch before female offspring
disperse to colonize new patches” (Waage
1985). On the other hand, the observed fe-
male biased sex ratio could merely be an
artifact of collecting techniques, assuming
that mating occurs in or near ant colonies
and females searching for new colonies are
the main dispersers. Thus, random sweep-
ing or flight intercept traps would produce
mostly females, whereas collections made
in ant colonies might yield a more balanced
sex ratio.
Family Paxylommatidae
Pachylommatoidae Foerster, 1862: 247.
Oldest family-group name (see Mason
1981 for discussion).
Hybrizon Fallen
Hybrizon Fallén, 1813, p. 19. No species.
Type-species: Hybrizon latebricola Nees,
1834. Monotypic, first included species
by Nees (1834:28).
Paxylomma de Brébisson, 1825: 23. Type-
species: Paxylomma buccatum de Bréb-
bison. Monotypic. Synonymy by Wes-
mael, 1835.
Plancus Curtis, 1833: 188. Type-species:
Plancus apicalis Curtis. Monotypic. Syn-
onymy by Stephens, 1835.
Eurypterna Foerster, 1862: 247. Type-
species: Paxylomma cremieri Romand.
Monotypic. Synonymy by Marshall, 1891.
Eupachylomma Ashmead, 1894: 58. Type-
species: Wesmaelia rileyi Ashmead. Orig-
inal designation. Synonymy by Watana-
be, 1935.
Ogkosoma Haupt, 1913: 52. Type-species:
Ogkosoma schwarzi Haupt. Monotypic.
Synonymy by Strand, 1914.
The names Paxyloma (Stephens 1835),
Paxylomme (Wesmael 1835), Paxyllomma
(Curtis 1837), Paxylloma (Blanchard 1840),
and Pachylomma (Ratzeburg 1848) are all
to be considered emendations of Paxylom-
ma (see Shenefelt 1969 and Mason 1981).
Because of the small size of these wasps
and the lack of a second recurrent vein in
the fore wing, Hybrizon will key to Braconi-
dae in most general textbooks with keys to
Hymenoptera families. In view of this, 7y-
brizon was included by Marsh et al. (1987)
in their identification manual for North
American genera of Braconidae. The genus
can be diagnosed by reference to couplet |
of that key and the associated figures. Adult
Hybrizon have a distinctive habitus (Fig. 3):
narrow head with bulging eyes and deep an-
terior tentorial pits (Figs. 6, 7), strongly
arched thorax, long spindly legs, and long
thin abdomen.
The two North American species of Hy-
brizon can be separated by the following
key.
Ocelli small, ocell-ocular distance at least equal to
diameter of lateral ocellus, often greater (Fig. 9);
first segment of radius in fore wing shorter than
first segment of discoideus and about ' length of
recurrent vein, branchial cell not as tall as disco-
cubital cell (Fig. 1); head, thorax and abdomen
usually entirely black rileyi (Ashmead)
Ocelli larger, ocell-ocular distance less than diam-
eter of lateral ocellus, often less than half (Fig. 8);
VOLUME 91, NUMBER |
disco-cubital
Figs. 1, 2.
first segment of radius equal to or longer than dis-
coideus and about *% length of recurrent vein; bra-
chial cell equal in height to disco-cubital cell (Fig.
2); head black, at least pronotum, mesopleuron
and base of abdominal terga 3 and 4 honey yellow,
sometimes thorax and abdomen extensively
marked with honey yellow . . flavocinctus (Ashmead)
Hybrizon rileyi (Ashmead)
Figs; 3,4:.7; 9
Wesmaelia rileyi Ashmead, 1899: 641. Ho-
lotype female in U.S. National Museum,
Washington, D.C.
Female. Length of body, 2-3 mm. Color:
head black, clypeus and mouthparts white;
antennal scape and pedicel yellow, flagellum
black; thorax black or dark brown, rarely
deep honey yellow; legs yellow with hind
femur, tibia, and coxa often light brown;
tegula yellow; abdomen black or dark brown,
rarely basal segments dark honey yellow.
Head: very weakly reticulate, smooth and
shining; ocellar-ocular distance equal to or
greater than diameter of lateral ocellus (Fig.
9): clypeus lengthened, apical margin well
below level of lower eye margin, malar space
slanted (Fig. 7); antenna with 11 flagello-
meres. Thorax: pro and mesothorax smooth
and shining; propodeum irregularly rugose,
without any indication of median longitu-
dinal carina. Abdomen: terga smooth and
shining, terga 1 and 2 sometimes weakly
striate at base (Fig. 4). Wings (Fig. 1): first
segment of radius shorter than first segment
31
Wings of Hybrizon species. |, H. rileyi(Ashmead). 2, H. flavocinctus (Ashmead) (scale = 0.5 mm).
of discoideus and about '2 length of recur-
rent vein, brachial cell not as tall as disco-
cubital cell.
Male. Essentially similar to female.
Type locality. UNITED STATES: Ox-
ford, Indiana.
Material examined. 241 99, 14 4¢ from
the following states and provinces: District
of Columbia, Georgia, Indiana, Iowa, Kan-
sas, Maine, Maryland, Michigan, New
Hampshire, New Jersey, New York, North
Carolina, Nova Scotia, Ontario, Pennsyl-
vania, Quebec, South Carolina, Virginia,
West Virginia, Wisconsin.
Biology. The type material is recorded as
being reared from Toxopotera (= Schiza-
phis) graminum, but this is probably not
correct. Three specimens from New Hamp-
shire are labelled “Attracted to disturbed
nest of Lasius alienus.”
This species is easily distinguished from
flavocinctus by its darker color, smaller
ocelli, and wing venation.
Hybrizon flavocinctus (Ashmead)
Figs. 2, 5, 6, 8
Eupachylomma flavocincta Ashmead, 1894:
59. Holotype female in U.S. National
Museum, Washington, D.C.
Female. Length of body, 3.5—-4 mm. Col-
or: head black, clypeus and mouth parts light
yellow; antennal scape and pedicel yellow,
flagellum black; prothorax honey yellow;
32 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Mj
a tt
Hybrizon species. 3, H. rileyi (Ashmead), habitus (scale = 2 mm). 4, H. rileyi, abdominal terga
1-2 (scale = 500 u). 5, H. flavocinctus (Ashmead), abdominal terga 1-2 (scale = 500 yu). 6, H. flavocinctus, face
Figs. 3-9.
(scale = 100 uw). 7, H. rileyi, face (scale
(scale = 200 x).
mesonotum dark brown or black, some-
times with yellow longitudinal lines; scu-
tellum yellow with brown spot at base:
mesopleuron varying from entirely brown
to yellow; propodeum dark brown; tegula
yellow; legs yellow, hind femur, tibia, and
coxa light brown; abdomen brown, terga 3
and 4 yellow at base. Head: reticulate and
200 uw). 8, H. flavocinctus, vertex (scale = 200 yu). 9, H. rileyi, vertex
dull; ocell-ocular distance less than diam-
eter of lateral ocellus, inner edge of each
ocellus margined by a scrobiculate groove
(Fig. 8); clypeus short, apical margin only
slightly below level of lower eye margin,
malar space nearly horizontal, eyes bulging
below (Fig. 6); antenna with 11 flagello-
meres. Thorax: pro- and mesothorax smooth
VOLUME 91, NUMBER 1
and shining; propodeum irregularly rugose,
often with a short median carina. Abdomen:
first and second terga usually distinctly
striate (Fig. 5), rest of terga smooth and
shining. Wings (Fig. 2): first segment of ra-
dius equal to or longer than discoideus and
about *% length of recurrent vein, brachial
cell about as tall as discocubital cell.
Male. Essentially as in female, occasion-
ally body mostly honey yellow.
Type locality. UNITED STATES: Wash-
ington, D.C.
Material examined. 137 99, 5 64 from the
following states and provinces: District of
Columbia, Maryland, Michigan, New York,
Ontario, Virginia, Wisconsin.
Biology. Unknown.
Prior to this study, the only authentically
determined specimen of flavocinctus was the
male holotype. The large number of female
specimens collected in Virginia were gen-
erally much darker in color than the holo-
type and I had thought them to be an un-
described species. After closer examination,
they agree morphologically with the holo-
type and I now consider them to be the
undescribed female of flavocinctus.
This species differs from ri/ey/ in its larger
size, larger ocelli, generally lighter body col-
or, and wing venation. It is also very similar
to the European buccatus (de Brébisson)
which is distinguished by its darker body
color, by having stronger sculpturing on the
head which is almost punctate, and by hav-
ing a few punctures on the mesonotum along
where the notauli would be.
ACKNOWLEDGMENTS
Scott Shaw, Museum of Comparative Zo-
ology, Harvard University, read the manu-
script and offered many helpful suggestions
for improvement, especially the sections on
classification and biology, for which I am
grateful. David Wahl, American Entomo-
logical Institute, Gainesville, Florida and
Robert Smiley, Systematic Entomology
Laboratory, Beltsville, Maryland also of-
fered useful comments on the manuscript.
33
Thanks also go to David Smith for oper-
ating the Malaise traps that produced the
specimens which motivated this study.
LITERATURE CITED
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1984. Essay on the phylogeny of Braconidae
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Ashmead, W. H. 1889 (1888). Descriptions of new
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1894. Notes on the family Pachylommato-
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Blanchard, C. E. 1840. Histoire nataurelle des ani-
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de Brébisson, L. A. 1825. Paxylomme, Paxylomma.
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1837. A guide to an arrangement of British
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Notes on the genus Paxylomma (Hym. Brac.), with
the description of a new species taken in Britain.
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Fallen, C. F. 1813. Spec. Nov. Hym. Disp. Meth., p.
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Foerster, A. 1862. Synopsis der Familien und Gat-
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Westwood, J O., ed., An Introduction to the
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Hamilton, W. D. 1967. Extraordinary sex ratios. Sci-
ence 156: 477-488.
Haupt, H. 1913. Beitrage zur Hymenopteran-Fauna
von Halle a. S. und Ungengend. Mitt. Entomol.
Ges. Halle 5-7: 47-62.
Marsh, P.M. 1963. A key to the Nearctic subfamilies
of the family Braconidae (Hymenoptera). Ann.
Entomol. Soc. Amer. 56: 522-527.
1971. Keys to the genera of the families Bra-
conidae, Aphidiidae, and Hybrizontidae (Hyme-
noptera). Ann. Entomol. Soc. Amer. 64: 841-850.
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1979. Family Braconidae, pp. 144-295. In
Krombein, K. V. et al., eds., Catalog of Hyme-
nopterain America North of Mexico. Washington,
D.C., Smithsonian Institution Press.
Marsh, P. M., S. R. Shaw, and R. A. Wharton. 1987.
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genera of the family Braconidae (Hymenoptera).
Mem. Entomol. Soc. Wash. 13, 98 pp.
Marshall, T. A. 1891. Les Braconides. Jn Andre, E.,
Species de Hyménoptéres d’Europe et d’Algerie,
vol. 5, 635 pp.
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PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 35-50
THE GENERA ATAENOGERA AND PHYCUS IN THE NEW WORLD
(DIPTERA: THEREVIDAE: PHYCINAE)
DONALD W. WEBB AND MICHAEL E. IRWIN
Illinois Natural History Survey, 607 East Peabody Drive, Champaign, Illinois 61820.
Abstract. —Two of the three species names available for the New World genus Ataenog-
era Krober are synonymized in this paper. The genus Phycus Walker, previously unknown
from the New World, is represented here by two new species. A key to the New World
species of these two genera is provided along with species descriptions and distributions.
Descriptions of the pupal stage of Ataenogera abdominalis and of the larva and pupa of
Phycus frommeri n. sp. are included.
Key Words:
distribution
In a revisionary monograph on the
Nearctic genera of Therevidae, Irwin and
Lyneborg (1981) described several new gen-
era, redescribed others, and assigned an un-
described therevid species from northwest-
ern Mexico and southern California to the
genus Phycus. They separated the genera
Phycus Walker and Ataenogera Krober on
the basis of several morphological charac-
teristics and suggested that species of Atae-
nogera have no close relatives in the Old
World other than Phycus. Subsequently,
Lyneborg (1983) characterized and critical-
ly evaluated the Old World species of Phy-
cus.
Because Phycus was unknown from the
New World prior to Irwin and Lyneborg’s
study, we undertook to compare that genus
to the closely related Ataenogera. The species
in these genera are similar in gross structure
and are placed together within the subfam-
ily Phycinae. Because Ataenogera and Phy-
cus are Clearly separated as a new clade from
such New World phycine genera as Heni-
comyia Coquillett on the one hand, and
Pherocera Cole, Schlingeria Irwin, and Par-
apherocera Irwin on the other (Irwin and
Diptera, Therevidae, Phycus, Ataenogera, New World, species descriptions,
Lyneborg 1981), we elected to describe and
diagnose their New World species here in a
single publication.
METHODS
The morphological terminology used for
the male terminalia was originally defined
and described by Lyneborg (1968) and sub-
sequently modified by Lyneborg (1972,
1976, 1978) and Irwin (1977a, b); that for
the female terminalia was defined and de-
scribed by Irwin (1976); that for the larva
and pupa follows Irwin (1972). Other mor-
phological features are described in termi-
nology generally accepted in the literature
on Diptera. The range for each measure-
ment and ratio is followed by the average;
the range for each setal count is followed by
the mode.
Each specimen was assigned a THER-
EVIDAE/M. E. IRWIN/SPECIMEN num-
ber attached to the specimen. This number
is used to associate the ecological and label
data with a given specimen and is printed
in italics in this paper. The data are incor-
porated into an automated data manage-
36 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ment system originally designed by Rauch
(1970).
To conserve space and include as much
information as possible about each speci-
men, a layout adopted from Irwin (1983) is
used in the “Specimens Examined” section
of each species. Many of the terms used
there are explained by Stuckenberg and Ir-
win (1973). The layout follows these typo-
graphical conventions:
1) Full capitals: LARGEST POLITICAL
UNIT (country, or state within the United
States).
2) Boldface type: intermediate political
unit (state or province outside the United
States or county within the United States)
and elevation expressed 1n m above sea level.
3) Roman or normal print: smallest po-
litical unit (city or town) and modifier of
that unit (distances in km, direction, and
subunits of that unit).
4) Collector names. Acronyms were used
for the following collectors: M. E. Irwin,
MEI; R. M. Worley, RMW;; S. I. Frommer,
SIF.
5) Numbers of specimens is followed by
the sex designation M for male or F for
female.
6) A semicolon terminates one series of
specimens and signals the beginning of the
next. Data not repeated in a subsequent se-
ries are the same as those of preceding se-
ries.
Depositories.—Paratypes of Phycus
frommeri n. sp. have been deposited in the
following museums: AMNH, American
Museum of Natural History, New York;
ANSP, Academy of Natural Sciences,
Philadelphia; ASU, Arizona State Univer-
sity, Tempe; AMS, Australian Museum,
Sydney; BMH, Bernice P. Bishop Museum,
Honolulu, Hawaii; BMNH, British Mu-
seum of Natural History, London; BYU,
Brigham Young University, Provo, Utah;
CAS, California Academy of Sciences, San
Francisco; CIS, California Insect Survey,
University of California, Berkeley; CMNH,
Field Museum of Natural History, Chicago;
CMP, Carnegie Museum, Pittsburgh; CNC,
Canadian National Collection, Ottawa;
CSDA, California State Department of Ag-
riculture, Sacramento; CSIRO, Common-
wealth Scientific Industrial Research Or-
ganization, Canberra, Australia; CSU,
Colorado State University, Fort Collins; CU,
Cornell University, Ithaca; DEI, Deutsches
Entomologische Institut, Berlin, East Ger-
many; DSIR, Department of Scientific and
Industrial Research, Nelson, New Zealand;
DZSA, Departamento de Zoologia Agricul-
tura, Sao Paulo, Brazil; EEA, Estacion Ex-
perimental Agronomica, Universidad de
Chile, Maipu; FSCA, Florida State Collec-
tion of Arthropods, Gainesville; IAS, Insti-
tute of Agricultural Sciences, Tokyo; IE, In-
stituto di Entomologia, Bologna, Italy; IML,
Instituto Miguel Lillo, Tucuman, Argen-
tina; INHS, Illinois Natural History Survey,
Champaign; INIA, Instituto Nacional de
Investigaciones Agricolas, Chapingo, Mex-
ico; IOC, Instituto Oswaldo Cruz, Rio de
Janeiro, Brazil; IRSN, Institut Royal des
Sciences Naturelle de Belgique, Brussels;
ISU, Iowa State University, Ames; ITM,
Instituto Technologico y de Estudios Super-
iores, Monterrey, Mexico; KSU, Kansas
State University, Manhattan; KUF, Kyu-
shu University, Fukuoka, Japan; LACM,
Natural History Museum of Los Angeles
County, Los Angeles; Museum of Compar-
ative Zoology, Harvard University, Cam-
bridge, Massachusetts; MEI, M. E. Irwin
Collection, MHN, Museo de Historia Nat-
ural Javier Prado, Lima, Peru; MMB, Mo-
ravske Museum, Brno, Czechoslovakia;
MNH, Musei Nationalis Hungarici, Buda-
pest, Hungary; MNHN, Museum National
d’Histoire Naturelle, Paris, France; MSU,
Michigan State University, East Lansing;
NCSU, North Carolina State University,
Raleigh; NMB, Naturhistorisches Museum,
Basel, Switzerland; NMP, Natal Museum,
Pietermaritzburg, South Africa; NMSU,
New Mexico State University, Las Cruces;
Nevada State Department of Agriculture,
Reno; OSM, Ohio State Museum, Colum-
VOLUME 91, NUMBER |
bus; OSU, Oregon State University, Cor-
vallis,; PAS, Polish Academy of Sciences,
Warsaw, Poland; RNHL, Rijkmuseum van
Natuurlijke Historie, Leiden, The Nether-
lands; SDAP, State Department of Agri-
culture, Harrisburg, Pennsylvania; SDCM,
San Diego County Museum, San Diego;
SJSC, San Jose State University, San Jose,
California; SMN, Staatlichen Museum fiir
Naturkunde, Stuttgart, West Germany;
SWRS, Southwestern Research Station
(AMNH), Portal, Arizona; TAM, Texas Ag-
ricultural and Mechanical University, Col-
lege Station, UA, University of Arizona,
Tucson; UAC, University of Alberta, Ed-
monton, Alberta, Canada; UBC, University
of British Columbia, Vancouver, Canada;
UCD, University of California, Davis;
UCM, University of Colorado Museum,
Boulder; UCR, University of California
Riverside; UCVM, Universidad Central de
Venezuela, Maracay, Venezuela; UG, Uni-
versity of Georgia, Athens; UI, University
of Idaho, Moscow; UK, University of Kan-
sas, Lawrence; UM, University of Minne-
sota, Saint Paul; UMA, University of Mich-
igan, Ann Arbor; UNLP, Museo de Ciecias
Naturales, Universidad Nacional de La Pla-
ta, La Plata, Argentina; USI, University of
Southern Illinois, Carbondale; USNM,
United States National Museum, Washing-
ton, D.C.; USSR, Zoological Institute USSR,
Leningrad; USU, Utah State University,
Logan; UTA, University of Texas, Austin;
UTI, University of Tel Aviv, Israel; UU,
University of Utah, Salt Lake City; UW,
University of Wisconsin, Madison; UZM,
Universitetets Zoologiske Museum, Copen-
hagen, Denmark; VNM, Naturhistorisches
Museum Vienna, Austria; WSU, Washing-
ton State University, Pullman; ZIB, Zool-
ogisches Institut, Berlin, West Germany;
ZSI, Zoological Survey of India Collection,
Calcutta, India. The paratypes of Phycus
frontalis n. sp. are deposited in the following
museums: AMNH, American Museum of
Natural History, New York; INHS, Illinois
Natural History Survey, Champaign;
37
USNM, United States National Museum,
Washington, D.C.
Key TO NEw WoRrLD SPECIES OF
ATAENOGERA AND PHYCUS
1. Maxillary palpus one-segmented (Fig. 3); eye
margins distinctly divergent from level of ocel-
lar tubercle toward genae (Fig. 2); discal cell
pointed basally (Fig. 4); sternites 5-8 in males,
5-7 in females (Fig. 10) with numerous lan-
ceolate setae ventrally; male genitalia with large,
subtriangular hypandnum (Fig. 8) oe
sult Sichags eater ans a Ataenogera abdominalis Krober
- Maxillary palpus two-segmented (Fig. 18); eye
margins divergent slightly from level of ocellar
tubercle toward genae (Fig. 17); discal cell trun-
cate basally (Fig. 19); sternites lacking lanceo-
late setae; male genitalia lacking hypandrium
or much reduced. (Phycus Walker) .......... 2
2. Eyes separated dorsally by distance greater than
width of ocellar tubercle (Fig. 17); width of
frons at level of lateral ocelli greater than 0.5
times width at level of antennal bases; pos-
terolateral extensions of gonocoxites short,
thick in ventral view (Fig. 23)
: . Phycus frommeri n. sp.
- Eyes sepamited dorsally only by width of ocellar
tubercle (Fig. 35); width of frons at level of
lateral ocelli less than 0.5 times width at level
of antennal bases; posterolateral extensions of
gonocoxites narrow, attenuate in ventral view
(Fig. 39) Phycus frontalis n. sp.
Superficially, the genus Afaenogera 1s
morphologically similar to Phycus, the two
genera are however quite distinct. The best
distinguishing characteristics are: (a) max-
illary palpus two-segmented in Phycus, one-
segmented in Ataenogera; (b) fore coxae have
1-2 apical setae in Phycus, 4-6 apical setae
in Ataenogera; (c) ventral epandrial sclerite
fused anteriorly with epandrium in Phycus,
fused laterally with epandrium in Alaenog-
era; (d) hypandrium extremely small or ab-
sent in Phycus, large in Ataenogera; (e) ven-
tral apodeme of aedeagus composed of two
long, narrow rods in Phycus, very reduced,
appearing absent in Afaenogera.
Genus Ataenogera Krober
Ataenogera Krober (1914: 31):
(1932: 255). Type species:
Krober, by monotypy.
Malloch
A. abdominalis
Figs. 1-15. Ataenogera abdominalis.—1. Antenna (5747). 2. Head of male, frontal view (5747). 3. Maxillary
palpus, lateral view (5747). 4. Wing, dorsal view (5747). 5. Male tergite 8, epandrium, cerci, and ventral epandrial
sclerite, dorsal view (576/). 6. Male ventral epandrial sclerite, ventral view (576/). 7. Male gonocoxites and
gonostylus, dorsal view (576/). 8. Male gonocoxites and gonostylus, ventral view (576/). 9. Male aedeagus,
lateral view (5761). 10. Female abdomen, terminal segments, lateral view with enlargement of ventral lanceolate
setae (5752). 11. Female terminalia, dorsal view (5752). 12. Female terminalia, ventral view (5752). 13. Pupal
exuvium lateral view (5764). 14. Pupa exuvium, ventral view (left), dorsal view (right) (5764). 15. Frontal plate
(5764). Abbreviations: (AS) antennal sheath; (C) cercus; (DA) dorsal apodeme; (Dp) distiphallus; (DSp) dorsal
spiracle; (EA) ejaculatory apodeme; (Epa) epandrium; (F) furca; (Gs) gonostylus; (Gx) gonocoxites; (Hpa) hy-
pandrium; (S,) sternite 8; (T,) tergite 8; (T.) tergite 9; (VES) ventral epandrial sclerite. Scale = 0.1 mm, unless
otherwise indicated. 38
VOLUME 91, NUMBER |
Leptocera Kroéber (1928: 117). Type species:
L. gracilis Kréber by monotypy. Preoc-
cupied by Olivier (1813: 489).
Ziehenia Krober (1929: January: 434). New
name for Leptocera Krober.
Epileptocera Richards (1929: August: 171).
Unjustified new name for Leptocera
Krober.
Moderate-sized, slender flies.
Head.—Frons (Fig. 2) of both sexes at its
narrowest as wide as ocellar tubercle, to-
mentum brownish gray; setae sparse, dor-
solateral to antennal bases and in males in
two lateral rows ventral to ocellar tubercle;
male eyes dichoptic, eye margin diverges
distinctly from level of ocellar tubercle to-
wards gena, facets equal, setae absent; para-
facial lacking setae; head slightly protrudes
at antennal level; facial and genal callus ab-
sent; length of antenna |.3-2.0 times head
depth; antenna long (Fig. 1); scape cylin-
drical; pedicel subtriangular; flagellum
elongate, tapered apically, setae absent; style
terminal, two-segmented, with minute ter-
minal spine; clypeus concave, setae absent;
maxillary palpus one-segmented (Fig. 3).
Thorax.—np 2-3, sa 1, pa 1, dc 0-1, sc
1. Vittae absent; mesonotal setae moder-
ately long, subappressed; prosternum bare
in and around central depression; pleural
setae elongate, scattered on propleuron and
dorsal fourth of anepisternum. Wing (Fig.
4).—Ground color hyaline to pale brown,
clouding apparent near apex; veins brown;
pterostigma absent; setulae dorsal on R,;
subcostal cell and cell r, elongate, open; veins
R, and R, subequal; cell r, large, encloses
apex of wing, length 1.5—2.6 times width at
apex; veins M,, M,, and M, originate sep-
arately from apical margin of discal cell; cell
m, generally closed, petiolate; discal cell
acute basally; posterior cells 5; posterior
cubital cell closed with short petiole; m-cu/
r-m subequal. Legs.— Fore coxae moderate-
ly long, sparsely haired, bare on posterior
surface, with 4-6 stiff setae on anterior sur-
face; femora lacking stiff setae; tibiae with
several stiff apical setae.
39
Abdomen.—Slender, especially in male,
longer than wide, laterally compressed; dor-
sum convex, shining fuscous to black; setae
sparse, short, sternites 5-8 in males, 5-7 in
females (Fig. 10) with numerous lanceolate
setae ventrally. Male terminalia (Figs. 5-
9).—Tergite 8 (T;) wide, deeply constricted
medially. Sternite 8 large, simple. Epan-
drium (Epa), longer in midline than maxi-
mum width, posterior margin broadly
emarginate; cerci (C) extending posteriorly
beyond epandrium and ventral epandrial
sclerite; ventral epandrial sclerite, ventral
view (VES) setose, attached laterally to
epandrium with no membranous attach-
ment to aedeagus, posterior margin broadly
emarginate. Gonocoxites in ventral view not
united ventrally, attached anteriorly by nar-
row membrane; hypandrium (Hpa) broad,
subtriangular, large. Aedeagus with dorsal
apodeme laterally attached by a strongly
sclerotized bridge to basal section of para-
mere; distiphallus narrow, downwardly di-
rected; ventral apodeme lacking; ejacula-
tory apodeme (EA) large, apical fourth
expanded. Female terminalia (Figs. 11-
12).—Tergite 8 (T;) subrectangular, wider
than long, posterior margin broadly con-
cave. Tergite 9 (T,) suboval, separated me-
dially, with several strong setae along pos-
terior margin. Cerci (C) subtriangular,
membranous, with scattered setae. Sternite
8 (S,) large, as long as wide, with short me-
dian emargination on posterior margin.
Sternite 9 greatly modified, invaginated be-
neath sternite 8 to form internal furca (F).
Immature stages. — Characteristics of pu-
pal exuvium (Figs. 13-15) in description of
species.
Ataenogera abdominalis Krober
Ataenogera abdominalis Kroéber (1914: 31):
Malloch (1932: 255).
Leptocera gracilis Kréver (1928: 118). New
synonym.
Henicomyia brevicornis Bromley (1934:
361). New synonym.
Derivation of name. — abdomen (Latin) =
belly; a/is (Latin) = pertaining to.
40 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Diagnosis.—Ataenogera abdominalis is
the only recognized species in this genus.
The characteristics given in the key and de-
scription of the genus separate it from Phy-
cus. Examination of the type specimens of
Ataenogera abdominalis, Leptocera gracilis,
and Henicomyia brevicornis revealed no
distinctive characteristics that would justify
retaining them as separate species. Some
variation was noted in the color pattern of
the fore and middle femora but this appears
to represent clinal variation.
Description of male lectotype (5759).—
Body length 8.8 mm.
Head.— Head depth 0.84 mm. Ocellar tu-
bercle fuscous, subshiny to lightly tomen-
tose; ocellar setae black, moderately long,
scattered. Eyes fuscous. Frons fuscous, to-
mentum light gray, variable, with silver pile
lateral to antennal bases, setae (Fig. 2) white,
moderately long, dorsolateral to antennal
bases and dark brown, short, in lateral row
ventral to ocellar tubercle. Antenna (Fig. 1)
brown, pruinosity light gray, length of an-
tenna 1.9 times head depth; length of scape
0.40 mm, 2.9 times width, 4.0 times length
of pedicel, setae dark brown, short, with
several stiff setae ventrally; length of pedicel
0.10 mm, 0.7 times width, setae dark brown,
in apical band; length of flagellum 1.02 mm,
5.7 times width, 2.6 times length of scape;
length of basal stylomere 0.02 mm, length
of apical stylomere 0.06 mm. Parafacial
narrow, tomentum dense silver. Genal setae
white, elongate, scattered. Maxillary palpus
(Fig. 3) dark brown, pruinosity light gray,
length 0.52 mm, 5.2 times width, setae white
to pale yellow, elongate, scattered. Label-
lum black, pruinosity light gray; setae fus-
cous, short, scattered. Postocular setae white,
elongate, abundant ventrally becoming
black, short, appressed dorsally.
Thorax.—np 2, sa 1, pa 1, dc 0, sc 1. Dark
brown in ground color, tomentum dark gray;
setae white, moderately long, subappressed.
Postpronotal lobe concolorous with thorax;
setae white, moderately long, scattered.
Pleuron fuscous, tomentum dense gray,
anepisternum with ventral three-fourths
glossy. Pleural setae white, elongate, scat-
tered on propleuron, scattered on dorsal
fourth of anepisternum, absent on remain-
ing pleural sclerites. Scutellum fuscous, to-
mentum dark gray. Postnotum and latero-
tergite dark brown, subshiny to pruinosity
light gray; setae on laterotergite white, elon-
gate, abundant. Wing (Fig. 4).—Length 5.7
mm, 3.8 times width. Membrane hyaline.
Pterostigma dark brown, narrow. Halter
dark brown. Legs.—Dark brown, tomen-
tum dark gray, concolorous. Anterior tu-
bercle on hind coxa round, fuscous, apical
half pale yellow.
Abdomen.—Dark brown, subshiny, ter-
gites 2-4 with narrow, white posterior mar-
gin; setae black, short, appressed on dark
areas, white, moderately long on white pos-
terior margin, sternites 5-8 with dark brown
lanceolate setae (Fig. 10). Male termi-
nalia.—Characteristics given in description
of genus.
Variation in males (N = 10).— Body length
6.3-8.8, 7.9 mm. Head depth 0.60--0.84,
0.70 mm. The frons varies in having the
tomentum entire to the ventral third glossy.
Length of antenna |.8—2.2, 2.0 times head
depth; length of scape 0.32-0.46, 0.40 mm,
2.9-4.2, 3.6 times width, 3.2-5.3, 4.4 times
length of pedicel; length of pedicel 0.08-
0.10, 0.09 mm, 0.7-0.8, 0.8 times width;
length of flagellum 0.74—-1.02, 0.84 mm, 4.1-
5.7, 4.9 times width, 1.6-2.6, 2.1 times
length of scape; length of basal stylomere
0.02 mm, length of apical stylomere 0.06-
0.10, 0.08 mm. Length of maxillary palpus
0.36-0.52, 0.44 mm, 5.2-7.3, 5.9 times
width. Notopleural setae 2-3, 2. Dorsocen-
tral setae 0-1, 0. Wing length 4.5-5.7, 5.0
mm, 3.5-3.8, 3.7 times width. A cline exists
in the color pattern of the fore and middle
femora from dark brown in specimens from
Argentina, Paraguay, and southern Brazil to
dark yellow in specimens from Mexico.
Female.—Similar to male with following
exceptions (N = 10). Body length 7.5-8.6,
7.9 mm. Head depth 0.84-0.98, 0.92 mm.
VOLUME 91, NUMBER 1
The frons shows variation similar to male.
Length of antenna |.3-1.5, 1.4 times head
depth; length of scape 0.44—-0.50, 0.47 mm,
3.7-5.0, 4.2 times width, 4.8-6.0, 5.4 times
length of pedicel; length of pedicel 0.08-
0.10, 0.08 mm, 0.67-0.83, 0.75 times width;
length of flagellum 0.70-0.72, 0.71 mm, 3.9-
4.5, 4.3 times width, 1.4-1.6, 1.5 times
length of scape; length of basal stylomere
0.02-0.04, 0.03 mm, length of apical sty-
lomere 0.06-0.12, 0.08 mm. Fore and mid-
dle femora show variation similar to males.
Characteristics of terminalia (Figs. 1 1-12)
given in description of genus.
Seasonal activity.—In the specimens ex-
amined, adults were taken throughout the
year with no difference in the collecting pe-
riod between Central America and southern
South America. Females (25) were collected
2.8 times more often than males (9).
Pupa (Figs. 13-15).—Length 8.0 mm (N
= 1), width 1.5 mm. Alar process not spi-
nose. Labial sheath broad (Fig. 15), truncate
apically, not bisecting proboscial sheath.
Length of antennal sheath 0.56 mm, length
of subapical spine 0.08 mm. Thoracic spi-
racle tapered apically, apex truncate, length
0.20 mm, 2.0 times width. Posterior spines
elongate, not divergent apically, length 0.44
mm. Dorsal spines on abdominal segments
I-VII in transverse row on posterior third
of segment. Abdominal spiracles on seg-
ments I-VII.
Type material.—A syntypic series of 3 males
of Ataenogera abdominalis Kréber is in the
Museum fiir Naturkunde der Humboldt-
Universitat zu Berlin. A male of this series,
herein designated the lectotype, was col-
lected at San Bernardino, Paraguay, in IV,
by K. Fiebrig S. V. The holotype female (not
male as stated in the description by Bromley
1934) of Henicomyia brevicornis Bromley
(American Museum of Natural History) was
collected at Kartabo, British Guiana, on May
2, 1924. A syntypic series of one male and
one female of Leptocera gracilis Krober is
in the Zoologische Institut, Halle. The male
of this series, herein designated the lecto-
41
type, was collected at Parana, Brazil, in De-
cember.
Distribution.—The range of Ataenogera
abdominalis, a widespread Neotropical
species, extends from southern Mexico to
Argentina and Uruguay.
Specimens examined (37).—ARGEN-
TINA: Tucuman: Amaicha, |6-XI-1966, L.
A. Stange, 1M. Mendoza: Potrerillos, 6-I-
1927, F. and M. Edwards, | F. La Rioja: La
Rioja, 1928, 1F. Catamarca: 17 km NW
Chumbicha, 1143 m, 25-XII-1971, C. A.
Pearson, |F. BOLIVIA: Beni: Rio Itenez
opposite Costa Marques, Brazil, 1—3-IX-
1964, J. K. Bouseman and J. Lussenkop,
1F; 4-6-IX-1964, 1M; 30-31-VIII-1964, 1F;
Rio Itenez, Pampa de Meio, | 1-13-IX-1964,
J. K. Bouseman and J. Lussenkop, IF.
BRAZIL: Parana: 1912, 1M 1F; Para, Bak-
er, 1F; Santa Catarina: Nova Teutonia,
27°8', 52°23’, 16-XII-1947, F. Plaumann,
1M; I-1945, 1F; 27°11’, 52°23’, 18-XII-
1959, F. Plaumann, IF. BRITISH
GUIANA: Kartabo, 20-V-1924, 1F. COS-
TA RICA: Guanacaste: La Pacifica, 4 km
NW Canas [Las Canas], 1-9-IV-1974, P. A.
Opler, 1M 4F; 29-III-1974, 1F. EL SAL-
VADOR: Quezaltepeque, 2-XI-1977, MEI,
1F. MEXICO: Morelos: 11.7 km S Yaute-
pec, 17-VIII-1962, N. L. Marston, 2F;
Puebla: 4.8 km NW Petlalcingo, 3-1V-1962,
F. D. Parker, 1F. PANAMA: Canal Zone,
Ancon, 4-V-1926, C. T. Greene, 1F; 9-IV-
1926, 1F. PARAGUAY: San Bernardino,
IV, K. Fiebrig, 1M; 5-IV, 1M; -/-/-, 1M.
TRINIDAD: St. Augustine, V-1959, F. D.
Barrett, 1F. URUGUAY: Minas: Arassu-
ahy, X-1929, Thieman, 1F. VENEZUELA:
Barinitas: 10 km SE Barinitas, 1-III-1986,
R. B. Miller, 1M.
Genus Phycus Walker
Phycus Walker (1850: 1): Irwin and Lyne-
borg (1981: 260); Lyneborg (1978: 212).
Type species: Xylophagus canescens
Walker (1848: 129) by monotypy (= X)-
lophagus brunneus Wiedemann, 1824:
19).
42 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Caenophanes Loew (1874: 415): Lyneborg
(1978: 212). Type species: C. insignis
Loew (1874: 415) by monotypy. Preoc-
cupied by Caenophanes Foerster (1862:
236).
Caenophanomyia Bezzi (1902: 191): Lyne-
borg (1978: 212). New name for Caeno-
phanes Loew (1874: 415).
Paraphycus Becker (1923: 62): Lyneborg
(1978: 212). Type species: Phycus nitidus
Wulp (1897: 137) by original designation.
Caenophaniella Séguy (1941: 112): Lyne-
borg (1978: 212). Type species: C. nigra
Séguy (1941: 112) by original designation.
Moderate-sized, slender flies.
Head.—Frons of both sexes at its narrow-
est from as wide as to more than twice as
wide as ocellar tubercle (Figs. 17, 35), wider
in female than in male of same species, with
shining black areas and tomentose areas,
setae sparse, scanty, or absent; male eyes
dichoptic, eye margins only slightly diver-
gent from level of ocellar tubercle toward
genae (Figs. 17, 35), facets equal, setae ab-
sent; parafacial lacking setae; head slightly
to distinctly protruding at antennal level;
facial and genal calli absent; antenna (Fig.
16) elongate, length 1.4-1.9 times head
depth; scape cylindrical; pedicel subtrian-
gular; flagellum elongate, tapered apically,
setae absent, ratio of lengths of flagellum to
scape differs greatly among species; flagellar
style terminal, two-segmented, with minute
terminal spine; clypeus with surface con-
cave, setae absent; maxillary palpus two-
segmented (Fig. 18), basal segment cylin-
drical, apical segment oval or lobate with
large, apical sensory pit.
Thorax.—np 1-2, sa 1, pa 1, de 0-1, sc
1; vittae absent; mesonotal setae short, uni-
form, erect; prosternum bare in and around
central depression; pleural setae very short,
sparse. Wing (Fig. 19).—Pterostigma ab-
sent; setulae dorsal on R,; subcostal cell and
cell r, elongate, open; veins R, and R, sub-
equal; cell r, large, enclosed apex of wing,
m, usually closed, petiolate; discal cell
trunctae basally; m-cu/r-m subequal. Legs. —
Fore coxae elongate, sparsely setose, with 3
stiff apical macrosetae; middle coxae bare
on posterior surface; femora without setae;
tibiae with short setae.
Abdomen.—Slender, especially in male,
laterally compressed toward apex; dorsum
convex, shining black or reddish brown; se-
tae sparse and short; male and female lack
lanceolate setae ventrally. Male terminalia
(Figs. 20-24).—Tergite 8 (Ts) comparative-
ly wide and deeply to moderately constrict-
ed medially. Sternite 8 large and simple.
Epandrium (Epa) simple, without posterior
incision, may be shorter, equal, or longer in
midline than maximum width; cerci (C) free,
well sclerotized, extending posteriorly be-
yond epandrium and ventral epandrial
sclerite. Ventral epandrial sclerite, ventral
view (VES) large, well sclerotized, setose,
extending to anterior margin of epandrium
with membranous attachment to dorsal
apodeme of aedeagus. Hypandrium absent
in New World species. Gonocoxites (Gx)
broad, heavily sclerotized; separated ven-
tromedially. Gonostylus (Gs) large, thick,
extending slightly beyond apex of gonocox-
ite. Aedeagus laterally attached by a strong-
ly sclerotized bridge to midsection of par-
amere; distiphallus (Dp) forms a fine,
downwardly directed tube; dorsal apodeme
(DA) broad; ventral apodeme (VA) shaped
as two elongate extensions; ejaculatory apo-
deme (EA) large; paramere composed of a
distal process, which curves in inward and
downward approaching base of gonostylus,
and a stout basal apodeme. Female termi-
nalia (Figs. 25, 26).—Tergite 8 (T,) broad,
broadly concave along posterior margin.
Tergite 9 (T,) subtriangular, separated me-
dially with several strong setae along pos-
terior margin. Tergite 10 bilobed, membra-
nous. Cerci (C) membranous, simple, with
scattered setae. Sternite 8 (S,) large, as long
as wide, with short median incision on pos-
VOLUME 91, NUMBER 1 43
21
Figs. 16-26. Phycus frommeri.—16. Antenna (5446). 17. Head of male, frontal view (5/69). 18. Maxillary
palpus, lateral view (5446). 19. Wing, dorsal view (5/69). 20. Male tergite 8, epandrium, cerci, and ventral
epandrial sclerite, dorsal view (5446). 21. Male ventral epandrial sclerite, ventral view (5446). 22. Male gono-
coxites and gonostylus, dorsal view (5446). 23. Male gonocoxites and gonostylus, ventral view (5446). 24. Male
aedeagus, lateral view (5446). 25. Female terminalia, dorsal view (5730). 26. Female terminalia, ventral view
(5730). Abbreviations: (ApSg) apical segment; (BaSg) basal segment; (C) cercus; (DA) dorsal apodeme; (Dp)
distiphallus; (EA) ejaculatory apodeme; (Epa) epandrium; (F) furca; (Gs) gonostylus; (Gx) gonocoxites; (S,) sternite
8; (Stp) stipes; (T,) tergite 8; (T,) tergite 9; (T,,) tergite 10; (VA) ventral apodeme; (VES) ventral epandrial
sclerite. Scale = 0.1 mm, unless otherwise indicated.
terior margin. Sternite 9 greatly modified, Washingtonia palms or other vegetation in
invaginated beneath sternite 8 to form in- desert canyon bottoms (Irwin, personal ob-
ternal furca (F). servation).
Several Old World species of Phycus have
been observed walking along dead and fal-
len tree trunks 1n riverine habitats. The New
World species from western North America Derivation of name: this species 1s named
have been observed walking on rocks under in honor of Saul I. Frommer, University of
Phycus frommeri Webb and Irwin,
NEw SPECIES
44 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
California, Riverside, who collected most
of the specimens.
Diagnosis.—P. frommeri can be distin-
guished from P. frontalis by the following
combination of characteristics: eyes (Fig. 17)
separated by a distance greater than the
width of the ocellar tubercle; width of frons
at level of lateral ocelli greater than 0.5 times
width at level of antennal bases; postero-
lateral extensions of gonocoxites in ventral
view (Fig. 23) short, thick.
Description of male holotype (5/69).—
Length (excluding antenna) 6.7 mm.
Head.—Ocellar tubercle black, tomen-
tum light gray; ocellar setae black, short,
appressed. Eyes dark brown, separated dor-
sally by distance greater than width of ocel-
lar tubercle (as in Fig. 17). Frons black, to-
mentum light gray, converging slightly at
vertex (as in Fig. 17), setae black, short,
erect, in mediolateral row. Antenna (as in
Fig. 16) dark brown, tomentum light gray;
length of scape 0.32 mm, 2.3 times width,
2.3 times length of pedicel, setae black, scat-
tered, erect, with several larger macrosetae
ventrally; length of pedicel 0.14 mm, 1.0
times width, setae black, short, stiff} length
of flagellum 0.78 mm, 4.9 times width, 2.4
times length of scape; length of basal sty-
lomere 0.02 mm, length of apical stylomere
0.10 mm. Parafacial narrow, dark brown,
tomentum silver. Genal setae white, elon-
gate. Clypeus brown, tomentum silver.
Maxillary palpus (as in Fig. 18) dark brown,
tomentum silver; length of basal segment
0.34 mm, 5.4 times width, setae black, elon-
gate; length of apical segment 0.10 mm, 1.7
times width, 0.29 times length of basal seg-
ment, setae shorter than those of basal seg-
ment. Labellum dark brown, tomentum light
gray; setae along ventral margin brown,
moderately long. Postocular setae white,
fine, abundant ventrally becoming black,
stiff, erect toward vertex.
Thorax.— Black, tomentum light silver,
setae white, fine, scattered; np 2, sa 1, pa 1,
de 0, sc 1. Postpronotal lobe concolorous
with thorax; setae white, fine, scattered.
Propleuron, anepisternum, katepisternum,
meron dark brown, tomentum silver; ane-
pimeron fuscous, glossy. Pleural setae white
to pale yellow, abundant on propleuron, and
scattered over anepisternum; absent on re-
maining pleural sclerites. Scutellum black,
tomentum silver; setae white to pale yellow,
scattered. Postnotum and laterotergite dark
brown to black, tomentum silver; setae on
laterotergite white to silver, elongate, abun-
dant. Wing (as in Fig. 19).—Length 5.2 mm,
width 1.8 mm, length 2.9 times width. Wing
membrane hyaline with apical third pale
smoky brown. Halter dark brown, tomen-
tum light gray. Legs.—Coxae dark brown,
tomentum silver; femora, tibiae and tarsi
dark brown. Coxae with 3 stiff, brown apical
setae.
Abdomen. — Dark brown, subshiny; setae
dark brown, short, appressed with pale yel-
low, elongate setae laterally on tergite 1. Male
terminalia (as in Figs. 20—24).—Tergite 8
(T,) as wide as epandrium, deeply concave
medially. Epandrium (Epa) as long as wide
medially; cerci broad (C), rounded apically;
ventral epandrial sclerite (VES), ventral
view, deeply incised medially on apical
margin. Gonocoxites (Gx) in dorsal view
broad, heavily sclerotized; in ventral view
separated medially. Gonostylus (Gs) large,
thick, extending slightly beyond apex of
gonocoxites.
Variation in males (N = 10).— Body length
(excluding antenna) 5.7—7.3, 6.6 mm. Head
depth 0.76-0.84, 0.81 mm. Length of an-
tenna 1.5-1.9, 1.7 times head depth; length
of scape 0.30-0.40, 0.34 mm, 2.3-3.0, 2.5
times width, 2.3— 2.9, 2.5 length of pedicel;
length of pedicel 0.12—-0.14, 0.13 mm, 0.8-
1.2, 1.0 times width; length of flagellum
0.76-0.92, 0.83 mm, 4.7-5.8, 5.0 times
width, 1.7-2.5, 2.0 times length of scape;
length of basal stylomere 0.02 mm, length
of apical stylomere 0.10 mm. Length of bas-
al segment of maxillary palpus 0.34—0.40,
0.38 mm, 4.5—5.7, 5.0 times width; length
of apical segment 0.10-0.14, 0.12 m, 1.2-
1.8, 1.5 times width, 0.26-0.39, 0.31 times
VOLUME 91, NUMBER 1
length of basal segment. Wing length 4.5—
6.8, 4.9 mm, 2.8-3.0, 2.9 times width. Vari-
ation in the color pattern was found in one
specimen (5329) where the anepisternum,
anepimeron, meron, metepleuron, middle
and hind coxae, and entire abdomen were
pale brown and subshiny.
Female.—Similar to male with frontal se-
tae scattered.
Female terminalia (Figs. 25, 26).—Char-
acteristics given in description of genus.
Variation in females (N = 10).—Body
length (excluding antenna) 8.2—9.6, 8.8 mm.
Head depth 0.94—0.96, 0.95 mm. Length of
antenna 1|.4—-1.6, 1.5 head depth; length of
scape 0.40-0.46, 0.43 mm, 2.5-2.9, 2.7
times width, 2.0-2.9, 2.4 times length of
pedicel; length of pedicel 0.16-0.20, 0.18
mm, |.0-1.3, 1.2 times width; length of fla-
gellum 0.68-0.80, 0.74 mm, 4.3-5.0, 4.6
times width, 1.1-1.3, 1.2 times length of
scape; length of basal stylomere 0.02 mm,
length of apical stylomere 0.10 mm. Length
of basal segment of maxillary palpus 0.42-
0.46, 0.45 mm, 5.3-5.8, 5.7 times width;
length of apical segment 0.14 mm, 1.8 times
width, 3.0—3.3, 3.2 times length of basal seg-
ment. Wing length 6.2—6.5, 6.4 mm, length
2.0-2.2, 2.0 times width.
Ecology.—The majority of specimens
were collected in Malaise traps situated in
canyon washes. Individual specimens were
collected on rocks and under Washingtonia
palms, with a single male collected on Er-
iogonum fasciculatum polifolium (5602). In
the collections examined, a distinct differ-
ence was noted in the pattern between tim-
ing of adults collected in Mexico and in Cal-
ifornia. In Mexico, 92% of the adults were
collected between 12 March and 5 May,
while 8% were collected between 23 August
and 8 November; 77% of the adults were
collected during April. In California, adults
were collected between 25 April and 29 July;
the majority of specimens were collected
between early May and mid June. Overall,
males (405) were collected 1.7 times more
frequently than females (243). Figure 27
45
No. of Adults
1970
Adults
No, of
go 15 31 15 30 15
APRIL MAY JUNE JULY
Fig. 27. Phycus frommert.—Flight periodicity in
Deep Canyon, Riverside County, California, in 1969
and 1970 as measured in a single Malaise trap.
shows the variation in flight periodicity at
Deep Canyon, Riverside County, Califor-
nia, during 1969 and 1970. Two female lar-
vae of Phycus frommeri were collected by
R. B. Miller and L. A. Stange, one (5620)
from under small conglomerates of fine soil
at the base of a tree in a shaded gully and
the other (56/9) in loose soil under boul-
ders, both in the state of Colima, Mexico.
These were reared to adults in our labora-
tory.
Larva.— Head (Figs. 28-31) length 0.42
mm; metacephalic rod (MR) elongate, pos-
terior third clavate, length 0.86 mm, 2.0
times length of head. Prothoracic spiracle
(Fig. 30). Posterior spiracle (Fig. 31).
Pupa (Figs. 32-34).— Length 7.7-9.1 mm
(N = 2), width 1.7-2.0 mm. Alar process
not spinose. Labial sheath broad, truncate
apically, not bisecting proboscial sheath.
Length of antennal sheath 0.54 mm, length
of subapical spine 0.14 mm. Length of tho-
racic spiracle 0.38 mm, tapered apically,
apex truncate. Posterior spines elongate, not
divergent apically, length 0.50 mm. Dorsal
spines on abdominal segments I-VII and
46 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ye eae MR
ae - KE
MP i: s 28 — Sa
ANT
a a iI
A , Sus ) AS G / EN
vee / K Sa © ] t
/ XS} Clay
SEN Qe
|. ‘ \) 33 \\ —— 10
a ) Bert t\ i ° , \\ wy —
ry é s by | 3 \I %
29 | ( ca lh \{ LAY
| : KT N 4 K
4h KK \ 32 j De pw \
a) \ i \ fp’ us Ne y
MR—+ = \ \\ h [! yl S il ; } 34
\ ? x pi | Xx ull
WF. NM 4%
| Re ad ka
[ ? « Nelo
| A 31 \ \V/
Figs. 28-34. Phycus frommeri larva and pupa exuvia (56/9)—28. Larval head capsule, lateral view. 29.
Larval head capsule, dorsal view (right), ventral view (left). 30. Prothoracic spiracle of larva, lateral view. 31.
Posterior spiracle of larva, posterior view. 32. Pupal exuvium, lateral view. 33. Pupal exuvium, dorsal view
(right), ventral view (left). 34. Frontal plate of pupal exuvium. Abbreviations: (ANT) antenna; (AS) antennal
sheath; (DSp) dorsal spiracle; (M) mandible; (MP) maxillary palpus; (MR) metacephalic rod. Scale = 0.1 mm,
unless otherwise stated.
ventral spines on segments I-VII fine, not
fused basally. Abdominal spiracles on seg-
ments I-VII moderately thick, tubular.
Distribution. — The range of Phycus from-
meri extends from southern California to
the southern tip of Baja California Sur and
along the northwestern border of Sonora,
Mexico southward to the state of Colima.
Specimens examined (661).— Holotype:
male, Irwin specimen number 5/69, CAS
type no. 15741 (on permanent loan to the
California Academy of Sciences from the
University of California, Riverside); Cali-
fornia, Riverside County, P. L. Boyd Desert
Research Center, 3.5 mi S Palm Desert,
marker #57, 18-—23-V-1970, S. I. Frommer,
in a Malaise trap. Paratypes as follows:
UNITED STATES—CALIFORNIA: Riy-
erside Co.: P. L. Boyd Desert Deep Canyon
Research Center, 5.6 km S Palm Desert, 6-
13-VI-1969, SIF, 14 M 13 F; 6-8-V-1970,
SIF, 1 M; 15-18-V-1970, SIF, 23 M 5 F;
18-23-V-1970, SIF, 50 M 20 F; 27-V-1-VI-
1970, SIF, 35 M 15 F; 13-18-VI-1969, SIF
RMW, 13 M 10 F; 18-19-VI-1969, SIF
RMW, 2M 2F; 19-20-VI-1969, SIFRMW,
1M 1 F; 20-24-VI-1969, SIF RMW, 4 M
1 F; 8-VII-1969, SIF RMW, 1 F; 4-6-V-
1970, SIF RMW, | F; 26-IV-3-V-1970, SIF
RMW, | M; 8-12-V-1970, SIF RMW, 7 M
2 F; 11-13-V-1970, SIF RMW, 5 M; 13-
15-V-1970, SIF RMW, 8 M 2 F; 23-25-V-
1970, SIF RMW, 16 M 18 F; 25-26-V-1970,
SIF RMW, 25 M 17 F,; 1-4-VI-1970, SIF
RMW, 8 M4 F; 21-29-V-1973, A. B. Tabet,
2 M; 5-13-VI-1973, A. B. Tabet, 7 M 4 F;
24-V-1969, MEI SIF, 44 M 12 F; 8-VI-1965,
MEI, | M 1 F; 16-V-1969, MEI, 1 M 1 F;
18-V-1969, MEI, 8 M3 F; 21-V-1969, MEI,
11 M 2 F; 13-20-VI-1973, A. B. Tabet, 5
VOLUME 91, NUMBER 1
M; 21-VI-1962, EIS, 1 F; 19-V-1964, MEI,
1 M; 11-VI-1965, MEI, | F; 9-16-V-1973,
A. B. Tabet, 1 M; 5 mi W Sage, 3-VII-1963,
P. D. Hurd, | F; 2-VII-1963, E. I. Schlinger,
1 F; Idyllwild, 27-VI-1956, M.S. Wasbauer,
1 M; Carrizo Creek, 30-VI-1964, E. I.
Schlinger, | F; Massacre Canyon, 300 yds
up from Highway 79, 29-VII-1964, MEI, 1
F. San Diego Co.: Culp Canyon, | 2-VI-1958,
E. I. Schlinger, 12 F; Borego, 1-V-1946, J.
S. Perry, | M; 9.7 km E San Diego, 26-VI-
1963, H. L. Griffin, | F, R. L. Langston, |
F; San Vicente Res., 229 m, 16-VI-1965,
MEI, 1 F. MEXICO—Baja California Sur:
100 km NW La Paz, Arroyo Guadalupe,
107 m, 20-IV-1968, MEI, 2 M; 14.5 km S
Loreto, 17-[V-1968, MEI, 12 M 1 F; 4.8
km E San Ignacio, 171 m, 14-IV-1968, MEI,
1 F; 3 km E La Burrera, 515 m, 2-3-IX-
1977, J. L. Fisher and R. L. Westcott, | M;
2.5 km E La Burrera, 549 m, E. M. and J.
L. Fisher, | M: 6.4 km NW Don Pancho,
1-VII-1964, MEI, 1 F; Las Barracas, ca. 30
km E Santiago, 25-3 1-III-1982, P. DeBach,
1 M; 1-6-IV-1982, P. DeBach, 2 M IF; 13-
18-IV-1982, P. DeBach, 1 F; 19-24-IV-
1982, P. DeBach, | M; 25-30-IV-1982, P.
DeBach, | M; 7-12-V-1982, P. DeBach, 4
M 2 F. Sinaloa: 86.9 km S Culiacan, 164.6
m, 23-I1V-1969, MEI, | M; “Las Escondi-
das,” 106.2 km N Mazatlan, 137 m, 22-IV-
1968, MEI, 16 M 25 F. Colima: Los Tem-
panes, 12-III-1985, R. B. Miller and L. A.
Stange, | F reared from larva; Rio Salado,
7 km S Colima, 14-III-1985, | F reared
from larva.
Phycus frontalis Webb and Irwin,
NEw SPECIES
Derivation of name: front (Latin) = brow;
alis (Latin) = pertaining to.
Diagnosis.—P. frontalis can be distin-
guished from P. frommeri by the following
combination of characteristics: eyes sepa-
rated dorsally only by width of ocellar tu-
bercle (Fig. 35); frons distinctly convergent
towards vertex (Fig. 35), width at level of
lateral ocelli less than 0.5 times width at
47
level of antennal bases; posterolateral ex-
tensions of gonocoxites in ventral view nar-
row, attenuate (Fig. 39).
Description of male holotype (5/79).—
Length (excluding antenna) 6.8 mm.
Head.—Ocellar tubercle fuscous, tomen-
tum light gray; ocellar setae fuscous, mod-
erately long, scattered. Eyes fuscous, sepa-
rated medially by distance equal to width
of ocellar tubercle (Fig. 35). Frons fuscous,
tomentum light gray, dense silver around
antennal bases; width at level of ocellar tu-
bercle 0.40 times width at level of antennal
bases (Fig. 35); setae pale yellow, moder-
ately long, in mediolateral row; callus ab-
sent. Antenna as in P. frommeri (Fig. 16),
fuscous, tomentum light gray; scape, length
0.34 mm, 2.8 times width, 2.8 times length
of pedicel, setae fuscous, short, subap-
pressed with several large, stiff macrosetae
ventrally; pedicel, length 0.12 mm, 1.0 times
width, setae fuscous, short, suberect; flagel-
lum, length 0.82 mm, 4.6 times width, 2.4
times length of scape; basal stylomere, length
0.04 mm, apical stylomere, length 0.10 mm,
apical spine minute. Parafacial fuscous, to-
mentum dense silver. Genal setae white to
silver, elongate, abundant. Clypeus fuscous,
tomentum dense silver. Maxillary palpus as
in P. frommeri (Fig. 18), dark brown, to-
mentum light gray; length of basal segment
0.34 mm, 4.3 times width, setae dark brown,
moderately long, erect, on ventral surface;
length of apical segment 0.14 mm, 0.41
times length of basal segment, setae sparse.
Labellum dark brown, tomentum light gray;
setae brown, moderately long, scattered.
Postocular setae white to silver, elongate,
abundant on ventral half, with numerous
black, stiff setae dorsally.
Thorax.—Fuscous, tomentum light gray;
setae white to silver, moderately long, scat-
tered; np 2, sa 1, pa 1, dc 0, sc 1. Postprono-
tal lobe concolorous with thorax; setae white
to silver, moderately long, scattered. Pro-
pleuron, anepisternum, katepisternum,
meron fuscous, tomentum silver, anepi-
meron glossy. Pleural setae white to silver,
48 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
\
37 \
Figs. 35-40.
Phycus frontalis (5180). 35. Head of male, frontal view. 36. Male epandrium, cerci, and ventral
epandrial plate, dorsal view. 37. Male ventral epandrial plate, ventral view. 38. Male gonocoxites and gonostylus,
dorsal view. 39. Male gonocoxites and gonostylus, ventral view. 40. Male aedeagus, lateral view. Scale = 0.1
mm, unless otherwise stated.
elongate, abundant on propleuron, scat-
tered on anepisternum, on ventral third of
katepisternum, absent on anepimeron and
meron. Scutellum fuscous, tomentum light
gray; setae pale yellow, moderately long,
scattered. Postnotum and laterotergite fus-
cous, tomentum light gray; setae on later-
otergite white to silver, elongate, abundant.
Wing as in P. frommeri (Fig. 19).—Length
5.0 mm, width 1.8 mm, length 2.8 times
width. Halter fuscous, tomentum light gray.
Legs.— Dark brown, coxae with tomentum
silver; anterior tubercle on hind coxa dark
brown, apical half pale.
Abdomen.—Fuscous, subshiny; setae
fuscous, short, appressed, mixed with pale
yellow, elongate, suberect setae. Male Ter-
minalia (as in Figs. 36-40).—Tergite 8 sim-
ilar to P. frommeri. Epandrium rectangular,
1.25 times longer than wide, posterior mar-
gin truncate; cerci lobate; ventral epandrial
sclerite, ventral view, with dark brown se-
tae. Gonocoxites in ventral view with pos-
terolateral extension narrow, attenuate.
Variation in males (N = 2).— Body length
(excluding antenna) 6.8-6.9, 6.8 mm. Head
depth 0.78-0.92, 0.85 mm. Antenna, length
1.1-1.5, 1.3 times head depth; scape, length
0.32-0.34, 0.33 mm, 2.3-2.8, 2.6 times
width, 2.8—3.2, 3.0 times length of pedicel;
pedicel, length 0.10-0.12, 0.11 mm. Max-
illary palpus, basal segment length 0.30-
0.34, 0.32 mm, 3.8—-4.3, 4.1 times width;
apical segment length 0.10-0.14, 0.12 mm,
1.3-1.8, 1.6 times width, 0.33-0.41, 0.37
times length of basal segment. Wing length
5.0-5.5, 5.3 mm, 2.8-3.1, 3.0 times width.
Female.—Similar to male. Female Ter-
minalia.—As in P. frommeri (Figs. 25, 26).
Variation in females (N = 2).—Body
length (excluding antenna) 7.4 mm. Head
depth 1.10-1.18, 1.14 mm. Antenna length
VOLUME 91, NUMBER |
1.4 times head depth; scape, length 0.41-
0.44, 0.43 mm, 2.6-2.8, 2.7 times width,
2.43.0, 2.7 times length of pedicel; pedicel,
length 0.14-0.18, 0.16 mm, 0.8-1.0, 0.9
times width; flagellum, length 0.84 mm, 4.7
times width, 2.0 times length of scape; basal
stylomere length 0.04 mm, apical stylomere
length 0.10 mm. Maxillary palpus, basal
segment length 0.40-0.52, 0.46 mm, 5.0-
5.2, 5.1 times width; apical segment length
0.16-0.20, 0.18 mm, 1.4-2.0, 1.6 times
width; 0.38—0.40, 0.39 times length of basal
segment. Wing length 6.7—7.7, 7.2 mm, 3.0-
3.1, 3.15 times width.
Distribution. — The range of Phycus fron-
talis extends from Yucatan, Mexico to Cos-
ta Rica.
Specimens examined (4).— Holotype:
male, Irwin specimen number 5/79
(AMNB), Mexico, Yucatan, Chichén Itza,
VI-29. Paratypes as follows: MEXICO—
Yucatan: Chichén Itza, VI-1929, | M 1 F.
COSTA RICA—La Suiza, 1924, P. Schild,
] F.
ACKNOWLEDGMENTS
We thank Drs. G. L. Godfrey, L. M. Page,
and W. E. LaBerge for reviewing this manu-
script and Audrey Hodgins for editorial
comments. We also thank the following in-
stitutions and their curators or former cu-
rators for the loan of material relevant to
this study: American Museum of Natural
History, P. Wygodzinsky; British Museum
(Natural History), K. V. G. Smith; Califor-
nia Department of Food and Agriculture,
M. S. Wasbauer; California Insect Survey,
F. R. Cole; Canadian National Collection,
H. J. Teskey; Cornell University, L. L. Pe-
chuman; Instituto Miguel Lillo, Tucuman,
Argentina, M. L. de Grosso; Kansas State
University, D. H. Blocker; Naturhisto-
risches Museum, Wien, R. Lichtenberg; E.
I. Schlinger Collection; U.S. National Mu-
seum of Natural History, W. W. Wirth; C.
W. O’Brien Collection; University of Cali-
fornia, Davis, R. O. Schuster; University of
California, Riverside, S. I. Frommer; Zool-
49
ogisches Institut, Halle, Professor Hiising;
Zoologisches Museum aus der Humboldt-
Universitat zu Berlin, H. Schumann.
Support for this study was supplied in
part by the Illinois Natural History Survey
and the University of Illinois at Urbana-
Champaign. Additional support was pro-
vided to M. E. Irwin by the National De-
fense Education Act Fellowship, the Uni-
versity of California Regents Patent Fund,
the University of California Dry Lands In-
stitute, and the University of California
Deep Canyon Research Fund.
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PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 51-54
A MID-SUMMER COMPARISON OF SIZES AND GROWTH
RATES AMONG NYMPHS OF THREE SYMPATRIC MANTIDS
(MANTODEA: MANTIDAE) IN TWO OLD-FIELD HABITATS
L. E. HURD AND R. M. EISENBERG
Ecology Program, School of Life Sciences, University of Delaware, Newark, Delaware
19716.
Abstract.—We compared three species of mantid nymphs in two field habitats (sites
CHRY and AG), in two censuses during mid-summer 1986. Tenodera sinensis (Saussure)
nymphs exhibited no difference between sites, either in size of nymphs, or in rate of
maturation. Both 7. angustipennis and M. religiosa were significantly larger in site CHRY
than in site AG in August, although no such difference had been evident in July. More
individuals of these two species had also matured in CHRY than in AG by the August
census. These data suggest that 7. angustipennis and M. religiosa were more food limited
at site AG than at site CHRY during the time just prior to maturation. These results are
consistent with the hypothesis that 7enodera sinensis is more of a habitat generalist than
the other two species.
Key Words:
ators
Mantids are generalist predators subject
to food limitation which varies in impor-
tance during a growing season, as well as
between habitats and years (Matsura et al.
1975, Hurd et al. 1978, Hurd and Eisenberg
1984, Eisenberg et al. 1981). Food limita-
tions on newly hatched nymphs can retard
development and decrease size of the im-
ago, which in turn reduces fecundity (E1-
senberg et al. 1981, Matsura and Marooka
1983, Hurd and Eisenberg 1984, Hurd and
Rathet 1986). Therefore, comparing sizes of
mantids collected at the same time from
different habitats could provide a relative
measure of the differences between habitats
in terms of food limitation and fitness for a
given species. We can then ask the same
question of any number of mantid species
found together in the same habitat to dis-
cern whether the resource level in a given
Mantodea, Mantidae, Tenodera, Mantis, food limitation, sympatry, pred-
field is qualitatively the same for each
species.
Most ecological work on mantids has
concentrated on a single species in a single
habitat during either the first month of life
or adulthood. Rathet and Hurd (1983) stud-
ied growth rates and habitat placement
within a single field site, of nymphs of three
species which commonly occur together in
old fields in northern Delaware: Tenodera
sinensis (Saussure), 7. angustipennis (Saus-
sure), Mantis religiosa (Linnaeus). These
three morphologically similar species hatch
at different times, differ in body size (7.
sinensis > T. angustipennis > M. religiosa),
and/or inhabit different levels of foliage
within a field. These niche differences sug-
gest to us that there may be differences in
resource utilization among these species. In
an exploratory study designed to detect both
52 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
MANTID LENGTH (mm)
hes
Ten. aA a a
50 0 50
ee (%)
Fig. |. Sizes of mantid nymphs in two habitats on
19 July 1986. Open bars represent frequency of nymphs
of various sizes found at site AG; shaded bars are for
site CHRY. Ts = Tenodera sinensis, Ta = T. angus-
tipennis, Mr = Mantis religiosa. One-way ANOVA: F
= 24.945: df = 5, 72; P < 0.001. LSD comparisons
indicate 7. sinensis is significantly larger than the other
two species; no differences between sites.
site and interspecific differences, we com-
pared sizes and growth rates of these three
mantid species in two old fields during the
last nymphal stages.
MATERIALS AND METHODS
The two old-field habitats in our study
are in Newark, New Castle County, Dela-
ware. They are approximately 3 km apart
and have different vegetation. One field, site
AG, is located on the experimental farm of
the School of Agriculture, University of
Delaware, and is dominated by a dense
ground cover of timothy (Ph/eum pratense)
and Canada bluegrass (Poa compressa) with
patches of goldenrod (Solidago spp.) and
thistle (Cirsium spp.). The other field, site
CHRY, is adjacent to the Chrysler plant in
Newark, and is dominated by goldenrod
with a ground cover of grasses, chiefly tim-
othy.
We censused mantids in both habitats on
100 5 Ts
904
i Eaey
MANTID LENGTH (mm)
=
40 S|
30
Ttolahalalstetapalctetelfqtetalislalml alata heliatn lite lam
50 0 50 0 50 0 50
FREQUENCY (%)
Fig. 2. Sizes of mantid nymphs in two habitats on
13 August 1986. Open bars represent frequency of
nymphs of various sizes found at site AG; shaded bars
are for site CHRY. Ts = Tenodera sinensis, Ta = T.
angustipennis, Mr = Mantis religiosa. One-way AN-
OVA: F = 25.714; df = 5, 84; P < 0.001. LSD com-
parisons indicate 7. sinensis is significantly larger than
the other two species; site differences indicated by as-
terisks.
19 July and 13 August 1986, which brack-
eted the last month of nymphal life for these
species. Individuals were hand caught, iden-
tified, measured for length (front of head to
tip of abdomen), and then released at the
point of capture. Sex was recorded for those
mantids which had matured by the second
census. Lengths were compared among
species and habitats on each census date
with one-way ANOVA and least signifcant
difference post hoc comparison using Stat-
graphics (STSC, version 2.1).
RESULTS AND DISCUSSION
Tenodera sinensis nymphs were larger
than the other two species at both sites in
July (one-way ANOVA, F = 24.945; df =
5, 72; P < 0.001), and M. religiosa was not
significantly different in size from 7. an-
gustipennis (Fig. 1). This interspecific dif-
ference was expected from earlier results
VOLUME 91, NUMBER 1
Table 1.
53
Numbers of individuals, % adult and adult sex ratio for three mantid species in two habitats (AG
and CHRY) in July and August of 1986. Mr = Mantis religiosa, Ts = Tenodera sinensis, Ta = T. angustipennis.
Species Site July N August V % Adult M:F
AG 12 1 75 Ee
MECH ETOSe CHRY 20 20 100 10:10
T. angustipennis AG ay Y -
CHRY 23 8 25 2:0
T. sinensis AG 2 2 oy
CHRY 26 22 59 7:6
(Rathet and Hurd 1983) and published size
differences for adults (Gurney 1950, Helfer
1963). There were no differences in size be-
tween sites for any species at this time. Ap-
parently, then, the sites did not differ in food
limitation from egg hatch until mid-July.
There were no adults of any species present
in either field on 19 July.
The August census revealed a number of
differences which were not apparent in July
(Fig. 2). Tenodera sinensis was stil signifi-
cantly larger than the other two species at
both sites (one-way ANOVA, F = 25.714;
df = 5, 84; P < 0.001), and did not exhibit
a difference between sites in size or per-
centage of adults (Table 1). However, both
M. religiosa and T. angustipennis were sig-
nificantly larger at site CHRY than at site
AG (Fig. 2), and more had matured at site
CHRY (Table 1). The size differences could
not be attributed solely to a greater number
of adults at CHRY because even the largest
adults at AG were smaller than the largest
adults at CHRY for both species. These data
suggest that there was greater food limita-
tion during the intervening month at site
AG for these two species than at site CHRY.
Some adult M. religiosaat CHRY were con-
siderably larger (75-80 mm) than previ-
ously published records for this species
(about 65 mm) (Gurney 1950, Helfer 1963).
The sizes of the other two species (Fig. 2)
were consistent with the literature.
Eisenberg et al. (1981) hypothesized that
mantids could rely on insects which forage
on late summer flowers (e.g. goldenrod) for
a major portion of their prey when insect
biomass declines in the rest of the field. Hurd
(1989) found that adult female 7. sinensis
on flowers gained significantly more weight
and deposited oothecae with more eggs (i.e.
had greater fitness) than those on plants not
in flower. We have found mostly 7. sinensis
on these flowers, which may explain why
they apparently were not as food limited at
site AG as the other two species. Thus, while
all three species might rely on the same prey
early in the season, M. religiosa and T. an-
gustipennis could be better indicators of
habitat quality in terms of resident arthro-
pod prey availability than 7. sinensis late
in the season. A detailed examination of the
diets of these species in the field is needed
to test these ideas.
ACKNOWLEDGMENTS
We thank Cynthia Kendall for help in the
field, and two anonymous referees for very
thorough reviews of the manuscript. This
work was supported by NSF Grant BSR
8506181, and is contribution #1 11 from the
Ecology Program, University of Delaware.
LITERATURE CITED
Eisenberg, R. M., L. E. Hurd, and J. A. Bartley. 1981.
Ecological consequences of food limitation for adult
mantids (Tenodera aridifolia sinensis Saussure).
Am. Midl. Nat. 106: 209-218.
Gurney, A. B. 1950. Praying mantids of the United
States, native and introduced. Smithsonian Inst.
Annu. Rep. 1950: 339-362.
Helfer, J. R. 1963. How to Know the Grasshoppers,
Cockroaches, and Their Allies. Dubuque, Iowa,
W. C. Brown. 353 pp.
Hurd, L. E. 1989. On the importance of late season
54 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
flowers to the fitness of an insect predator, Ten-
odera sinensis Saussure (Orthoptera: Mantidae),
in an old field community. Entomologist (U.K.).
Hurd, L. E.and R. M. Eisenberg. 1984. Experimental
density manipulations of the predator Tenodera
sinensis (Orthoptera: Mantidae) in an old-field
community. I. Mortality, development and dis-
persal of juvenile mantids. J. Anim. Ecol. 53: 269-
281.
Hurd, L. E., R. M. Eisenberg, and J. O. Washburn.
1978. Effects of experimentally manipulated den-
sity on field populations of the Chinese mantis
(Tenodera aridifolia sinensis Saussure). Am. Midl.
Nat. 99: 58-64.
Hurd, L. E. and |. H. Rathet. 1986. Functional re-
sponse and success in juvenile mantids. Ecology
67: 163-167.
Matsura, T., T. Inoue, and Y. Hosomi. 1975. Eco-
logical studies of a mantid, Paratenodera angus-
tipennis de Saussure I. Evaluation of the feeding
condition in natural habitats. Res. Pop. Ecol. (To-
kyo) 17: 64-76.
Matsura, T. and K. Marooka. 1983. Influence of prey
density on fecundity in a mantis, Paratenodera
angustipennis (S.). Oecologia 56: 306-312.
Rathet, 1. H. and L. E. Hurd. 1983. Ecological rela-
tionships among three co-occurring mantids,
Tenodera sinensis (Saussure), T. angustipennis
(Saussure), and Mantis religiosa (Linnaeus). Am.
Midl. Nat. 110: 240-248.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, p. 54
NOTE
Ixodes downsi (Acari:
Ixodes downsi Kohls was originally de-
scribed from Aripo Cave, Trinidad, on the
basis of a male, female, and three nymphs
found on the wall of the cave, and one larva
found on a bat, Anoura g. geoffroyi Gray in
the cave (Kohls 1957. Proc. Entomol. Soc.
Wash. 59: 257-264). On the basis of this
scant information, Kohls (op. cit.) specu-
lated that it might be a bat tick although he
noted that oil birds, Steatornis caripensis
Humboldt, nested in the cave and a large
rat was also seen (in the cave). J. downsi has
not been recorded since the original descrip-
tion.
We report a second locality for the species
at a considerable distance from the type lo-
cality. An engorged female was collected
from the throat of a young, fully feathered
S. caripensis found on the floor of a cave
near Tingo Maria, Department Huanuco,
Peru, on 11 February 1975 by Baker. Bats
were heard in the cave but none were seen
and several unidentified parrots flew from
the cave. This locality is approximately 2600
km southwest of the type locality, on the
Ixodidae) from Peru
opposite side of South America. The finding
of a specimen on an oil bird only confuses
the issue as to whether /. downsi is a bat
tick or a bird tick. There are no known
species of /xodes commonly found on both
birds and bats, and additional collecting
from hosts will be necessary before the true
host relationships can be ascertained.
The length and width measurements of
the engorged female from Peru, compared
with those of the unengorged allotype fe-
male in parentheses, were 7.22 (2.53) mm
and 4.07 (1.87) mm, respectively. In all oth-
er respects the Peruvian specimen agrees
with the original description.
The specimen will be deposited in the
Florida State Collection of Arthropods,
Florida Department of Agriculture and
Consumer Services, Gainesville, Florida.
Nixon Wilson, Department of Biology,
University of Northern Iowa, Cedar Falls,
Iowa 50614; W. Wilson Baker, Tall Tim-
bers Research Station, Rt. 1, Box 678, Tal-
lahassee, Florida 32312.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 55-58
TEMPORAL DISTRIBUTION OF HATCHING TIMES IN THREE
SYMPATRIC MANTIDS (MANTODEA: MANTIDAE) WITH
IMPLICATIONS FOR NICHE SEPARATION AND COEXISTENCE
L. E. HurpD AND R. M. EISENBERG
Ecology Program, School of Life Sciences, University of Delaware, Newark, Delaware
19716.
Abstract. —Mantids are members of a guild of generalist predators in arthropod com-
munities. Three species commonly coexist in old fields: Tenodera sinensis Saussure, T.
angustipennis Saussure, and Mantis religiosa Linnaeus. Egg hatch occurs early in the spring,
producing high predator density when prey are in short supply. First instar nymphs of
the two Tenodera species, which are most similar in size and occupy the same vegetational
stratum in the field, are most divergent in timing of egg hatch. Nymphs of 7. sinensis
hatch first, and enter the second instar as 7. angustipennis are beginning to hatch. The
timing of egg hatch for Mantis religiosa, which is much smaller than the other two and
occupies a different portion of the vegetation, is intermediate. Therefore nymphs of the
three species are not the same body size at the same time; body size determines size of
prey which can be captured most efficiently. We suggest that the temporal disparity in
egg hatch may be one mechanism for coexistence among these species by reducing niche
overlap when resources are critically limiting.
Key Words:
patry
Praying mantids (Mantodea: Mantidae)
are members of a guild (sensu Root 1967)
of generalist predators which inhabit ar-
thropod communities. This guild includes
other taxa, notably spiders, and all members
have in common a bitrophic niche; they oc-
cupy both the third and fourth trophic levels
by virtue of feeding on other predators as
well as on herbivores. Thus, the relationship
among guild members is complicated by the
potential for both interspecific competition
and mutual predation. Added to this is the
propensity for cannibalism in some groups.
Therefore, interspecific coexistence in this
guild, especially among very similar species,
may depend upon factors which mitigate a
variety of antagonistic interactions.
Three similar species of mantids com-
Mantodea, Mantidae, Tenodera, Mantis, bitrophic niche, predators, sym-
monly co-occur in northern Delaware. Te-
nodera sinensis Saussure, the largest of the
three, was originally introduced from the
orient near Philadelphia, Pennsylvania, in
1896 (Laurent 1898), and has occurred in
Delaware at least since the 1930s (Jones
1933). Its congener, 7. angustipennis Saus-
sure, was found in Maryland in 1926 (Gur-
ney 1950), and had become established in
Delaware by 1930 (Jones 1933). Mantis re-
ligiosa Linnaeus, the smallest of the three,
was introduced into the northeastern United
States in 1899 (Gurney 1950); its duration
of residence in Delaware is uncertain (Rath-
et and Hurd 1983). All three species are
sympatric in Asia (Jing-jin et al. 1981).
We have been studying various popula-
tions of mantids, especially 7. sinensis, in
56 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
northern Delaware over the past decade (E1-
senberg and Hurd 1977, Hurd et al. 1978,
Eisenberg et al. 1981, Rathet and Hurd 1983,
Hurd and Eisenberg 1984a, b, Hurd 1985,
Hurd and Rathet 1986, Hurd 1988). We
have observed that all three species can be
found in the same general habitat: succes-
sional old fields. Emergence from oothecae
takes place in early spring when arthropod
biomass is very low. In fact, mantid bio-
mass at egg hatch is apt to be much greater
than biomass of all other arthropods com-
bined, so nymphs are likely to be severely
food limited (Hurd and Eisenberg 1984a,
b). It is therefore reasonable to ask: how do
the niches of these similar predators differ
to allow them to coexist in a limited envi-
ronment? Rathet and Hurd (1983) found
that both Tenodera species occupy the same
vegetational stratum (30-60 cm_ foliage
height), whereas M. religiosa is generally
found lower in the vegetation (5-25 cm).
Therefore, M. religiosa may escape preda-
tion from the two larger species, although
this does not rule out interspecific compe-
tition. Rathet and Hurd (1983) also found
that appearance of first instar nymphs in the
field was not simultaneous among the three
species, and that as a consequence there was
a consistent interspecific size difference
throughout the season. Mantid nymphs of
different sizes have different optimal prey
sizes based upon capture efficiency (Bartley
1983). In the present study we ask whether
there is an intrinsic difference in timing of
egg hatch which could be another mecha-
nism which reduces niche overlap in this
guild of generalist predators.
MATERIALS AND METHODS
We collected oothecae of 7. sinensis, T.
angustipennis, and M. religiosa from two
old fields in Newark, New Castle County,
Delaware, on 10 February 1986. The col-
lection sites are approximately 3 km apart,
and represent somewhat different vegeta-
tional assemblages. One field is located on
the experimental farm property of the School
of Agriculture, University of Delaware (=
site AG), and dominated by a dense ground
cover of timothy (Phleum pratense) and
blue-stemmed grass (Poa compressa) with
patches of goldenrod (Solidago spp.) and
thistle (Cirsium canadensis). The other site
is adjacent to the Chrysler plant in Newark
(= site CHRY), and is dominated by gold-
enrod with a sparse ground cover of grasses,
chiefly timothy.
Twelve oothecae of each mantid species
from each of the two field sites were re-
turned to the laboratory and immediately
placed in separate containers and incubated
at 25°C and ambient light. When eggs
hatched, newly emerged nymphs were re-
moved from the containers and killed by
freezing, after which they were counted. A
random sample of 20-30 nymphs from each
group was measured for body length (tip
of abdomen to front of head).
RESULTS
Tenodera sinensis began hatching after 18
days of incubation, and hatching continued
for 11 days from oothecae collected from
both sites, with a few nymphs emerging from
AG oothecae as late as day 33 (Fig. 1). The
pattern of hatching was sporadic with no
clear difference between field sites. In only
two cases did nymphs emerge on two or
more consecutive days from the same
ootheca, which is consistent with results
from Eisenberg and Hurd (1977). Mean size
of nymphs from site CHRY (9.7 mm, SD
= (0.37) was somewhat larger than from site
AG (9.4 mm, SD = 0.40) (t = —2.84, df=
48, P < 0.01).
Tenodera angustipennis did not start
hatching until 7. s7nensis had finished, and
duration of hatch was six days (Fig. 1). The
pattern of hatching was nearly identical for
eggs from both sites. Only one ootheca ex-
hibited hatching on two consecutive days.
Mean body length of nymphs (8.9 mm, SD
= (0.20) was not different between sites.
Mantis religiosa was the only one of the
three species to exhibit a marked site dif-
VOLUME 91, NUMBER 1
ference in hatching pattern (Fig. |). Hatch-
ing time was intermediate, and overlapped
with both Tenodera species. Unlike Teno-
dera spp., all but one ootheca produced
nymphs on consecutive days, as many as
five days in a row. As with 7. sinensis,
nymphs from site CHRY were larger (6.1
mm, SD = 0.31) than nymphs from site AG
(mean = 5.8 mm, SD = 0.20) (t = —3.78,
df = 58, P < 0.001).
DISCUSSION
We found that under identical laboratory
conditions of a “common garden” experi-
ment, eggs of 7. sinensis hatch before those
of the other two species. This is in agree-
ment with field observations of first ap-
pearance of nymphs (Rathet and Hurd
1983), and indicates an intrinsic difference
between species. Matsura (pers. comm.) has
also observed that 7. sinensis hatches before
T. angustipennis in Japanese fields.
A potential significance of this phenolog-
ical difference is a reduction in overlap
among species with regard to optimal prey
size (Bartley 1983), which could reduce
competition for limited food early in the
growing season: by the time 7. angustipen-
nis and M. religiosa begin to hatch, the ear-
liest 7. sinensis nymphs would be entering
the second instar (Hurd and Eisenberg
1984a, Hurd and Rathet 1986). This is like-
ly to be more important for the two Teno-
dera species, since they are similar in size
at emergence and occupy the same vegeta-
tional stratum, than for M. religiosa which
occupies a different portion of the vegeta-
tion (Rathet and Hurd 1983) and is much
smaller. The fact that the timing of egg hatch
in M. religiosa overlaps the other two species
may be further indication that interaction
between these genera is less important than
between species of Tenodera. Of course the
question remains as to whether the tem-
poral differences in hatching arose from his-
toric interspecific competition, or represent
a fortuitous preadaptation to living in the
same habitat.
57
Mantis religiosa
40 Tenodera angustipennis
PERCENT TOTAL EGGS HATCHING
no Tenodera sinenis
18 20 25 30 35 40
DAYS OF INCUBATION
Fig. 1. Timing and pattern of egg hatch, as a percent
of total eggs hatching for each species from oothecae
collected at two field sites: AG = shaded bars, CHRY
= open bars.
In instances where food is severely lim-
iting, the size difference between second in-
star nymphs of 7. sinensis and first instar
T. angustipennis would confer an additional
benefit on the former species, that of being
able to use its congener (and later-hatching
conspecifics) as a food source. We have ob-
served in the laboratory that intra-instar
predation between and within these con-
generic nymphs is relatively uncommon, as
is the case with M. religiosa according to
Roeder (1936). However, we have also ob-
served that the difference in size between
successive instars is sufficiently great to pro-
voke much cannibalism and interspecific
predation (Hurd 1988). This may help to
explain why we have never found 7. an-
gustipennis to be abundant in fields wherein
T. sinensis was well-established.
We found two indications of differences
between field sites: the difference in hatch-
ing patterns for M. religiosa, and the larger
nymphs at site CHRY for both this species
and 7. sinensis. The difference in nymph
58 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
size may well be an indication of differences
in feeding conditions for adult mantids dur-
ing the previous season. Eisenberg et al.
(1981) found that food limitation among
adults plays an important role in biomass
of oothecae; egg size may also be affected.
This would indicate that adults of these two
species were more food limited at site AG
than at site CHRY. We are currently testing
this.
Another possibility is that the size differ-
ences represent ecotypes for one or both
species. This remains to be tested, as does
the question of what is the gain in fitness
by hatching at a larger size? One possibility
is a decline in time of development, al-
though 7. sinensis, at least, is flexible in this
regard (Hurd and Rathet 1986). We cannot
as yet offer a satisfactory explanation for the
difference in hatching patterns for /. reli-
giosa (unless ecotypic); however, roughly the
same pattern difference was observed the
following year (unpublished data). The
question of the significance of multiple con-
secutive-day hatching of eggs in this species
also remains to be answered.
ACKNOWLEDGMENTS
This work was supported by NSF Grant
BSR 8506181. This is contribution #110
from the Ecology Program, University of
Delaware.
LITERATURE CITED
Bartley, J. A. 1983. Prey selection and capture by the
Chinese mantid (7enodera sinensis Saussure).
Ph.D. dissertation, University of Delaware, New-
ark, DE.
Eisenberg, R. M. and L. E. Hurd. 1977. An ecological
study of the emergence characteristics for egg cases
ofthe Chinese mantis (Tenodera aridifolia sinensis
Saussure). Am. Midl. Nat. 97: 478-482.
Eisenberg, R. M., L. E. Hurd, and J. A. Bartley. 1981.
Ecological consequences of food limitation for adult
mantids (7enodera aridifolia sinensis Saussure).
Am. Midl. Nat. 106: 209-218.
Gurney, A. B. 1950. Praying mantids of the United
States, native and introduced. Smithson. Inst.
Annu. Rep., 339-362.
Hurd, L. E. 1985. Ecological considerations of man-
tids as biocontrol agents. Antenna, Bull. Royal
Entomol. Soc. London 9: 19-22.
1988. Consequences of divergent egg phe-
nology to predation and coexistence in two sym-
patric, congeneric mantids (Orthoptera: Manti-
dae). Oecologia 76: 547-550.
Hurd, L. E. and R. M. Eisenberg. 1984a. Experi-
mental density manipulations of the predator
Tenodera sinensis (Orthoptera: Mantidae) in an
old-field community. I. Mortality, development
and dispersal of juvenile mantids. J. Anim. Ecol.
53: 269-281.
1984b. Experimental density manipulations
of the predator 7enodera sinensis (Orthoptera:
Mantidae) in an old-field community. II. The in-
fluence of mantids on arthropod community struc-
ture. J. Anim. Ecol. 53: 955-967.
Hurd, L. E., R. M. Eisenberg, and J. O. Washburn.
1978. Effects of experimentally manipulated den-
sity on field populations of the Chinese mantis
(Tenodera aridifolia sinensis Saussure). Am. Mid.
Nat. 99: 58-64.
Hurd, L. E. and I. H. Rathet. 1986. Functional re-
sponse and success in juvenile mantids. Ecology
67: 163-167.
Jing-jin, Y., X. Chong-hua, Y. De-fu, and L. Ying-mei.
1981. Studies on the bionomics of six mantids
from China. Coll. Res. Papers, Chinese Acad. For-
estry 2: 67-74.
Jones, F. M. 1933. Another oriental mantis well es-
tablished in the U.S. (Tenodera angustipennis
Saussure). Entomol. News 44: 1-3.
Laurent, P. 1898. A species of Orthoptera. Entomol.
News 9: 144-145.
Rathet, 1. H. and L. E. Hurd. 1983. Ecological rela-
tionships among three co-occurring mantids,
Tenodera sinensis Saussure), 7. angustipennis
(Saussure), and Mantis religiosa (Linnaeus). Am.
Midl. Nat. 110: 240-248.
Roeder, K. D. 1936. Raising the praying mantis for
experimental purposes. Science 83: 582-583.
Root, R. B. 1967. The niche exploitation pattern of
the blue-gray gnatcatcher. Ecol. Monogr. 37: 95-
124.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 59-65
NEW WORLD SPECIES OF HOLCOPELTE AND IONYMPHA
(HYMENOPTERA: EULOPHIDAE), WITH DESCRIPTIONS OF
TWO NEW SPECIES
CHRISTER HANSSON
Department of Systematic Zoology, Lund University, Helgonavagen 3, S-223 62 Lund,
Sweden.
Abstract.—The genera Holcopelte Forster and Jonympha Graham (Hymenoptera: Eu-
lophidae) are for the first time recorded from the Americas. The American Holcopelte
comprises two new species: H. americana from Canada and the United States, and H.
huggerti from the United States, Ecuador and Peru. Females and males are described for
both species. Jonympha is represented by two species in the Americas: J. carne (Walker)
from Canada, Ecuador and Peru, and /. ochus (Walker) from Canada. The hosts are
unknown for all four species.
Key Words:
Forster (1856) described Holcopelte to in-
clude Elachistus obscurus (Forster, 1841).
Later, he described H/. fulvipes (1861), which
was later synonymized with H. obscura by
Boucek & Askew (1968). Thomson (1878)
included Holcopelte as a species-group in
the genus Derostenus; he also described D.
sulciscuta which was later transferred to
Holcopelte (Graham, 1959). Ashmead
(1894) described six species of Holcopelte
from the Island of St. Vincent. Later (1904),
he synonymized Holcopelte with Horisme-
nus Walker. Erdés (1958) described Horis-
menus lenticeps from Hungary, which was
later transferred to Holcopelte (Boucek &
Askew, 1968). Graham (1959) resurrected
Holcopelte and presented a key to the two
British species. Boucek (1969) described
Holcopelte stelteri from Germany and gave
a key to the four European species.
The six species described from St. Vin-
cent by Ashmead were Horismenus, and
prior to this paper no records of Holcopelte
from the Americas have been published.
Since only the females are known in the
Eulophidae, Holcopelte, Ionvmpha, New World, taxonomy
European species, this is the first time male
Holcopelte are described. Hosts are only
known for two of the European species (ob-
scura, Stelteri), both have been reared from
gallmidges (Diptera, Cecidomyiidae) (Bou-
éek & Askew 1968, Boucek 1969).
Ionympha was described by Graham
(1959) to include Entedon carne and E.
ochus; both species were originally de-
scribed by Walker (1839). No additional
species have been described. The genus was
only known from Europe. Hosts are un-
known for both species.
Abbreviations used in the text are: HE =
height of an eye; MO = width of mouth
opening; MS = malar space; OOL = dis-
tance between one posterior ocellus and eye;
POL = distance between posterior ocelli;
POO = distance between posterior ocelli and
occipital margin; WH = width of head (dor-
sal view); WT = width of thorax across
shoulders. Abbreviations of museums and
private collections are as follows: BMNH =
British Museum (Natural History), London,
England; CH = collection of the author;
60 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
CNC = Canadian National Collections, Ot-
tawa, Canada; LUZM = Lund University
Zoological Museum, Lund, Sweden; USNM
= National Museum of Natural History,
Washington, D.C., USA.
Genus Holcopelte Forster
Holcopelte Forster, 1856: 78. Type-species:
Elachistus obscurus Forster, 1841: 40, by
original designation.
Diagnosis.— Head lenticular; face, frons
and occiput smooth and polished; interan-
tennal elevation high and clearly delimited
(Figs. 3, 5, 7, 9); mandibles four-dentate
(Fig. 7); antenna with two small, discoid
anelli; notaular depressions distinct, long
and narrow (Fig. 1); thorax with a median
furrow extending from posterior mesoscu-
tum to anterior scutellum (Fig. 1); post-
marginal vein shorter than stigmal vein;
male four basal flagellar segments with a
single whorl of long hairs at the base of each
segment, fifth segment with a basal whorl
and with scattered hairs in front of whorl
(Figs. 4, 8).
Remarks.—The monophyly of Holco-
pelte is shown through the following syn-
apomorphies: 1) face, frons and occiput
smooth and polished; 2) notaular depres-
sions distinct, long and narrow, and clearly
delimited from remaining mesoscutum; 3)
thorax with a median furrow extending from
posterior mesoscutum to anterior scutel-
lum.
The American species differ from Euro-
pean species by having a pale petiole and
female forecoxa brown (European species
with petiole dark and all coxae with same
color in female).
KEY TO THE NEw WORLD SPECIES OF
HOLCOPELTE
l= Females\. =. 2
— Males
Antennal scrobes joining before reaching ver-
tical line of frontal fork (Fig. 3); scape 5.0 x as
long as wide with 6-7 setae along ventral edge
(Fig. 2); reticulation on median thoracic dor-
sum engraved (slightly raised on the sides and
i)
along hind part of scutellum); cubital hair-line
of forewing almost straight (Fig. 1); propodeum
sculptured and with 4 plicae (Fig. 1); petiole
conical without a dorsal shield
La hee shad siviscere katie H. americana new species
- Antennal scrobes never meet (Fig. 7); scape
8.0 x as long as wide with 3-5 setae along ven-
tral edge (Fig. 6); thoracic dorsum with raised
reticulation; cubital hair-line of forewing
strongly sinuate below speculum (Fig. 12); pro-
podeum smooth to almost smooth with 2 pli-
cae (Fig. 11); petiole quadrangular (shape
transverse to slightly elongate) with a dorsal
shield that covers petiolar foramen (Figs. 10,
0 |) ee ee Ae . H. huggerti new species
3. Petiole small and conical, without a dorsal
shield (like Fig. 1); cubital hair-line of forewing
almost straight (like Fig. 1); scape apically ex-
panded (Fig. 4); flagellum infuscate, segments
210=2S: x asslongas wide s,s saeco
. H. americana new species
- Petiole distinctly elongate (1. 5-3.0 x as long as
wide) with a dorsal shield (like Figs. 10, 11);
cubital hair-line of forewing strongly sinuate
below speculum (like Fig. 12); scape distinctly
narrowing at apex (Fig. 8); flagellum pale, seg-
ments 3.5 (apical segment 5.0) as long as
wide and slightly narrowed medially
. H. huggerti new species
Holcopelte americana,
NEw SPECIES
Figs. 1-5
Diagnosis.—Petiole small and conical,
without a dorsal shield; cubital hair-line of
forewing almost straight; reticulation on
median thoracic dorsum engraved; propo-
deum with four plicae; ratio width/length
of dorsellum = 2.4/1.0. Female: antennal
scrobes joining before reaching vertical line
of frontal fork; scape 5.0 as long as wide
with 6-7 setae along ventral edge. Male:
scape apically expanded; flagellum infus-
cate; flagellar segments 2.0-2.5 x as long as
wide; head transverse in frontal view.
Description.— Female: Scape yellowish-
white, more or less infuscate in apical part,
remaining antenna brown. Face, frons and
occiput golden-green, occasionally bluish.
Thorax varying from blackish to brownish,
usually also with a slight golden-green tinge
VOLUME 91, NUMBER | 61
er a
wi
PG hc %
yes
Sess : ‘
Figs. 1-9. Holcopelte spp. 1, ° H. americana new species, habitus. 2, Same, lateral view of head. 3, Same,
frontal view of head. 4, ¢ H. americana, lateral view of head. 5, Same, frontal view of head. 6, 2? H. huggerti
new species, lateral view of head. 7, Same, frontal view of head. 8, ¢ H. huggerti, lateral view of head. 9, Same,
frontal view of head. Scale representing 0.50 mm.
62 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 10-14. 10-12, 2 Holcopelte huggerti new species. 10, Side view of petiole. 11, Dorsal view of propodeum
and petiole. 12, Base of forewing. 13-14, 2 Jonympha carne (Walker). 13, Frontal view of head. 14, Left mandible.
Scale to the left representing 0.25 mm (Fig. 10), and 0.50 mm (Figs. 11-13); scale to the nght representing 0.10
mm (Fig. 14).
(lacking in some specimens). Forecoxa
brown, mid- and hindcoxae pale (usually
whitish, but occasionally yellowish), re-
maining parts of legs varying from yellowish
to predominantly infuscate. Wings hyaline
or weakly infuscate, veins pale. Petiole yel-
lowish-white. Gaster with same color as
thorax. Length of body: 0.9-1.3 mm. Fla-
gellum with all five segments free, segment
I 1.5-2 x, Il about 2 x, If] and IV 2.3-2.5 x,
and V about 3 x as long as wide. Ratios HE/
MS/MO = 4.2/1.0/3.0. Malar space 1.5 x
as wide as width of scape in widest part.
Frontal fork V-shaped. Inner orbit of eye
with one row of setae. Ratios POL/OOL/
POO = 8.4/3.6/1.0. Entire occipital margin
with a fine and sharp edge, extending from
eye to eye. Ratio WH/WT = 1.3/1.0. Pro-
notal collar without transverse carina. An-
terior part of mesoscutum with reticulation
slightly raised above the surface, with trans-
verse meshes. Median and posterior meso-
scutum with finer and engraved reticula-
tion, meshes isodiametric or slightly
elongate. Notaular depressions clearly de-
limited from remaining mesoscutum, nar-
row and extending along posterior half of
mesoscutum. Midlobe of mesoscutum with
a fine furrow, usually extending along pos-
terior-median half of mesoscutum, but oc-
casionally shorter. Scutellum with fine and
engraved reticulation, slightly raised along
sides and hind part. Meshes slightly to very
elognate in median part, isodiametric along
sides and hind part. Furrow on median
mesoscutum continuing on scutellum, ex-
tension varying from anterior to entire scu-
tellum. Dorsellum 2.4 as wide as long,
with irregular sculpture and usually divided
into three concave areas. Forewing round-
ed, hindwing with apex varying from round-
ed to sharply pointed. The shape of apex of
hindwing is linked to the size of the speci-
men: small specimens have a pointed
apex,while large specimens have a rounded
apex. Speculum developed and closed be-
low, cubital hair-line almost straight. Ratios
length of marginal/postmarginal/stigmal
veins = 7.1/1.0/1.2. Anterior part of pro-
podeum with a relatively wide transverse
furrow, extending between stigmata. With
two pair of plicae, outer pair separating pro-
podeal callus from propodeum, inner pair
situated half way between outer plica and
median propodeum (these are occasionally
bifurcate in posterior part), with or without
a median carina. With a pair of carinae as-
cending from upper corners of petiolar fo-
ramen, reaching half way up on propodeum.
Propodeal surface usually with weak reticu-
lation and/or other weak sculpture. Pro-
podeal callus with two setae. Petiolar fora-
men triangular to rounded in shape. Petiole
conical, at most as long as wide, usually
VOLUME 91, NUMBER 1
slightly transverse. Gaster ovate and mod-
erately acuminate posteriorly, about 1.2 x
as long as thorax + propodeum.
Male: Color like female, except all coxae
brown and head with brighter color. Length
of body: | mm. Scape apically expanded,
flagellar segments I-III about 2 x, [V and V
about 2.5 x as long as wide. Malar space as
wide as width of scape in widest part. Ratios
HE/MS/MO = 3.4/1.0/2.6, POL/OOL/
POO = 14.0/6.0/1.0, WH/WT = 1.5/1.0.
Gaster slightly longer than thorax + pro-
podeum. Otherwise as in female.
Type material.—Holotype 2 labelled:
“USA: West Virginia, Greenbrier Co., 10
miles E. Richwood, Summit Lake,
29.vu1.1983, leg. L. Huggert,” in LUZM.
Paratypes: 2 2 with same label as holotype;
1 9 “USA: West Virginia, Greenbrier Co.,
Richwood, Summit Lake, 27.vu1.1983, leg.
L. Huggert’’; 2 2, 1 6 “USA: West Virginia,
Pocahontas Co., Falls of Hills Creek,
22.vii.1983, leg. L. Huggert’; 3 2 “USA:
West Virginia, Pocahontas Co., Falls of Hills
Creek, Monongahela Forest, 26.vii.1983,
leg. L. Huggert”’; | 6 “USA: West Virginia,
Pocahontas Co., Dogway Road, Monon-
gahela Forest, 26.vi1.1983, leg. L. Huggert”’;
1 2 **Canada: Quebec, Bouchette, Lac Rod-
dick, 12.1x.1982, leg. L. Huggert.”” These
paratypes deposited: 3 2, 1 6 in CH, 5 9,
1 6 in LUZM, 1 2 in USNM; 1 ¢
**N.S.C.B.H.N.P., Skyline Trail, 25.vii.
1983,” “Birch PG 635802, B.R.I. Survey”;
1 2 “Prince Edward I., Sand Banks Prov.
Pks., 25.vii.1982, L. Masner”’; 1 2 “St. Law-
rence Is. Nat. Park, Ontario, Thwartway Is.,”
“19 vii.1976, L. Masner, Code 4212-7”; fi-
nal three paratypes in CNC.
Distribution.— Canada (Nova Scotia,
Ontario, Prince Edward Island, Quebec) and
the United States (West Virginia).
Holcopelte huggerti, New SPECIES
Figs. 6-12
Diagnosis. —Petiole quadrate (2), or 1.5—
3.0 as long as wide (4) with a thin dorsal
shield that covers petiolar foramen; cubital
63
hair-line strongly sinuate below speculum;
reticulation on thoracic dorsum raised;
midlobe of mesoscutum separated from
scutellum by a narrow transverse furrow;
propodeum with two plicae; ratio width/
length of dorsellum = 5.9/1.0. Female: an-
tennal scrobes never meet; scape 8.0 as
long as wide, with 3-5 setae along ventral
edge. Male: scape narrowed apically; flagel-
lum pale; flagellar segments 3.5 x (apical
segment 5.0 x) as long as wide and slightly
narrowed in median part; head about as high
as wide in frontal view.
Description.— Female: Scape yellowish-
white, remaining antenna infuscate. Frons
and occiput dark brown with a weak golden
or golden-green tinge. Thorax dark brown
with a golden tinge. Forecoxa brown, mid-
and hindcoxae pale (usually white, but oc-
casionally yellowish), remaining parts of
fore- and midlegs usually more or less in-
fuscate (especially femora), hindleg usually
predominantly pale. Wings hyaline with pale
veins. Petiole yellowish-white. Gaster with
same color as thorax. Length of body: 0.9-
1.3 mm. Scape narrow with 3-5 setae along
ventral edge. All five flagellar segments free,
segment I about 2.8 x, HH, II and V about
3.8, and IV about 3.4 as long as wide.
Ratios HE/MS/MO = 4.2/1.0/2.4. Malar
space about 2.5 x as wide as width of scape
in widest part. Frontal fork V-shaped. Inner
orbit of eye with one row of setae. Ratios
POL/OOL/POO = 13.0/10.0/1.0. Entire
occipital margin with a fine and sharp edge,
extending from eye to eye. Ratio WH/WT
= 1.5/1.0. Pronotal collar without trans-
verse carina. Mesoscutum and scutellum
with fine but distinctly raised reticulation,
stronger along sides and hind part of scu-
tellum. Meshes elongate in median part of
both mesoscutum and scutellum, otherwise
isodiametric. Median furrow on_ thorax
weak, extending from posterior ' of meso-
scutum to anterior 3 of scutellum, some-
times absent on mesoscutum. Notaular
depressions like americana. Midlobe of
mesoscutum separated from scutellum by a
narrow furrow. Dorsellum short, 5.9 as
64 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
wide as long, with surface convex. Shape of
wings like americana. Speculum present and
closed below, cubital hair-line strongly sin-
uate below speculum. Ratios length of mar-
ginal/postmarginal/stigmal veins = 7.4/1.0/
1.3. Anterior part of propodeum with a rela-
tively wide transverse furrow, extending be-
tween stigmata. Only one pair of plicae pres-
ent, situated in level with stigmata. Also
with a pair of carinae ascending from upper
corners of petiolar foramen, reaching about
half way up on propodeum. Propodeal sur-
face otherwise smooth. Propodeal callus
with two setae. Petiolar foramen quadrate,
upper margin rounded. Petiole quadrate,
with a thin dorsal shield anteriorly, shield
covering petiolar foramen. Gaster ovate,
more acuminate than americana posteriorly,
about 1.3 as long as thorax + propodeum.
Male: Color like female, except entire an-
tenna pale and all coxae infuscate. Length
of body: 1.1-1.4 mm. Scape narrowed api-
cally. Flagellar segments I-IV about 3.5 x,
and V about 5.0 as long as wide. Malar
space 1.2x as wide as width of scape in
widest part. Ratios HE/MS/MO = 2.9/1.0/
2.0, POL/OOL/POO = 15.0/7.2/1.0, WH/
WT = 1.3/1.0. Median furrow on thorax
extending along entire scutellum in some
specimens. Petiole 1.5—3.0 x as long as wide.
Gaster slightly longer than thorax + propo-
deum. Otherwise as in female.
Type material.—Holotype @ labelled:
“USA: West Virginia, Greenbrier Co., 10
miles E. Richwood, Summit Lake, 29.vu1.
1983, leg. L. Huggert” in LUZM. Paratypes:
2 2 with same label as holotype; | ° “USA:
West Virginia, Pocahontas Co., Falls of Hills
Creek, Monongahela Forest, 26.vu.1983,
leg. L. Huggert”; 1 @ “Peru: Huanucu, To-
cache, 2.11.1984, leg. L. Huggert”’; 1 2 “‘Peru:
Cusco, Machu Pichu, 21.xii.1983, leg. L.
Huggert”; 2 2 6 6 “Peru: Cusco, Aqua Cal-
iente, 28.x1i.1983, leg. L. Huggert’’; 1 2 “Ec-
uador: Napo, Lumbaqui, 10-1 1.111.1983, leg.
L. Huggert”; 1 9 “Ecuador: Pichin, Rio Pal-
enque, 4.ii.1983, leg. L. Huggert”; 2 2 “Ec-
uador: Pichin, Puerto Quito, 2.111.1983, leg.
L. Huggert’’; 4 2 2 6 in CH, 623 6in LUZM,
1?1¢6in USNM.
Distribution.—The United States (West
Virginia), Ecuador and Peru.
Genus Jonympha Graham
Tonympha Graham, 1959: 199. Type-
species: Entedon ochus Walker, 1839: 21,
by original designation.
Diagnosis.— Mandibles multidentate,
long and narrow (Fig. 14); antennal scrobes
never meet (Fig. 13); frons below fork with
fine reticulation at least in some places; an-
tenna with two small and discoid anelli; male
flagellar segments with hairs placed evenly
(not only a basal whorl as in Holcopelte),
notaular depressions shallow and not de-
limited from remaining mesoscutum; pro-
podeum smooth and shiny without plicae;
male gaster with a pale subbasal spot.
Remarks.-The monophyly of Jonympha
is shown through the following synapo-
morphies: 1) mandibles long and narrow; 2)
male gaster with a pale subbasal spot.
For identification and description of the
species see Graham (1959: 199-200). The
key in Graham can be supplemented with
the following character: female /. carne with
2-5 setae on propodeal callus, 7-9 setae in
ochus.
Tonympha carne (Walker)
Figs. 13-14
Entedon carne Walker, 1839: 123.
Ionympha carne (Walker), Graham, 1959:
200.
Material. —CANADA: | 2 Ontario, Ron-
deau Prov. Park, 26.vili. 1982. ECUADOR:
1 ° Pichin, Tinalandia, 800 m, 7.11.1983.
PERU: 1 2 Cusco, Ollantaytambo, 19.xi.
1983: 1 2 Cusco, Machu Pichu, 21.xii.1983.
Allleg. L. Huggert (1 2 in CH, 3 ?in LUZM).
Type material of E. carne in BMNH (not
seen).
Remarks. — Most European specimens of
this species have dark coxae and remaining
parts of legs predominantly infuscate. Three
VOLUME 91, NUMBER 1
of the females from the Americas have mid-
and hindcoxae and remaining hindleg pale.
However, there are some European speci-
mens with hindcoxa pale, and one of the
females from Peru has all legs entirely in-
fuscate. The infuscation of the legs frequent-
ly shows a high degree of intraspecific vari-
ation in many species of Eulophidae and I
do not regard this difference in color be-
tween European and American specimens
as a species character. American specimens
of I. carne have, on the average, a weaker
reticulation on the thoracic dorsum than
European specimens. The midlobe of meso-
scutum is usually partly, and the scutellum
is predominantly, smooth and polished in
American specimens. However, in some
species of Eulophidae with a Holarctic dis-
tribution, e.g. Chrysocharis prodice (Walk-
er), American specimens tend to have a
weaker reticulation on the thoracic dorsum
(Hansson 1987). This tendency becomes
even more apparent in specimens from the
southern Nearctic region.
Distribution.—West Palearctic (Boucek
& Askew 1968), Canada (Ontario), Ecuador
and Peru. Jonympha carne was previously
not recorded from the Americas.
Tonympha ochus (Walker)
Entedon ochus Walker, 1839: 21.
Ionympha ochus (Walker), Graham, 1959:
200.
Material CANADA: 1 @ British Co-
lumbia, N. Vancouver, 31.viii.1960, S. M.
Clark (CNC). This specimen agrees well with
European specimens. Type material of /.
ochus in BMNH (not seen).
Distribution.—West Palearctic (Boucek
& Askew 1968) and Canada (British Colum-
65
bia). Jonympha ochus was previously not
recorded from the Nearctic Region.
ACKNOWLEDGMENTS
I am particularly grateful to Lars Huggert
(Lund, Sweden) who collected the majority
of the material forming the base of this pa-
per. My thanks also to R. Danielsson
(LUZM), M. E. Schauff (USNM) and C. M.
Yoshimoto (CNC) for loan of material.
LITERATURE CITED
Ashmead, W. H. 1894. Report on the parasitic Cy-
nipidae, part of the Braconidae, the Ichneumon-
idae, the Proctotrypidae, and part of the Chalcidi-
dae. Part II. J. Linn. Soc. (Zool.) 25: 108-188.
1904. Classification of the chalcid flies or the
superfamily Chalcidoidea, with descriptions of new
species in the Carnegie Museum, collected in South
America by Herbert H. Smith. Mem. Carneg. Mus.
1(-IX): 225-551.
Bouéek, Z. 1969. Descriptive and taxonomic notes
on ten, mainly new, species of West Palearctic
Eulophidae (Hymenoptera). Acta Entomol. Mus.
Natl. Pragae 38: 525-543.
Boucek, Z. and R. R. Askew. 1968. Palearctic Eu-
lophidae (excl. Tetrastichinae). Index of Ento-
mophagous Insects. Le Francois, Paris. 254 pp.
Erdos, J. 1958. Eulophidae in Hungaria recenter de-
tectae. Acta Zool. Hung. 3: 205-223.
Forster, A. 1841. Beitrage zur Monographie der
Pteromalinen Nees. Aachen. 46 pp.
1856. Hymenopterologische Studien. II.
Chalcidiae und Proctotrupii. Aachen. 152 pp.
. 1861. Ein Tagin den Hochalpen. Progr. Real-
schule Aachen fur 1860/61. 44 pp.
Graham, M. W. R. de V. 1959. Keys to the British
genera and species of Elachertinae, Eulophinae,
Entedontinae and Euderinae (Hym., Chalcidoi-
dea). Trans. Soc. Br. Entomol. 13: 169-204.
Hansson, C. 1987. Revision of the New World species
of Chrysocharis Forster (Hymenoptera: Eulophi-
dae). Entomol. Scand. Suppl. 31: 1-87.
Thomson, C. G. 1878. Hymenoptera Scandinaviae.
Vol. 5. Lund. 307 pp.
Walker, F. 1839. Monographia Chalciditum. I. Lon-
don. 333 pp.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 66-70
NATURALLY OCCURRING HOST SITES FOR
XYLOPHILIC CECIDOMYIIDAE (DIPTERA)
Emity A. Rock AND DALE JACKSON
(EAR) Wayne General and Technical College, 10470 Smucker Road, Orrville, Ohio
44667; (DJ) Department of Biology, The University of Akron, Ohio 44325.
Abstract. — Existing data on host sites for xylophilic Cecidomyiidae larval development
are derived from observations of sawn trees and logs. Field studies of hardwood trees at
three sites in northeast Ohio show that strong winds and animal activity break live branches
of suitable size and thus expose vessels of =75 u which are used for larval development.
Such damaged branches can support large populations of midges. Larvae develop in both
the proximal and distal sides of the break. Branches remain suitable for oviposition and
larval development for one to several months, depending on branch diameter and weather
conditions.
Key Words:
Xylophilic Cecidomyiidae use freshly ex-
posed vessels of hardwoods as a larval hab-
itat. Relationship between host selection and
vessel diameter has been previously re-
ported (Rock and Jackson 1985, 1986). As
with that of other investigators (Kieffer 1900,
Brues 1922), our initial encounter with these
cecidomyiids began when we observed fe-
males swarming on the cut surfaces of logs
and stumps of trees exposed in logging op-
erations. Although we also used cut logs
during earlier studies, we speculated on the
availability of such niches under natural
conditions. Our current investigation ex-
amines 1) the role of strong winds and an-
imal activity in exposing larval niches, 2)
utilization of vessels in live branches that
remain on the tree, 3) the number of larvae
that one branch can support and 4) the length
of time after exposure that a branch remains
suitable for use.
METHODS
Storm and squirrel damage.—Two sites
in northeast Ohio were monitored for two
wood loving, tree damage, squirrel activity
years (1985 and 1986) for live, broken
branches. One site was a small island of
approximately 7500 square meters located
near a lake shore with five mature oak
(Quercus alba L.) and one mature ash (Frax-
inus americana) trees (known xylophilic
cecidomyiid hosts (Rock and Jackson
1986)). The other was part of a large sub-
urban garden with a study area of 5625
square meters with five mature oak and three
mature ash trees. The areas below the trees
were mowed regularly to facilitate collection
of fallen twigs and branches. We recorded
the diameter of only the live branches be-
cause previous studies have shown that dead
wood is not a suitable larval habitat. Some
larger branches were kept at the field site to
monitor their use by cecidomyids. The
suitability of the branches for larval devel-
opment was verified by subsequent collec-
tion of larvae from the branch ends.
Squirrels were active at the field sites and
numerous live host tree branches were col-
lected that squirrels had severed by chew-
ing. Branches broken by squirrels were
VOLUME 91, NUMBER 1 67
YO —-———— ——— ay
i
70
60
f 50 ie
;
4 40
;
a 30
8
fs
B 20
10
SB,
0 rae ieee oe cele —
es rr ae Fer ee rt Fe a eee 5 [a east
0.0 0.5 15 2.0 2.5 3.0 15°50 55 60 65 70 7.5 8.0 85 9.0 9.5 100
BRANCH DIAMETER (inm)
Fig. 1. Number and size of live branches broken from hardwood hosts at test sites. Branches <2 mm d not
included in collection.
readily identified by teeth marks. Branch
diameters were recorded.
Suitability of proximal section of broken
branches.—In earlier studies, we had for
convenience used only the severed branch
sections. The current study investigates the
use of the intact basal parts as their use would
double the number of potential larval de-
velopment sites.
We cut a total of fifty-nine branches rang-
ing from 13 mm d to 30 mm d from ash,
oak and elm trees at field sites in Akron and
Orrville, Ohio. The severed branches were
placed at the base of the tree to confirm the
presence of females. Branch ends on the trees
were left exposed for ten days to allow fe-
males to oviposit. After this time, the ends
were covered by a plastic cup; the branch
and cup were then covered with a nylon bag
to hold the cup in place. Rain wetting the
branches stimulated the emergence of lar-
vae. We recorded the number of larvae that
fell into the cups without identifying them
to species. The branches were uncovered for
three days to allow for possible further ovi-
position; the cups were then replaced. This
procedure was repeated for up to seven
weeks.
Potential larval yield from small branch-
es.—We had previously determined that
host branches of 6 mm or greater in di-
ameter usually contain xylem vessels of =>75
u d, which are suitable for larval develop-
ment (Rock and Jackson 1986), but never
recorded the number of midges that emerged
from indvidual twigs. Twenty sassafras
(Sassafras albidum (Nutt.) Nees) branches
with diameters varying from 3 mm to 20
mm were offered to a population of 300
caged Xylodiplosis longistylus Gagné fe-
males for one week. The branches were held
at room temperature in plastic bags for 14
68 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
NUMBER OF BRANCHES COLLECTED
40 45 5.0 5.5 60 65 70 75 80 85 9.0
BRANCH DIAMETER (mm)
Fig. 2.
days. Each was then soaked in a separate
container of water, and the number of larvae
emerging from both exposed ends was re-
corded.
Period of suitability of cut branches for
larval development.—To establish if there
was a connection between length of use of
a severed branch and branch diameter, we
monitored larval emergence in branch sam-
ples of various diameters from the field sites.
Unidentified xylophilic larvae emerged from
vessels in approximately 14 days after ovi-
position if the branch was soaked in water
(Rock and Jackson 1985). When a wood
sample ceased to yield larvae, we assumed
it had lost its attractiveness to females ap-
proximately two weeks earlier. We then
compared duration of attractiveness to
sample diameter.
RESULTS AND DISCUSSION
Strong winds create a source of suitable
Oviposition sites. Although the majority of
Number and size of live branches broken from host trees by squirrel activity.
the live branches broken from host trees
during an entire summer are <5 mm d (Fig.
1), high winds break off some branches of
6 mm d or greater, which contain suitably
sized vessels. Branches with diameters rang-
ing from 15 mm to 35 mm were occasion-
ally broken throughout the summer. At oth-
er locations we observed major damage due
to lightning and heavy snow that also cre-
ated sources of exposed vessels. As dis-
cussed below, the availability of larger
branches is very important as they can sup-
port several generations of gall midges each
summer.
Squirrels expose the ends of live branches
when they feed and build their nests (Short-
en 1954). Although most of the branches
are less than 6 mm d (Fig. 2), some larger
branches are severed and can serve as suit-
able sites for midge development. Gray
squirrels (Sciurus nigra) are known to build
nests in many of the hardwoods that are also
hosts to xylophilic midges. Uhlig (1955), in
VOLUME 91, NUMBER 1
NUMBER OF LARVAE COLLECTED
(Thousands)
69
20.0
9.0 10.0
BRANCH DIAMETER (mm)
Fig. 3.
an extensive study of the gray squirrel in
West Virginia, noted the use of oak, elm,
and hickory, and less frequently willow,
walnut, and sassafras, in nest building. Wal-
nut and oak trees at our field sites contained
squirrel nests. Leaf nests built of twigs and
leaves from the tree in which they are lo-
cated have been noted from early spring
throughout the summer months. Juvenile
squirrels are the primary leaf nest builders,
and the actual number of nests is directly
proportional to the rearing success of the
spring and summer litters (Uhlig 1955). One
juvenile may build more than one nest
throughout the summer and early fall. Thus,
with several juvenile squirrels in an area, a
supply of exposed branches exists for sev-
eral months.
Fig. 3 summarizes a laboratory study, us-
ing sassafras, which shows a direct correla-
tion between branch diameter and the num-
ber of larvae collected. We previously
determined that larvae require vessels of
Relationship of number of larvae collected to branch diameter: Sassafras albidum.
=75 ud for development and that it is pos-
sible to predict the number of potential lar-
val habitats based on the number of vessels/
mm? in the sapwood. The 6 to 8 mm d
branches are suitable for larval develop-
ment but each yielded only a few larvae.
However, the 20 mm d sassafras sample had
approximately 1850 vessels =75 » d/mm?
in each end, and it yielded over 1700 larvae.
Preliminary field data indicate that actual
utilization of vessels is much less than in
laboratory samples. A 12 mm dash branch,
for example, yielded 6 larvae, and a 50 mm
d branch yielded 100 larvae during 12 days
of field collecting. In natural conditions,
small, broken branches are available to the
flies throughout the summer and can sup-
port small overlapping populations. Al-
though large branches are available less fre-
quently, they permit rapid population
increases and provide larval niches for ex-
tended periods of time.
The live basal sections of severed branch-
70 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Relationship of host sample diameter to
length of attractiveness.
Pengtn
c8)
Attrac-
Diameter tiveness
Host Plant (cm)* (Weeks)
Juglans nigra L. 35 15
Fraxinus americana 22 10
Carya ovata (Mill.) K. Koch 21 12
Quercus alba L. 14 14
Ulmus americana L. 12 14
Populus deltoides Bartr. 12 10
U. americana 11 1
Salix babylonica Marsh 9 8
Sassafras albidum (Nutt.) Nees 7 8
F. americana 5 8
F. americana 2to4 8
F. americana 1 to 2 6
F. americana 0.6 to | 4
* All samples 25 to 35 cm long.
es are attractive to females and suitable for
larval development. Larvae emerged from
most of the attached broken branches, the
largest number collected at one time being
30 from a 30 mm ash branch. No larvae
emerged after the sixth week of exposure,
and we assumed branches had ceased to at-
tract female midges.
The time period during which midges uti-
lize a cut branch (Table 1) is directly pro-
portional to its diameter. As the life cycle
of most xylophilic species is approximately
4 weeks, during the summer in northeast
Ohio it is possible for one 35 cm d log to
support three generations of gall midges.
Major factors that limit the period of at-
tractiveness are speed of decay and rainfall.
General observations over several summers
indicate that abnormally wet periods pro-
mote fungal growth on exposed branch ends
and so reduce the length of time that a log
is suitable. Weather records, however, 1n-
dicate that summers with increased shower
activity are associated with more frequent
episodes of high winds which lead to an
increase in the number of severed branches
(Robert Thompson, personal communica-
tion, National Weather Service, North Can-
ton, Ohio 1988).
LITERATURE CITED
Brues, C. T. 1922. Some hymenopterous parasites of
lignicolous Itonididae. Proc. Am. Acad. of Arts
and Sci. 57: 263-287.
Kieffer, J. J. 1900. Monographie des Cecidomyides
d’Europe et d’Algérie. Ann. Soc. Entomol. Fr. 69:
181-472 and pls. 15-44.
Rock, E. and D. Jackson. 1985. The biology of xy-
lophilic Cecidomyiidae (Diptera). Proc. Entomol.
Soc. Wash. 87: 135-141.
1986. Host selection in xylophilic Cecido-
myiidae (Diptera). Proc. Entomol. Soc. Wash. 88:
316-319.
Shorten, M. 1954. Squirrels. Collins, London. 212
pp.
Thompson, R. 1988. Personal communication. Na-
tional Weather Service, North Canton, Ohio.
Uhlig, H.G. 1955. The gray squirrel: Its life history,
ecology, and population characteristics in West
Virginia. Pittman-Robertson Project 31-R. Con-
servation Commission of West Virginia. 175 pp.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 71-80
TWO NEW TERRESTRIAL ISOPODA (ONISCIDEA) FROM
CORALLINE CAYS OF VENEZUELA’S CARIBBEAN COAST
Maurizio G. PAOLETTI AND BENJAMIN R. STINNER
(MGP) Department of Biology, University of Padova, 35100-Padova, Italy, Visiting
Professor, Department of Entomology, The Ohio State University; (BRS) Department of
Entomology, The Ohio State University, Wooster, Ohio 44691.
Abstract.—Two terrestrial, halophilous isopods, Metastenoniscus neotropicalis n. sp.
(Stenoniscidae) and Armadilloniscus caraibicus n. sp. (Scyphacidae) are described from
coralline cays of Venezuela’s Caribbean coast. Buchnerillo litoralis Verh. and Stenoniscus
pleonalis Aubert and Dollfus are reported from the Caribbean region.
Key Words:
Isopoda, Oniscidea, Neotropical, Caribbean, Metastenoniscus neotropicalis
n. sp., Armadilloniscus caraibicus n. sp., Buchnerillo litoralis, Stenoniscus
pleonalis
Knowledge of the Venezuelan terrestrial
isopoda fauna is reported in a few post-
worldwar papers: Brian 1957, Vandel 1952,
1968, 1972, Andersson 1960, Mulaik 1960,
Strinati 1971, Schultz 1971, 1983, 1984,
following the classic studies of Van Name
1936, 1940, 1942. We have followed in gen-
eral the terminology proposed by Holdich
1984, Holdich et al. 1984. Our work is
mostly based on SEM (Scanning Electron
Microscopy).
The two new species that we describe be-
long to the halophilous seacoast fauna. The
tergites covered by longitudinal ribs suggest
that these species belong to the eco-mor-
phological category of creepers (Schmalfuss
1984, Paoletti 1987).
ONISCOIDEA
Stenoniscidae
Metastenoniscus neotropicalis n. sp.
Type locality. Coralline key (Cayo) of Pla-
yuela, Parque Morrocoy, estado Falcon,
Venezuela. 28 females and 12 males were
collected in soil litter of Coccolobis uvifera
(L) Jacq. (Polygonaceae), by M. G. Paoletti
on January 1, 1986 (Paoletti 1988).
Male holotype, allotype and paratypes are
located in the M. G. Paoletti collection; 3
female paratypes are deposited in the Mu-
seo Zoologico of the Padova University; 3
female paratypes are located in the general-
collection of Instituto Museo de Zoologia
Agricola, Universitad Central de Venezue-
la, Maracay, Aragua, Venezuela.
Diagnosis. This new species is similar to
Metastenoniscus osellai Taiti and Ferrara,
1981. The smaller body is more cylindrical
and holds less enlarged epimera. The telson
is shorter and three-lobed.
Dimensions: length males: 1.4-1.86 mm,
mean 1.57 mm; width males: 0.42-0.56 mm,
mean 0.50; length females: 1.48-—2.42 mm,
mean 2.14 mm; width females: 0.48-0.84
mm, mean 0.71 mm (Figs. 1, 2).
Body ornamentation: consists mainly of
subcircular plaques (Figs. 4A, C, D); body
shape subparallel, costulated, light violet
colored; pereonites I-IV without medial rib
(Figs. 3A, C).
72 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
0.57274
0.532 4
E
E 0492
B=
mo)
= a
0.452 4 y -13.8493+01428-x
r= 0.2107
p-01942
a
0412
T T T if T
136 150 1.64 178 192
Length (mm)
Fig. 1. Regression for male dimensions of Metas-
tenoniscus neotropicalis n. sp.
Head: with three posterior main tubercles
and two median main tubercles (Figs. 3A,
B, C); eye with 34 ommatidia; antenna with
flagellum consisting of two articles; the sec-
ond article bears on its foreword-facing sur-
face three aesthetascs (Figs. 3E, F). Anten-
nule two-articulated with three apical
aesthetascs, the exterior occasionally bro-
ken at the base (fig. 4E).
Telson: short and distinctly three-lobed
(Figs. 3C, D; 4B, D).
Uropods: the basis is longer than wide
(Fig. 4B), the exopod bears distally a spike
of 4—5 aesthetascs (Figs. 4D, F); the endo-
pod much longer than exopod (Figs. 4C, D),
thickened in the middle, end with a spike
of three elements (Fig. 4C).
Pleopods: male endopod of first and sec-
ond pair are little differentiated (Fig. 5).
Affinities. M. neotropicalis n. sp. is dis-
tinct from Metastenoniscus osellai Taiti and
Ferrara, 1981, described from Bali by the
following features:
1. females and males are smaller;
2. epimeral appendages less developed and
body smaller;
3. head with a different arrangement of tu-
bercles especially in the posterior part:
five on M. osellai and three on M. neo-
tropicalis n. sp.;
4. pereonites I and II with only a hind ves-
tige of medial rib; in M. osellai the per-
0.86 4
y --0.1586 + 03349-x
r= 0.8204
p< 0.0001 a 48
0.76 4
0.66 5
Width (mm
0.56 5
046 5
‘lt Tig
144 1.70 196 2.22 248
Length (mm)
Fig. 2. Regression on female dimensions of Me-
tastenoniscus neotropicalis n. sp.
eonite from | to 4 has medial costa well
developed (Taiti and Ferrara 1981 Figs.
lGand 2©@):
. telson shorter and distinctly three-lobed;
6. uropod exopods shorter and with wider
basis.
Nn
Habitat. Found under Coccolobis uvifera
(L.) Jacq. (Polygonaceae) (uva de plaja) lit-
ter, eating decayed litter tissue (Paoletti 1987
Figs. 7D, G, H). This isopod was not yet
found in the intertidal zone but strictly in
the interior part of the coralline cays and
above the upper tidal level.
Distribution. Discovery of M. neotropi-
calis n. sp. considerably enlarges the known
geographic range of the family Stenonisci-
dae. In fact Metastenoniscus is now repre-
sented not only in the Oriental region (Bali
and the Andaman Islands) but also in a Pan-
tropical belt including the Caribbean region.
That they only now have been discovered
in the Caribbean region is probably due to
their small size.
Discussion. Stenoniscus pleonalis Aubert
and Dollfus, 1890 (sensu Vandel 1962) is
easily distinguishable from Metastenonis-
cus oSellai (Taiti and Ferrara, 1981) and from
M. neotropicalis n. sp. It was reported only
rarely beyond the Mediterranean region and
in the Neotropical region only twice (Vandel
1968, Schultz 1972). We collected Stenon-
iscus pleonalis Aubert and Dollfus (sensu
VOLUME 91, NUMBER 1
Fig. 3.
ee ee
Metastenoniscus neotropicalis n. sp. Female: A dorsal view, B head frontal view, C lateral view, D
73
100umS
—e
2
pleon and telson, E and F second antennal flagellum article from dorsal and ventral views.
Vandel 1962) in Florida Cays, Tavernier
Creek, on rocky beach: in the soil, under
Posidonia and on coconut drift in the beach,
M. G. Paoletti, October 26, 1987. This lat-
ter observation is interesting. Under the
Posidonia drift in the same location was col-
lected one female of Buchnerillo litoralis
Verhoeff, 1943, which represents a first re-
cord for the Neotropical region (Schultz and
Johnson 1984), and Vandeloscia culebre
(Moore, 1901), Ty/los niveus Budde-Lund.
and Armadilloniscus ellipticus (Harger,
1878).
ONISCOIDEA
Scyphacidae
Armadilloniscus caraibicus n. sp.
Type locality. Coralline cays (Cayos) of
Parque Morrocoy, estado Falcon, Venezue-
la: Cayo Sombrero, one gravid female
31.X11.1985; Cayo Cico, 8 males, 16 fe-
males, 7 gravid females, 3.1.1986 were col-
lected by M. G. Paoletti. Isopods were col-
lected in the intertidal zone under coralline
rocks lying on the sandy beach at Cayo Cico;
at Cayo Sombrero the gravid female was
74 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
NW
Aho
>, aoe | TRS:
ORION
Fig. 4.
Metastenoniscus neotropicalis n. sp. Female: A I and II pereonites, B ventral view of hind pereon,
pleon and uropods, C uropodal endopods, D below view of pleon and uropods, E second article of antennulae
with aesthetascs, F uropodal exopod.
found among small woody debris on the
coarse sandy beach (Paoletti 1988).
Male holotype, allotype and paratypes are
located in M. G. Paoletti collection; 3 fe-
male paratypes are deposited in the Museo
Zoologico of the Padova University; 3 fe-
male paratypes are deposited in the general-
collection of Instituto Museo de Zoologia
Agricola, Universitad Central de Venezue-
la, Maracay, Aragua, Venezuela.
Diagnosis. Body elliptical, dull brown,
covered with prominent ridges of tubercles
on the head, forming ribs on the pereon and
pleon (Figs. 10, 11). Ornamentation con-
sists of circular plaques (Figs. 11F, H) and
digitiform trichomes.
VOLUME 91, NUMBER |
Fig: 5:
Metastenoniscus neotropicalis n. sp. Male, A and B pleopod I endopods, C pleopods I exopod, D
pleopods II endopod. Armadilloniscus caraibicus n. sp. Male, E pleopods I, F pleopods II, G VII male pereopods.
Dimensions: length males: |.92—2.46 mm,
mean 2.27; width males: 0.84-1.26 mm,
mean 1.05 (Fig. 6); length non gravid fe-
males: 1.25—3.9 mm, mean 2.34; width non
gravid females: 0.75-1.92 mm, mean 1.095
(Fig. 7); length gravid females: 2.85—3.36
1.30 4
y - 0.0437 + 0.4449-x
= 0.37
ASI p= 0.45
4
E
E
= 1064
2
Ss
0944
a
0.824
T a af = a5 | T. st
189 2.04 219 2.34 249
Length (mm)
Fig.6. Armadilloniscus caraibicus n. sp. Regression
of male dimensions.
mm, mean 3.0; width gravid females: 1.35—
1.56 mm, mean 1.44 (Fig. 8); 6-9 pulli per
gravid female were counted (Fig. 9).
Head: with lateral lobes truncated at tip
and median lobe pointed, with three main
tubercles in the posterior and two in the
23-4
y = 0.0302 *0.4546:x
ol 0.84
1
a9) P<0.000 ‘
E
E
= 154
Sl
=
11
a
07-4
a =| aE T a i
14 19 27 35 43
Length (mm)
Fig. 7. Armadilloniscus caraibicus n. sp. Regression
of non gravid female dimensions.
76 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1584 : 3.38 4 > -
a
a y = 2.1322+01178-x
r= 0.564
a
152 < 3244 p-00912
D
2 2
E ®
E 2
- 4.464 ® 310-4
= c
3 5
‘ 0.7228 +0,2364 =
140 4 y & 296-4
r?- 0.205 5
ee p= 0.397
a a
1.344 2824
T T T 7 T
2.82 296 3.10 3.24 3.38
Length (mm) Pulli per female
Fig. 8. Armadilloniscus caraibicus n. sp. Regression Fig.9. Armadilloniscus caraibicus n. sp. Regression
of gravid female dimensions. of gravid female length and pulli per female.
BD he -
100 pm
Fig. 10. Armadilloniscus caraibicus n. sp. Female: A and B pleon; C and D head and pereon.
VOLUME 91, NUMBER |
Fig. 11. Armadilloniscus caraibicus n. sp. Female: A-D body surface in different perspectives; E head; F
pereonite ornamentation; G and H pleon and peculiar tubercle ornamentation.
anterior part (Figs. 1OC, D, 11A, C, D, E). (Figs. 12C, D); antennule characteristically
Eye with 4—5 visible ommatidia; antenna, three-articulated and apically bifurcated; it
without enlargements, holds four flagellar bears aesthetascs (5 on specimens exam-
articles, the second with three and the third ined) (Figs. 12A, B).
with two aesthetascs on the inferior surface Body ornamentation: pereon with four
77
78 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
: c ;
» Fs,
Fig. 12. Armadilloniscus caraibicus n. sp. Female A-F. Male G and H. A and B antennules; C and D antennal
four-article flagellum in dorsal and ventral view; E forelegs; F maxillipeds; G and H male pleopods I.
VOLUME 91, NUMBER 1
main medial ridges and four shallow lateral
ridges of tubercles (Figs. 10C, 1 1 A-E); pleon
supporting two main ridges of tubercles
(Figs. 1OA, B; 11B, D, G).
Uropods: endopod somewhat longer than
basis, holding a distal spike of 3-4 aesthe-
tascs (Figs. 10B, 11G). Male pereopods I
and VII slightly modified (Fig. SG). Pleo-
pods: male pleopods vary little from other
members of the genus (Figs. SE-F).
Affinities. The development of tubercle
ribs on the head, pereon and pleon are the
features of Armadilloniscus caraibicus n. sp.
by which it can easily be differentiated from
other species (Arcangeli 1957, Schultz 1972,
1977, Garthwaite et al. 1985). From the
more tuberculated A. coronacapitalis Men-
zies, this new species is distinguishable for
its smaller size, less tuberculation, differing
sculpture on the head, the shape of pleopod
I endopod, antennae, and antennules. It can
be separated from Armadilloniscus quadri-
cornis Vandel, 1971, 1973 by different head
structure and ornamentation and by its
smaller size.
Habitat. The specimens were found un-
der coralline rocks on the sandy beach in
the intertidal zone. Only one specimen was
found between woody debris on the beach.
The intertidal habitat seems to be the pre-
ferred habitat of the genus 4rmadilloniscus.
Distribution and Discussion. Armadil-
loniscus caraibicus n. sp. was found on cor-
alline cays (cayos) of Parque Morrocoy,
Venezuela which extends southward in the
Neotropical region the distribution of Ar-
madilloniscus. A revision of the described
species and of Scyphacidae genera 1s needed
to have a better taxonomic and zoogeo-
graphic understanding of the group. At pres-
ent little is known about the phylogenetic
relationships of the Scyphacidae and other
neotropical isopods.
ACKNOWLEDGMENTS
Weare indebted to Carlos Bordon for field
collection and discussion. A. L. Dreon, C.
Furlan, U. Arezzini, C. Friso assisted with
79
the figures and photographs, C. Britton and
F. Ferrara with bibliographical research. F.
Purrington revised the text. The Ministero
Italiano della Pubblica Istruzione provided
financial support.
LITERATURE CITED
Andersson, A. 1960. South American isopods in the
collection of the Swedish State Museum of Natural
History. Arkiv for Zool. 12: 537-569.
Arcangeli, A. 1957. Il genere Armadilloniscus Ulj. €
gli Scyphacidae. Atti Acc. Sc. Torino 91: 1-30.
Brian, A. 1957. Descrizione di Neosanfilippia vene-
zuelana n. gen., n. sp. di Isopodo terrestre trog-
lobio. Ann. Mus. Civ. St. Nat. Genova 69: 352-
360.
Ferrara, F. and S. Taiti. 1981. Isopodi terrestri delle
isole Adamane. Boll. Mus. Civ. St. Nat., Verona
8: 459-492.
Garthwaite, R. L., F. G., Hochberg, and C. Sassaman.
1985. The occurrence and distribution of terres-
trial isopods on Santa Cruz Island with prelimi-
nary data for the other California islands. Bull.
Southern California Acad. Sci. 84: 23-37.
Holdich, D. M. 1984. The cuticular surface of wood-
lice: A search for receptors. The biology of terres-
trial isopods, S. L. Sutton and D. M. Holdich, eds.,
Zool. Soc. London Symp. 53: 9-48.
Holdich, D. M., R. J., Lincoln, and J. P., Ellis. 1984.
The biology of terrestrial isopods: Terminology
and classification. The biology of terrestrial iso-
pods, S. L. Sutton and D. M. Holdich, eds., Zool.
Soc. London, Symp. 53: 1-6.
Mulaik, S. B. 1960. Contribucion al conocimiento de
los Isopodos terrestres de Mexico. Revista de la
Soc. Mexicana de Hist. Natural 21: 79-220.
Paoletti, M. G. 1988. Life strategies of isopods and
“soil invertebrates” in neotropical Venezuela.
Monit. Zool. It., Mon. Ser. (In press.)
Schmalfuss, H. 1984. Eco-morphological strategies
in terrestrial isopods. The biology of terrestrial
isopods, L. S. Sutton and D. M. Holdich, eds.,
Zool. Soc. London Symp. 53: 49-63.
Schultz,G. A. 1971. A review of species of the family
Scyphacidae in the New World. Proc. Biol. Soc.
Wash. 84: 477-488.
1972. Ecology and systematics of terrestrial
isopod crustaceans from Bermuda. Crustaceana,
Supp. 3, pp. 79-99.
. 1977. Terrestrial isopod crustaceans from St.
Catherines Island, Georgia. Georgia J. Sci. 35: 151-
158.
1983. Disposition of three species of Onis-
coidea from Western Atlantic seashores. Proc. Biol.
Soc. Wash. 96: 440-451.
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Oniscoidea from Belize, Central America. J. Nat.
Hist. 19: 3-14.
Schultz, G. A. and C. Johnson. 1984. Terrestrial iso-
pod crustaceans from Florida. J. Crust. Biol. 4:
154-171.
Strinati, P. 1971. Recherches biospeleologiques en
Amerique du Sud. Ann. de Speleologie 26: 439-
450.
Taiti, S. and F. Ferrara. 1981. Metastenoniscus osellai
genere e nuova specie di isopodo terrestre dell’ isola
di Bali. Boll. Mus. Civ. St. Nat., Verona 8: 443-
452.
Vandel, A. 1952. Etude des Isopodes terrestres re-
coltes au Venezuela par le dr. G. Marcuzzi. Mem.
Mus. Civ. St. Nat. Verona 3: 59-203.
1962. Faune de France Isopodes terrestres.
Lechevalier, Paris.
1968. I. Isopodes terrestres. Mission zoolo-
1984. Three new and five other species of
gique belge aux iles Galapagos et en Ecuador (N.
et J. Leleup, 1964-1965). Resultats scientifiques.
Premiere partie 1: 35-168.
. 1971. Les Isopodes terrestres des iles Rennell
et Bellona. The natural history of the Rennell Is-
land, T. Wolff, ed., 6: 139-153.
. 1972. Les Isopodes terrestres de la Colombie.
Studies on the Neotropical Fauna 7: 147-172.
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Zool. verhandelingen, Leiden 125: 1-160.
Van Name, W. G. 1936. The American land and
fresh-water isopod Crustacea. Bull. Amer. Mus.
Nat. Hist. 71: 1-535.
1940. A supplement to the American land
and fresh-water isopod Crustacea. Bull. Amer. Mus.
Nat. Hist. 77: 109-142.
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Mus. Nat. Hist. 80: 299-329.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 81-87
ANTS ASSOCIATED WITH A COLEOPTEROUS LEAF-BUD
GALL ON XYLOPIA AROMATICA (ANNONACEAE)
G. WILSON FERNANDES, WILLIAM J. BOECKLEN, ROGERIO P. MARTINS,
AND ALEXANDRE G. CASTRO
(GWF, WJB) Box 5640, Department of Biological Sciences, Northern Arizona Uni-
versity, Flagstaff, Arizona 86011; RPM, AGC Departamento de Biologia Geral, Caixa
Postal 2486, ICB/Universidade Federal de Minas Gerais, 30.000-Belo Horizonte-MG,
Brazil.
Abstract.—We describe the ant fauna (11 species divided into four genera) associated
with a coleopterous leaf-bud gall on Xylopia aromatica (Annonaceae) in Minas Gerais,
Brazil. Ant occupied galls are significantly larger than are unoccupied galls, and the vari-
ance of gall diameters for occupied galls is significantly smaller than is the variance for
all galls. There is a significant positive relationship between ant colony size and gall
diameter. The six most common species of ants may partition galls according to gall size,
as there are significant differences in the diameters of occupied galls among species. At
the community level, patterns of co-occurrence of ants are indistinguishable from those
expected under a random assortment model. Individual ant species do exhibit non-random
patterns of co-occurrence.
Key Words:
Annonaceae, ants, Brazil, coleopterous gall, community ecology, habitat
selection, insect galls, Minas Gerais, resource partitioning, Xy/opia aro-
matica
Plant galls represent an important re-
source for many species other than the gall
formers. Owing to their localized concen-
tration of nutritive tissues and their marked
succulence, galls provide favourable breed-
ing sites for a variety of species (Brandhorst
1962, Mani 1964, Shorthouse 1973, Yu-
kawa 1983). Galls protect their inhabitants
from inclement weather (Felt 1940, Uhler
1951, Sandlant 1979; but see Baust et al.
1979) and natural enemies (Askew 1961,
1980, but see Price et al. 1986, 1987). Use
of galls ranges from species that open galls
simply to prey upon gall formers and in-
quilines to those that depend exclusively on
gall tissues for food and shelter.
Beauvisage (1883, cited in Mani 1964)
applied the term “‘locatari” to species, other
than the gall formers, associated with insect
galls. Mani (1964) divides the locatari into
33 categories according to their ecological
niches. The locatari often represent a vast
fauna with the most numerous categories
consisting of parasites and predators. For
example, Stegagno (1904) reported 177
species associated with cynipid galls on
Quercus in Italy; of these, 138 are parasites
and predators.
Species that inhabit galls after emergence
of gall formers and inquilines are the ‘“‘suc-
cessori’’ (Mani 1964, Yukawa 1983). Most
of these are plant-nesting ants and myr-
mecophilous insects, such as aphids and
coccids. Others include mites, spiders,
thrips, collembola, bees, and wasps (re-
viewed by Mani 1964). The most common
ant genera associated with old galls are:
Camponotus, Cataulacus, Crematogaster,
82 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
GERAIS
Sao Paulo
we
=
2h omine
Rio de Janeiro
Fig. 1.
Colobopsis, Lasius, Leptothorax, and Olop-
sis (Walsh 1864, Patton 1879, Wheeler 1910,
Ping 1920, Sturtevant 1925, Mani 1964,
Torossian 1971a, b). Despite their impor-
tance and dominance within the successori,
little is known about the ecologies of these
ants other than compendiums of ant species
associated with particular species of gall for-
mers (e.g. Brandhorst 1962, Espadaler and
Nieves 1983).
Here, we describe the ant fauna associ-
ated with a coleopterous leaf-bud gall on
Xylopia aromatica (Annonaceae). We de-
scribe patterns in distribution and abun-
dance of the ant fauna and document ele-
ments of habitat selection and resource
partitioning.
MATERIAL AND METHODS
One of us (G.W.F.) collected galls from
the Ecological Station of Parapitinga, Trés
Marias, Minas Gerais, Brazil during one
sample period in mid-May, 1984. The sta-
tion is a continental island located in a man-
Location of the Ecological Station of Parapitinga in Trés Marias, Minas Gerais, Brasil.
made lake (Trés Marias Lake) between lat-
itude 18°-19° south and longitude 45°-46°
west (Fig. 1).
The host tree, Xylopia aromatica, oc-
curred in a gallery forest along the northwest
border of the island. Only three individuals
of approximately six meters tall bearing galls
were observed in the area surveyed. The
galls are induced by an unidentified species
of curculionid (Coleoptera). Gall formers
held as vouchers were kept in the author’s
collection. The galls are spherical and gla-
brous and occur on leaf-buds (Fig. 2). They
are green when occupied by the gall former,
but turn brown after its emergence. All galls
which were on trees (n = 114) were collected
and subsequently measured and dissected
in the laboratory. Galls that fall on the forest
floor are also utilized by ants; however they
are not included here because of insufficient
sample size.
In this paper, we use the word “colony”
as any group of ants, composed of workers
and larvae (eggs, queen, and pupae if pres-
VOLUME 91, NUMBER I
e."
Fig. 2.
21.5 mm.
ent), occupying a single gall. In addition, we
defined those galls in which we found only
workers as “groups of workers.”
STATISTICAL ANALYSES
We compared the mean diameter of galls
that were occupied by ants with the mean
diameter of unoccupied galls using one-way
83
Coleopterous leaf-bud galls found on Xylopia aromatica. Gall diameter ranged from 5.0 to
analysis of variance. We also compared the
variance of occupied galls with the variance
of all galls (including occupied galls) with a
Chi-square test (Sokal and Rohlf 1969, page
175). We assumed that our collection con-
sisted of the entire population of galls on
the island and that the set of occupied galls
was a sample from that population.
84
Table |.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ant species associated with coleopterous leaf-bud galls on Xylopia aromatica (Annonaceae).
(2) Individuals
Gall Diameter
Ant Species Number of Colonies Adult Larvae/Pupae (2) mm
Azteca bicolor l 1.0 - 14.5
Azteca sp. 17 28.1 8.9 16.4
Leptothorax wilda 13 5.4 4.6 14.7
Leptothorax sp. | 45.0 37.0 17.0
Pseudomyrmex flavidus 4 0.8 5:5 16.4
Pseudomyrmex sp. A 1 1.0 = 12:5
Pseudomyrmex sp. B 1 1.0 — 15.0
Zacryptocerus pallens 7 2.0 - 12.9
Zacryptocerus pusillus 8 51-3 12.5 15.9
Zacryptocerus sp. A 11 S27, DD) 14.9
Zacryptocerus sp. B l 4.0 4.0 10.0
We compared the mean diameters of oc-
cupied galls among the six most common
species of ants by one-way analysis of vari-
ance.
We estimated the relationship between the
logarithm of colony size (number of indi-
viduals, of all castes, in a gall) and gall di-
ameter through linear regression.
Finally, we examined patterns of occur-
rence within galls. We compared the num-
ber of galls that contained 0, 1, 2, and 3
species of ants with those expected under a
Poisson distribution. We estimated A, the
Poisson parameter, from the sample. We
compared the observed distribution to the
expected distribution with a Chi-square
goodness-of-fit test.
RESULTS
We collected 114 galls of which 52 (45.6%)
were occupied by ants. Eleven species of
ants were represented in the sample; the
most common were Azteca sp. which oc-
cupied 17 galls, Leptothorax wilda which
occupied 13 galls, and Zacryptocerus sp. “A”
which occupied 11 galls (Table 1). Five
species were represented by a single indi-
vidual. “Colony sizes” ranged from three
individuals to a colony of Zacryptocerus
pusillus which contained 129 adults and lar-
vae.
The internal structure of the galls was
highly modified in some cases, unmodified
in others (Fig. 3). Modification is defined as
any internal architectural departure, such as
tunnel and gall wall holes, from the usual
spheroid larval chamber. In particular, galls
inhabited by Azteca sp. and Zacryptocerus
pusillus exhibited extensive modifications
when compared to unmodified galls. Galls
inhabited by Pseudomyrmex flavidus were
unmodified. Modified galls typically con-
tained larger colonies than did unmodified
galls.
Ants typically occupied large galls (Fig.
4). The mean diameter of occupied galls,
15.6 mm, was significantly larger than the
mean diameter of unoccupied galls, 13.6 mm
(Fi.112 = 12.8, P < 0.001). The variance of
diameters of occupied galls was significantly
smaller than would be expected if ants were
selecting galls at random (x?,,; = 72.1, P <
0.005).
There was a significantly positive linear
relationship between the logarithm of col-
ony size (number of individuals) and gall
diameter (Fig. 5).
The six more common species of ants fur-
ther partitioned the subset of occupied galls
according to gall size; mean gall diameters
of occupied galls were significantly different
among species (F; 5, = 2.74, P < 0.05). Az-
teca sp. typically occupied the largest galls
(% = 16.4 mm), Zacryptocerus pallens the
smallest (¥ = 12.9 mm).
We collected 65 colonies distributed
VOLUME 91, NUMBER 1
Fig. 3.
cm
Cross-sections of galls showing internal modifications by ants. Gall (A) contained Azteca sp., (B)
Leptothorax sp., (C) Zacryptocerus pallens, (D) Zacryptocerus pusillus, and (E) Pseudomyrmex flavidus.
among | 14 galls. We estimated X, the Pois-
son parameter, as 65/114 = 0.57. The ob-
served distribution of colonies among galls
closely resembled the expected under a
Poisson distribution (Table 2). The good-
ness-of-fit test indicated no significant dif-
ALL DIAME TEF MM
Fig. 4. Relative frequency histograms of gall di-
ameter for the entire set of galls (foreground) and for
the subset of galls that were occupied by ants (back-
ground).
ference between the observed and expected
distributions (x7, = 2.51, P > 0.10).
Individual species did exhibit frequencies
of co-occurrence that appeared different
from those expected by chance alone (Table
3). For example, Zacryptocerus pusillus
<p) eile umaice
a | e e
<< = on - e
=) e fc
(ea) © eco
— e e e
> $ aoe e
a ae ne at ce e e
= ee, ry - <
(©) ee enn 0 e
oO
a
T 2 Ts
12.0 15C 18.0 PAK
GALL DIAMETER
Fig. 5. Relationship between the logarithm of col-
ony size (number of individuals) and gall diameter.
The relationship is Y = —0.929 + 0.122X (F, ,; = 20.1,
P < 0.001; R? = 0.24).
86 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Distribution of colonies among galls. Ex-
pected values are based on a Poisson distribution with
dA = 0.57.
Colonies/Gall Observed Expected
0 62 64.5
1 42 36.8
2 7 10.5
3 3 2.0
never jointly occupied a gall, while approx-
imately 4 such co-occurrences would be ex-
pected at random. Azteca sp. and Lepto-
thorax wilda were roughly half as likely to
co-inhabit a gall as chance alone would pre-
dict. On the other hand, Zacryptocerus pal-
/ens exhibited a slight tendency to share galls.
DISCUSSION
Galls represent an important resource for
this ant community, and ants exploit this
resource in a non-random fashion. Ants se-
lect significantly larger galls, over a narrow-
er range of sizes, than chance alone would
predict. Both the difference in means and
the difference in variance indicate habitat
selection. There may be strong selective
pressures for this habitat selection as indi-
cated by the significant positive relationship
between colony size and gall size. Selection
of larger galls by ants may be due to several
reasons, among them increase in queen fit-
ness. However, more work is called for to
observe the selection and the use of larger
versus smaller galls.
It is not clear whether galls are a limiting
resource to these ants. On the other hand,
the most common species of ants exhibited
resource partitioning. This is a necessary
condition for stable coexistence of multi-
species assemblages exploiting similar, lim-
iting resources predicted by Lotka-Volterra
based analyses of community dynamics
(May 1973, Schoener 1974). Of course, nei-
ther observation by itself is conclusive. Un-
occupied galls may be the result of interfer-
ence competition or priority effects (sensu
Torres 1984). Significant differences among
Table 3. Patterns of co-occurrences among ant
species.
Joint Occurrences
Species Observed Expected
Azteca bicolor l 0.58
Azteca sp. 4 Tad
Leptothorax wilda 3 5.93
Leptothorax sp. 0 0.58
Pseudomyrmex flavidus 3 2B)
Pseudomyrmex sp. A 1 0.58
Pseudomyrmex sp. B 0 0.58
Zacryptocerus pallens 5 3.54
Zacryptocerus pusillus 0 3.99
Zacryptocerus sp. A 5 5.19
Zacryptocerus sp. B 0 0.58
species in the diameters of occupied galls
may not be the result of competitively in-
duced resource partitioning, but rather,
species’ idiosyncratic responses to their en-
vironment (James et al. 1984). Correlative
data provide notoriously weak inferences
regarding mechanisms (Brady 1979); ex-
periments are required to establish causa-
tion.
Patterns of species co-occurrence at the
community level are indistinguishable from
those predicted from a random assortment
model. However at the constituent level,
individual species deviate from expecta-
tion. In particular, Zacryptocerus pusillus
never shares a gall, although approximately
four joint occurrences are predicted. Fowler
et al. (1985) provide evidence that Z. pusil-
/us is interspecifically territorial. Whether
deviations from chance for the other species
are biologically significant is unclear. Un-
fortunately, little is known about the ecol-
ogies of these ants.
Gall-inhabiting ants may provide an ideal
system to examine the determinants of
community organization in arboreal ants.
Galls are a discrete resource, amenable to
experimental manipulation in time and
space.
ACKNOWLEDGMENTS
We thank Hélcio R. Pimenta and Ivo das
Chagas for field assistance. We also thank
VOLUME 91, NUMBER 1
A. Mabelis and W. W. Benson for infor-
mation on ants, H. Castanheira and L. M.
Araujo for identification of ants, L. Ki-
noshita for identification of Xy/opia aro-
matica, and W. E. Clark for identification
of the gall former insect. In addition, we
thank the comments of H. Larew and an
anonymous reviewer on an early draft of
this manuscript. The Departamento de
Biologia Geral, ICB, Universidade Federal
de Minas Gerais and the staff of Codevasf-
Trés Marias, MG provided logistical sup-
port. This work was supported by a spe-
cialization fellowship from SEMA/CNPg (n°
10.6193/84) to G. W. Fernandes.
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PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 88-111
REVISION OF THE ORNATUS SPECIES GROUP OF THE GENUS
ANTHONOMUS GERMAR (COLEOPTERA: CURCULIONIDAE)
WAYNE E. CLARK AND Horace R. BURKE
(WEC) Department of Entomology and Alabama Agricultural Experiment Station, Au-
burn University, Alabama 36849; (HRB) Department of Entomology, Texas A&M Uni-
versity, College Station, Texas 77843.
Abstract.—The seven species in the Anthonomus ornatus species group are A. ornatus
Blanchard, A. signatipennis Blanchard, A. chilicola Clark, new species, A. kuscheli Clark,
new species, 4. araucanus Clark, new species, A. blanchardi Clark, new species, and A.
berberidis Clark, new species. These are hypothesized to constitute a monophyletic group
on the basis of morphological characters of the adult weevils, their common association
with plants in the genus Berberis (Berberidaceae) and their restricted distributions in
southern Chile and Argentina. Other characters determined to be apomorphic by com-
parisons with other anthonomines were used to produce a phylogeny of the species in the
A. ornatus group. Characters diagnostic of the group and of each of the species are pre-
sented, along with illustrations and a key to adults of the species. The larval and pupal
stages of A. ornatus and A. kuscheli are also described and illustrated.
Key Words:
The Anthonomus ornatus group includes
A, ornatus Blanchard, A. signatipennis Blan-
chard, and five heretofore undescribed
species. The species occur in southern Chile
and Argentina at least as far south as Isla
Navarino (55°S) in the Chilean province of
Magallanes northward to the province of
Coquimbo (30°S). Known hosts of the
species are plants in the genus Berberis (Ber-
beridaceae). The members of the A. ornatus
group appear to be the only representatives
of the subfamily Anthonominae in Chile.
This fact is noteworthy because the antho-
nomines are rich and diverse in the re-
mainder of the Neotropical Region (O’Brien
and Wibmer 1982, Wibmer and O’Brien
1986). It seems likely that the sister group
of the 4. ornatus group will be found among
the Nearctic or Neotropical species, but hy-
potheses of such relationships cannot be
formulated until these species are more
Anthonomus, Anthonominae, PAUP, phylogeny, Berberis, Chile, Argentina
completely known taxonomically. Mean-
while, the descriptions, illustrations and keys
presented in this paper should facilitate
identification of the species in the 4. ornatus
group.
MATERIALS AND METHODS
Specimens of 785 adults, including the
types of the previously described species,
were examined. These were from the col-
lections of the following individuals and in-
stitutions (letter codens identify the collec-
tions in the text):
AMNH The American Museum of Natural
History, New York City, New
York, USA (L. H. Herman, Jr.);
CACA_ Collection of Allan C. Ashworth,
Fargo, North Dakota, USA;
CCBM_ Collection of C. Bordon, Maracay,
Venezuela;
VOLUME 91, NUMBER 1
CWOB Collection of C. W. O’Brien, Tal-
lahassee, Florida, USA;
Collection of H. and A. Howden,
Ottawa, Ontario, Canada;
Museum of Comparative Zoology,
Cambridge, Massachusetts, USA
(S. R. Shaw);
Muséum National d’Histoire Nat-
urelle, Paris, France (H. Perrin);
Museo Nacional de Historia Nat-
ural, Santiago, Chile (M. Elgueta
1D
Museu de Zoologia, Universidade
de Sao Paulo, Sao Paulo, Brazil (U.
R. Martins);
Division of Scientific and Indus-
trial Research, Auckland, New
Zealand (G. Kuschel);
Texas A&M University, College
Station, Texas, USA (H. R. Burke);
National Museum of Natural His-
tory, Washington, D.C., USA (D.
R. Whitehead).
HAHC
MCZC
MNHN
MNNC
MZSP
NZAC
TAMU
USNM
Measurements of adult specimens were
made with an ocular micrometer in a dis-
secting microscope as follows: total length
from anterior margin of eye to elytral apex
in lateral view; width across elytra at widest
point; length of pronotum, dorsally, from
anterior to posterior margins; length of ros-
trum from anteroventral margin of eye to
apex, across arc, in lateral view; length of
distal portion of rostrum from antennal in-
sertion to apex in lateral view; width of frons
at narrowest point between eyes; width of
base of rostrum just distad of eyes in dorsal
view; and width of pro- and metafemora,
in anterior view, excluding the ventral teeth.
The range and, in parentheses, the mean
and sample size of each measurement are
given for each species.
Descriptions of larvae are based on full-
grown specimens. The terminology follows
Anderson (1947) and Thomas (1957). Where
differences in terminology exist between the
two systems, the Thomas terms are placed
in parentheses following those of Anderson.
89
Terminology of pupal characters follows
Burke (1968).
THE ANTHONOMUS ORNATUS GROUP
Adults of the seven species in the A. or-
natus group are about average in size for the
genus (specimens examined range from 1.88
to 4.00 mm in length), somewhat elongate
and slender in body form, without signifi-
cant elytral prominences, and with relative-
ly slender profemora armed with a mod-
erately large ventral tooth and a smaller,
distal tooth. They are distinguished by their
contrasting patterns of white, pallid to dark
ferruginous and fuscous scales that cover
corresponding areas of pallid or dark integ-
ument (Figs. 1-12). No other anthonomines
examined have exactly this same pattern.
Adults of the species also have an elongate,
flat, distal endophallic sclerite (Figs. 19-30).
They are further characterized as follows:
Head: vertex with narrow ferruginous
scales, frons with broader, more pallid scales;
eyes strongly convex, slightly to promi-
nently elevated, separated by distance 0.6-
0.9 x width of rostrum at base; antennal
funiculus with 7 articles. Rostrum: slender;
proximal portion with well-developed ca-
rinae and sulci; finely setose. Prothorax:
pronotum densely, coarsely punctate, with
narrow, attenuate, pallid to dark ferrugi-
nous scales and broader, more pallid scales
in narrow middorsal vitta; pleuron with
small anteromedian patch of pallid, broad
scales that is feebly developed in small spec-
imens. E/ytra: humeri not prominent; sides
subparallel. Pygidium: not impressed or sul-
cate. Legs: pro- and mesocoxae with dense,
broad, pallid scales that are weakly devel-
oped in small specimens; profemur slender,
ca. 1.1-1.2 x stouter than metafemur, with
small, conical, acute ventral tooth and
smaller distal tooth; metafemur with small
ventral tooth and minute distal tooth; pro-
tibial uncus slender, slightly curved. Geni-
talia: male tegmen with long parameres.
Distribution (Figs. 42-45). The species in
the 4. ornatus group are known mainly from
90 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. Habitus of adults (lines represent | mm): 1) A. signatipennis, male, Quinta Pittet, Magallanes,
Chile, lateral view; 2) the same, dorsal view; 3) A. ornatus, male, Rio El Ganso, Seno de Otway, Magallanes,
Chile, lateral view; 4) the same, dorsal view; 5) 4. kuscheli, female, Chepu, Chiloé, Chile, lateral view; 6) the
same, dorsal view.
the Valdivian and Magellanic forest regions
which, according to Kuschel (1960), extend
south of the 36th parallel beyond the south-
ernmost portion of the continent to Isla
Navarino at the 55th parallel. Blanchard
(1851: 387) stated that both A. signatipennis
and A. ornatus were found in “la provincia
de Coquimbo.” These records, along with
records of A. signatipennis from the Chilean
provinces of Valparaiso and Santiago (Figs.
42, 43), extend the limits of the group sig-
nificantly north of the Valdivian Forest into
more arid regions. A possible explanation
for this extension is found in the observa-
tion by Davis (1986) that in parts of the
Coquimban Desert Province, moisture from
coastal fog supports remnants of the Val-
divian forest. Davis listed several southern
VOLUME 91, NUMBER |
91
Figs. 7-12.
lateral view; 8) the same, dorsal view; 9) A. chilicola, male, 20 km. E Manzanar, Malleco, Chile, lateral view;
10) the same, dorsal view; 11) 4. berberidis, male, Frutillar, Llanquihue, Chile, lateral view; 12) the same, dorsal
view.
temperate plant species (not including Ber-
beris, the known hosts of the members of
the A. ornatus group), as occurring in these
moist refugia.
Plant associations. Six of the 7 species of
the 4. ornatus group are known to be as-
sociated with one or more of the following
Berberidaceae: Berberis buxifolia Lamarck;
Habitus of adults (lines represent | mm): 7) 4. araucanus, male, Laguna Malleco, Malleco, Chile,
B. chilensis Hort. ex C. Koch; B. darwini
Hooker; and B. ilicifolia Forster in Comm.
The two species for which the immatures
are known, A. ornatus and A. kuscheli, de-
velop in flower buds. The site of develop-
ment of the remaining species is not known.
Label data indicate that adults of A. sig-
natipennis and A. ornatus have been col-
92 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
18
17
Figs. 13-16. Pygidium, adult female, dorsal view (line represents 0.25 mm): 13) A. signatipennis, Quinta
Pittet, Magallanes, Chile; 14) 4. ornatus, Lago Frio, Aisén, Chile; 15) A. kuscheli, Panquipulli, Valdivia, Chile;
16) A. araucanus, Laguna Laja, Bio-Bio, Chile.
Figs. 17-18. Abdomen, adult female, ventral view (line represents 0.25 mm): 17) A. ornatus, Lago Frio,
Aisén, Chile; 18) 4. blanchardi, El Coigo, Curic6, Chile.
lected together, on the same date, at several velop in flower buds, but the developmental
localities. One of these records indicates that site of A. signatipennis is unknown. Adults
specimens of the two species were taken on of the two species have also been taken at
the same day on B. buxifolia. Larvae ofone the same locality on B. ilicifolia, but one
of the species, 4. ornatus, are known to de- day apart. Adult 4. blanchardi have been
= 29 30
Figs. 19-30. Aedeagus, adult male (line represents 0.5 mm): 19) 4. signatipennis, Estancia Fenton, Magal-
lanes, Chile, dorsal view; 20) the same, lateral view; 21) 4. ornatus, Trapatrapa, Concepcion, dorsal view; 22)
the same, lateral view; 23) A. ornatus, Lago Frio, Aisén, dorsal view; 24) the same, lateral view; 25) 4. blanchardi,
El Coigo, Curico, Chile, dorsal view; 26) the same, lateral view; 27) A. kuscheli, Pichinahuel, Malleco, Chile,
dorsal view; 28) 4. araucanus, 20 km. E Manzanar, Malleco, Chile, dorsal view; 29) 4. chilicola, 20 km. E
Manzanar, Malleco, Chile, dorsal view; 30) 4. berberidis, Frutillar, Llanquihue, Chile, dorsal view.
93
94 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 31-37.
Figs. 38-41.
Third instar larva of Anthonomus kuscheli. 31) lateral view; 32) head capsule, frontal view; 33)
labium and right maxilla, ventral view; 34) epipharynx; 35) clypeus and labrum; 36) abdominal segments 3 and
4, dorsal view; 37) mandible. Line accompanying Fig. 31 = 2 mm; Fig. 32 = 0.5 mm; Fig. 33 = 0.25 mm; other
figures greatly enlarged.
Pupa of Anthonomus ornatus. 38) head, rostrum and ventral view of prothorax; 39) prothorax,
dorsal view; 40) terminal abdominal segments, dorsal view; 41) 9th abdominal segment; lateral view. Line
accompanying Fig. 38 = | mm; Fig. 39 = 1 mm; Fig. 40 = 0.5 mm.
collected on B. buxifolia, but not at the same
locality as A. signatipennis and A. ornatus.
Adults of another pair of species, 4. kuscheli
and A. berberidis, have been collected on B.
darwini at the same time and locality.
Discussion. No observed characters of
adults of the members of the A. ornatus
group can be cited with confidence as evi-
dence of a sister group relationship with any
anthonomine group (Clark 1987a, b, c, 1988,
Clark and Burke 1985, 1986a, b, c, Clark
and Martins 1987). The larvae of two of the
species, A. ornatus and A. kuscheli, are
known. These larvae trace to A. pomorum
L. and A. nebulosus LeConte in a key to
larvae of members of the subfamily (Ahmad
and Burke 1972), but there is no other in-
dication that these species are very closely
related. Larvae of A. kuscheli are distinct in
possessing three rather than four epipharyn-
Fig. 42.
Map of southern South America showing the distribution of 4. signatipennis
Fig. 43. Map of southern South America showing the distribution of 4. ornatus
Fig. 44. Map of southern South America showing the distributions of 4. kuscheli (circles) and A. chilicola
(square).
Fig. 45. Map of southern South America showing the distributions of 4. blanchardi (circles), A. araucanus
(triangles) and A. berberidis (squares).
95
96 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
geal sensory pores. Larvae of Coccotorus
scutellaris (LeConte) are the only other an-
thonomines known to have only three epi-
pharyngeal sensory pores, but available in-
formation does not indicate that this species
is closely related to the 4. ornatus group.
The pupae of members of the 4. ornatus
group have a single process on the 9th ab-
dominal segment. This structure, however,
is also shared with several other, apparently
unrelated anthonomines, including Tachy-
pterellus quadrigibbus Say, Pseudanthono-
mus validus Dietz, Anthonomus albopilosus
Dietz, A. stupulosus Champion, A. griseis-
quamis Champion, A. unipustulatus Cham-
pion and 4. nubiloides Fall (Burke 1968).
The association of the species in the 4.
ornatus group with plants in the genus Ber-
beris is biogeographically significant. Guil-
lermo Kuschel (pers. comm.), speaking from
years of experience collecting weevils and
other insects in Chile and neighboring coun-
tries, noted that the 4. ornatus group mem-
bers were found exclusively on Berberis. He
also insisted that he “frequently and quite
thoroughly” checked Berberis plants all over
northern Chile, Peru and Bolivia, but found
no anthonomines on those plants there. The
+500 species of Berberis are distributed
throughout the north temperate zones and
southward, mainly at higher altitudes, in
tropical regions of Africa, Asia and South
America (Cronquist 1981: 130, Good 1964:
80-81). As noted by Cabrera and Willink
(1973: 98), species of Berberis are abundant
in the Valdivian and Magellanic forests of
southern Chile and adjacent portions of Ar-
gentina. Raven (1963: 155) listed Berberis
among several genera that used the moun-
tains to extend their distribution from the
northern hemisphere to the south. No other
anthonomines are known to have hosts in
the family Berberidaceae (Burke [1976] list-
ed plant families known to contain
anthonomine hosts).
It is fairly certain that the members of the
A. ornatus group are the only anthonomines
to be found in Chile. No others are listed
by Wibmer and O’Brien (1986) in their
checklist of South American Curculionidae.
Schenkling and Marshall (1934) did list two
other Chilean Curculionidae, Anthonomus
variabilis Philipi and A. australis Philipi, as
anthonomines, but Kuschel (1950: 17) de-
termined that these are actually members
of the genus Rhopalomerus Blanchard in the
subfamily Eugnominae.
Key TO ADULTS OF THE SPECIES OF
ANTHONOMUS IN THE A. ORNATUS GROUP
1. Elytra (Figs. 1, 2) with oblique, alternating light
and dark fasciae, without posterolateral mac-
ulae; elytral interstria 2 with discrete postero-
median pallid line bounded anteriorly and
posteriorly by short, dark portions (Figs. 1, 2)
Baer? signatipennis
1’. Elytra with large, posterolateral maculae (Figs.
3-6, 9, 10) (feebly developed in some speci-
mens, Figs. 7, 8, 11, 12); elytral interstria 2
without discrete posteromedian pallid seg-
ment bounded anteriorly and posteriorly by
Short;:dark portions’; sacs. scene oe ieeee 2
Sutural elytral interstriae with acute, overlap-
ping apical extensions; metatibia of male with
ventral margin strongly concave in distal 7,
with enlarged, excavated apical mucro; aedea-
gus asymmetrical (Figs. 29, 30)
‘. Sutural elytral interstriae without apical ex-
tensions; metatibia of male with ventral mar-
gin not or only slightly concave in distal *,
apical mucro small; aedeagus symmetrical
(Figs. 19-28)
3. Sternum 5 of male with large ‘posteromedian
prominence (Fig. 11); pallid annuli around
posterolateral elytral maculae weakly devel-
oped (Fig. 12) . a eee TESS eee berberidis
3’. Sternum 5 of male einen posteromedian
prominence; pallid annuli around postero-
lateral maculae distinct (Fig. 10) ...... chilicola
4. Elytra with posterolateral maculae completely
surrounded dorsally by pallid annuli (Figs. 5,
6); pygidium of female with subapical prom-
inences (Fig. 15); body slender, somewhat flat-
tened dorsally (Fig. 5) ................ kuscheli
4’. Elytra with posterolateral maculae incom-
pletely surrounded dorsally by pallid annuli
(Figs. 3, 4), or macula feebly developed (Figs.
7, 8); pygidium of female with subapical prom-
inences weakly developed (Fig. 14) or absent;
body less slender, less flattened (Figs. 3,7) .. 5
5. Elytral pattern distinct (Figs. 3, 4); sternum 5
of female with shallow emarginations (Fig. 18)
or deep incisions (Fig. 17) on each side of me-
th
i)
VOLUME 91, NUMBER |
dian prominence; aedeagus with apicolateral
prominences (Figs. 21, 23, 25); pygidium of
female without apicomedian prominence (Fig.
wn
oy
=
pss
=
=
uo}
S
5
=
oO
p
a
Leo}
a
oO
<
fa)
°
b> a
(3s
ie
oOo.
Se
Gus
ia}
Oo.
>
oS:
<4
prominent midbasal macula (Figs. 7, 8); ster-
num 5 of female without emarginations or in-
cisions; apex of aedeagus narrowed to apex,
without apicolateral projections (Fig. 28); py-
gidium of female with small apicomedian
prominence (Fig. 16) . . araucanus
6. Sternum 5 of female with deep posteromedian
incisions (Fig. 17); aedeagus (Figs. 21, 23) about
as broad at broadly rounded apex as at base,
with acutely pointed apicolateral prominences
low emarginations on each side of small, blunt,
posteromarginal prominence; aedeagus (Fig.
25) narrower at apex than at base, apex slightly
distended, with blunt, weakly developed api-
colateral prominences . ... blanchardi
Anthonomus signatipennis Blanchard
igs. 15.2, 13, 19; 20).42
Anthonomus signatipennis Blanchard 1851:
387. Holotype. Chile, “provincia de Co-
quimbo,” male, labelled with a green disc
and [67/ 7] [TYPE] [MUSEUM PARIS/
CHILI/ CL. GAY 1849] [Anthonomus/
signatipennis/ Type Blanchard/ H. Perrin
det. 19] (MNHN). Blackwelder 1947: 839.
Schenkling and Marshall 1934: 59. Wib-
mer and O’Brien 1986: 204.
Recognition (Figs. 1, 2). Adults of A. sig-
natipennis are distinguished by the follow-
ing combination of characters:
Elytra (Figs. 1, 2) without posterolateral
maculae; interstria 2 with posteromedian
pallid portion bounded anteriorly and
posteriorly with short dark portions; in-
terstriae 3, 5, 7 and 9 with long postero-
median dark portion contrasting with
short dark portions on interstriae 4, 6 and
8; aedeagus (Figs. 19, 20) with broad, blunt
apicomedian prominence.
They are easily distinguished from adults of
the other members of the A. ornatus group
by the distinct elytral pattern. They lack the
97
large posterolateral elytral maculae char-
acteristic of adults of the other members of
the group. Distinctive are the prominent
posthumeral patch of pallid scales, the
oblique discal fascia of pallid scales extend-
ing across interstriae 1-4, and the oblique,
alternating dark and pallid elytral fasciae.
They are most likely to be confused with
adult 4. ornatus but, in addition to the dif-
ferent elytral pattern (cf. Figs. 1—4), the fe-
male sternum 5 lacks posteromarginal
emarginations and the male aedeagus lacks
acute subapicolateral prominences (cf. Figs.
19S 23):
Adult male. Length: 3.28-3.64 mm (x =
3.40, n = 10). Width: 1.46-1.60 mm (x =
1.53,n = 10). Rostrum: length 1.30-1.51 x
(x = 1.40, n = 10) pronotal length; distal
portion finely punctate, length 18-30% (x =
23,n = 10) of total rostral length. Prothorax:
pronotum with pallid scales in broad sub-
apical fascia and in posterolateral vittae.
Elytra: interstriae 3 and 5 slightly elevated
at base and on disc; integument and scales
dark on humeri, on basal, elevated portions
of interstriae 3 and 5, on discal portions of
interstriae 1-3, in oblique, irregular, an-
teromedian and posteromedian fasciae, and
on apices of interstriae 4-6; pallid integu-
ment and scales predominant in small
oblique patches extending across interstriae
1-3, in large, rectangular patch behind hu-
meri, in narrow, oblique, anteromedian fas-
cia that extends posteriorly from interstria
5 to suture, in broader, oblique postero-
median fascia, and on declivity. Abdomen:
posteromedian portion of sternum 5 slightly
depressed behind transverse impression.
Legs: protibia with slight ventral marginal
prominence at proximal 3; metatibia nar-
row, ventral margin slightly, broadly con-
vex in middle *3; metatibial mucro short,
straight, acute, not excavated.
Adult female. Length: 3.40-3.72 mm (x
= 3.52, n = 10). Width: 1.56-1.66 mm (x
= 1.61,n = 10). Rostrum: length 1.25-1.47
x (X¥ = 1.36, n = 10) pronotal length; distal
portion subcylindrical, shallowly punctate,
98 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
glabrous, length 22-29% (* = 25, n = 10)
of total rostral length. Pygidium (Fig. 13):
apical margin rounded, without subapical
ridge. Abdomen: sternum 5 without median
emargination. Legs: metatibial mucro mi-
nute.
Distribution (Fig. 42). The 336 adult
specimens of A. signatipennis examined are
from the following localities. ARGEN-
TINA. Neuquén: San Martin de los Andes.
Rio Negro: Norquinc6é. Santa Cruz: Cana-
don Leon; Lago Argentina. CHILE. Aisén:
Chile Chico; Coyhaique; 10 km. N Puerto
Ibanez. Bio-Bio: Los Barros, Laguna Laja;
Volcan Copahue. Cautin: Cherquenco.
Chiloé: Castro; Chepu; Dalcahue. Concep-
cidn: Salto de Laja. Llanquihue: Lepihue, W
of Puerto Montt; Puerto Varas; Los Muer-
mos; Maullin; Puerto Montt. Magallanes:
Estancia Cameron; Estancia Fenton; La-
guna Amarga; Puerto Williams; Punta Are-
nas; Quinta Pittet; Rio Santa Maria. Mal-
leco: Lonquimay; Licura; Laguna Malleco;
19 km. E Manzanar; 20 km. E Manzanar;
Pichinahuel, Cordillera Nahuelbuta; 38 km.
SE Victoria. Nuble: Las Cabras, Cordillera
Chillan; Shangrila, 75 km. E Chillan. Osor-
no: Laguna la Copa, Parque Nacional de
Puyehue. Santiago: Melocoton; 4 km. W
Portillo; Quinta San Ramon. Talca: 5 km.
W Molino. Valparaiso: Algarrobo. The
specimens examined were collected during
the months of October through February
and in April.
Plant associations. Adults of A. signati-
pennis were collected on Berberis buxifolia
at Chepu, Chiloé, Quinta Pittet and Puerto
Williams, Magallanes; on B. chilensis at Sal-
to de Laja, Concepcion and Algarrobo, Val-
paraiso; on B. darwinii at Laguna la Copa,
Osorno, and on B. ilicifolia at Chile Chico,
Aisén, Chile.
Anthonomus ornatus Blanchard
Figs. 3, 4, 14, 17, 21-24, 38-41, 43
Anthonomus ornatus Blanchard 1851: 387.
Holotype. Chile, female, “provincia de
Coquimbo,” labelled with green disc and
[15/ 43] [illegible handwritten label] [An-
thonomus/ ornatus.] [TYPE] [MUSEUM
PARIS/ CHILE/ Gay 1843] (MNHN).
Kuschel 1950: 17 (= biplagiatus Fair-
maire, not Redtenbacher [redtenbacheri
Blackwelder]). Ashworth and Hoganson
1987. Schenkling and Marshall 1934: 58.
Wibmer and O’Brien 1986: 204 (= bipla-
giatus Fairmaire, not Redtenbacher [red-
tenbacheri Blackwelder)).
Anthonomus biplagiatus Fairmaire 1884:
503-504. Holotype. Chile: ‘“Punta-Are-
na,” female [223] [TYPE] [MUSEUM
PARIS/ SANTA-CRUZ/ LEBRUN
1883] [anthonomus/ biplagiatus/ Fairm]
(MNHN). Kuschel 1950: 17. Wibmer and
O’Brien 1986: 204.
Anthonomus redtenbacheri Blackwelder
1947: 839. Replacement name for A. bi-
plagiatus Fairmaire (1884) not Redten-
bacher (1867).
Recognition (Figs. 3, 4). Adults of A. or-
natus are distinguished by the following
combination of characters:
Elytra (Figs. 3, 4) with large posterolateral
maculae; pallid dorsal annuli around
maculae broken dorsomedially by dark
posteromedian portion on interstria 2;
sutural elytral interstriae without apical
extensions; sternum 5 of female (Fig. 17)
with deep apicomedian emarginations on
each side of slender median projection;
aedeagus (Figs. 21-24) symmetrical, with
rounded apex and acute lateral promi-
nences; pygidium of female (Fig. 14) with
weakly developed subapical promi-
nences.
They are distinguished from adults of A.
signatipennis by the characters discussed
under the latter. The elytral patterns are dis-
tinct in all specimens of A. ornatus exam-
ined but there is considerable variation in
the ratio of dark and pallid portions. In
some, the pallid portions are extensive and
clothed wtih predominantly white scales,
with dark fuscous portions correspondingly
VOLUME 91, NUMBER 1
limited primarily to the elevated basal and
discal portions of interstria 3 and to the
edges of the posterolateral maculae. In oth-
ers, the pallid portions are more extensive,
the pallid scales are mostly ferruginous, and
dark integument and fuscous scales are ex-
tensive on the basal portions of interstriae
2-5, on the median portion of the disc, on
the posterolateral maculae and on the de-
clivity. The specimen illustrated (Figs. 3, 4)
is somewhat intermediate between the ex-
tremes. There is also variation in the shape
of the aedeagus. At one extreme the sides,
seen in dorsal view (Fig. 21), are subparallel
behind the apicolateral projections and the
lateral plates are only slightly distended dor-
sally, as seen in lateral view (Fig. 22). At the
other extreme, the sides are broadly con-
stricted in the distal *4 in dorsal view (Fig.
23) and the lateral plates are correspond-
ingly distended dorsally in lateral view (Fig.
24). This variation 1s not geographical; both
extremes are exhibited in specimens from
Trapatrapa, Concepcion, which do not dif-
fer otherwise.
Adult male. Length: 2.56-3.32 mm (x =
2.90, n = 10). Width: 1.16-1.60 mm (x =
1.39, n = 10). Rostrum: length 1.40-1.70 x
(x = 1.60, n = 10) pronotal length; distal
portion finely punctate, length 23-32% (x =
28, n= 10) of total rostral length. Prothorax:
pronotum without distinct subapical fascia
and dorsolateral vittae of pallid scales. E/y-
tra: interstria 3 with slightly elevated basal
and discal portions; dark integument and
scales predominant on humeri, on basal el-
evated portions of interstriae 3 and 5 and
on discal portions of interstriae 1-3, form-
ing irregular edges of posterolateral mac-
ulae, and on apices of interstriae 4—6; pallid
integument and scales predominant on
posthumeral portions, in broad annuli
around edges of posterolateral maculae, in
median portion of that macula, and on de-
clivity. Abdomen: posteromedian portion of
sternum 5 slightly depressed behind trans-
verse impression. Legs: protibia with slight
midventral marginal prominence; metatib-
eye)
la narrow, straight, with slight ventral mar-
ginal prominence; metatibial mucro large,
curved, excavated.
Adult female. Length: 2.48-3.32 mm (x
= 2.98, n = 10). Width: 1.12-1.58 mm (x
= 1.41, n = 10). Rostrum: length 1.59-1.76
x (X = 1.68, n = 10) pronotal length; distal
portion slender, shining, glabrous, length 29-
37% (X = 35, n = 10) of total rostral length.
Legs: metatibial mucro minute.
Larva (2 specimens from Frutillar, Llan-
quihue, Chile, collected November 2, 1983,
by G. Kuschel, ex flower buds Berberis buxi-
folia). The third instar larva of A. ornatus
resembles that of 4. kuscheli in size and
other characters except as follows: it is more
strongly curved; the ninth abdominal seg-
ment is not as long; and there are four epi-
pharyngeal sensory pores instead of three as
in A. kuscheli. The significance of these ap-
parent differences will have to await the
availability of additional specimens.
Pupa (5 specimens from Frutillar, Llan-
quihue, Chile, collected November 2, 1983,
by G. Kuschel ex flower buds of Berberis
buxifolia). Body: length 3.8-4.2 mm (n =
5). Head (Fig. 38): frontal setae straight to
slightly curved; each borne on summit of
small, rounded tubercle; pair separated by
distance greater than length of a seta. Su-
praorbital setae absent. One pair of fine ba-
sirostral (interocular) setae; each about '2
length of frontal seta. One pair of minute
sessile distirostral setae. Pronotum (Fig. 39):
pronotal setae straight to feebly curved; se-
tae on anterior margin slightly longer and
stouter than posterior setae. Anteromedian
setae each borne subapically on anterior face
of conical, sharply pointed tubercle; tuber-
cles separated by distance equal to ca. width
of tubercle at base. Three pairs of antero-
lateral setae; each borne on summit of slight
to distinct tubercle; tubercle of anterolateral
3 often taller than others; setae on each side
of pronotum in straight to feebly curved
line. Posteromedian setae each borne at or
near apex of subconical to acutely pointed
tubercle; tubercles separated by distance
100
slightly greater than length of a seta. Three
pairs of posterolateral setae; arranged in
curved line on each side of pronotum; each
borne on summit of small, rounded tuber-
cle. Mesonotum: three pairs of straight
mesonotal setae; each borne on summit of
rounded tubercle or seta, innermost oc-
casionally subapical on small, acutely point-
ed tubercle. Metanotum: three pairs of
straight to curved metanotal setae; more
widely spaced than mesonotals; each borne
on summit of rounded tubercle; metanotal
2 usually located closer to 3 than to 1. Ab-
domen (Fig. 40): three pairs of discotergal
setae and occasionally with additional se-
taless tubercles on some terminal segments;
setae each borne on summit of low tubercle
on first two abdominal terga, remaining ter-
ga with seta located at base of sharply point-
ed tubercles that become progressively larg-
er posteriorly. Laterotergal setae 1 and 2
present on each of first 8 terga; seta 1 mi-
nute, borne at base of small, sharply pointed
tubercle on all terga; seta 2 borne subapi-
cally on sharply pointed tubercle, slightly
curved, longer than discotergal setae. An-
teronotal setae absent. Spiracles well de-
veloped on first 5 abdominal segments, fee-
bly developed on segment 6, absent on
others. Laterosternal and sub-laterosternal
setae absent. Segment 9 bearing a single
sharply pointed process which in side view
(Fig. 41) is turned slightly upward apically;
minute seta borne on each side of segment
ca. midway between apex and base of seg-
ment.
Distribution (Fig. 43). The 235 adult
specimens of A. ornatus examined are from
the following localities: ARGENTINA.
Neuquén: San Martin de los Andes. Rio Ne-
gro: Colonia Catedral; Norquinco; Rio Foy-
el. Santa Cruz: Canadon Leon. Tierra del
Fuego: Bahia San Sebastian, Punta de Are-
nas. CHILE. Aisén: Chile Chico; Coy-
haique; Lago Escondido; Lago Frio; 10 km.
N Puerto Ibanez; Quellon. Bio-Bio: Los
Barros, Laguna Laja. Chiloé: Chepu. Con-
cepcion: Trapatrapa. Linares: Parral. Llan-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
quihue: Frutillar; Rio Penon, Maullin. Ma-
gallanes: Cerro Castillo; Dos Lagunas;
Estancia Cameron; Estancia Canelo; Estan-
cia Fenton; Isla Navarino; Laguna Azul and
Ultima Esperanza, Parque Nacional Torres
del Paine; Laguna Los Robles; Quinta Pit-
tet; Puerto Williams; Punta Arenas; Rusfin;
Rio Tres Pasos; Rio el Ganso, Seno de Ot-
way. Malleco: Icalma; La Fusta; 20 km. E
Manzanar; Marimenuco; Cordillera Na-
huelbuta: Villa Portales. Nuble: Cordillera
Chillan, Las Cabras, Las Trancas, 70 km. E
Chillan; El Marchant; Recinto; 4.5 km. SE
Recinto. Osorno: Antillanca; 2 km. SW Vn.
Casablanca, Parque Nacional de Puyehue.
The specimens examined were collected
during the months of October through Feb-
ruary and in April and August. Only one of
the specimens on which the reference to A.
ornatus by Ashworth and Hoganson (1987:
887) is based is that species. The others are
A. signatipennis, A. berberidis and A. kus-
cheli.
Plant associations. Specimens of A. or-
natus were collected on Berberis buxifolia
at Chepu, Chiloé, Frutillar, Llanquihue, and
Quinta Pittet and Puerto Williams, Maga-
llanes; on Berberis ilicifolia at Chile Chico,
Aisén, and Puerto Williams, Magallanes;
and on Berberis sp. at Trapatrapa, Concep-
cion, and Laguna Azul and Ultima Espe-
ranza, Magallanes, Chile.
Anthonomus blanchardi Clark,
NEw SPECIES
Figs. 18, 25, 26, 45
Type specimens. Holotype: Chile, male
[CHILE/ El Coigo/ Cord. Curico/ Oct-Nov
1959] [G. Kuschel/ Collection] [Entomol-
ogy/ Division/ D.S.I.R./ New Zealand]
(NZAC). Paratypes: Chile, 1 male, 2 females
[CHILE/ El Coigo/ Cord. Curico/ Nov—Dec
1959] [G. Kuschel/ Collection] [Entomol-
ogy/ Division/ D.S.I.R./ New Zealand]; |
female [Santiago/ Farellones/ 2200 m/ 10
Dec 1950] [G. Kuschel/ Berberis/ buxifolia]
[Coll./ Kuschel] [Entomology/ Division/
D.S.I.R./ New Zealand]; 1 female [Chile:
VOLUME 91, NUMBER 1
8600'/ Farellones/ P. Santiago/ XII-25-
1968] [under dung L &/ C. W. O’Brien]; |
female [Chile Santiago/ Penalolén/ 25-I-
1975/ Coll: M. Beeche]; 1 female [Altos de
Vilches/ Talca A280 mts./ 25—26-XI-1970/
J. Solvericens]. Total paratypes, 7 (CWOB,
MNNC, NZAC).
Recognition. Adults of A. blanchardi are
distinguished by the following combination
of characters:
Elytra (cf. Figs. 3, 4) with large postero-
lateral maculae; pallid dorsal annuli
around macula broken medially by dark
posteromedian portion on interstria 2;
sutural elytral interstriae without apical
extensions; sternum 5 of female (Fig. 18)
with shallow apicomedian emarginations
on each side of short median prominence;
aedeagus (Fig. 25, 26) symmetrical, nar-
rowed in distal '2, with rounded apex and
short, acute apicolateral prominences;
pygidium of female (cf. Fig. 14) with
weakly developed subapical promi-
nences.
They are distinguished from adults of the
closely allied 4. ornatus by characters of the
male genitalia and of sternum 5 of the fe-
male. The aedeagus of A. blanchardi (Figs.
25, 26) is constricted and narrowed in the
distal 7 to the apex. The apex itself is slight-
ly distended, rather than being rounded as
in A. ornatus (cf. Figs. 21, 23, 25). It is also
narrower than the apex in that species and
has less well-developed apicolateral prom-
inences. The median posteromarginal
prominence on sternum 5 of the female is
bounded on each side by shallow emargi-
nations rather than being deeply incised as
in A. ornatus (cf. Figs. 17, 18).
Adult male. Length: 2.48-2.68 mm (xX =
2.58, n = 2). Width: 1.30-1.34 mm (Xx =
1.32, n = 2). Rostrum: length 1.43-1.55 x
(x = 1.49, n = 2) pronotal length; distal
portion finely punctate, length 32-35% (x =
34, n = 2) of total rostral length. Prothorax:
pronotum without distinct subapical fascia
and dorsolateral vittae of pallid scales. E/y-
101
tra: interstria 3 with slightly elevated basal
and discal portions; dark integument and
scales predominant on humeri, on basal el-
evated portions of interstriae 3 and 5 and
on discal portions of interstriae 1-3, form-
ing irregular edges of posterolateral mac-
ulae, and on apices of interstriae 4—6; pallid
integument and scales predominant on
posthumeral portions, in broad annul
around edges of posterolateral maculae, in
median portion of macula, and on declivity.
Abdomen: posteromedian portion of ster-
num 5 slightly depressed behind transverse
impression. Legs: protibia with slight mid-
ventral marginal prominence; metatibia
narrow, straight, with slight ventral margin-
al prominence; metatibial mucro large,
curved, excavated.
Adult female. Length: 2.48-3.16 mm (x
= 2.81, n = 6). Width: 1.10-1.50 mm (x =
1.34, n = 6). Rostrum: length 1.54-1.69 x
(X = 1.62, n = 6) pronotal length; distal
portion slender, shining, glabrous, length 30-
40% (X = 35, n = 6) of total rostral length.
Legs: metatibial mucro minute.
Distribution (Fig. 45). Anthonomus blan-
chardi is known from the type series from
the following localities. CHILE. Curico: El
Coigo, Cordillera Curico. Santiago: Farel-
lones; Penalolén. Ta/ca: Altos de Vilches.
Plant associations. The paratype of 4.
blanchardi from Farellones, Santiago, Chile,
was collected on Berberis buxifolia.
Etymology. This species is named for
Charles Emile Blanchard (1819-1900) in
honor of his contribution to the study of the
A. ornatus group.
Anthonomus kuscheli Clark,
NEw SPECIES
Figs. 5, 6, 15, 27, 31-37, 44
Type specimens. Holotype: Chile, male
(CHILE/ Pichinahuel/ Cord. Nahuelbuta/
1-31 Jan 1959] [G. Kuschel/ Collection]
[Entomology/ Division/ D.S.I.R./ New
Zealand] (NZAC). Paratypes: Argentina, |
female [Argentina/ Rio Negro/ L. Nahuel-
huapi/ Llao Llao] [24 Nov 1950/ W. Witt-
102 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mer] [Coll. Kuschel] [Entomology/ Divi-
sion/ D.S.I.R./ New Zealand]; 1 male
{Argentina/ R. Negro/ L. Nahuelhuapi] [1
Mar 1949/ W. Wittmer] [Coll. Kuschel]
[Entomology/ Division/ D.S.I.R./ New
Zealand]. Chile, 1 male [CHILE] [Ento-
mology/ Division/ D.S.I.R./ New Zealand];
3 males [Aysen/ Rio Correntoso/ 22-I-1971/
F. Silva G.] [Ex. Berberis/ buxifolia]; 1 male,
2 females [CHILE: P./ Aisen, 7 km./ W.
Coyhaique/ I-21-1968] [at night L. &/ C.
W. O’Brien]; 1 female [CHILE: P./ Aisen,
15 km./ W. Coyhaique/ I-21-1968] [Collec-
tors: L &/ C. W. O’Brien]; | male [Bio-Bio/
Pemehue/ R. Renaico] [12 Jan 1946/ G.
Kuschel] [Coll./ Kuschel] [Entomology/ Di-
vision/ D.S.I.R./ New Zealand]; 1 male
[male symbol] [Cautin/ Vn. Llaima/ 17 Sep
1951/ M. Codoceo] [Coll./ Kuschel] [Ento-
mology/ Division/ D.S.I.R./ New Zealand];
1 male, | female |CHILE/ Chiloé/ Chepu/
22.10.58] [s/ Berberis/ darwini] [Coll./ Kus-
chel] [Entomology/ Division/ D.S.I.R./ New
Zealand]; | male, | female [CHILE/ Chiloé/
Chepu/ 23.9.54] [Coll./ Kuschel] [Ento-
mology/ Division/ D.S.I.R./ New Zealand];
6 males, 7 females [CHILE, Llanquihue/
Frutillar/ 31 October 1983/ G. Kuschel]
[Berberis/ darwini/ em. 9 Nov 1983]; 1 fe-
male [CHILE/ Frutillar/ 27 Feb 1950/ G.
Kuschel] [G. Kuschel/ Collection] [Ento-
mology/ Division/ D.S.1.R./ New Zealand];
1 male [CHILE/ Pichinahuel/ Cord. Na-
huelbuta/ 1-31 Jan 1959] [G. Kuschel/ Col-
lection] [Entomology/ Division/ D.S.I.R./
New Zealand]; 2 males [CHILI/ Pichina-
huel 1200 m/ 14-18 Feb 1956] [G. Kuschel/
Collection] [Entomology/ Division/ D.S.I.R./
New Zealand]; 12 males, 17 females
(CHILE: 19 km./ E. Manzanar/ P. Malleco/
XI-3-1967] [Collectors: L &/ C. W. O’Brien];
1 female [Chile: 20 km./ SE Victoria/ P.
Malleco/ XI-3-1967] [Collectors: L &/ C.
W. O’Brien] [compared/ with type] [An-
thonomus/ ornatus/ Blanch/ det COB 1971];
2 males, 2 females [Chile: 20 km./ SE. Vic-
toria/ P. Malleco/ XI-3-1967] [Collectors:
L&/C.W. O’Brien]; | male, | female [Chile:
38 km./ SE. Victoria/ P. Malleco/ XI-3-
1967] [Collectors: L &/ C. W. O’Brien]; 1
female [CHILE: Laguna de Copa/ P. N. de
Puyehue, Osorno Pro./ Site 20, El. 520 m.
7-XII-77/ Valdivian Rain Forest/ Ash-
worth, Hoganson, Mooers] [on Berberis
darwinii] [Anthonomus/ ornatus/ Blan-
chard]; 14 males, 8 females [CHILE: La-
guna Espejo/ P. N. de Puyehue, Osorno Pro./
Site 17, El. 520 m. 21-XII-77/ Valdivian
Rain Forest/ Ashworth, Hoganson, Mooers]
{on Berberis darwinii] [Anthonomus/ or-
natus/ Blanchard]; | male, 2 females [CHI-
LI/ Panquipulli/ Valdivia/ 12 Jan 1944] [G.
Kuschel/ Collection] [Entomology/ Divi-
sion / D.S.I.R./ New Zealand]. Total para-
types, 95; (CACA, CWOB, MNNC, NZAC,
TAMU).
Recognition (Figs. 5, 6). Adults of 4. kus-
cheli are distinguished by the following
combination of characters:
Elytra (Figs. 5, 6) with posterolateral mac-
ulae; pallid dorsal annuli around maculae
uninterrupted by dark portion on inter-
stria 2; sutural elytral interstriae without
apical extensions; aedeagus (Fig. 27) sym-
metrical, narrowed apically, with slight
subapicolateral prominences; pygidium
of female (Fig. 15) with subapical prom-
inences; sternum 5 of female with small
apicomedian prominence; endophallus
(Fig. 27) with two dentate median scler-
ites in addition to distal sclerite.
They resemble A. ornatus but are more
elongate in body form and have the discal
area of the elytra more flattened so that the
basal and discal elevations of interstria 3
are not so distinct. In addition, the pallid
annuli around the dorsal portions of the
posterolateral elytral maculae are complete
in A. kuscheli (Figs. 5, 6), not interrupted
by a dark portion of interstria 2 as in 4.
ornatus (Figs. 3, 4). The anterior margin of
the macula is also less irregular in A. kus-
cheli. Additional differences are found in the
male genitalia and in the female abdomen
and pygidium. The aedeagus of 4. kuscheli
VOLUME 91, NUMBER |
is strongly narrowed toward the apex and
the apicolateral prominences are weakly de-
veloped (Fig. 27). The deep posteromedian
incisions of sternum 5 of the female A. or-
natus are replaced in A. kuscheli by a small
apicomedian prominence. The female py-
gidium has large subapical prominences in
A kuscheli (Fig. 15), whereas the promi-
nences are weakly developed in 4. ornatus
(Fig. 14).
Adult male. Length: 2.72-3.00 mm (
2.86, n = 10). Width: 1.22-1.
1.26, n = 10). Rostrum: length 1.31-1.59 x
(x = 1.46, n = 10) pronotal length; length
of distal portion 24-33% (X = 28, n = 10)
of total rostral length. Prothorax: pronotum
without distinct subapical fascia and dor-
solateral vittae of pallid scales. E/ytra: in-
terstria 3 with slightly elevated basal por-
tion, discal elevation feebly developed; dark
integument and scales predominant on bas-
al and discal elevations of interstria 3 and
around edges of large, posteromedian mac-
ulae; pallid integument and scales predom-
inant on sides behind humeri and in com-
plete annuli around dorsal portions of
posterolateral maculae. Abdomen: postero-
median portion of sternum 5 slightly de-
pressed behind transverse impression. Legs:
protibia with slight ventral marginal prom-
inence at proximal '3; metatibia narrow,
straight, with slight ventral marginal prom-
inence at proximal 4; metatibial mucro
large, curved, excavated.
Adult female. Length: 2.60-3.12 mm (x
= 2.88, n = 10). Width: 1.12-1.40 mm (x
= 1.26,n = 10). Rostrum: length 1.38-1.61
x (¥ = 1.49, n = 10) pronotal length; distal
portion subcylindrical, shallowly punctate,
glabrous, length 29-38% (x = 33, n = 10)
total rostral length. Abdomen: sternum 5
with small apicomedian prominence. Legs:
metatibial mucro short, curved.
Larva (8 specimens from Frutillar, Llan-
quihue, Chile, collected October 31, 1983,
by G. Kuschel, ex flower buds of Berberis
darwint). Body (Fig. 31): curved; length 3.1—
5.4 mm (n = 8). Asperities rounded to sub-
103
conical, fairly evenly distributed over body.
Color dingy white, except for brownish pig-
mented pronotal area. Head (Fig. 32): dark
brown; width of head capsule 0.51-0.54 mm
(n = 8). Basal article of antenna bearing
elongate-oval accessory appendage and three
short, stout setae, one of which is longer
than others. Endocarina (endocarinal line)
distinct, slightly longer than '2 length of
frons. Epicranial suture (coronal suture) less
than '2 length of head capsule. Four pairs
of frontal setae; setae | and 3 short, seta 4
ca. 2x length of 3, seta 5 distinctly longer
than 4; seta 3 located dorsolaterad of 4. Two
pairs of frontal sensilla, one pair in front of
frontal setae | and other pair located dor-
solaterad of setae 3. Five pairs of dorsal
epicranial setae; setae 1, 4 and 5 slender,
long, ca. equal in length; setae 2 and 3 nearly
equal in length, distinctly shorter than other
dorsal epicranial setae; seta 3 remote from
frontal suture; seta 4 closer to frontal suture
than seta 1. Four pairs of minute, peglike
posterior epicranial setae that tend to be
arranged in a slightly curved line; the lower
seta of series located directly above or
slightly mesad of dorsal epicranial seta 2.
Three pairs of posterior epicranial sensilla;
one pair near vertex; one pair dorsad and
mesad of dorsal epicranial seta 1; one pair
about halfway between dorsal epicranial se-
tae 4 and 5. Lateral epicranial seta | short,
about 4 length of seta 2. Ventral epicranial
setae 1 and 2 ca. equal in length. Clypeus
(Fig. 35) with an oblique narrow, nonpig-
mented area on each side. Clypeal setae
slender; seta | slightly longer and some-
times located closer to anterior margin of
frons than 2. One pair of clypeal sensilla
that are located closer to seta | than to 2.
Three pairs of labral setae: setae | and 2 ca.
same length; seta 3 shorter. Epipharynx (Fig.
34) (epipharyngeal lining) bearing two pairs
of anteromedian setae of ca. same length.
Three pairs of stout anterolateral setae pres-
ent. Labral rods (tormae) stout, converging
posteriorly. Three epipharyngeal sensory
pores arranged in single cluster. Two pairs
104
of stout median epipharyngeal spines pres-
ent between labral rods (tormae). Mandible
(Fig. 37) with two well-defined teeth; oc-
casionally outermost tooth considerably re-
duced; two slender setae ca. same length;
one sensillum. Maxillary palps (Fig. 33) with
apical article shghtly longer than basal ar-
ticle, bearing several minute papillae at apex
and sensillum; basal article with short seta
and two sensilla. Stipial setae 1, 3 and 4
long, seta | slightly longer than other two;
seta 2 much shorter than others. Mala (la-
cinial lobe) bearing | 1 stout setae, 6 ventral
(ventral lacinial) and 5 dorsal (dorsal lacin-
ial). Labial palpus consists of one segment
that bears several small papillae at apex and
sensillum. Premental sclerite with long pos-
terior process. One pair of long premental
(prelabial) setae present. Three pairs of glos-
sal setae of ca. equal size. Postmental setae
(postlabial) 1 and 3 each shorter than 2.
Thorax (Fig. 31) (setae described on one
side of body only): pronotum bearing 8 long
setae and two short ones. Five postdorsal
setae present; setae 2 and 5 shorter than
others. Two spiracular (alar) setae present;
one several times longer than other. Epi-
pleural (dorsopleural) lobe bearing one long
seta. Two long pleural (ventropleural) setae
on prothorax; one seta on each of meso- and
metathorax. Three long pedal (laterosternal)
setae. Sternal (mediosternal) seta minute.
Abdomen (Fig. 31): posterior margins of
most or all of first seven abdominal seg-
ments emarginate dorsally (Fig. 36). Ab-
dominal segments |—7 bearing three distinct
dorsal folds. Prodorsum (fold I) of segments
1-8 with short setae. Five postdorsal (setae
of abdominal segment, fold III) setae pres-
ent; setae 1, 3 and 5 longer than 2 and 4.
Two spiracular (alar) setae; seta 2 ca. 3 x
longer than |. Epipleurum (dorsopleurum)
bearing one long and one short seta. One
short pleural (ventropleural) seta and pedal
setae ca. same length. Two minute pedal
setae present. Anus subterminal; surround-
ed by four lobes, each bearing minute seta.
Seven pairs of minute setae borne on seg-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ment 9. Spiracles bicameral; air tubes with
6 annul.
Pupa (5 specimens from Frutillar, Llan-
quihue, collected October 31, 1983, by G.
Kuschel, ex flower buds of Berberis dar-
wini). The pupa of A. kuscheli is similar in
form and size to A. ornatus except the pos-
teromedian pronotal tubercles tend to be
larger and the setae are borne apically rather
than subapically as usual with the latter
species. The best distinguishing character
appears to be the discotergal setae on terga
6, 7 and 8 that are borne on the summits
of rounded tubercles rather than subapically
on acutely pointed tubercles as in 4. orna-
tus.
Distribution (Fig. 44). The type series of
A. kuscheli consists of adult specimens from
the following localities. ARGENTINA. Rio
Negro: Llao Llao, Lago Nahuel Huapi.
CHILE. Aisén: 7 km. W Coyhaique, Rio
Correntoso. Bio-Bio: Pemehue. Cautin:
Volcan Llaima. Chiloé: Chepu. Llanquihue:
Frutillar. Malleco: 19 km. E Manzanar,
Cordillera Nahuelbuta, Pichinahuel, 20 and
38 km. SE Victoria. Valdivia: Panquipulli.
Plant associations. Specimens of A. kus-
cheli were collected on Berberis darwini at
Chepu, Chiloe, and Frutillar, Llanquihue;
and on B. buxifolia at Rio Correntoso, Aisén,
Chile. Larvae and pupae were taken from
flower buds of B. darwini.
Etymology. This species is named in hon-
or of Guillermo Kuschel to commemorate
his contributions to the understanding of
the weevil fauna of Chile and as a token of
appreciation for help extended in the prep-
aration of this paper.
Anthonomus araucanus Clark,
New SPECIES
Figs. 7, 8, 16, 28, 45
Type specimens. Holotype: Chile, male
{(CHILE/ Laguna Laja/ Los Barros 1500 m/
14 Jan 1948] [on/ Berberis] [G. Kuschel/
Collection] [Entomology/ Division/ D.S.I.R./
New Zealand] (NZAC). Paratypes: Chile, 2
females [CHILE/ Laguna Laja/ Los Barros
VOLUME 91, NUMBER 1
1500 m/ 14 Jan 1948] [on/ Berberis] [G.
Kuschel/ Collection] [Entomology/ Divi-
sion/ D.S.I.R./ New Zealand]; 2 males [Bio-
Bio/ Cord. Pemehue/ 1500 m] [16 Jan 1944/
G. Kuschel] [Coll./ Kuschel] [Entomology/
Division/ D.S.I.R./ New Zealand]; | female
[Bio-Bio/ La. Laja/ Los Barros/ 1500 m] [13
Jan 1945/ G. Kuschel] [Coll./ Kuschel]
[Entomology/ Division/ D.S.I.R./ New
Zealand]; | male, 4 females [Chile: 19 km/
E. Manzanar/ P. Malleco/ XI-3-1967] [Col-
lectors L &/ C. W. O’Brien]; 2 males
[CHILE, Malleco Prov./ 20 km E Manza-
nar/ 1100 m 19-21.xii./ 1976 H. F. Howden]
[beating]; 4 males [Malleco/ L. Malleco/
1100 m] [22 Jan 1946/ G. Kuschel] [Coll./
Kuschel] [Entomology/ Division/ D.S.1.R./
New Zealand]; 1 female [Chile. 12 km./ W.
Curicautin (sic)/ P. Malleco/ XI-3-1967]
[Collectors L &/ C. W. O’Brien]; 3 males,
3 females [Chile: 20 km./ SE. Victoria/ P.
Malleco/ XI-3-1967]. Total paratypes, 23;
(CWOB, HAHC, NZAC, TAMU).
Recognition. Adults of A. araucanus are
distinguished by the following combination
of characters:
Elytra (Figs. 7, 8) with prominent mid-
basal macula; sutural elytral interstriae
without apical extensions; posterolateral
elytral maculae weakly developed (Figs.
7, 8), pallid annuli around maculae bro-
ken by dark portion on posteromedian
portion of interstria 2; rostrum of female
short, relatively stout; pygidium of female
(Fig. 16) with small apicomedian prom-
inence; aedeagus (Fig. 28) symmetrical,
narrowed to bluntly rounded apex; en-
dophallus (Fig. 28) with two dentate me-
dian sclerites in addition to distal sclerite.
They are similar in appearance to adults of
A. berberidis. They are relatively small. In
addition, the elytral patterns, with the ex-
ception of the prominent, dark midbasal
macula, are feebly developed in both species
(cf. Figs. 7, 8, 11, 12). The two are easily
distinguished; 4. araucanus lacks the apical
extensions of the sutural elytral interstriae
105
and the large prominence on sternum 5 of
the male. In addition, adults of A. araucanus
have the aedeagus symmetrical rather than
asymmetrical as in adult 4. berberidis (cf.
Figs. 28, 30).
Adult male. Length: 1.88-2.52 mm (x =
2.36, n = 10). Width: 0.84—-1.22 mm (x =
1.12,n = 10). Rostrum: length 1.33-1.64 x
(x = 1.44, n = 10) pronotal length; length
of distal portion 23-32% (X = 27, n = 10)
of total rostral length. Prothorax: pronotum
without subapical fascia and dorsolateral
vittae of pallid scales, middorsal vitta in-
distinct. E/ytra: interstria 3 with slight basal
and discal elevations; dark integument and
scales predominant in large, midbasal mac-
ula and forming edges of weakly developed
posterolateral maculae. 4bdomen: sternum
5 with slight apicomedian prominence. Legs:
protibia with slight ventral marginal prom-
inence at proximal '4; metatibia narrow,
ventral margin slightly curved in distal 74,
with slight prominence at proximal '4; me-
tatibial mucro large, curved, excavated.
Adult female. Length: 2.00-2.44 mm (x
= 2.30, n = 10). Width: 1.02-1.22 mm (x
= 1.08, n = 10). Rostrum: length 1.00-1.52
x (X = 1.37, n = 10) pronotal length; distal
portion subcylindrical, shallowly punctate,
glabrous, length 25-34% (x = 30, n = 10)
total rostral length. 4bdomen: sternum 5
with posterior margin straight. Legs: meta-
tibia straight, narrow at apex; metatibial
mucro short, straight.
Distribution (Fig. 45). The type series of
A, araucanus consists of adult specimens
from the following localities. CHILE. Bio-
Bio: Laguna Laja, Cordillera Pemehue.
Malleco: 12 km. W Curacautin; 20 km. E
Manzanar; Laguna Malleco; 20 km. SE Vic-
toria.
Plant associations. The holotype and two
paratypes were collected on Berberis sp. at
Los Barros and Laguna Laja, Bio-Bio, Chile.
Etymology. The name of this weevil is
derived from that of the Araucanians and
emphasizes the fact that it occurs in the heart
of their country and that of the Araucaria.
106
Anthonomus chilicola Clark,
NEw SPECIES
Figs. 9, 10, 29, 44
Type specimens. Holotype: Chile, male
[CHILE, Malleco Prov./ 20 km E Manza-
nar/ 1100 m 19-21.x11./ 1976 H. F. How-
den] [beating] (HAHC). Paratypes: Chile, 3
males [CHILE, Malleco Prov./ 20 km E
Manzanar/ 1100 m 19-21.xi1./ 1976 H. F.
Howden] [beating] (HAHC).
Recognition (Figs. 9, 10). Adults of A.
chilicola are distinguished by the following
combination of characters:
Sutural elytral interstriae with acute,
overlapping apical extensions; with pos-
terolateral elytral maculae (Figs. 9, 10);
pallid annuli around dorsal portions of
maculae extending across part of short
posteromedian dark portion on interstria
2; metatibia of male with ventral margin
strongly concave in distal 7, apical mucro
enlarged, excavated; aedeagus (Fig. 29)
asymmetrical, constricted medially, with
slight subapicolateral prominences.
Like adults of A. kuscheli, adult A. chilicola
are more slender than those of A. ornatus
(cf. Figs. 4, 10). The elytral pattern in A.
chilicola is essentially the same as in A. or-
natus, but the pallid annuli around the pos-
terolateral maculae are partially interrupted
by enroachment of dark integument and
scales on interstria 2 (Fig. 10). Adult 4.
chilicola are further distinguished from
adults of both of these species, however, by
the apical extensions of the sutural elytral
interstriae, by the male metatibia which is
strongly, broadly concave on the ventral
margin and has a much larger, more strong-
ly excavated metatibial mucro, and by the
asymmetrical aedeagus. These are all char-
acters adult 4. chilicola share with adult A.
berberidis, which are distinguished by the
less distinct elytral pattern (cf. Figs. 9-12),
the large posterolateral prominence on ster-
num 5 of the male, and by differences in the
aedeagus (cf. Figs. 29, 30).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Adult male. Length; 3.40-4.00 mm (x =
3.75, n = 4). Width: 1.40-1.64 mm (x =
1.57, n = 4). Rostrum: length 1.47-1.67 x
(x = 1.58, n = 4) pronotal length; length of
distal portion 21-31% (X = 25, n = 4) of
total rostral length. Prothorax: pronotum
without distinct subapical fascia and dor-
solateral vittae of pallid scales. E/ytra: in-
terstria 3 with slightly elevated basal and
discal portions; dark integument and scales
predominant on humeri and on basal and
discal elevations of interstria 3, less dense
on basal portions of interstriae 2, 4 and 5
and on discal portion of 2, forming broad,
irregular edges of posteromedian maculae;
pallid integument and scales predominant
behind humeri and in oblique fascia ex-
tending posteriorly to posteromedian por-
tion of interstria 2 then curving posteriorly
to apices of interstriae 4-6. Abdomen: pos-
teromedian portion of sternum 5 slightly
depressed. Legs: procoxae with dense, broad,
pallid scales; protibia with slight ventral
marginal prominence at proximal '3; meta-
tibia narrow, slightly curved, with slight
ventral marginal prominence at proximal
Ys; metatibial mucro large, curved, exca-
vated.
Adult female. Unknown.
Distribution (Fig. 44). The type series of
A. chilicola consists of adult specimens from
the following locality. CHILE. Malleco: 20
km. E Manzanar.
Plant associations. Unknown.
Etymology. The name of this species is
derived from that of its native country, em-
phasizing the unique austral distribution of
the group to which it belongs.
Anthonomus berberidis Clark,
NEw SPECIES
Figs.) lil 125 30745
Type specimens. Holotype: Chile, male
[CHILE/ Frutillar/ 14 Dec 1943/ G. Kus-
chel] [G. Kuschel/ Collection] [Entomolo-
gy/ Division/ D.S.I.R./ New Zealand]
(NZAC). Paratypes: ARGENTINA, | male,
1 female [ARGENTINA/ El Bolson, Rio/
VOLUME 91, NUMBER 1
Negro/ [X-20-1963/ A. Kovacs]. Chile, |
male [Chili] [Deyr.]; 2 males, 4 females
[CHILE: 19 km. S. of Ancud,/ Is. Chiloe/
Site C2. El. 120 m. 14-II-79/ Valdivian Rain
Forest/ Ashworth, Hoganson, Gordon] [on
Berberis darwin] [Anthonomus/ ornatus/
Blanchard]; 5 males, 6 females [CHILE/
Chiloé/ Chepu/ 22.10.58] [s/ Berberis/ dar-
wini] [Coll./ Kuschel] [Entomology/ Divi-
sion/ D.S.I.R./ New Zealand]; 3 males, 2
females [Chile: 9 km./ E. Chepu/ Chiloe Is/
II-4-1968] [at night L. &/ C. W. O’Brien];
2 females [Chile: 13/ km. S. Castro/ Is Chi-
loe/ II-5-1968] [Collectors: L &/ C. W.
O’Brien]; 1 female [Dalcahue/ Chiloe,
CHILE]; | female [CHILE/ Dalcahue/ Chi-
loe/ 10-20 Nov 1957] [G. Kuschel/ Collec-
tion] [Entomology/ Division/ D.S.I.R./ New
Zealand]; 1 male [Quellon-Chi/ loe I-1955/
J.S. Vargas]; | female [CHILE: Chiloé/ Dal-
cahue/ 17-31.1.1962/ Luis Pena] [H. & A.
Howden/ Collection]; | male, | female [Pto
Montt/ 15.1.71]; 1 female [Chile: 12 km.
N./ Puerto Montt/ P. Llanquihue/ II-8-
1968] [at night L. &/ C. W. O’Brien]; 3 males,
5 females [CHILE/ Llanquihue/ Frutillar/
13.9.54] [Coll./ Kuschel] [Entomology/ Di-
vision/ D.S.I.R./ New Zealand]; | male, 3
females [CHILE, 2 km./ S. Frutillar/ P.
Llanquihue/ 280’ II-3-1968] [Collectors: L
&/C.W. O’Brien]; | male, | female [CHILE/
Llanquihue/ Frutillar/ 23.9.54] [Coll./ Kus-
chel] [Entomology/ Division/ D.S.I.R./ New
Zealand]; 3 males, 2 females [CHILE/ Llan-
quihue/ Frutillar/ 24.2.50] [Coll./ Kuschel]
[Entomology/ Division/ D.S.I.R./ New
Zealand]; 1 male, 3 females [CHILE/ Fru-
tillar/ 14 Dec 1943/ G. Kuschel] [G. Kus-
chel/ Collection] [Entomology/ Division/
D.S.1.R./ New Zealand]; 1 male [CHILE/
Frutillar/ 28 Dec 1944/ G. Kuschel] [G.
Kuschel/ Collection] [Entomology/ Divi-
sion/ D.S.1I.R./ New Zealand]; | male, 7 fe-
males [CHILE/ Frutillar/ 27 Feb 1950/ G.
Kuschel] [G. Kuschel/ Collection] [Ento-
mology/ Division/ D.S.I.R./ New Zealand];
2 males, 1 female [MAULLIN a/ Las Que-
mas/ Llanquihue/ 17,-Dic.1972/ Coll: L. E.
107
Pena]; | male [CHILE/ Volcan Osorno/
4.3.50/ Kuschel leg.] [Coll./ Kuschel] [Ento-
mology/ Division/ D.S.I.R./ New Zealand].
1 male [CHILE: Laguna de Copa/ P. N. de
Puyehue, Osorno Pro./ Site 20, El. 520 m.
7-XII-77/ Valdivian Rain Forest/ Ash-
worth, Hoganson, Mooers] [on Berberis
darwinii] [Anthonomus/ ornatus/ Blan-
chard]; 3 males, 2 females [CHILE: Laguna
Espejo/ P. N. de Puyehue, Osorno Pro./ Site
17, El. 520 m. 21-XII-77/ Valdivian Rain
Forest/ Ashworth, Hoganson, Mooers] [on
Berberis darwinii] [Anthonomus/ ornatus/
Blanchard]; 3 males, 8 females [CHILE: La-
guna Espejo/ P. N. de Puyehue, Osorno Pro./
Site 17A. El. 520 m. 10-I-79/ Valdivian Rain
Forest/ A. C. Ashworth, J. W. Hoganson]
{on Berberis sp.] [Anthonomus/ ornatus/
Blanchard]; Total paratypes, 87; (AMNH,
CACA, HAHC, MNNC, MCZC, NZAC,
TAMU).
Recognition (Figs. 11, 12). Adults of 4.
berberidis are distinguished by the following
combination of characters:
Sutural elytral interstriae with acute,
overlapping apical extensions; postero-
lateral elytral maculae feebly developed
(Figs. 11, 12), surrounded dorsally by
complete pallid annuli; sternum 5 of male
with large posteromedian prominence;
metatibia of male with ventral margin
strongly concave in distal 74, with en-
larged, excavated, apical mucro; pygid-
ium of female with small apicomedian
prominence; aedeagus (Fig. 30) asym-
metrical, constricted medially, with blunt
subapicolateral prominences.
These, and adults of the closely related A.
chilicola have the aedeagus similarly asym-
metrical (cf. Figs. 29, 30), similarly modi-
fied male metatibiae with the ventral mar-
ginal concavity, enlarged, strongly excavated
apical mucrones, and similar extension of
the extreme apices of the sutural elytral in-
terstriae. This latter feature is even more
pronounced in adult 4. berberidis where the
108
extension of the right elytron overlaps that
of the left one. The elytral pattern of light
and dark integument and scales is also sim-
ilar in adult 4. chilicola and A. berberidis,
but is weakly developed in adult 4. berber-
idis, which have a prominent midbasal
macula (cf. Figs. 9, 12). The large apico-
median prominence on sternum 5 of the
male is unique to adults of A. berberidis.
Adult male. Length: 2.52-3.04 mm (x =
2.83, n = 10). Width: 1.12-1.44 mm (% =
1.28, n = 10). Rostrum: length 1.02-1.48 x
(X = 1.32, n = 10) pronotal length; length
of distal portion 22-31% (¥ = 25, n = 10)
of total rostral length. Prothorax: pronotum
without subapical fascia and dorsolateral
vittae of pallid scales; middorsal vitta in-
distinct. E/ytra: interstria 3 without distinct
basal and discal elevations; dark integu-
ment and scales predominant in large, mid-
basal macula and forming edges of weakly
developed posterolateral maculae. Legs:
protibia with distinct ventral marginal
prominence at proximal '/3; metatibia nar-
row, ventral margin strongly curved in dis-
tal 74, with distinct prominence at proximal
Y4; metatibial mucro large, curved, exca-
vated.
Adult female. Length: 2.16-3.16 mm (x
= 2.87, n = 10). Width: 0.92-1.36 mm (x
= 1.26, n = 10). Rostrum: length 0.87-1.40
x (X = 1.28, n = 10) pronotal length; distal
portion subcylindrical, shallowly punctate,
glabrous, length 26-33% (xX = 29, n = 10)
total rostral length. Abdomen: sternum 5
with posterior margin straight. Legs: meta-
tibia straight, narrow at apex, apical mucro
short, straight.
Distribution (Fig. 45). The type series of
A. berberidis consists of adult specimens
from the following localities. ARGEN-
TINA. Rio Negro: El Bolson. CHILE. Chi-
loé: 19 km. S Ancud; 13 km. S Castro; Che-
pu; Dalcahue; Puerto Montt; 12 km. N
Puerto Montt; Quellon. Lianquihue: Frutil-
lar; 2 km. S Frutillar; Maullin. Osorno: La-
guna la Copa and Laguna Espejo, Parque
Nacional de Puyehue; Volcan Osorno.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Plant associations. Specimens of A. ber-
beridis were collected on Berberis darwini
19 km. S Ancud and at Chepu, Chiloé, and
at Laguna la Copa and Laguna Espejo, Par-
que Nacional de Puyehue, Osorno, Chile.
Label data indicate that adults have been
collected in January, February, March, Sep-
tember, October, November and Decem-
ber.
Etymology. The name of this species is
derived from that of the plant genus with
which it and several other 4. ornatus group
members are associated.
PHYLOGENY
The classification of the subfamily An-
thonominae has been called “chaotic”
(Burke 1976). Some 500, mostly New World,
named species are assigned to the subfam-
ily, but recent revisionary studies (Clark
1987a, b, c, 1988, Clark and Burke 1985,
1986a, b, c) indicate that the actual number
of species may be twice that. Within this
unwieldy mass of species, some apparently
natural groupings do present themselves, but
it is not often possible to identify the sister
groups of these. This is the case with the A.
ornatus group. The group seemed by pre-
liminary inspection to be natural, but iden-
tification of the sister group remains for fu-
ture studies. Thus, in order to apply the
outgroup method of comparison to identify
apomorphic characters in the 4. ornatus
group, it was necessary to attempt to iden-
tify characters unique to the group by ex-
amining a broad, though incomplete, array
of anthonomine taxa. Further studies may
reveal that some of these characters are more
widely distributed.
The most detailed comparisons included
those of characters of the 4. ornatus group
with those of anthonomines examined in
connection with ongoing revisionary stud-
ies. These include the species in the An-
thonomus albolineatus, furcatus, grandis,
gularis, unipustulatus, and venustus groups,
the species of the Anthonomus subgenera
Anthomorphus, Anthonomocyllus and An-
VOLUME 91, NUMBER |
109
107,12
a
signatipennis
ornatus
ace
blanchardi
2,4,7,8-
kuscheli
ee |
3,5,9-——]
Fig. 46.
thonomorphus, the genera Atractomerus,
Loncophorus, Omogonus and Pseudantho-
nomus, and the types of most of the named
Neotropical Anthonominae. Numerous
Nearctic and Old World species have also
been examined, but not in as much detail.
There is no indication that any of the species
examined should be placed in the 4. ornatus
group, nor are there indications that any of
them is the sister group of the 4. ornatus
group.
As a result of the comparisons made, 13
characters (listed below) were determined
to be unique to adults of the 4. ornatus
group. The distribution of these characters
among the seven species in the group is de-
picted in Table 1. Presence 1s indicated by
a score of “1,” absence by a score of “0,”
“missing,” because the female of 4. chili-
araucanus
Phylogenetic tree depicting hypothesized relationships of the species of the A. ornatus group.
cola is unknown, by a score of 9. These
characters were analyzed using the PAUP
computer programs developed by Swofford
(1985), on an IBP Personal Computer to
determine the most parsimonious branch-
ing pattern consistent with this distribution
of characters. Analysis of the 13 characters,
all given equal weight, with the ALLTREES
option which searches all possible trees,
produced the phylogenetic tree depicted in
Fig. 46. The characters, with comments on
their distributions under the constraints of
the accepted solution, are listed below.
1) Elytral integument with markedly con-
trasting patterns of pallid and dark por-
tions clothed with correspondingly pal-
lid and dark scales ranging from white
through pallid to dark ferruginous to
Table 1. Data matrix for the phylogenetic tree in Fig. 46.
Characters
0 0 0 0) 0 0 0 0 0 1 ] 1 ]
| 2 3 4 5 6 7 8 9 0 I 2 3
Outgroup 0 0 0 0 0 0 0 0 0 0 0 0 (0)
signatipennis 1 0 0 0 0 l 0 0 0 0 0 0 0
ornatus l 1 0 1 0 I 0 i 0) l l ] 0
blanchardi 1 1 0 1 0 1 0 1 0 1 ] 1 (0)
kuscheli 1 l 0 1 0 1 1 l 0 l ] 0 0
araucanus l 1 0 0 0 1 1 1 0) 0 0 0 l
chilicola 1 1 1 1 l l 0 l 1 9 9 9 9
] 1 1 1 1 0 0 1
berberidis
110 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
fuscous (Figs. 1-12). Although con-
trasting patterns of light and dark in-
tegument and light and dark scales are
not unique to the A. ornatus group, the
patterns are similar in the species in the
group and do not seem to be exactly
duplicated in any of the other anthon-
omines.
2) Elytra with large, posterolateral mac-
ulae (Figs. 3-12). These are weakly de-
veloped in A. araucanus and A. ber-
beridis (Figs. 7, 8, 11, 12).
3) Sutural elytral interstriae with acute,
overlapping apical extensions.
4) Aedeagus with apicolateral promi-
nences (Figs. 21, 23, 25, 27, 29). These
are assumed to have been lost in 4.
araucanus (Fig. 28) and A. berberidis
(Fig. 30).
5) Aedeagus asymmetrical (Figs. 29, 30).
6) Endophallus with an elongate, flat, dis-
tal sclerite (Figs. 19-30).
7) Endophallus with two dentate median
sclerites (Figs. 27, 28). These are as-
sumed to have been secondarily lost in
the ancestor of A. ornatus and A. blan-
chardi and in that of A. chilicola and A.
berberidis.
8) Parameres of tegmen of male genitalia
close together or fused at base.
9) Metatibia of male with ventral margin
strongly concave in distal 74, with en-
larged, excavated, apical mucro.
10) Pygidium of female with subapical
prominences (Figs. 14, 15). These are
weakly developed and assumed to be
obsolescent in 4. ornatus (Fig. 14) and
A. blanchardi.
11) Sternum 5 of female with apicome-
dian prominence. This is a simple
prominence in 4. kuscheli, a promi-
nence bounded by shallow emargina-
tions in A. blanchardi (Fig. 18), a long
prominence bounded on each side by
deep incisions in 4. ornatus (Fig. 17).
12) Sternum 5 of female incised or emar-
ginate (Figs. 17, 18).
13) Pygidium of female with apicomedian
prominence (Fig. 16).
ACKNOWLEDGMENTS
Thanks are extended to the individuals
and institutions listed in the Materials and
Methods section for arranging loans of the
specimens used in this study. Guillermo
Kuschel initially suggested the group as suit-
able for revision, provided information and
other valuable assistance, along with a crit-
ical review of the manuscript. Charles W.
O’Brien and Guillermo Wibmer also re-
viewed the manuscript. This paper is pub-
lished as Alabama Agricultural Experiment
Station Journal Series No. 17-881496P and
Texas Agricultural Experiment Station T.A.
23182.
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weevil tribe Anthonomini (Coleoptera: Curculion-
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73-88.
1987b. Revision of the Nearctic species of
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nus Anthomorphus Weise (Coleoptera: Curculion-
idae). Quaest. Entomol. 23: 317-364.
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1988. The species of Anthonomus in the al-
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thonomus Germar (Coleoptera: Curculionidae).
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1986a. Revision of the gu/aris species group
of the genus Anthonomus Germar (Coleoptera:
Curculionidae). Coleopts Bull. 40(1): 1-26.
1986b. Phylogeny of the species of the An-
thonomus subgenus Anthonomorphus Dietz, with
discussion of relationships with Anthonomus
grandis Boheman (Coleoptera: Curculionidae). J.
Kans. Entomol. Soc. 59(3): 508-516.
1986c. A new neotropical species of Anthon-
omus Germar (Coleoptera: Curculionidae) asso-
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11
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PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 112-123
A REVIEW OF THE WORLD SPECIES OF NOTOGLYPTUS MASI
(HYMENOPTERA: PTEROMALIDAE)
Steve L. HEYDON
Department of Entomology, NHB; Mail Stop 165, Smithsonian Institution, Washing-
ton, D.C. 20560.
Abstract. — Until the present review, Notog/yptus Masi was a monotypic genus containing
the single species N. virescens Masi, which was known from throughout the Old World.
This species is redescribed, and its known distribution is extended to the Nearctic region.
Four new species of Nofoglyptus from the New World are described: N. bidentatus, N.
luteicrus, N. nesiotes and N. tzeltales. Notoglyptus is redescribed, its relationships to other
genera of the Miscogasterinae are discussed, and a key is given for distinguishing the
known world species. The host(s) of species in this genus are still unknown.
Key Words:
taxonomy
Masi (1917) described the genus Noto-
glyptus and included two species, N. vires-
cens Masi from the Seychelles Islands, and
N. niger Masi from Italy. He designated the
latter species as the type-species of the ge-
nus. Boucek (1976) synonymized N. niger
with N. virescens. This cosmopolitan species
was by then known to occur throughout
southern Europe, across to India, and down
into southern Africa (Graham 1969, Boucek
1976). I have seen specimens from Japan
and a single male from North America.
During the course of sorting collections for
a continuing study of the Nearctic misco-
gasterine Pteromalidae, I came across spec-
imens belonging to four undescribed species
of Notoglyptus from the New World—N. bi-
dentatus n. sp. from South America, N. tzel-
tales n. sp. from Central America and north-
ern South America, N. /uteicrus n. sp. from
the continental Nearctic region south to
Venezuela, and N. nesiotes n. sp. from the
Caribbean and coastal southeastern U.S.
Graham (1969) placed Notog/yptus in the
Sphegigasterini (Miscogasterinae), probably
Hymenoptera, Pteromalidae, Notoglyptus, Nearctic, Palearctic, Neotropics,
because the elongate T1 is found in other
genera he placed there such as Crypto-
prymna Forster and Novitzkyanus Boucek.
Genera with an elongate T1 occur sporad-
ically throughout the Pteromalidae, and I
feel its common occurrence in these three
genera is due to convergence (Heydon 1988).
The closest relatives of Notoglyptus are
among genera related to Halticoptera Spi-
nola and Thinodytes Graham. These genera
all share a similar unique propodeal struc-
ture—the median panels of the propodeum
are reticulately sculptured or smooth and
have a distinct median carina and plicae
connected posteriorly by a W-shaped carina
(compare Figs. 12 and 13). This carina is
most distinctly developed in N. bidentatus
and N. fzeltales, the most morphologically
generalized Notoglyptus species. The ter-
minal two segments of the maxillary palps
of male Halticoptera are lamellately ex-
panded, and this character state also occurs
in Notoglyptus luteicrus (Fig. 11). However,
using this character state as direct evidence
for a close relationship between Halticop-
VOLUME 91, NUMBER 1
teraand Notoglyptus is complicated because
N. luteicrus is one of the more morpholog-
ically derived members of the genus; this
character state is not present in the more
primitive extant species of Notog/yptus.
The characters defining Notoglyptus are
reviewed in the Discussion section follow-
ing the generic description below. Noto-
glyptus may have evolved in South America
since that is where the most primitive species
(N. bidentatus and N. tzeltales) are found.
MATERIALS AND METHODS
This study is based on examination of 67
specimens from the museums whose acro-
nyms are given 1n the Acknowledgment sec-
tion below. Type depositions are given in
parentheses in the appropriate section un-
der each species description. Terminology
in this paper generally follows that of Gra-
ham (1969), except that club is used instead
of clava and the gastral terga are numbered
1-7 starting with the basal tergite of the gas-
ter. The following abbreviations are used:
the multiporous plate sensillae are MPP
sensillae, the lower ocular line is LOcL, the
antennal funicular segments are F1, F2,.. ,
F6, and the gastral terga are T1, T2,.. ,T7.
The units of measurement given in the de-
scriptions can be converted to millimeters
by multiplying by 0.02.
Notoglyptus Masi
Notoglyptus Masi, 1917: 181. Type species
N. niger Masi (original designation). Ga-
han and Fagan, 1923: 98. Peck, Boucek,
and Hoffer, 1964: 36 (key). Graham, 1969:
124 (key), p. 140. Boucek, 1976: 15.
Dzhanokmen, 1978: 77 (key), p. 80. Fa-
rooqi and Subba Rao, 1985: 259, 310F,
310G; 1986: 295.
Description. Color: Body ranges from
black to metallic green or blue; legs metallic
or entirely nonmetallic. Structure: Head
ovate (Fig. 7) or triangular (Fig. 1) in an-
terior view; clypeus with anterior margin
straight or produced (Fig. 7), bidentate in
113
N. bidentatus (Fig. 1); genal concavities well
developed, extending 2 to 5 malar length.
Antenna inserted near or below LOcL; with
2 anelli, 6 funicular segments, and 3-seg-
mented club; MPP sensillae in 1 row on
each funicular segment (Figs. 3, 5); club often
with micropilosity on terminal segment(s)
(Figs. 3, 5), apical spine not present. Meso-
soma with pronotum reduced, much nar-
rower and depressed relative to mesoscu-
tum (Figs. 8, 10, 12), horizontal collar
developed in N. bidentatus (Fig. 8), N. tzel-
tales, and N. virescens but not present in N.
luteicrus (Fig. 10) and N. nesiotes (Fig. 12),
this collar bordered anteriorly by weak to
strong anterior transverse carina; mesoscu-
tum with notauli complete, groovelike (Figs.
8, 10, 12); upper epimeron smooth; scutel-
lum with scutoscutellar sulcus foveolate
medially in N. nesiotes (Fig. 12) and N. vi-
rescens, frenal sulcus distinct (Figs. 8, 10,
12), frenum weakly coriaceous or smooth;
propodeum with median panels alveolate or
smooth, plicae and median carina distinct,
connected posteriorly by W-shaped carina
(Fig. 9). Wing hyaline; relative lengths of
veins as submarginal > marginal > post-
marginal > stigmal; basal cell and vein se-
tate (Figs. 2, 4) except in N. tzeltales (Fig.
6); fore wing sometimes with distinct ad-
marginal setae. Petiole quadrate to elongate,
granulate to alveolate; median carina some-
times present; basal flange present. Gaster
ovate; Tl nearly concealing succeeding ter-
ga, hind margin straight; hypopygium ex-
tending nearly to tip of gaster. Male max-
illary palps pale in all species except N.
bidentatus, terminal two segments lamel-
lately expanded in N. /uteicrus (Fig. 11).
Discussion. The following combination
of character states will reliably distinguish
Notoglyptus: the presence of distinct genal
concavities; a 13-segmented antenna; com-
plete, groovelike notauli (Figs. 8, 10, 12);
the propodeum with the median carina and
plicae distinct, and connected posteriorly by
W-shaped carina (Fig. 9); the petiole about
as long as wide, with a basal ventral flange;
114
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-7.
(ww
(exe)
1-2, Notoglyptus bidentatus n. sp., male. 1, Anterior view of head. 2, Fore wing. 3, Notoglyptus
luteicrus n. sp., female. 3. Antenna. 4—5, Notoglyptus nesiotes n. sp., female. 4, Fore wing. 5, Antenna. 6-7,
Notoglyptus tzeltales n. sp., female. 6, Fore wing. 7, Anterior view of head.
Tl almost completely covering the entire
length of gaster, with its hind margin entire;
and the hypopygium extending to the tip of
the gaster. An autapomorphy defining No-
toglyptus is the reduction in size of the
pronotum relative to the rest of the meso-
soma. This character 1s easily seen in com-
parison with other closely related misco-
gasterine genera such as Halticoptera
(compare Figs. 8, 10, 12 with 13). The
unique fovea in the center of the scutellum
sO prominent in the type species is clearly
ofno more than specific value when looking
at this genus on a worldwide basis.
The species of Notoglyptus can be divided
into two distinct species-groups. Notoglyp-
tus bidentatus and N. tzeltales comprise the
first group which is characterized by a large
body size, the body and legs with extensive
bright metallic coloration, the anterior mar-
gin of the clypeus produced (Figs. 1, 7), the
toruli located below the LOcL (Figs. 1, 7),
the female antennal club with no micropi-
losity, the fore wing lacking any distinct ad-
marginal setae, and the petiole without a
median carina. Except possibly for the low
insertion of the antennae, these characters
states are all plesiomorphic compared with
those defining the other species-group. No-
toglyptus luteicrus, N. nesiotes, and N. vi-
rescens form the second species-group.
These species are relatively small, have the
VOLUME 91, NUMBER 1
Figs. 8-13. 8-9, Notoglyptus bidentatus n. sp., male. 8, Dorsolateral view of mesosoma. 9, Frenum, dorsellum,
and propodeum. 10-11, Notoglyptus luteicrus n. sp., male. 10, Dorsolateral view of mesosoma. 11, Maxillary
palps. 12, Notoglyptus nesiotes n. sp., female. 12, Dorsolateral view of mesosoma. 13, Halticoptera sp., female.
13, Dorsolateral view of mesosoma. Scale bar = 0.1 mm.
body color very dark, the legs (including the antennal club with micropilosity, the fore
coxae) yellow and nonmetallic, the anterior wing with distinct admarginal setae, and the
margin of the clypeus nearly straight, the petiole with a median carina.
toruli located above the LOcL, the female Despite these obvious morphological dif-
116 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ferences, the species I have placed together
in Notoglyptus belong together as a mono-
phyletic unit within the Miscogasterinae on
the basis on the reduced size of the prono-
tum, and with respect to closely related gen-
era such as Halticoptera, because of the
elongate first gastral tergite and hypopyg-
ium. Division of the two species-groups into
separate genera would result in the species-
group containing Notoglyptus bidentatus and
N. tzeltales being paraphyletic. Further, the
important character of the presence or ab-
sence of the horizontal pronotal collar cuts
across the species-groups as I have outlined
them here. Notoglyptus virescens has a hor-
izontal collar as do N. bidentatus and N.
tzeltales; yet, in the characters separating
the species-groups, N. nigrescens belongs
with N. /uteicrus and N. nesiotes.
Key TO WORLD SPECIES OF
NoOTOGLYPTUS Masi
1. Pronotum lacking horizontal collar, sloping
downward immediately from anterior margin
of mesoscutum (Figs. 10, 12) ;
— Pronotum with short horizontal collar sepa-
rated from declivitous neck by a weak to
strongly developed transverse carina (Fig. 8) .. 3.
2. Scutellum with a pair of diverging foveae at
base (Fig. 12). Dorsellum cariniform. Female
antenna with funicular segments 5—6 quadrate
to elongate (Fig. 5). (West Indies and coastal
southeastern United States) nesiotes Heydon
— Scutellum with scutoscutellar sulcus a contin-
uous deep furrow, not foveolate (Fig. 10). Dor-
sellum bandlike, length nearly half that of fre-
num. Female antenna with funicular segments
5-6 transverse (Fig. 3). (Continental North
America south to Venezuela) luteicrus Heydon
3. Scutellum with discal fovea. Torulus above
lower ocular line. Coxae and femora yellow,
nonmetallic .. virescens Masi
— Scutellum smoothly convex (Fig. 8). Torulus
below lower ocular line (Figs. 1, 7). Coxae and
femora blue or green, metallic. (Neotropical) 4.
4. Head triangular in anterior view; anterior mar-
gin of clypeus bidentate (Fig. 1). Basal cell of
fore wing setate (Fig. 2). Body length 2 mm or
more. (South America) bidentatus Heydon
— Head ovate in anterior view; anterior margin
of clypeus truncate (Fig. 7). Basal cell of fore
wing bare (Fig. 6). Body length 1.8 mm or less.
(Central America and northern South Ameri-
ca) tzeltales Heydon
io)
Notoglyptus bidentatus,
New SPECIES
Figs. 1-2, 8-9
Holotype, female. Description. Color:
Head dark green; mesosoma mostly dark
green with strong coppery reflections on
dorsum and upper epimeron; pronotum,
propodeum, petiole, gaster bluish black.
Antenna with scape, pedicel dark green; fla-
gellum black. Maxillary palp brown. Legs
with coxae dark blue; fore, middle trochan-
ters orange-yellow, hind trochanter brown;
femora blue-green, orange-yellow basally
and apically; tibiae orange-yellow basally,
remainder orange-brown with weak metal-
lic reflections medially; tarsi dark brown.
Wing veins brown.
Sculpture: Clypeus, gena finely coria-
ceous; head finely alveolate otherwise.
Mesosoma (Fig. 8) with middle lobe of
mesoscutum alveolate; side lobes, scutel-
lum, frenum (weakly) finely coriaceous;
dorsellum smooth; median panels of pro-
podeum alveolate (Fig. 8); petiole finely al-
veolate, alveoli 2x as long as wide; gastral
terga smooth.
Structure: Body length 2.4 mm. Head (Fig.
1) triangular in anterior view, width 1.4 x
height (46:34), 2.2 x length (46:21); clypeus
separated from face by deep furrowlike ca-
rina, anterior margin produced and biden-
tate; genal concavity shallow, extending 1
malar distance; frons regularly concave be-
tween eyes; eye height 1.4 length (16.5:
12.0), 1.1 malar length (16.5:15.0), eye
length 1.5 temple length (12:8); ratio of
MOD, OOL, POL, LOL as 3:11:8:4. An-
tenna with torulus one inside diameter be-
neath LOcL; combined length of pedicel and
flagellum 0.85 head width (39:46); rela-
tive lengths of scape, pedicel, anelli, Fl-6,
club as 22.0:7.0:3.0:4.0:4.5:4.0:4.0:4.0:3.5:
7.0; widths of Fl, F6, club as 3:5:5; second
anellus 2x as large as first; club without
micropilosity. Mesosoma (Fig. 8) length
1.6 width (59:36); collar with weak an-
terior transverse carina; scutellum regularly
rounded, scutoscutellar sulcus continuous
VOLUME 91, NUMBER 1
furrow; dorsellum bandlike, length equal to
frenal length; spiracles ovate, <1 x own di-
ameter from anterior margin of propodeum;:
nucha bordered by carina anteriorly. Fore
wing (Fig. 2) length 2.3 width (145:63);
ratio of submarginal, marginal, postmargin-
al, stigmal vein lengths as 50.0:30.0:24.0:
13.5; costal cell densely setate; basal cell
setate to base along anterior margin; spe-
culum closed posteriorly; no distinct ad-
marginal setae present. Petiole broadening
apically, length 1.8 x apical width (15.0:8.5);
lacking median carina. Gaster length 1.4 x
width (42:30); deep, height 1 x width (30:
30).
Allotype, male. Color: Similar to holo-
type female except front of head blue-green
with violet reflections, anelli and funicular
segments tan ventrally. Structure: Body
length 2.4 mm. Antenna with combined
length of pedicel and flagellum 0.93 x head
width (40:43); lengths of scape, pedicel,
anelli, Fl-6, club as 20:5:3:4:4:4:4:4:4:9;
widths of Fl, F6, club as 3.5:4.0:4.0; funic-
ular segments tend to be hemispherical.
Maxillary palps slender. Petiole length 2.2 x
width (15.5:7.0). Gaster length 1.3 x width
(67-28):
Variation. The body length of the para-
type males ranges between 2.1 and 2.8 mm.
Though all the paratype males were col-
lected from approximately the same area,
they show considerable variation in color.
Most are basically green like the allotype,
but one is blue, and several of the others
have greater or smaller areas of the head
and mesosoma purple. One male has a pet-
iole only 1.4 times as long as wide; the pet-
iole of the other males ranges between 1.8
and 2.3 as long as wide. The basal cell
varies from totally setate, as in the holotype,
to two specimens which have only a couple
of rows of setae along the anterior margin
of the basal cell.
Discussion. Unique features of this species
are the triangular head (Fig. 1), the broad
and deep antennal scrobes, the long malar
distance, the bidentate clypeus (Fig. 1), the
117
deep sulcus around the clypeus (Fig. 1), and
the dark maxillary palps. Notoglyptus bi-
dentatus exhibits a few primitive character
states not found in congeners. It has a dis-
tinct pronotal collar (Fig. 8) (a character state
shared with N. tze/tales and N. virescens), a
bidentate clypeus, and dark maxillary palps.
The extensively setate wings (Fig. 2) may
also be primitive. However, setate wings are
common in miscogasterine species living at
high elevations or latitudes, so it is difficult
to say in this case whether the setate wings
of N. bidentatus are primitive or an adap-
tation to the mountainous habitats where
this species occurs.
Etymology. The specific name is from the
Latin word bidentatus, meaning two-
toothed, and refers to the unique bidentate
state of the anterior margin of the clypeus
in this species.
Biology. Nothing is known of the host(s)
of this species.
Type material. The holotype, allotype, and
5 male paratypes (all CNC) were collected
in Ecuador in the elfin forest at 3800 meters
along the Quito-Baeza Road on | March
1979 by W. Mason. Four additional para-
types (CNC, INHS, USNM) were collected
as follows: Ecuador. Napo (4100 m, Quito-
Baeza Road), 24:-II- 1983, 1 4, Paruma (4200
m, Quito-Baeza Road), 14-17-II- 1982, 3 4.
Notoglyptus luteicrus,
New SPECIES
Figs. 3, 10-11
Holotype, female. Description. Color:
Head, mesosoma, petiole black, except the
following blue: clypeus; pairs of diffuse spots
extending anteriorly from lateral ocelli; ver-
tex along orbits; lateral parts of pronotum
and mesoscutum, frenum, dorsellum, pro-
podeum; pleural regions posteriodorsally.
Gaster brown, Tl with bluish reflections.
Antenna with scape yellow-brown, darker
in apical 4: pedicel brown; flagellum dark
brown. Mandibles yellow, teeth reddish yel-
low. Maxillary palps yellow. Legs yellow,
pretarsi dark brown. Wing veins pale brown.
118
Sculpture: Clypeus smooth; remainder of
head, mesoscutum (Fig. 10), scutellum del-
icately alveolate; frenum coriaceous (Fig.
10); dorsellum smooth; median panels of
propodeum alveolate, pattern very weak in
center of each panel: petiole granulate; gas-
tral terga smooth.
Structure: Body length 1.4 mm. Head
ovate in anterior view, width 1.3 height
(27:21), 2.2 x length (27:12); clypeus set off
from face by obscure sulcus, anterior mar-
gin straight; genal concavity extending half-
way to eye; antennal scrobes shallow; eye
height 2.4 length (13.0:5.5), 2.2 malar
length (13:6), length 4.9 x temple length (9.5:
2.0); ratio of MOD, OOL, POL, LOL as 1.5:
5:6.5:3. Antenna (Fig. 3) with torulus one
outside diameter above LOcL; combined
length of pedicel and flagellum 0.93 x head
width (25:27); relative lengths of scape, ped-
icel, anelli, Fl-6, club as 13.0:4.0:1.0:2.0:
2.5:2.0:2.0:2.0:1.5:8.0; widths of Fl, F6,
club as 2.0:3.5:4.0; anelli subequal in size;
micropilosity completely covering ventral
side of terminal segment. Mesosoma length
1.4 width (28:20); collar undeveloped:
scutellum regularly rounded, scutoscutellar
sulcus continuous furrow; dorsellum length
‘> frenal length; spiracles round, | x own
diameter from anterior margin of propo-
deum; nucha acarinate anteriorly. Fore wing
length 3.1 width (65:21); ratio of sub-
marginal, marginal, postmarginal, stigmal
vein lengths as 20:11:9:6; costal cell with
single complete row of setae; basal vein with
row of 4 setae; basal cell with | seta on left
wing; speculum open posteriorly; | row of
distinct admarginal setae present. Petiole
length 1.1 x width (4.5:4.0); median carina
present. Gaster ovate, length 1.4 width
(29:21); height 0.86 x width (18:21).
Allotype, male. Description. Color: Sim-
ilar to holotype except reflections of frenum,
dorsellum, propodeum green. Structure:
body length 1.4 mm. Antenna with com-
bined length of pedicel and flagellum 1.2 x
head width (31.0:26.5); relative lengths of
scape, pedicel, anelli, F1-6, club as 14:3:1:
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
3:3:3:3:3:3:9; widths of Fl, F6, club as 2:3:
3. Maxillary palps (Fig. 11) with 2 apical
segments lamellately expanded. Petiole
length 1.2 x width (6:5). Gaster length 1.4 x
width (22:16).
Variation. Body size in the females ranges
from 1.0 to 1.5 mm; in the males, 1.2 to
1.4 mm. The patches of metallic coloration
on the head and mesosoma vary among
specimens in extent, intensity, and color.
The color varies from blue, as in the ho-
lotype, to dark green. The specimens from
the eastern U.S. and Canada usually have
the scape brown only at the apex, but those
from Mexico and the western U.S. and Can-
ada have the scape mostly brown. The setal
patterns of the wings are variable, but the
wings generally resemble that in Fig. 4.
Discussion. The unique feature of this
species is the lamellately expanded male
maxillary palps (Fig. 11). Within its species-
group, Notoglyptus luteicrus resembles N.
nesiotes in lacking any horizontal collar on
the pronotum (Fig. 10) and a patch of mi-
cropilosity on the apical segment of the fe-
male club (Fig. 3). Notoglyptus luteicrus has
spots of metallic coloration located in sim-
ilar places to those in N. nesiotes but the
boundaries of the spots on the head are dif-
fuse in N. /uteicrus and sharp in N. nesiotes.
Etymology. The specific name comes from
the Latin words /uteus, meaning yellow, and
crus, meaning leg, and refers to the yellow
legs of this species.
Biology. Nothing is known of the host(s)
of this species, but specimens have been
collected from alfalfa, soybeans, Baccharis
L. (Compositae), and the crucifers Sisym-
brium altissimum L. and Descurainia so-
phia (L.) P. B. Webb.
Type material. The holotype (INHS) was
collected from the railroad siding at the end
of Gerty Drive on the South Farms of the
University of Illinois, Champaign County,
Illinois, on 25 June 1982, by S. L. Heydon.
The allotype (INHS) came from the same
locality, but was collected 21 August 1981.
Eighteen additional paratypes were collect-
VOLUME 91, NUMBER |
ed as follows (AMNH, CNC, INHS, SEC,
USNM): Canada. ALBERTA: Lethbridge,
5-VII-1956, (swept from brome) 2 2; Lost
River, Onefour (10 mi. WNW Wild Horse),
3-VI-1956, 1 2. ONTARIO: 13 mi. N Belle-
ville, 27-V-1970, 1 °. United States. CAL-
IFORNIA: Rancho Santa Fe, 14-1-1959,
(alfalfa field) 1 9. COLORADO: Fort Col-
lins, V- 1894, (alfalfa) 1 6. FLORIDA: Col-
lier Seminole State Park (Collier Co.), 25-
26:V-1978, 1 2; Fort Ogden, 8-IV-1952, 1
2. IDAHO: Hansen, 29: V- 1930, (Sisymbri-
um altissimum and Descurainia sophia) |
9. INDIANA: Hovey Lake (Posey Co.), 3-
VI-1981, 1 6. ILLINOIS: Dixon Springs Ag-
ricultural Research Station (Pope Co.), 29-
31-VII-1980, 1 6. LOUISIANA: Cameron
Parish, 4:-VIII-1969, (soybeans) | @.
MARYLAND: Patuxent Research Center
(Prince Georges Co.), 1- VII: 1982, 12. NEW
JERSEY: Ramsey, 31-VII-1918, 1 9. NEW
MEXICO: Elmendorf, 21-VII-1936, 1 2.
UTAH: Richfield, 18-V-1954, 1 9; Utah
Lake (Utah Co.), 1 2. Mexico. VERACRUZ:
Jalapa, HI-IV-1965, 1 2. Venezuela. POR-
TUGUESA: 10 km N Biscucuy, 9: VI-1981
(sweeping Baccharis), 1 6. Country? San Ra-
fael Jicoltepec, 1 2, 3 6.
Notoglyptus nesiotes,
NEw SPECIES
Figs. 4-5, 12
Holotype, female. Description. Color:
Head, mesosoma, petiole black except pro-
podeum metallic coppery; inner orbit along
vertex, area between median and lateral
ocelli, pronotum and mesoscutum laterally,
posteriodorsal pleural region metallic green.
Gaster dark reddish brown with coppery re-
flections dorsally. Antenna with scape, anel-
li yellow; pedicel, flagellum brown. Man-
dibles yellow, teeth reddish yellow.
Maxillary palps yellow. Legs yellow, pre-
tarsi black. Wing veins yellow, parastigma
darker.
Sculpture: Body sculpture weak, face co-
riaceous, mesoscutum (Fig. 12) weakly al-
119
veolate mesally, petiole granulate, body
nearly smooth otherwise.
Structure: Body length 1.4 mm. Head
ovate in anterior view, width 1.4 height
(22:16), 2.2 x length (22:10); clypeus sepa-
rated from face by obscure sulcus, anterior
margin straight; antennal scrobes shallow;
genal concavity weakly developed, extend-
ing only '4 of way to eye; eye height 1.2 x
length (9.0:7.5), 2.0 x malar length (9.0:4.5),
eye length 3.8 x temple length (7.5:2.0); ra-
tio of MOD, OOL, POL, LOL as 1.5:5:5.5:
3. Antenna (Fig. 5) with torulus | own
outside diameter above LOcL; combined
length of flagellum and pedicel 1.0 head
width (23:22): relative lengths of scape, ped-
icel, anelli, Fl-6, club as 10.5:3.5:1.0:2.0:
2.0:2.0:2.0:2.5:2.0:6.0; widths of Fl, F6,
club as 1.5:2.0:2.0; anelli subequal in size;
club with patch of micropilosity down ven-
tral side of apical segment. Mesosoma (Fig.
12) length 1.7 width (26:15); collar not
developed; scutellum uniformly convex,
scutoscutellar sulcus with pair of contiguous
diverging foveae mesally; dorsellum cari-
nalike; spiracles 1.5 x own diameter from
anterior margin of propodeum, nucha bor-
dered by carina anteriorly. Fore wing (Fig.
4) length 2.7 x width (48:18); submarginal,
marginal, postmarginal, stigmal vein lengths
as 17.0:11.5:7.0:5.0; costal cell with one
complete row of setae plus a few others dis-
tally; basal vein setate; speculum open pos-
teriorly; distinct row of admarginal setae
present. Petiole length 0.88 width (3.5:
4.0); median carina present. Gaster ovate,
length 1.4 width (25:18); height 1.0 x
width (18:18); T2-7 protruding from be-
neath T1.
Male unknown.
Variation. The specimens in the type se-
ries from Isla Mona are all of a rather uni-
form size, coloration, and morphology. The
number of setae along the basal vein varies
from just a couple of setae to a row extend-
ing down the length of the basal cell and
curving basally down the cubital vein. About
half the specimens have a few setae distally
120
in the basal cell. The paratype female from
Sapelo Island, Georgia, lacks the metallic
patches on the head and is more distinctly
sculptured than the series from Isla Mona.
Its body sculpturing resembles that given
above for N. /uteicrus. It also resembles N.
luteicrus in lacking the distinct carina along
the anterior margin of the nucha. However,
it has long terminal funicular segments of
the antenna, a foveolate scutoscutellar sul-
cus (Fig. 12), and a cariniform dorsellum,
features which are all diagnostic for N. /u-
teicrus. Collection of specimens from a wid-
er geographic range may help make sense
of the morphological divergence between the
populations from Puerto Rico and those of
the continental U.S.
Discussion. Unique features of this species
are the weak body sculpture (Fig. 12) and
the cariniform dorsellum (Fig. 12). Noto-
glyptus nesiotes and N. virescens both have
terminal funicular segments of the antenna
quadrate or elongate and a foveolate scu-
toscutellar sulcus, but N. nesiotes lacks the
fovea on the disc of the scutellum charac-
teristic of N. virescens.
Etymology. The species name is derived
from the Greek word nesiotes, meaning in-
sular, and refers to the island distribution
of this species.
Biology. The host(s) of this species are
unknown. The paratype female from Sapelo
Island, Georgia, was collected on Spartina
Schreber (Gramineae).
Type material. Holotype (USNM) and 8
paratype females (CNC, USNM) were col-
lected on Isla Mona, Puerto Rico, in August
1944, by H. A. Beatty. One additional para-
type female was collected as follows: United
States. GEORGIA: Sapelo Island (Mcin-
tosh Co.), 10-I1X-1963 (on Spartina).
Notoglyptus tzeltales,
New SPECIES
Figs. 6-7
Holotype, female. Description. Color:
Front of head dull dark green; vertex, dor-
sum of mesosoma blue-green; propodeum,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
pleural regions, coxae green; petiole bluish
black; gaster dark reddish brown with strong
bluish reflections. Antenna with scape blue-
green; remainder brown, pedicel with weak
metallic reflections. Mandibles brownish
yellow; teeth pale brownish red. Maxillary
palps cream-colored. Legs with most of
femora brown with weak metallic reflec-
tions; tibiae brownish yellow, slightly dark-
er mesally; basal tarsal segment brownish
yellow, rest of tarsi darkening distally tll
pretarsus black. Wing veins reddish brown.
Sculpture: Clypeus granulate; head,
mesosoma, scutellum regularly alveolate;
frenum, dorsellum, propodeum smooth;
petiole finely strigulate dorsally; gaster
smooth.
Structure: Body length 1.6 mm. Head (Fig.
7) broadly ovate in anterior view, width
1.3x height (32:24), 2.1 length (32:15);
clypeus separated from face by distinct sul-
cus, anterior margin produced but truncate;
antennal scrobes shallow; genal concavity
extending 2 way to eye; eye height 1.3 x
length (14:11), 1.8 malar length (14:8),
length 3.7 temple length (11:3); ratio of
MOD, OOL, POL, LOL as 2.5:6:6:3. An-
tenna with torulus located just below LOcL;
combined length of pedicel and flagellum
0.72 head width (23:32); relative lengths
of scape, pedicel, anelli, Fl—-6, club as 14.0:
4.5:1.5:2.0:2.0:2.0:2.0:2.0:2.0:5.0; relative
widths of Fl, F6, club as 2.0:3.0:3.5, all
funicular segments except Fl transverse;
anelli subequal in size; club lacking ventral
patch of micropilosity. Mesosoma length
1.4 width (34:24); collar developed, an-
terior edge rounded; scutellum regularly
convex, scutoscutellar sulcus continuous
furrow; dorsellum length about ' frenal
length; spiracles ovate, <1 x own diameter
from anterior margin of propodeum; carina
bordering nucha anteriorly. Fore wing (Fig.
6) length 2.2 width (71:33); ratio of sub-
marginal, marginal, postmarginal, stigmal
vein lengths as 28:18:8.5:6; costal cell with
one complete row of setae and couple of
others distally; basal cell and vein bare;
VOLUME 91, NUMBER |
speculum open posteriorly; no distinct ad-
marginal setae. Petiole length 1.6 width
(8:5); lacking median carina. Gaster ovate,
length 1.7 x width (30:18); deep, height | x
maximum width (18:18).
Allotype, male. Color: Pattern similar to
holotype but antenna beyond basal half of
pedicel nonmetallic, brownish yellow; club
slightly darker on outer side. Body length
1.5 mm (critical point-dried). Antenna with
combined length of pedicel and flagellum
0.73 x head width (20.5:28.0); relative
lengths of scape, pedicel, anelli, F1-6, club
as 12:3:1:2:2:2:2:2:2:5: relative widths of F1,
F6, club as 2:3:3; funicular segments cylin-
drical. Maxillary palps slender. Petiole
length 2.0 x width. Gaster length 1.6 x width
(31:20).
Variation. The color of N. tzeltales varies
greatly over its geographic range. The fe-
male from Huetamo de Nunez is almost
black except for the dorsum of the meso-
soma which is dark green, and the antennal
flagellum is yellow beyond the basal half of
the pedicel. The male has the face violet,
the remainder of the head and mesosoma
green, the pedicel to F6 yellow, and the club
brown. Females of the series from Vene-
zuela have the body greenish black and the
antennal flagellum brown. In the males, one
paratype has antennal coloration like that
of the allotype, while the other has the fla-
gellum brown on the outer face and pale on
the inner face. The males from Venezuela
also have the tip of the middle tibia dark-
ened. The Venezuela series was collected
into alcohol and then critical point-dried.
This process can sometimes alter the colors
of specimens. Whether the color differences
in N. tzeltales are caused by environmental
influences or genetic differences needs study.
Discussion. Unique features of N. tzel-
tales are the bare basal cell and vein (Fig.
6). This species most closely resembles N.
bidentatus in having metallic legs and the
anterior margin of the clypeus produced;
although in N. tze/tales (Fig. 7), the clypeus
does not have the median emargination
121
present in N. bidentatus (Fig. 1). The two
species are reliably distinguished by the
characters given in the key.
Etymology. The specific name of this
species comes from the Tzeltales Indians
who live in the area of Mexico around San
Cristobal de las Casas.
Biology. The host(s) of this species is un-
known.
Type material. The holotype (CNC) is
from San Cristobal de las Casas, Mexico,
and was collected 1-3 June 1969. The al-
lotype male and 8 female and 2 male para-
types (USNM) were collected in Venezuela,
10 km north of Biscucuy, on 9 June 1981,
by E. E. Grissell, while sweeping Baccharis
(Compositae). Ten female and | male para-
types were collected as follows (CNC,
USNM): Costa Rica. Cartago (1500 m),
VIII-1980, 1 9; Heredia, 10-VIII-1975, 2
2. Mexico. CHIAPAS: San Cristobal de las
Casas, 27-1V-1969, 2 2, 12-VI-1969, 1 9.
MICHOACAN: Huetamo de Nunez, 7: III:
1972, 1 2, 1 6. QUERETARO: 10 mi. E San
Juan Del Rio, 30-VII-1954, 2 ¢. Guate-
mala. GUATEMALA: Guatemala City, IX-
LOS 9S lS:
Notoglyptus virescens Masi
Notoglyptus virescens Masi, 1917: 181-183.
Gahan and Fagan, 1923: 98. Boucek,
1961: 67. Graham, 1969: 140. Boucek,
1976: 15 (synonymy). Boucek, Subba Rao,
and Farooqi, 1978: 448. Farooqi and
Subba Rao, 1986: 295. Holotype, 8,
BMNH Hym. 5.873 (examined).
Notoglyptus niger Masi, 1917: 181. Erdés,
1948: 38. Boucek, 1961: 67. Peck et al.,
1964: 36. Boucek, 1976: 15 (synonymy).
Farooqi and Menon, 1973: 55. Boucek,
1977: 46. Dzhanokmen, 1978: 80. Bou-
cek et al., 1978: 448.
The following description of N. virescens
is based on a pair of specimens on loan to
me from the BMNH.
Female. Description. Color: Head, meso-
soma, petiole dark green; gaster brown. An-
tenna with scape yellow, slightly more brown
122
apically; remainder brown, pedicel pale
ventrally. Mandible yellow, teeth reddish
brown. Maxillary palps yellow. Legs yellow.
Wing veins yellowish brown.
Sculpture: Clypeus smooth, remainder of
head delicately alveolate; mesoscutum
roughly alveolate medially, becoming co-
riaceous laterally; scutellum coriaceous;
frenum, dorsellum smooth; median panels
of propodeum alveolate, sculpturing weak
in center of panels; petiole alveolate; gaster
smooth.
Structure: Body length 1.3 mm. Head
ovate in anterior view, width 1.3 x height
(23.5:18.0), 2.0 length (23.5:11.5); clyp-
eus separated from face by obscure sulcus,
anterior margin nearly straight, slightly re-
flexed; antennal scrobes shallow; genal con-
cavity extending ' way to eye; eye height
1.4 length (11:8), 2.8 malar length (11:
4), length 2.7 x temple length (8:3); ratio of
MOD, OOL, POL, LOL as 2.0:4.5:6.0:3.0.
Antenna with torulus 1x own diameter
above LOcL; combined length of pedicel
and flagellum 1.1 x head width (25.0:23.5):
relative lengths of scape, pedicel, anelli, Fl—
6, sclubvas -11:0:3.0; 10:210:2'522-5:2:5:225:
2.5:7.0; relative widths of Fl, F6, club as
1.75:2.0:2.0; anelli subequal in size; micro-
pilosity in line down ventral side of terminal
two segments of club. Mesosoma length
1.5 width (28:19); horizontal collar de-
veloped, anterior edge carinate; scutellum
with distinct discal fovea, scutoscutellar sul-
cus foveolate medially; dorsellum length '/
frenal length; propodeum with spiracles cir-
cular, 1.5 own diameter from anterior
margin of propodeum; nucha not bordered
by carina anteriorly. Fore wing length 2.5 x
width (57:23); ratio of submarginal, mar-
ginal, postmarginal, stigmal vein lengths as
20:12:9:5; costal cell with | complete setal
row basally and a couple others apically;
basal cell with apical quarter setate; spe-
culum closed posteriorly; distinct admar-
ginal setae present. Petiole length 1.2 x width
(5:4); with weak median carina. Gaster
length 1.3 width (21:16); height 1.1 x
width (17:16).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Male. Color: Similar to female except face
with face bluish, dorsum of mesosoma
greenish. Structure: Body length 1.0 mm.
Antenna with combined length of pedicel
and flagellum 1.5 x head width (31:21); rel-
ative lengths of scape, pedicel, anelli, F1—6,
club as 10.0:2.5:1.0:3.0:3.0:3.0:3.0:3.5:3.0:
8.0; relative widths of Fl, F6, club as 2:2:
2. Maxillary palps slender. Gaster length
1.5 width (27:18).
Variation. The male from Ontario is a
little larger (body length 1.3 mm), its scape
is mostly brown, its legs are more amber,
and its petiole lacks the median carina; but
agrees otherwise with the above description.
Discussion. Unique features possessed by
this species are the fovea on the disc of the
scutellum and the club having a line of mi-
cropilosity down the ventral side. This
species 1s intermediate in morphology with-
in the genus since it has the developed hor-
izontal pronotal collar like N. bidentatus and
N. tzeltales, but has the straight clypeal mar-
gin, the toruli located above the LOcL, the
antennal club with micropilosity, yellow legs,
admarginal setae, and a median carina on
the petiole like N. /uteicrus and N. nesiotes.
Distribution. This species is found from
southern Europe to India, Japan, and south-
ern Africa (Boucek 1976, 1977). There is a
male in the Canadian National Collection
from Mount Pakenham, Ontario.
Material Examined (BMNH, CNC). In-
dia. Delhi, IMRI, 1-IV-1977, (on grass) 1
2. Zimbabwe. Salisbury, | ¢. Japan. KYO-
TO: Kibune, 6-VIII-1980, 1 ¢@. Canada.
ONTARIO: Mount Pakenham, 11-VI-
1969, 1 6.
ACKNOWLEDGMENTS
I wish to thank the following persons for
the loan of material: M. Favreau, American
Museum of Natural History (AMNH), New
York, NY; Dr. J. S. Noyes, British Museum
of Natural History (BMNH), London; Dr.
G. P. Gibson, Canadian National Collec-
tion (CNC), Ottawa, ON; Dr. W. E. La-
Berge, Illinois Natural History Survey,
Champaign, IL; Dr. G. W. Byers, Snow En-
VOLUME 91, NUMBER 1
tomological Collection (SEC), Lawrence, KS;
and Dr. E. E. Grissell, United States Na-
tional Museum (USNM), Washington, D.C.
I would like to express my appreciation to
Shirely McClellan and Drs. W. E. LaBerge
and George Godfrey (INHS), and two anon-
ymous readers for reviewing this paper. I
would like to thank Drs. W. E. LaBerge and
E. E. Grissell for help and encouragement,
J. Sherrod (INHS) for assistance with the
illustrations, the staff at the Center of Elec-
tron Microscopy at the University of Illinois
at Urbana-Champaign for use of the facil-
ities, and Molly Scott (INHS) for preparing
the plates.
LITERATURE CITED
Boucek, Z. 1961. Beitrage zur Kenntnis der Ptero-
maliden-fauna von Mitteleuropa, mit Beschrei-
bungen neuer arten und Gattungen (Hymenop-
tera). Acta Entomol. Mus. Natn. Pragae 34: 55-
95.
1976. African Pteromalidae (Hymenoptera):
New taxa, synonymies and combinations. J. Ento-
mol. Soc. South. Afr. 39: 9-31.
1977. A faunistic review of the Yugoslavian
Chalcidoidea (Parasitic Hymenoptera). Acta
Entomol. Jugosl. 13 (Suppl.): 1-145.
Boucek, Z., B. R. Subba Rao, and S. I. Farooqi. 1978.
A preliminary review of Pteromalidae (Hymenop-
tera) of India and adjacent countries. Orient. In-
sects 12: 433-468.
Dzhanokmen, K. A. 1978. [Identification of the 1n-
sects of the European part of the USSR. Vol. 3.
Hymenoptera. Second part. Pteromalidae.] Opre-
deliteli Faune USSR, No. 120: 57-228.
Erdos, J. 1948. Additamenta ad cognitionem faunae
Chalcidoidarum (Hymenoptera) in alveo Carpa-
thorum. IV. 11. Pteromalidae. Frag. Faun. Hung.
11: 36-51.
Farooqi, S. l.and M.G. R. Menon. 1973. New com-
bination proposed for a pteromalid species Oxy-
sychus sauteri (Masi) comb. n. (= Dinarmus sau-
teri Masi). Entomol. Newslet. 3: 55.
Farooqi, S. I. and B. R. Subba Rao. 1985. Family
Pteromalidae, pp. 254-263. Jn Subba Rao, B. R.
and Hayat, M., eds., The Chalcidoidea (Insecta:
Hymenoptera) of India and the adjacent countries.
Part I. Reviews of families and keys to families
and genera. Orient. Insects 19: 163-310.
. 1986. Family Pteromalidae, pp. 279-306. Jn
Subba Rao, B. R. and Hayat, M., eds., The Chal-
cidoidea (Insecta: Hymenoptera) of India and the
adjacent countries. Part II. A catalogue of Chal-
cidoidea of India and the adjacent countries. Ori-
ent. Insects 20: 1-430.
Gahan, A. B. and M. Fagan. 1923. The type species
of the genera of Chalcidoidea or chalcid-flies. U.S.
Natn. Mus. Bull. 124: 1-173.
Graham, M. W.R. de V. 1969. The Pteromalidae of
northwestern Europe (Hymenoptera, Chalcidoi-
dea). Bull. Brit. Mus. (Nat. Hist.) Entomol. Suppl.
16: 1-908.
Heydon, S. L. 1988. The Sphegigasterini: A cladistic
analysis and generic classification with reviews of
selected genera (Hymenoptera: Pteromalidae).
Ph.D. Thesis, Univ. of Illinois at Urbana—Cham-
paign, Urbana. 167 pp.
Masi, L. 1917. Chalcididae of the Seychelles Islands.
Novit. Zool. 24: 121-230.
Peck, O., Z. Boucek, and A. Hoffer. 1964. Keys to
the Chalcidoidea of Czechoslovakia. Mem. Ento-
mol. Soc. Canada 34: 1-120.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, p. 124
Book REVIEW
Bees of the New Genus Ctenoceratina i”
Africa South of the Sahara (Hymenoptera:
Apoidea). By Howell V. Daly. University
of California Publications in Entomology
108: 1-69. February 1988. University of
California Press.
Once again, Howell Daly has prepared an
excellent taxonomic revision. This mono-
graph of the new genus Ctenoceratina Daly
and Moure (tribe Ceratinini) encompasses
13 species of small carpenter bees indige-
nous to subsaharan Africa. Included are a
new subgenus, Simioceratina Daly and
Moure, and three new species, Ctenocera-
tina malindiae Daly, C. paulyi Daly, and C.
rossi Daly. A key, synonymies, descriptions,
distributions (precise, but unfortunately
without locality maps), and some biological
data including parasites are provided.
The publication’s format provides a clear,
easily followed explanation of how and why
various taxonomic decisions were made.
Thus. the text will prove useful to both ex-
perienced and novice hymenopterists. The
high quality illustrations include SEM pho-
tographs (Figs. 1-8) and line drawings (Figs.
9-50). One minor complaint is that figure
numbering (particularly of facing plates pp.
64-65) is not consistently from left to right
or from top to bottom. The left to right
convention seems most easily followed.
Dr. Daly’s thorough approach to this study
is evidenced by his efforts to assess vari-
ability. Through rearing, careful observa-
tion of live material, and collection at select
localities, he documented striking allome-
tric differences among nest mates, and
among, individuals within local popula-
tions. (Such differences undoubtedly led
earlier taxonomists to distinguish some 55
species.) When large series were available,
an effort was also made to assess variation
between populations. I appreciated Dr. Da-
ly’s treatment of species displaying complex
geographic variation. Rather than designat-
ing formal subspecies, informal “forms”
were discussed for these polytypic species.
I also appreciated the clarity provided by
his “similar species” discussions contained
within the species descriptions.
A critical test of the key was never made
since our African material was still on loan
to Dr. Daly. However, the key was easily
read and the illustrations made interpreta-
tion appear straight forward. One point of
potential confusion that should be noted is
his use of the term “scale.” The genus name,
Ctenoceratina, was selected because of the
comb-like rows of enlarged setae (‘‘scales’’)
found on the apical edges of terga and sterna
2-4 and frequently 5. While not incorrect,
this usage might momentarily baffle the un-
suspecting bee worker who dives into the
key without previously reading the intro-
duction.
Reviewing this work whetted my appetite
for future investigations of Ctenoceratina.
We are indebted to Howell for providing
yet another valuable contribution.
Beth B. Norden, Department of Ento-
mology, Smithsonian Institution, National
Museum of Natural History—105, Wash-
ington, D.C. 20560.
—
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, p. 125
Book REVIEW
Control of Arthropod Pests of Livestock:
A Review of Technology. By Roger O.
Drummond, John E. George, and Sidney
E. Kunz. 1988. CRC Press, Inc., Boca Ra-
ton. Florida, 245 pp. Price US $125.00.
This book reviews a large number of pa-
pers published in the U.S. and Canada con-
cerning the materials and methods which
have been used to control arthropod pests
of livestock. It is principally a summary of
the history of chemical treatments. It is not
a manual of pest control techniques, nor is
it a treatise on the evolution and/or failures
of chemical insecticides or acaracides which
have been used against pests of livestock.
The book covers the entire spectrum of ar-
thropod pests of livestock. The material re-
viewed on the biology and significance of
each pest ranges from one to two pages and
the review of control technologies used
against each pest ranges from three to six-
teen pages.
The first 15 chapters are arranged by com-
modity, i.e. cattle, horses, sheep and goats,
swine and poultry, and each chapter is sub-
divided into sections on individual pests.
The biology and importance of each pest is
discussed briefly, followed by a two-to-sev-
en page review of control efforts, and each
chapter is summed up by an Overview and
Current Technology section. The overview
sections are well done and provide a concise
and valuable statement of current control
methods for each pest group.
Chapters 16 through 18 review the con-
trol efforts which have been used against
pests that breed in poultry manure, against
ticks and chiggers and against flies in live-
stock manure and around structures. These
chapters are well arranged but the five-to-
nine pages devoted to each subject neces-
sarily limit the number of papers and tech-
nologies reviewed.
Chapter 19 provides a 4-page review of
pest resistance to insecticides and is prob-
ably the most disappointing chapter in the
book, particularly in view of the emphasis
that the authors place on the historical usage
of chemicals as insecticides and acaracides.
Chapter 20 provides a nice review of the
major sterile insect release programs (6
pages) and chapter 21 reviews the status of
biocontrol technology (5 pages).
Chapter 22 provides an intriguing pre-
view of the possibilities of developing the
resistance of livestock to arthropod pests (5
pages).
There is a 4-page appendix which lists the
trade and chemical names of many of the
insecticides and acaracides mentioned in the
book, but no addresses are listed for the
manufacturers of the chemicals.
Although a large number of references are
listed for each pest, the list is, of necessity,
incomplete and even some bibliographies
(1.e. West, West and Peters) are not men-
tioned. There are no lists of the published
bibliographies or reviews of literature per-
taining to an individual pest or commodity
except for the sterile insect technique for
screwworms.
There are no charts, chronological lists of
control methods (except for cattle grubs), or
analyses of the histories of the development
of resistance of pests to chemicals.
In summary, this is an ambitious book
which meets the stated objectives of the au-
thors to provide a history of control tech-
nologies. It should be of interest to anyone
who is interested in the history of livestock
pest control methods and should be valu-
able as a background source to anyone who
is just entering the livestock pest control
field. In my opinion, however, it is grossly
overpriced at U.S. $125.00.
Lawrence G. Pickens, Livestock Insects
Laboratory, Livestock and Poultry Sciences
Institute, USDA-ARS, Beltsville, Maryland
20705.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 126-127
Book REVIEW
Ecological Methods with Particular Ref-
erence to the Study of Insect Populations.
T. R. E. Southwood. 1978. Second re-
vised edition. Chapman and Hall, Lon-
don, New York; xxiv + 524 pp. $32.50
Softcover.
This second edition, now available in pa-
perback, is a thoroughly updated version of
the first edition from 1966. Some chapters
have been completely or largely rewritten
and substantial new information has been
added. Most impressively, and vividly il-
lustrating the rapid growth of available eco-
logical information, close to a thousand new
references have been included. The book
presents a synthesis of ““methods that are
peculiar to the ecologist,” and especially
those aimed at the study of insect popula-
tions. The usage, however, is much broader
as techniques generally applied to the study
of earthworms, mollusks, nematodes,
planktonic organisms, and several non-in-
sect arthropods have been included. The ap-
proach is mainly syn-ecological and only
little attention has been paid to the habitat
although the importance of spatial and tem-
poral heterogeneity as well as predictability
is stressed. The first chapter is introductory
and discusses classification of population
estimates, errors and confidence, and chap-
ter 2 deals with the sampling program and
the description of population dispersion.
Chapters 3-8 are mainly aimed at the ques-
tion of how to obtain an absolute population
estimate and contain marking techniques,
capture-recapture methods, and sampling
procedures. Chapters 9-11 deal with de-
mographical aspects. In chapter 12 systems
analysis and modelling are briefly discussed,
and the realm of community ecology is
reached in chapter 13 which deals with di-
versity, species packing and habitat. The last
chapter discusses productivity and energy
budgets.
‘Ecological Methods’ spans the gap be-
tween suction traps made from homestyle
vacuum cleaners and, for example, the “‘lin-
ear dioristical [systems] model with alge-
braically defined transfer functions.” To at-
tain this, a lot of information has to be
presented in a fairly condensed state, but
Southwood manages to do this in a very
coherent and readable style.
Sampling techniques are discussed thor-
oughly and their strengths and weaknesses
with regard to their ease, the samples they
produce and the financial resources they de-
mand are stressed. Much attention is paid
to possible sources of errors, whether stem-
ming from the equipment proper or from
the heterogeneity and unpredictability of
living organisms or their habitats. Basic
mathematical and statistical theory is not
discussed at length but is still presented in
a very clear form and with ample references
for those interested in its derivation.
It is, perhaps, somewhat ironical that a
book devoted to the study of populations
does not provide an explicit discussion on
how to define a population as a spatiotem-
poral entity. Largely left unanswered are
questions like: How are geographical limits
defined?, which constituing features
emerge?, and, what are the temporal di-
mensions?
A decade has passed since the 1978 edi-
tion, and methods and theory in insect ecol-
ogy have kept evolving. Emerging fields as
tropical rainforest canopy ecology have de-
veloped largely new techniques, and much
new information is available on patterns of
population growth, e.g. how to predict in-
sect pest outbreaks using integrated trap-
ping with various combinations of phero-
mones, kairomones, colors, and host-mimic
lures. The need for precision and critical
cost/benefit considerations, however, re-
mains important, and it is still imperative
to be able to evaluate the limitations and
VOLUME 91, NUMBER |
possible biases of the techniques involved
in any study. Therefore, the book still stands
out as a must for anyone teaching the prac-
tical aspects of insect (or animal) ecology,
and as an extremely valuable manual for all
who need to investigate animal populations,
127
whether as a research ecologist, a conser-
vationist or an applied entomologist.
Thomas Pape, Zoological Museum,
Universitetsparken 15, DK-2100 Copenha-
gen, Denmark.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, p. 127
NOTE
Rediscovery of the Ant Gnamptogenys hartmani
(Hymenoptera: Formicidae) in Eastern Texas
Wheeler (1915. Bull. Amer. Mus. Nat.
Hist. 34: 390) described Gnamptogenys
hartmani from a single worker collected in
Huntsville, Walker Co., eastern Texas. This
record seemed anomalous for a genus oth-
erwise confined to tropical environments
(Brown. 1961. Psyche 68: 69). Brown (op
cit) suggested that the Texas record was
either a locality error or an adventive spec-
imen taken from bananas (it also occurs in
Honduras). There are no ecological data as-
sociated with the specimen. Brown (op cit)
concluded that the presence of Gnamptoge-
nys in Texas remained to be convincingly
demonstrated.
We have collected a second worker spec-
imen from Texas, Brazos Co., 10 km N of
Kurten, about 60 km west of the type lo-
cality. The specimen was collected in a pit-
fall trap in an open grassy area located 30
meters from dense post oak (Quercus spp.)
woods on 5 Aug. 1987 (trap was in field for
24 h). Although we did extensive pitfall
trapping, baiting, general collecting and
berlese funnel sampling of the litter in the
study area, we have not collected any ad-
ditional specimens. We have not collected
this species in several other similar areas in
eastern Texas, using similar methods. This
specimen was collected in a disturbed area
(mowed and occasionally grazed by cattle),
but the ant was definitely not associated with
any introduced products. These data, in ad-
dition to the collection of this species in
Louisiana (Echols. 1964. Ann. Entomol. Soc.
Amer. 57: 137) clearly demonstrate that this
species is a rarely collected member of the
fauna of the United States. This species preys
on the brood of the ant Trachymyrmex sep-
tentrionalis (McCook) (Echols op cit), which
is common in the area. The ant, Afta texana
(Buckley), 1s also common and may serve
as a prey species.
It is gratifying that this interesting ele-
ment of our native ant fauna has withstood
the onslaught of the spread of the imported
fire ant (Solenopsis invicta Buren), which
eliminates many native ant species. The ant
was collected in an area of sandy soil where
the density of the imported fire ant is rel-
atively low (115 mounds/ha). The specimen
is in the collection of WPM, field number
9676.
Alex Mintzer suggested the study area, O.
L. Tate gave us permission to use the area,
Mark Strain, Shelley Stonecipher and Cecil
Pinder assisted with the field work. The re-
search was supported by the Texas Depart-
ment of Agriculture. This report is ap-
proved as Number TA 23642 of the Texas
Agricultural Experiment Station.
Wiliam P. MacKay and S. Bradleigh
Vinson, Department of Entomology, Texas
A&M University, College Station, Texas
77843.
PROC. ENTOMOL. SOC. WASH.
91(1), 1989, pp. 128-132
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of
WASHINGTON
Volume 90
OFFICERS FOR THE YEAR 1988
President F. Eugene Wood
President-Elect F. Christian Thompson
Recording Secretary Richard G. Robbins
Corresponding Secretary John M. Kingsolver
Treasurer Norman E. Woodley
Editor Hiram G. Larew
Custodian Anne M. Wieber
Program Chairman Warren E. Steiner
Membership Chairman Geoffrey B. White
Delegate to The Washington Academy of Sciences Manya B. Stoetzel
Published by the Society
WASHINGTON, D.C.
1988
TABLE OF CONTENTS, VOLUME 90
ARTICLES
BAKER, G. T.—See CHAN, W. P.
BAO, N.—See ROBINSON, W. H.
BARNES, J. K.—Notes on the biology and immature stages of Poecilographa decora (Loew)
(Diptera: Sclomyzidae) ... ie eas SR, HE aoe TEP eee te 474
BENNETT, F. D.—See PALMER, W. A.
BIN, F.—See JOHNSON, N. F.
BOLDT, P. E., W. WOODS, and T. O. ROBBINS—Phytophagous insect fauna of Baccharis
sarothroides Gray (Asteraceae) in Arizona and New Mexico ..............-..00+++e000- 207
BURGER, J. F.—A new genus and two new species of Pangoniini (Diptera: Tabanidae) of
zoogeographic interest from Sabah, Malaysia ............... ooh abi ratictey Necteratay Sher sabe eR
CARLSON, R. W.—See DREA, J. J.
CARPENTER, J. M. and M. C. DAY—Nomenclatural notes on Polistinae (Hymenoptera:
VOLUME 91, NUMBER 1
CAVE, R. D.—See NORTON, R. A.
CHAN, W. P., G. T. BAKER, and M. M. ELLSBURY —Sensilla on the larvae of four Hypera
Species:(Coleopteras Curculionidae)! Ma. as.2s as see sa aaiandendltea) pee e eee eene enous
CHEMSAK, J. A. and C. FELLER—New species of Gamniivatac fom Twin Cays, Belize
(Coleoptera) tee create Sete ice re tte ee ne Ne nnn ccs este Gt ee Ores
CHESSER, R. K.—See VAN DEN BUSSCHE, R. A.
CLEMENT, S. L. and T. MIMMOCCHI— Occurrence of selected flower head insects of Cen-
taurea solstitialisin Italyand Greece... 02.0 en ees cae eea seas wees snnecass
CLEMENT, S. L., T. MIMMOCCHI, R. SOBHIAN, and P. DUNN—Host Seahah) ofadule
Eustenopus hirtus (Waltl) (Coleoptera: Curculionidae), a potential biological control agent of
yellow starthistle, Centaurea solstitialis L. (Asteraceae, Cardueae) .............
DAVIS, D. R. and E. G. MILSTREY — Description and biology of Acrolophus phalcen ives
idoptera: Tineidae), a new moth commensal from gopher tortoise burrows in Florida . .
DAY, M. C.—See CARPENTER, J. M.
DIATLOFF, G. and W. A. PALMER—The host specificity and biology of Aristotelia ivae
Busch (Gelechiidae) and Lorita baccharivora Pogue (Tortricidae), two microlepidoptera se-
lected as biological control agents for Baccharis halimfolia (Asteraceae) in Australia ......
DREA, J. J. and R. W. CARLSON — Establishment of Cybocephalus sp. (Coleoptera: Nitidu-
lidae) from Korea on Unaspis euonymi (Homoptera: Diaspididae) in the eastern United
SS TALES Me ehyn saiegs ys ere ca oy ase een cys ehastoreentearaxeGter oc is tieer nich orca ia Nes atlas eraaiara te eee Mapa Stee
DROOZ, A. T. and H. H. NEUNZIG—Notes on the biology of two Phycitines (Lepidoptera:
Pyralidae) associated with Toumeyella pini (Homoptera: Coccidae) on pine ..............
DUNN, P.—See CLEMENT, S. L.
ELLSBURY, M. M.—See CHAN, W. P.
EMERSON, K. C. and R. D. PRICE—A new species of Haematomyzus (Mallophaga: Hae-
matomyzidae) off the bushpig, Potamochoerus porcus, from Ethiopia, with comments on lice
FOUN GON GpIGS ears ee teleaie ces oer ite cae atone stern tk RE Sereno ete tas ain
FELLER, C.—See CHEMSAK, J. A.
GITTINS, A. R.—See TRIPLETT, D. C.
GOEDEN, R. D.—Gall formation by the capitulum-infesting fruit fly, Tephritis stigmatica
(Di pterassle pi mit dae) peer ceere anys 2 eee ee Me ae Se ee See ere ke eee
GOTT, K. M.—See NEAL, J. W., Jr.
GRISSELL, E. E.—See HEYDON, S. L.
HALSTEAD, J. A.—Belaspidia longicauda, new species, the first Nearctic Belaspidia (Hy-
menoptera: Chalcididae) ............. bens Se SORE ae
HANNA, R. B.—See LANE, M. A.
HANSON, P. E. and J. C. MILLER—Notes on the biology of Caenocephus aldrichi Bradley
(ivimenopteras Gephidae)i mes: sacrsce sian = ce ateniere ite cic caectace eins cuits Gas qgrmicuets
HANSSON, C.—A revision of the genus Me. ot oeRAIS anid a review of det genus ‘Grain
(Eivmenopteras lop hidae)) cen emis estes ici cic nreiaeiier n= one ree crete oa
HARMON, J. D. and M. H. ROSS— Effects of aalathion and diazinon exposure on feniale
German cockroaches (Dictyoptera: Blattellidae) and their oothecae ............
HEYDON, S. L.—A review of the Nearctic species of Cryptoprymna Forster, with the decane
tion of a new genus, Polstonia (Hymenoptera: Pteromalidae) ...........................
HEYDON, S. L. and E. E. GRISSELL—A review of Nearctic Mes rismus Walker and Toxeuma
Walker (Hymenoptera: Chalcidoidea: Pteromalidae) .......... Cer igioat SuoaeActacr aero eee
HOEBEKE, E. R.—See WHEELER, A. G., Jr.
HUANG, Y. M.—Aedes (Stegomyia) josiahae, a new species of the simpsoni subgroup (Diptera:
Culicidae) . RA Tere
JOHNSON, N. F. ec. of Australian ‘Telenominae (Hiymenopter: Scelionidae) of A. Pp.
Dodd and A. A. Girault Ne a Rie eC ee eee rere
JOHNSON, N. F. and F. BIN— Tele. OTRAS egies EVER BTETe: Sedionidae) associnied with
the eggs of Zygaenidae (Lepidoptera)
269
179
47
501
164
458
307
44
338
7
87
130 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
JOHNSON, W. T.—See SPECKER, D. R.
KONDRATIEFF, B. C.—See STARK, B. P.
KURCZEWSKI, E. J.—See KURCZEWSKI, F. E.
KURCZEWSKI, F. E., E. J. KURCZEWSKI, and M. G. SPOFFORD—Nesting behavior of
Aporinellus wheeleri Bequaert and A. taeniolatus (Dalla Torre) (Hymenoptera: Pompilidae) 294
KURCZEWSKI, F. E.—See LANE, M. A.
LANE, M. A., F. E. KURCZEWSKI, and R. B. HANNA—Antennal sensilla and setae of
Evagetes parvus: (Hymenoptera; Pompilidae)) trae a ccie « cassieisteisiec cle acess edwin 428
MacDONALD, J. F.—New synonyms pertaining to Chelifera and generic key for North Amer-
ican Hemerodromiunae!(DipterasEmpididae)i- mere. ss: > 2 ee ceeeie eee celia haters 98
MALDONADO-CAPRILES, J.—See SANTIAGO-BLAY, J. A.
MARSHALL, S. A. and D. J.S. MONTAGNES— Ceroptera longicauda, a second North Amer-
ican species in the kleptoparasitic genus Ceroptera Macquart (Diptera: Sphaeroceridae) ... 189
MATHIS, W. N.—First record of the genus Procanace Hendel from North America, with the
descriptioniofianew:species:(DipteraxCanacidae) wath. certerct-clatsm cee ose «te ceils cies clea 329
MATHIS, W. N.—First record of the shore-fly genus Placopsidella Kertész from North America
(DipterazEphydridae)) «<:.5 c<cipcyhe Godctanetoeres oar atta aie dich eoa cits eee aie ae weerrerameue ale 334
MATHIS, W. N. and T. ZATWARNICKI-—Studies of the systematics of the shore-fly tribe
Dagini (DipterasEphydridae)| +10 ers sake shel sersapverd ah eee aici Stata Sasteyacid een. teeies 106
MILLER, J. C.—See HANSON, P. E.
MILSTREY, E. F.—See DAVIS, D. R.
MIMMOCCHI, R.—See CLEMENT, S. L.
MONTAGNES, D. J. S.—See MARSHALL, S. A.
NAKAHARA, S.—Generic reassignments of North American species currently assigned to the
genus Sericothrips Haliday (Thysanoptera: Thripidae) ...................-.e eee e eee eee 480
NEAL, J. W., Jr.—Unusual oviposition behavior on evergreen azalea by the Andromeda lace
bug Stephanitis takeyai (Drake and Maa) (Heteroptera: Tingidae) ...................... 52;
NEAL, J. W., Jr. and K. M. GOTT—Evidence for multivoltinism in Prodiplosis platani Gagné
(Diptera: Cecidomyiidae), a leaf curl midge of American sycamore ..................... 201
NEUNZIG, H. H.—See DROOZ, A. T.
NORTON, R. A., W. C. WELBOURN, and R. D. CAVE—First records of Erythraeidae
parasitic on oribatid mites (Acari, Prostigmata: Acari, Orbatida) ...................00.. 407
OSWALD, J. D.—A review of the South Pacific genus Austromegalomus Esben-Petersen (Neu-
roptera: Hemerobiidae) with a description of a new species from Rapa .................. 55
PALMER, W. A. and F. D. BENNETT—The phytophagous insect fauna associated with
Baccharisjnalimifolia W. intheseastern United'States 2.02... aca8).d4-ce ects hoc eee ok 216
PALMER, W. A.—See DIATLOFF, G.
PETERSON, B. V., M. VARGAS, V., and J. RAMIREZ-PEREZ—Simulium (Hemicnetha)
hieroglyphicum (Diptera: Simuliidae), a new black fly species from Costa Rica ........... 76
PETERSON, B. V.—See RAMIREZ-PEREZ, J.
POGUE, M. G.—Revision of the genus Lorita Busck (Lepidoptera: Tortricidae: Cochylini),
witht ardescniptionnofaynew SPECIES: s,s saa) aes bain stasitbecens, Saaee NIEMAN = eee cuter setae eae 440
PRICE, R. D.—See EMERSON, K. C.
QUICKE, D. L. J.—Digonogastra: The correct name for Nearctic [phiaulax of authors (Hy-
menoptera: Braconidae) ....... age, Sues RUM Nee Utha cd Marotenedn cd okt erat ai ccesd aoe acer naar eran 196
QUICKE, D. L. J.—See WHARTON, R. A.
RAMIREZ-PEREZ, J., B. V. PETERSON, and M. VARGAS, V.—Mayacnephia salasi (Dip-
tera: Simuliidae), a new black fly species from Costa Rica ........60..0ccs0eeesecee ees 66
RAMIREZ-PEREZ, J.—See PETERSON, B. V.
REEVES, R. M.—Distribution and habitat comparisons for Carabodes collected from conifer
branches with descriptions of brevis Banks and higginsi n. sp. (Acari: Orabatida: Cara-
{206 ) Cs F:1) e ae SR SE scre rts teh eee eee ECs ORR Od AiR A CN ey Oh 373
ROBBINS, R. K.—Comparative morphology of the butterfly foreleg coxa and trochanter (Lep-
idoptera)andlitssystéematics implications) 22. <.2¢00< act owes ngs wile ome pulses siete stelerste 133
VOLUME 91, NUMBER 1 131
ROBBINS, R. K.— Male foretarsal variation in Lycaenidae and Riodinidae, and the systematic
placement of Styx infernalis (Lepidoptera) ............00 00000 eee 356
ROBBINS, T. O.—See BOLDT, P. E.
ROBINSON, W. H. and N. BAO—The pest status of Periplaneta fuliginosa (Serville) (Dic-
Ly OpterasBlatidac) imi Chin ayer se etarevysreeyeleie eucnsereerecreeketena = aie cletele slcle aiaatsiacvemelalcrs 401
ROSS, M. H.—See HARMON, J. D.
SANTIAGO-BLAY, J. A. and J. MALDONADO-CAPRILES— Observations on the true bugs
Emesa tenerrima, a possible spider mimic, and Ghilianella borincana (Hemiptera: Redu-
wildae:JEMmesinac) Mom PUCKtOMRIGO) a2. .ccus anes sss bose ccee sey se vent ase a oe Reni Ayah 369
SITES, R. W.—See ZACK, R. S.
SOBHIAN, R.—See CLEMENT, S. L.
SPECKER, D. R. and W. T. JOHNSON—Biology and immature stages of the rhododendron
gall midge, Clinodiplosis rhododendri Felt (Diptera: Cecidomyiidae) ................... 343
SPOFFORD, M. G.—See KURCZEWSKI, F. E.
STAINES, C. L., Jr.—A review of the species of Acritispa Uhmann (Coleoptera: Chrysomelidae:
FATS Pinae)) seve caer eeces ceeicle ste et as rors eset cece re mons Mme eee rare Sacineee eunse eee Henne nesenttes 193
STARK, B. P., S. W. SZCZYTKO, and B. C. KONDRATIEFF—The Cultus decisus complex
of eastern North America (Plecoptera: Perlodidae) ................. 00. .c cece eee eee eee 91
STRICKMAN, D.—Redescription of the holotype of Culex (Culex) peus Speiser and taxonomy
of Culex (Culex) stigmatosoma Dyar and Thriambus Dyar (Diptera: Culicidae) ....... 484
SZCZYTKO, S. W.—See STARK, B. P.
TRIPLETT, D. C. and A. R. GITTINS—Nesting, mating and foraging habits of Melissodes
(Melissodes) tepida tepida Cresson in Idaho (Hymenoptera: Anthophoridae) ............. 462
TURNER, W. J.—Lectotype designation for E mpis chichimeca Wheeler and Melander (Diptera:
Ema pididae) yqeegarreruart atic aecnche ce astd cetera cetcicecasiteck Tele MeN ee Goch een Geto ee 62
VAN DEN BUSSCHE, R. A., M. R. WILLIG, R. K. CHESSER, and R. B. WAIDE—Genetic
variation and systematics of four taxa of neotropical walking sticks (Phasmatodea: Phas-
TM ATIGAG) aren, eee eis etre er reat oa eee ert RTO ETN EISISLE cad Ruane eel tes aut beereeae ee eaeiee 422
M. VARGAS, V.—See PETERSON, B. V.
M. VARGAS, V.—See RAMIREZ-PEREZ, J.
WAIDE, R. B.—See VAN DEN BUSSCHE, R. A.
WELBOURN, W. C.—See NORTON, R. A.
WHARTON, R. A. and D. L. J. QUICKE—A new species of Bracon (Hymenoptera: Bracon-
idae) parasitic on Eoreuma loftini (Dyar) (Lepidoptera: Pyralidae) ...................... 288
WHEELER, A. G., Jr., and E. R. HOEBEKE—Apterona helix (Lepidoptera: Psychidae), a
Palearctic bagworm moth in North America: New distribution records, seasonal history, and
NOS tpPlAN tS sere ewes crete eres sete ene ae tents aetna aaah ete erase SN ATaard “cetronescte 20
WHITFIELD, J. B.—Taxonomic notes on Rhyssalini and Rhysipolini (Hymenoptera: Bracon-
idae) with)first, Nearctic records. of three’eenera, 232. .ce. -sceesa ees dee eee cdctneene nue 471
WILKERSON, R. C.—Notes and redescriptions of some Anopheles series Arribalzagia holo-
types (Diptera: Culicidae) in the British Museum (Natural History) ..................... 411
WILLIG, M. R.—See VAN DEN BUSSCHE, R. A.
WOODS, W.—See BOLDT, P. E.
ZACK, R. S. and R. W. SITES—A new species of Donaceus Cresson (Diptera: Ephydridae)
licohe TI ETERS EL rs eka acs Ais oo a Sean te ie 5 Meine tae Ole a ey ee en Aa ee 101
ZATWARNICKI, T.—See MATHIS, W. N.
NOTES
BRUNNER, J. F.—See RATHMAN, R. J.
DAVIDSON, R. L.—Note on the habitat of Prerostichus (Pseudomaseus) tenuis (Casey) (Co-
leoptera: Carabidae) with six new state records ...................0.. Prete aoe A . 260
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
GAGNE, R. J.—A polynomial Riley name in Cecidomyiidae (Diptera) and implications of
Suchinames:for:Cynipidae (Elymenoptera)s «cms «.- peteletsderetertels erey lamin ieee 394
HALSTEAD, J. A.—A gynandromorph of Hockeria rubra (Ashmead) (Hymenoptera: Chalcid-
110 Vo) ee ee PR Rr AR a Sh OE CR A ee pant aa ae Hees eh WR ichigst 258
HULBERT, S. J.—See RATHMAN, R. J.
McCAFFERTY, W. P.—Neotype designation for Raptoheptagenia cruentata (Walsh) (Ephem-
eroptéranHeptageniiGdae) erences e crerstac = a Share eater eee aN oon eee eed ee ee 97
NAKAHARA, S.—A new synonym and revised status in Apterothrips (Thysanoptera: Thrip-
106 £212) le CRS er Ro SPS ony Gon aa aN Pe Pim es AS oe i Ge ie Got os Bnd etn o thn 3 'o Og 508
PAVUK, D. M. and R. N. WILLIAMS—A sesiid host record for Pterocormus chasmodops
(Hymenopteraziichneumonidae)}. ..:.acclaa-tereeretd vce ester tecmceslore seamen aretetoteketeese erates area 105
PLAKIDAS, J. D.—The newly discovered spring crown gall of Asphondylia rudbeckiaecon-
spicua (Diptera: Cecidomyiidae) on Rudbeckia laciniata (Asteraceae) in Pennsylvania .... 393
RATHMAN, R. J., J. F. BRUNNER, and S. J. HULBERT—Feeding by Medetera species
(Diptera: Dolichopodidae) on aphids and eriophyid mites on apple, Ma/us domestica (Ro-
SACEAC) ar eters gS vasane: o: arbi: aratebneicelavaterbeatleaeane Ra Wlap cud rasta Ree Mia Tree TOE oe ene eee $10
STOETZEL, M. B.—See VOEGTLIN, Dz.
VOEGTLIN, D. and M. B. STOETZEL—Hyadaphis tataricae (Homoptera: Aphididae): 10
years;afteritsintroductionsinto' North America’? 20.4206 .n\e ste et eal eeel-ie eee eee eeiers 256
WILLIAMS, R. N.—See PAVUK, D. M.
BOOK REVIEWS
AMRINE ). WsJrs—Bees and UNCIMRCEPers acacia cas tise eons csetieis aiarsieis ecient elerenetee 262
ARNAUD, P. H., Jr.—The Bombyliidae of Deep Canyon .........0 0c. ccc eect ene ee 124
GARROLE.: F.—The Behavioral E-COloeyiOfp Ants, occ ccc. tie ctece a, setae sage weseree ainregetererandte 396
FLINT, O. S., Jr.— Revision of the Caddisfly Genus Psilotreta (Trichoptera: Odontoceridae) .._ 400
FOOTE, B. A.—Manual of Nearctic Diptera, Volume 2 .... 0... ccc ccc ccc cece eee eees 125
HODGES, R. W.—Sphingidae Mundi (Hawkmoths of the World)... .....0000 0000000 es 127
KERANS Si Es— Une dxOdid TiCKS OPO SANAQY sleucc:s cass orsyestt sesershe ola oheieuasele waraenistule teres cater 398
KINGSOLVER, J. M.— The Metallic Wood-boring Beetles of Canada and Alaska: Coleoptera:
BUDE OS tIG ACI sce. oo. ac os agert MeO ETS Tes Sian eee STR a ee EET ER CEE TE TOC ete 397
NORTON, R. A. Be eaien Mitescand Human Welfare’ 2. on cs ccna sinicen iis oct eet 524
SMILEY, R. L.—Historical Perspective and Current World Status of the Tomato Russet Mite
(Acari: Eriophyidae) Brack, aah NS RR oh ner asain Ate 8: wee ape mtd 3 BN ecm 122
OBITUARY
KROMBEIN, K. V. and P. M. MARSH—Carl Frederick William Muesebeck, 1894-1987 .... 513
MISCELLANEOUS
INSTRUCTION TO AUTHORS ROC HET EC Gee On Eee ee 129
NEW MEMBERS FOR 1987 . ; Thar Sy dtataa ation ratars bos GAS LeTORSeie SSIS IE CTO SEERA 263
SOCIETY MEEMINGS a saecpsete nets raed ietienis evi seemed Seer Ones Be rae eae 264, 529
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CONTENTS
(Continued from front cover)
WEBB, D. W. and M. E. IRWIN—Genera Ataenogera and Phycus in the New World (Diptera:
Therevidae® Phycinbe)) itary oe doors sesh ngvicdale tdsals Olam scene ba Ret een ts tae ter men terres 3a
NOTES
MAacKAY, W. P. and S. B. VINSON—Rediscovery of the ant Gnamptogenys hartmani (Hy-
menoptera: Formicidae) aniasterm: MEXaS) 0. «es rersra sti sane ue uc oneleis h-Pa te aad eee oy 127
WILSON, N. and W. W. BAKER—/xodes downsi (Acari: Ixodidae) from Peru ............. 54
BOOK REVIEWS
NORDEN, B. B.—Bees of the New Genus Ctenoceratinia in Africa South of the Sahara (Hy-
MeNOPLEVAHADOIAEA), wk dos oh AES ¢ decile NAT ORION Paice otha chee cei eee cpa eet enone 124
PAPE, T.—Ecological Methods with Particular Reference to the Study of Insect Populations,
SecondiRevised! EGitiom-> 5. % 45 ccls, wire om teat oe toesteteretete eed ercekel ries cic tanche aie Clea nee eeer a epee 126
PICKENS, L. G.— Control of Arthropod Pests of Livestock: A Review of Technology ......... 125
TABLE OF GONTENTIS,, VOLUMES SON Ness ete ctctsta vi oteiathebtnt ito kele sircte lel can atekcnrel < h ainiat etal tata 128
IL. 91 APRIL 1989 NO. 2
(ISSN 0013-8797)
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON
PUBLISHED
UAR--ER
OaTHSONE,
CONTENTS | ‘#2 Qeseee
APPEL, A. G. and R. C. SPONSLER—Water and temperature glatiog $ofshe-s iri tive
lophagous cockroach Cryptocercus punctulatus Scudder (Dictyopteraefryptocercidaey ... 153
ASQUITH, A. and D. ADAMSKI—Description of the predaceous larva of Pseudogaurax sig-
MAES eGew (I pteras GHIOLOpldae) pam. se daie,2 cin ae te a eis hates yokes clays aes S 2) le, ees le oho ears 185
DIETRICH, C. H.—Surface sculpturing of the abdominal integument of Megobracidae and other
AMICHEN OM MUNCH A ELOMODIEKA) I azrrir a arte gcpeyecctare) sfere stn spots ox eee ers MP ers peel oe Gaahia 143
GAGNE, R. J. and P. E. BOLDT—A new species of Neolasioptera (Diptera: Cecidomyiidae)
from Baccharis (Asteraceae) in southern United States and the Dominican Republic .... 169
GOEDEN, R. D.—Host plants of Neaspilota in California (Diptera: Tephritidae) ........... 164
HENRY, C. S.—The unique purring song of Chrysoperla comanche (Banks), a green lacewing
of the rufilabris species group (Neuroptera: Chrysopidae) ......................2220... 133
KELLEY, R. W.—New species of micro-caddisflies (Trichoptera: Hydroptilidae) from New
ealedOniass Vata CURanGUni ine op eee een ees neh Me AT anole aaah lero hae desea 190
KNISLEY, C. B., D. L. REEVES, and G. T. STEPHENS—Behavior and development of the
wasp Pterombrus rufiventris hyalinatus Krombein (Hymenoptera: Tiphiidae), a parasite of
larvaletiper beeties (Coleoptera: Cicindelidae)) 0.0 estore wet tne wan aicvers fies) sles Soares ct ayers 179
LASALLE, J.— Notes on the genus Zagrammosoma (Hymenoptera: Eulophidae) with description
DAMIEN DECICS Ae met: Parity si Malia take meio Sauer gi elcnis “Saft lavolefergrot a Sahat ots sual oe eke yh Sas 230
MACKAY, W. P. and S. B. VINSON—Two new ants of the genus Solenopsis (Diplorhoptrum)
fromeastern Texas (Hymenoptera: Formicidae) .. 2... 0.5.0 cde cca e ect e ese s st enenee 175
MILLER, T. D.—First Nearctic record of the genus Nordlanderia (Hymenoptera: Eucoilidae),
MILA eSCMOLONS OLEWO NEW SPECIES! ase... Rut b-cfe ire Sae) tues <tace Sty carck ae outs einen he’s 158 -
NICKLE, D. A. and M. S. COLLINS—Key to the Kalotermitidae of eastern United States with
AME MW VEOLEFINES LOMMEIONIGa: (ISOPterd) sf of. cde cyte oe ule tle ges-ae We ane slates seit ace st /4& 269
PURRINGTON, F. F. and D. H. STINNER—Wasp parasites of the burdock seed moth, Metz-
neria lappella Zeller (Gelechiidae): New host record for Bracon mellitor Say (Hymenoptera:
RS CCCSANCL tS) Mee Rea eat Ce Se MPM LEA cele hale «kay» ig chsltlenete Aerie. aid Boea shee 203
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1989
F. CHRISTIAN THOMPSON President WARREN E. STEINER, JR. Program Chairman
JEFFREY R. ALDRICH President-Elect GEOFFREY B. WHITE, Membership Chairman
RICHARD G. RosBINS, Recording Secretary ANNE M. WIEBER, Custodian
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NORMAN E. WoobDLey, 7reasurer
HirAM G. LArReEw, Editor
RoBERT D. GorbDon, Associate Editor
Publications Committee
REBECCA F. SURDICK T. J. HENRY GEORGE C. STEYSKAL
B. V. PETERSON
Honorary President
Curtis W. SABROSKY
Honorary Members
Louise M. RUSSELL ALAN STONE THEODORE L. BISSELL
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Title of Publication: Proceedings of the Entomological Society of Washington.
Frequency of Issue: Quarterly (January, April, July, October).
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PRINTED BY ALLEN PRESS, INC., LAWRENCE, KANSAS 66044, USA
THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 133-142
THE UNIQUE PURRING SONG OF CHRYSOPERLA COMANCHE
(BANKS), A GREEN LACEWING OF THE RUFILABRIS
SPECIES GROUP (NEUROPTERA: CHRYSOPIDAE)
CHARLES S. HENRY
Professor of Biology, Department of Ecology and Evolutionary Biology, Box U-43, 75
North Eagleville Road, The University of Connecticut, Storrs, Connecticut 06268.
Abstract.—The courtship song of Chrysoperla comanche, a common green lacewing
from western North America, 1s described and compared to that of its apparent closest
relative, C. rufilabris. It was found to consist of a single long (about 15 seconds) volley
of abdominal vibration, producing a substrate-borne signal that steadily increased in
amplitude from beginning to end. Its primary frequency averaged about 87 Hz at 27°C,
and was modulated in amplitude by simultaneous low frequency vibrations caused by
slower oscillations of the abdomen. Both sexes made identical calls, and each female
answered the male’s signal halfway through his song. Temperature affected the song’s
phenotype, so linear regression equations were used to compare features at any temper-
ature. Unique song characteristics shared between C. comanche and its sister species, C.
rufilabris, reflect their close relationship and phylogenetic distinctness from the Chryso-
perla carnea group.
Key Words:
The green lacewing taxon Chrysoperla
Steinmann has recently been accorded full
generic status (Séméria 1977), yet further
subdivision may be necessary to represent
the evolutionary divergence that has oc-
curred within its boundaries. Currently, the
genus is defined by morphology, bionomics,
and behavior: its members share a common
ground plan of male genitalia and wing ve-
nation, feed as adults principally on hon-
eydews rather than plant lice, overwinter as
diapausing adults, and must “sing” to one
another before mating (Seméria 1977, Ha-
gen and Tassan 1966, Sheldon and Mac-
Leod 1974, Henry and Johnson in press).
However, more careful scrutiny of genitalia
in particular indicates that at least two nat-
ural species groups of Chrysoperla can be
recognized (Agnew etal. 1981, Adams pers.
commun.). The better studied of these, the
Chrysopidae, Chrysoperla, courtship, acoustical, communication
carnea group (Tauber and Tauber, 1987),
includes the common North American
species C. plorabunda (Fitch), C. downesi
(Banks), and C. mohave (Banks). The other
group, knownas the rufilabris group, 1s char-
acterized by male genitalia that are unique
among Chrysoperla; its members are most
easily recognized by their relatively narrow,
apically pointed hind wings, and by the
bright red stripe usually present on the facial
area (genae) of the head. Chrysoperla rufi-
labris (Burmeister), C. harrisii (Fitch), C.
externa (Hagen), and C. comanche (Banks)
belong to this lineage.
Singing during courtship and mating Is a
characteristic of all Chrysoperla species, and
has been studied in both of the species
groups. These songs are actually bouts or
volleys of male or female abdominal quiv-
ering/jerking that vibrate leaves, twigs, or
134
conifer needles with substrate-borne, low
frequency signals (Henry 1979)—a_ phe-
nomenon known as tremulation. The sig-
nals display complex frequency and ampli-
tude modulation and are highly consistent
and unique within each species. Both sexes
sing, reciprocally exchanging their signals in
prolonged duets, and in most species, the
two sexes produce identical songs (Henry
1980a, b, c, 1985a). Within the recognized
and undescribed species of the carnea group
in North America and Europe, the songs
have been shown to have a species-isolating
effect, preventing hybridization between
otherwise interfertile cryptic species (Henry
1985a, b, 1986).
The rufilabris group of Chrysoperla may
also use vibrational calls in this way, but
only the common C. rufi/abris has been ana-
lyzed in any detail (Henry 1980a). Here, I
describe the peculiar tremulating songs of
the sister species of C. rufilabris, C. coman-
che, from two sites in California. This species
was originally described from Texas and is
found throughout southwestern North
America (Bickley and MacLeod 1956, Ag-
new et al. 1981). It is a common resident
of orchards and vineyards at low elevation
and occurs sympatrically with C. rufilabris
at many localities. Just as C. rufilabris has
song features not found in the carnea group,
the call of C. comanche 1s of most unusual
design and dynamics, reflecting and rein-
forcing the distinct phylogenetic position of
the Chrysoperla rufilabris lineage.
METHODS AND MATERIALS
I collected adults of Chrysoperla coman-
che at two sites in California, 220 km apart,
during late September, 1987. Twelve were
obtained in the early afternoon of 24 Sep-
tember, 1987, from the southern tip of Se-
quoia National Forest, two kilometers south
of Isabella Lake, Kern County, at an ele-
vation of about 1000 meters. The collecting
area was very dry and hot (34°C), with scat-
tered 5-10 meter-tall scrub oak (Quercus
dumosa Nuttall) and larger (to 20 meters)
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
digger pine (Pinus sabiniana Douglas). All
were flushed from the oaks by prodding the
branches with an insect net. Twenty-one
other adults were obtained from a citrus
grove just northeast of the town of Mentone,
San Bernardino County, at about 500 me-
ters elevation. These were collected at dusk
(26°C) by beating the branches of 8 meter-
tall lemon trees. Phillip Adams (California
State University, Fullerton) chose this sec-
ond site and assisted in procuring and iden-
tifying the insects there. All were mailed to
my laboratory in Connecticut. The speci-
mens from each population were main-
tained as a separate breeding colony, re-
ferred to as Isabella or Mentone. Adults were
supplied with a water and Wheast” diet (Ha-
gen and Tassan 1966, Henry 1983b). Eggs
were removed daily and larvae reared on
ether-killed Drosophila spp., as described in
Henry (various papers). Photoperiods of
17:7 daylight: darkness and temperatures
of 25 + 1°C resulted in rapid growth and
continual reproduction in the laboratory.
Several field-collected individuals were
tested for reproductive and singing behav-
ior, but the majority of tests were performed
on first and second laboratory generations.
The songs of six males and seven females
from Isabella Lake and three males and one
female from Mentone were analyzed in de-
tail. Of these, one male and two females
from Isabella Lake and one male and one
female from Mentone were original, field-
caught individuals.
Adult males showed courtship activity
within 24 hours of eclosion, and females
after three days. Therefore, tests were ini-
tiated on week-old individuals, and contin-
ued for about one month (lifespan averaged
140 days). During testing, each lacewing was
placed individually or with a partner in a
small cardboard coffee cup covered with
plastic wrap. Vibrations from abdominal
jerking were monitored from the plastic sur-
face with a piezoelectric transducer con-
nected to a Tektronix® digital oscilloscope,
amplifier/speaker, and cassette tape record-
VOLUME 91, NUMBER 2
er, as described in earlier papers (Henry
1985a, b).
For analysis, recorded signals were digi-
tized by a Cambridge Electronics Design
CED1401 12-bit A/D,D/A device and
transferred to an IBM PC-AT microcom-
puter. Multiple-slice fast-Fourier trans-
forms (FFT’s) and accurate measures of
elapsed time could then be performed on
the signals with the WATERFALL-® soft-
ware package of Cambridge Electronics De-
sign, Ltd. Frequency and time data were
keyed into the spreadsheet, Microsoft EX-
CEL®, for further analysis and reduction,
and could then be transferred (through AS-
CII and DIF file conversion) to programs
like Asyst Software Technologies’ ASYS-
TANT +® for statistical analysis or Jandel’s
SIGMAPLOT® and Advanced Graphics
Software’s SLIDEWRITE for graphing.
A lacewing song (= call) is a vibrational
entity that an individual produces once or
at irregular intervals. Each may consist of
one, or many, volleys (= bouts) of abdom-
inal vibration, and those volleys are orga-
nized into “shortest repeated units” (SRU’s),
composed of at least one but often several
volleys delivered with some consistent tem-
poral spacing. During a duet, each partici-
pant repeatedly answers its partner with the
SRU characteristic of the species. In C. co-
manche, the song is best described as a sin-
gle volley, so the terms song, call, volley,
bout, and SRU are synonyms.
Because the features of lacewing songs
vary with temperature, data were collected
at 20 to 28°C to calculate linear regression
equations. Results could then be normal-
ized to a standard temperature, 27°C, to
compare means and standard deviations
among different subsets of data or between
taxa. I measured eight major features (Ta-
bles 1, 2, 3, and Fig. 1) of the songs of C.
comanche, using 71 male and 89 female vol-
leys at 8 different temperatures. These fea-
tures were specifically chosen to facilitate
comparison with Chrysoperla rufilabris (Ta-
ble 3; Figs. 1, 2, 5; Henry 1980a). To com-
135
pare the slopes of the temperature regres-
sions of different song features or different
populations or taxa, Q,, (the factor by which
a feature changes over a 10°C rise in tem-
perature) was determined for each song
component. This factor is obtained by cal-
culating the expected value of the feature at
20 and 30°C from the linear regression and
dividing the larger value by the smaller (Ta-
ble 4).
Significant differences were recognized
where the means of two normally distrib-
uted samples differed from one another by
a two-tailed t-test using confidence limits of
99% or better. These are marked in the ta-
bles with triple asterisks (***). Values fol-
lowing a + sign are one standard deviation
of the mean.
Voucher specimens were deposited in the
entomological collection of the University
of Connecticut. Some were frozen at —90°C
for future protein and DNA analysis.
RESULTS
Mating in C. comanche, as in other lace-
wings of the genus Chrysoperla, was pre-
ceded by a long, well-defined courtship in
which the partners exchanged volleys of ab-
dominal vibration. These songs, usually
produced spontaneously and identically by
both males and females under a wide range
of temperature and light conditions, oc-
curred most often at 25°C or above and dur-
ing the last hour of daylight. Individuals
also sang in response to the call of another
insect, or to a recorded or synthesized song
of the correct structure and duration played
through a small (12 cm) loudspeaker. The
shortest repeated unit (SRU) of the song
consisted ofa single prolonged volley of ab-
dominal vibration, lasting about 15 seconds
at 27°C (Fig. 1; Tables 1, 2, 3). The song,
when heard through headphones, resem-
bled a long, drawn out “purr” that began at
the threshold of audibility and slowly built
to a crescendo, then diminished quickly at
its conclusion with one or two secondary
bursts of vibration. The fundamental or pri-
136 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
SONG (SRU) of CHRYSOPERLA COMANCHE
AMPLITUDE
—— ee rt
13.30
10.64
0.00 2.66 5332) 7.98
TIME, in SECONDS
Fig. 1. Digitized oscillograph of the shortest re-
peated unit (SRU or volley) of the call of Chrysoperla
comanche, showing principal parameters A—D defined
in text and referenced in tables. A, initial volley fre-
quencies; B, early middle volley frequencies; C, late
middle volley frequencies; D, end-of-volley frequen-
cles.
mary frequency of vibration averaged 86-
88 Hz during the major, central portion (B
and C of Fig. 1) of each volley. Each call
started (A) and ended (D) at slightly but
significantly lower frequencies of about 75
and 79 Hz, respectively (Table 3). In ad-
dition, the primary frequency was modu-
lated by a lower frequency, high amplitude
oscillation whose frequency similarly in-
20.8 degrees C
| {| \ |
qn alty Nl, Wha.
HH } NAAN ANAK
Ny
mein
+f} r
4 27 degrees C
\ ini \| \ l iK, \ i| i
)
AVANT Nn Hl iN WMI At Hil MA
Wy TA
0 100 200 300 400 500
TIME, in MILLISECONDS
AMPLITUDE
MV VW
4 1 : y a
|
! )
Fig. 2. Digitized oscillographs of half-second frag-
ments of songs produced at two different temperatures
by a solo female Chrysoperla comanche. Spikes within
a volley are strokes of the individual’s abdomen and
constitute the primary frequency of the call. The regular
variations in overall amplitude are caused by the mod-
ulating frequency.
305 Y = -0.925x + 39.68
] ‘ r2=0.11,N = 150
AL eS
3 ce
Sye204 ate T
a 71 een
n | | = e e
£15 | 1 =| |
re. ] t r. I
FS | | =
ie | —-
a ] e
a
5 if e
0+—__.—_—_+.¥ > re
ZONpe Zils G22 925uee 24S on 260 noe
TEMPERATURE, in °C
Fig. 3. Effect of temperature on volley duration for
all calls of all males and females of Chrysoperla co-
manche. The filled circle at each temperature is the
mean of at least 15 volleys delivered at that temper-
ature, and the error bars bracket one standard devia-
uon of the mean. The linear regression equation, its
squared variance, and its sample base are entered above
the line plot.
creased, leveled off, and then decreased dur-
ing the course of each volley. The mean of
this modulation frequency was about 21 Hz,
but started at approximately 14 Hz and
stopped at 17.5 Hz (Table 3). It was clear
from watching receptive lacewings at low
temperatures that this modulating oscilla-
tion corresponded to gross vertical move-
ments of the insect’s abdomen, while the
higher, primary frequency was produced by
a low-amplitude abdominal “buzzing” su-
perimposed on the low-frequency carrier
(Fig. 2). At high temperatures, primary and
modulating frequencies were both high
enough and of sufficiently low amplitude to
render the motions of the abdomen nearly
invisible.
Males and females sang identically, in
every respect (Tables 1, 2, 3). In 27 of 36
observed heterosexual duets, the female an-
swered a spontaneous call produced by the
male well before the male completed his
song. When the female finished, a few sec-
onds passed before the male initiated
another volley, and then the female again
began to sing partway through the male’s
song. Thus C. comanche duets were not
“polite,” in the sense of nonoverlapping.
In fact, the beats that were produced from
the acoustical interaction between two su-
VOLUME 91, NUMBER 2 137
Table 1. Principal characteristics of the songs of individual males of Chrysoperla comanche, normalized to
27°C. Means and standard deviations are tabulated, with the number of measured volleys entered parenthetically.
Averages and standard deviations of the means are shown in the bottom rows. The letters A to D refer to the
parts of the song specified in Figure 1.
Males
Abdominal Vibration Frequency (strokes/second) Modulation Frequency (maxima/second) a Sone
uration
Indiv Initial (A) Mid! (B) Mid2 (C) End (D) Initial (A) Mid (C) End (D) Kec)
1 — 91.63 90.82 88.14 12.84 22.79 20.76 18.16
= + 11.35 = 1037 + 2.20 - + 0.43 + 1.47 + 5.65
= (7) (7) (7) (1) (7) (5) (7)
2 = 90.38 87.15 85.01 15.34 22 e712. 21.20 21.60
~ C13 +'0:05 + 0.50 - + 0.45 + 0.00 + 4.19
= (4) (4) (4) (1) (4) (4) (4)
3 77.72 88.11 86.88 81.86 12.87 19.70 16.09 LS:
+ 4.16 + 2.41 + 3.00 lil + 1.64 + 1.02 + 1.44 +417
(4) (8) (9) (9) (4) (8) (6) (9)
4 77.56 86.64 85.55 81.10 8.35 20.85 17.09 17.99
+ 3.13 = 2.33 e121 = 2.69 - + O75 se SAT + 3.58
(12) (16) (16) (11) (1) (16) (2) (15)
5 70.16 89.56 87.48 81.03 _ 21.74 18.56 18.01
— a= 0)54/) + 0.47 + 4.32 ad 10:37 + 1.61 + 5.43
(1) (6) (6) (4) = (6) (3) (4)
6 727 90.86 87.79 77.01 = 20.54 = 16.29
+ 5.91 2:19 E52 ED. _ + 0.94 - + 6.00
(4) (5) (5) (4) = (5) = (5)
a 75.92 86.18 83.98 80.47 16.20 21:93 20.13 17.14
+ 0.00 + 3.40 + 1.78 + 0.76 + 4.24 + 0:39 + 3.02 #11052
(2) (7) (7) (7) (5) (7) (6) (7)
8 68.28 85.17 84.99 72.51 14.51 20.13 16.11 13.56
#275512 + 1.08 + 1.24 + 2.01 an PB Xo) + 0.68 += 1253 = 3.29
(11) (11) (11) (1) (5) (11) (11) (11)
9 713.92 89.71 85.63 81.09 — 22.25 18.33 18.06
+ 2.45 + 2.28 + 0.33 3.20) — + 0.43 + 1.42 + 4.84
(5) (6) (6) (6) = (6) (6) (6)
Avg: 73.68 88.69 86.70 80.91 13.35 21.41 18.53 16.88
SD: 3.66 2.27 1.99 4.42 2519 1.14 2.02 3.00
N: 7.00 9.00 9.00 9.00 6.00 9.00 8.00 9.00
perimposed tones of similar but not iden-
tical frequency were very audible over head-
phones during the overlapping phases of the
songs. One to ten of such sequences always
occurred before copulation. Just prior to
copulation, the distinctions between male
and female calls became unclear: in five ob-
served copulations, each partner sang with-
out pause during the final 15 seconds of
courtship, producing a cacophony of sub-
strate-borne noise. The last phase of court-
ship was also characterized by a five to ten
second period of wing bumping and mutual
abdominal tapping, as in C. rufilabris and
most other lacewing species.
Insects were tested and measured at a
number of different temperatures. Data for
three of the eight principal song features are
graphed against temperature in Figs. 3 and
4, with the linear regression equations placed
138 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Principal characteristics of the songs of individual females of Chrysoperla comanche, normalized
to 27°C. Means and standard deviations are tabulated, with the number of measured volleys entered paren-
thetically. Averages and standard deviations of the means are shown in the bottom rows. The letters A to D
refer to the parts of the song specified in Figure 1.
Females
Abdominal Vibration Frequency (strokes/second) Modulation Frequency (maxima/second) Bee
uration
Indiv Initial (A) Mid! (B) Mid2 (C) End (D) Initial (A) Mid (C) End (D) (seconds)
l = 89.40 88.50 85.75 = 22.03 = 17.01
_ + 1.96 213 + 1.40 - + 0.58 = 216.15
= (3) (3) (2) = (3) = (3)
2 78.59 86.90 85.69 82.04 13.14 21.49 16.64 7.69
+ 2.80 + 1°56 = (0195 ae SiS) De. #10533 seu D EDIT + 4.60
(5) (6) (6) (6) (3) (6) (6) (6)
3} 79.72 88.17 86.75 83.29 13.34 20.75 19.12 13.56
+ 3.46 + 1.60 + 1.02 = 2:96 + 3.54 + 1.34 + 2.10 = 21677
(4) (9) (9) (6) (2) (9) (4) (9)
4 76.49 86.50 84.89 76.10 14.23 D1R25 16.01 11.91
+ 2.94 ae) lal + 0.89 3533 + 2.87 + 0.35 + 1.63 223.85)
(16) (22) (22) (22 (6) (22) (22) (20)
5 72.58 86.85 85.31 78.46 15.34 21.27 18.36 16.48
+ 4,3] = 1k69 + 1.14 2513:35) +, 1.28 + 0.48 + 1.45 sey eDil,
(9) (18) (18) (18) (3) (18) (17) (17)
6 73.66 88.68 84.90 79.53 - 21.41 17.09 12.07
- + 175 + 1.60 12911 — + 0.41 + 0.42 + 6.05
() (4) (4) (4) = (4) (4) (4)
7 70.61 90.09 88.19 79.31 _ 19.76 16.75 16.39
+ 4.10 412225 = 133) 2.51 = + 0.32 + JT 2.611)
(5) (10) (10) (9) = (10) (9) (10)
8 81.00 87.42 84.52 78.10 12.56 21.56 17.33 11.39
+ 1.65 + 2.66 + 1.36 22199 + 1.81 + 0.74 "1216 + 6.62
(10) (17) (17) (15) (5) (15) (12) (14)
Avg 76.09 88.00 86.09 80.32 13.72 21.19 1733) 13.31
SD: 3.91 1.31 1:55 3.14 1.09 0.68 1.07 3:21
N 7.00 8.00 8.00 8.00 5.00 8.00 7.00 8.00
beside each graph. Volley/SRU duration
(Fig. 3) varied inversely with temperature,
but the correlation was not very good: the
regression coefficient, R*, was a low 0.11.
All primary and modulating frequency
components (Fig. 4) showed a direct, linear
temperature relationship, and those in the
middle portions of the call were tightly cor-
related with temperature changes (R? = 0.84
to 0.85). The Q,, values calculated from the
regression equations (Table 4) were statis-
tically equivalent for all frequency compo-
nents of the calls. Compared with C. plora-
bunda and Chrysopiella minora Banks,
however, pitch alterations of the songs of
C. comanche were less pronounced for a
given temperature change.
The temperature regression equations al-
lowed conversion of all measured values to
their equivalents at 27°C. Individual lace-
wings varied little in their normalized av-
erage values for most of the principal song
features (Tables 1 and 2). When different
populations of insects were compared (Ta-
ble 3), males were insignificantly different
from females, and Isabella Lake individuals
VOLUME 91, NUMBER 2
90+ @ = MID SONG, PRIMARY FREQ.
852 te a
80 i
N t s =
we 754 : Y = 2.13x + 28.48, r2=0.84, N=160
lu 4 se
7s } e.
er
> 244
o ; # = MID SONG, MODULATION FREQ
a 21-4 Theat ad a
so | 4
Ww 18
i
1 a Y = 1.13x — 9.43, r2=0.85, N=157
15 qa
rr ae
20 «21 22) 8235) G24 25) = 26) 27) 28
TEMPERATURE, in OC
Fig. 4. Effect of temperature on the primary and
modulation frequencies of the mid-volley portion of
the songs of all males and females of C. comanche. The
filled circle or triangle at each temperature is the mean
of at least 15 volleys delivered at that temperature, and
the error bars bracket one standard deviation of the
mean. The linear regression equation, its squared vari-
ance, and its sample base are entered below each line
plot.
were essentially identical to those from
Mentone. Only one call feature showed sig-
nificant differences between compared sam-
ples: Mid2 primary frequency in the Isabella
and Mentone populations differed by 2 Hz.
Song duration was the most variable of the
measured song parameters among individ-
uals (Tables 1 and 2).
DISCUSSION
The song of Chrysoperla comanche was
found to be unique among those that have
been described for eleven other species of
its genus. First, it is characterized by the
longest unbroken volley of any North
American lacewing, averaging nearly 15
seconds in duration, but occasionally ex-
ceeding 30 to 35 seconds. The species that
most nearly matches C. comanche in volley
length is its North American sister species,
C. rufilabris, with volleys of four to eight
seconds (Henry 1980a). Another species
with volleys of comparable length is the un-
described P3 sibling of C. plorabunda. Its
volleys, produced several per SRU, can be
as long as eight seconds in individuals from
the Kofa Mountains of Arizona (unpub-
139
/ i) _-100 200 300 400 z soo
Ps MILLISECONDS
AMPLITUDE
0.00 1.00 2.00 3.00 4.00 5.00 6.00
TIME, in SECONDS
Fig. 5. Digitized oscillograph of the shortest re-
peated unit (SRU or volley) of the male “long call” of
Chrysoperla rufilabris. Inset is a half-second section
taken from the middle of the volley, which shows elev-
en individual oscillations of the abdomen (black
smears). Note the absence of a clear frequency struc-
ture; the abdomen strikes the substrate and generates
high-frequency noise.
lished data). The only lacewing species that
seems to exceed C. comanche in volley
length is an undescribed relative of C. car-
nea from near Ticino in the southern Alps
of Europe. A volley in this species may last
for more than two minutes. However, this
case may not be strictly comparable, be-
cause each long bout of abdominal vibra-
tion in the Ticino form actually consists of
numerous, independent short volleys re-
peated about eight times per second (Henry
1983a).
A second remarkable feature of the C.
comanche call is the modulation of the pri-
mary abdominal vibration frequency by
regular, intense oscillations of lower fre-
quency. Some green lacewings that produce
a series of short volleys in rapid succession
approach but do not attain the condition
seen in C. comanche. For example, a sibling
species of C. downesi from the redwood for-
ests of California (Founder’s Grove) repeats
its short volleys at nearly 10 per second (un-
published data), and two sibling species of
C. carnea from different parts of the Alps
of Switzerland may attain volley repetition
rates of about eight per second (Henry 1983a,
140
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 3. Principal characteristics of the songs of Chrysoperla comanche, normalized to 27°C and averaged
for all measured volleys produced by males versus females (upper section of table) or produced by individuals
from Isabella Lake versus Mentone, California (lower section). The corresponding features of the songs of
Chrysoperla rufilabris are entered at the bottom of the table for comparison. Mean values and standard deviations
are tabulated, with the number of measured individuals (column 1) or volleys (all other columns) entered
parenthetically. Significant differences (between rows) at the 1% level are indicated by asterisks. N/a = not
applicable.
Abdominal Vibration Frequency (strokes/second) Modulation Frequency (maxima/second) Toone
rat
Initial (A) Mid! (B) Mid2 (C) End (D) Initial (A) Mid (C) End (D) (Geconds)
All Males: 73.66 88.06 86.41 80.40 14.21 21.19 18.21 16.36
(N = 9) SEEDEDO) =. 2494 + 2.35 + 5.24 S23" EZ SG NG
(39) (70) (71) (63) (17) (70) (43) (67)
All Females: 76.51 87.54 85.64 78.70 13.70 21.14 Les 13.36
(N = 8) + 4.57 + 2.14 Ea 1!6)/ + 3.68 22339, 101825) EO nA OF
(50) (89) (89) (82) (19) (87) (74) (83)
All Isabella: 75.67 88.06 86.42 *** 80.19 13.59 20.45 16.47 14.90
(N = 13) + 4.38 + 2.38 + 2.04 + 4.48 25 0H SE 122" 22:37 75108
(61) (118) (119) (106) (21) (118) (82) (112)
All Mentone: 74.37 86.94 84.75" 77.41 14.42 20.95 16.99 14.13
(N = 4) + 6:57 + 2.78 Si + 3.87 9317 10196 = 27) G83
(28) (41) (41) (39) (15) (39) (35) (38)
Totals: 75.26 87.77 85.98 79.44 13.94 21.16 93 14.70
(N = 17) tS 17) 9253 + 2.03 + 4.49 £2579" VEO], 2314 E558
(89) (159) (160) (145) (36) (SH) (UA), (150)
Chrysoperla n/a n/a n/a n/a — 18.20 = BHII6)
rufilabris: =e — +1.23
(N = 10, 28 C) _ (116) - (183)
1985a). In contrast, C. comanche’s signals
consist of long, continuous volleys, with no
pauses in primary abdominal vibration ac-
tivity between the amplitude peaks (Fig. 2).
Also, the rate or frequency of modulation,
about 21 Hz, is higher by a factor of two or
more than in species with discrete, rapidly
repeated volleys. The interaction of the pri-
mary and modulation frequencies at two
temperature extremes is illustrated in Fig. 2.
The universal pattern of singing within
the carnea group of Chrysoperla is the pro-
duction of identical SRU’s alternately by
each individual of a courting pair. Whether
these are single volleys or complex series of
volleys, those of one partner do not overlap
or acoustically interfere with those of the
other. In contrast, duets in C. comanche are
not “polite”; the female typically answers
the male halfway through his one-volley call,
and then both partners often sing simulta-
neously for several minutes before copulat-
ing. Similarly, in C. comanche’s closest
North American relative, C. rufilabris, the
male produces a song consisting of a single,
long volley of abdominal vibration, and the
receptive female also answers the male in
the middle of his volley (Henry 1980a).
Other basic song features are shared be-
tween C. comanche and C. rufilabris (com-
pare Figs. | and 5). In both species, the SRU
is an exceptionally long volley of abdominal
vibration that increases steadily in ampli-
tude to a crescendo near its end. Neither
species seems to repeat its long volleys at
any regular interval: the volley, SRU, and
song are all one and the same. And finally,
both species modulate the intensity of their
calls with very low frequency, high ampli-
tude oscillations of the abdomen, at ap-
VOLUME 91, NUMBER 2
141
Table 4. Q,, factors over the range of 20 to 30°C for each of the principal features of the songs of all individuals
of Chrysoperla comanche. Corresponding values for Chrysoperla plorabunda (Henry 1983b) and for Chrysopiella
minora (Henry and Johnson in press) are also tabulated. N/a = not applicable.
Abdominal Vibration Frequency (strokes/second)
Species Initial (A) Mid! (B) Mid2 (C)
Chrysoperla comanche 1.19 132 1.30
(N = 17)
Chrysoperla plorabunda 1.61 1.47 1.47
(N = 10)
Chrysopiella minora 1.80 1.81 1.81
(N = 15)
Modulation Frequency (maxima/second) Song
Duration
End (D) Initial (A) Mid (C) End (D) (seconds)
1.28 1.18 1.86 1.67 1.78
125 n/a n/a n/a 1.83
1.71 n/a n/a n/a Dali)
proximately the same rate: 21 Hz in C. co-
manche, and 17 Hz in C. rufilabris. These
shared characteristics probably reflect the
close phylogenetic relationship between the
two species, because such traits are not found
in the carnea group of Chrysoperla.
However, important differences also exist
between C. comanche and C. rufilabris. First,
C. rufilabris has sexually dimorphic calls,
which were not found in C. comanche. Male
C. rufilabris usually produce long volleys,
but females answer with much shorter bursts
of abdominal vibration and have not been
found to produce anything resembling the
male SRU (Fig. 5; see also fig. 1F and G in
Henry 1980a). Second, the song of C. rufi-
labris is devoid of any primary frequency
structure analogous to that of C. comanche;
only lower frequency oscillations, corre-
sponding to C. comanche’s modulation fre-
quency, are apparent (Fig. 5, inset). It is
possible that C. rufilabris, unlike other lace-
wings that tremulate, strikes the substrate
with its abdomen, because considerable 500
Hz noise is carried on the 17 Hz amplitude
peaks (Fig. 5, inset). Whatever the cause,
the songs of the two species are distinctive
and “sound” different from one another
through headphones, despite their several
shared characteristics.
Whether or not C. comanche and C. rufi-
labris can interbreed is unknown. The two
species overlap geographically, thus provid-
ing the potential for hybridization: C. rufi-
labris occurs commonly in fields and mead-
ows throughout most of North and Central
America, including the American Southwest
and California (Bram and Bickley 1963),
and C. comanche has been collected at many
sites in California, Arizona, New Mexico
and Texas (Bickley and MacLeod 1956).
Both species reproduce continually under
long daylight conditions, suggesting pro-
longed overlap in their breeding seasons in
areas of sympatry. Although the relevant
breeding experiments remain to be done,
the pronounced differences between the
songs of the two sister species probably pre-
clude successful interspecific courtship and
mating.
The phylogenetic position of another
common member of the rufi/abris group, C.
harrisii, is uncertain. It, too, sings during
courtship, but its song is a peculiar, pro-
longed session of untidy, almost random
vibrational bursts, and has not yet been
characterized in detail. Its dark green color,
conifer-associated habits, and unique male
genitalia (Bram and Bickley 1963) make it
easy to distinguish from C. rufilabris or C.
comanche, and its song has not been found
to exhibit any of the features shared by C.
rufilabris and C. comanche. This indicates
that the latter two species are more closely
related to each other than either is to C.
harrisii, and that C. harrisii may be the sister
group of the rufilabris-comanche lineage.
The reproductive behavior of the fourth rel-
ative of these species, C. externa, has not
been investigated.
142
My studies of C. comanche and its rela-
tives suggest that singing behavior has taken
a different evolutionary direction in the ru-
filabris group of Chrysoperla than it has in
the carnea group. It also provides evidence
for separating Chrysoperla into two genera,
to avoid the confusion of species groups. In
both groups, singing 1s essential to court-
ship, and is complex in most species, per-
haps because of the reinforcement of be-
havioral isolating “‘mechanisms’’ in
genetically similar, potentially interbreed-
ing taxa (Butlin 1987). However, polite duets
between partners have developed princi-
pally in the carnea group, while the three
members of the rufilabris group have ac-
quired long, overlapping calls that may in-
volve drumming on the substrate rather than
tremulation.
ACKNOWLEDGMENTS
This work was supported by National Sci-
ence Foundation award BSR-8508080,
Charles S. Henry Principal Investigator. I
thank my research assistant and wife, Julie
J. Henry, for help in maintaining living lace-
wings. Phillip Adams (California State Uni-
versity, Fullerton) helped greatly in the col-
lection and identification of specimens, but,
more importantly, provided me with the
conceptual background in chrysopid tax-
onomy and phylogeny that made this study
possible. Special thanks go to an anony-
mous reviewer, who worked hard to im-
prove the clarity of the manuscript.
LITERATURE CITED
Agnew, C. W., W. L. Sterling, and D. A. Dean. 1981.
Notes on the Chrysopidae and Hemerobiidae of
eastern Texas with keys for their identification.
Southwestern Entomol., suppl. 4: 1-20.
Bickley, W. E. and E. G. MacLeod. 1956. A synopsis
of the nearctic Chrysopidae with a key to the gen-
era (Neuroptera). Proc. Entomol. Soc. Wash. 58:
177-202.
Bram, R. A. and W. E. Bickley. 1963. The green
lacewings of the genus Chrysopa in Maryland
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(Neuroptera: Chrysopidae). Univ. Maryland Agric.
Exp. Stn., Bull. A-124: 1-18.
Butlin, R. K. 1987. Species, speciation, and rein-
forcement. Am. Nat. 130: 461-464.
Hagen, K. S. and R. L. Tassan. 1966. The influence
of protein hydrolysates of yeast and chemically
defined diet upon the fecundity of Chrysopa car-
nea Steph. Vest. csl. Spol. Zool. 30: 219-227.
Henry, C.S. 1979. Acoustical communication during
courtship and mating in the green lacewing Chry-
sopa carnea (Neuroptera: Chrysopidae). Ann.
Entomol. Soc. Am. 72: 68-79.
1980a. Acoustical communication in Chry-
sopa rufilabris (Neuroptera: Chrysopidae), a green
lacewing with two distinct calls. Proc. Entomol.
Soc. Wash. 82: 1-8.
1980b. The courtship call of Chrysopa dow-
nesi Banks (Neuroptera: Chrysopidae): Its evolu-
tionary significance. Psyche 86: 291-297.
1980c. The importance of low-frequency,
substrate-borne sounds in lacewing communica-
tion (Neuroptera: Chrysopidae). Ann. Entomol.
Soc. Am. 73: 617-621.
. 1983a. Acoustic recognition of sibling species
within the holarctic lacewing Chrysoperla carnea
(Neuroptera: Chrysopidae). Syst. Entomol. 8: 293-
301.
1983b. Temperature-induced changes in the
calls of the green lacewing, Chrysoperla plorabun-
da (Neuroptera: Chrysopidae). Psyche 90: 343-
360.
1985a. Sibling species, call differences, and
speciation in green lacewings (Neuroptera: Chry-
sopidae: Chrysoperla). Evolution 39: 965-984.
1985b. The proliferation of cryptic species
in Chrysoperla green lacewings through song di-
vergence. Fla. Entomol. 68: 18-38.
1986. Good vibrations. Natural History 95:
46-53.
Henry, C. S. and J. B. Johnson. In Press. Sexual sing-
ing in a non-chrysoperlan green lacewing, Chry-
sopiella minora Banks. Can. J. Zool.
Séméria, Y. 1977. Discussion de la validité taxono-
mique du sous-genre Chrysoperla Steinmann
(Planipennia, Chrysopidae). Nouv. Rev. Entomol.
7: 235-238.
Sheldon, J. K. and E.G. MacLeod. 1974. Studies on
the biology of the Chrysopidae. 5. The develop-
mental and reproductive maturation rates of Chry-
sopa carnea (Neuroptera, Chrysopidae). Entomol.
News 85: 159-169.
Tauber, C. A. and M. J. Tauber. 1987. Inheritance
of seasonal cycles in Chrysoperla (Insecta: Neu-
roptera). Genet. Res., Camb. 49: 215-223.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 143-152
SURFACE SCULPTURING OF THE ABDOMINAL INTEGUMENT OF
MEMBRACIDAE AND OTHER AUCHENORRHYNCHA (HOMOPTERA)
CHRISTOPHER H. DIETRICH
Department of Entomology, Box 7613, North Carolina State University, Raleigh, North
Carolina 27695-7613.
Abstract. —Scanning electron microscopic studies of the abdominal integument of au-
chenorrhynchous Homoptera indicate the presence of a variety of features including
sensilla, acanthae, and microtrichia, the variability of which cannot be resolved by the
dissecting microscope. Three classes of structures are recognized based on their celluar
components: |) multicellular processes (sensilla); 2) unicellular processes (acanthae); and
3) subcellular processes (microtrichia). Four kinds of sensilla were found: sensilla coelo-
conica, sensilla papillosa (previously undescribed), sensilla trichodea, and sensilla pla-
codea. In a survey of 46 genera representing 12 families (Eurymelidae, Cicadellidae,
Membracidae, Biturritiidae, Aetalionidae, Cercopidae, Aphrophoridae, Tibicinidae, Cixi-
idae, Delphacidae, Fulgoridae, and Flatidae), sensilla and non-sensory protuberances were
found on the non-genital abdominal terga of members of all auchenorrhynchous super-
families except the Fulgoroidea.
Key Words:
ing, sensilla
Examination of the abdominal integu-
ment of many auchenorrhynchous Homop-
tera by scanning electron microscopy re-
veals several fine-structural features of
potential interest to systematists. Previ-
ously, these features, which include various
types of sensilla and non-sensory protuber-
ances, have been neglected or viewed su-
perficially as textures in revisionary works.
The objectives of this paper are to describe
the common fine-structural features of in-
tegument sculpturing in the Membracidae
and related groups, to propose a nomencla-
ture for such features, and to discuss their
diversity and value for hypothesizing rela-
tionships among taxa.
Harris (1979) summarized the descrip-
tive terminology of insect integumental
sculpturing, proposing the use of relative
rather than absolute measures of size of the
Homoptera, Auchenorrhyncha, morphology, integument, surface sculptur-
individual elements of sculpturing (e.g.
punctate vs. puctulate). He further proposed
the prefixing of terms for sculpturing visible
only at magnifications greater than 100 x
with “micro-” (e.g. micropunctate). While
such terminology is useful at the descriptive
level, 1t can be misleading with regards to
homology if the elements of the sculpturing
have fine-structure themselves. For exam-
ple, the terms “punctate” and “foveate”
could refer to inornate depressions in the
integument as well as sensillar pores. There-
fore, some nomenclatural means for differ-
entiating integument textures based on the
fine structure of the individual elements is
needed if homologies among such structures
are to be taken into account.
The terminology used herein is derived
as much as possible from the hypothesized
functional and developmental aspects of the
144
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
brochosomes (b).
features described following Snodgrass’
(1935) classification of the cuticular features
of insects as updated by Richards and Rich-
ards (1979) and Zacharuk (1985).
Most detailed observations of the integ-
umental fine-structure of Auchenorrhyncha
have been restricted to the mouthparts (e.g.
Backus and McLean 1983), antennae (e.g.
Lewis and Marshall 1970, Bourgoin 1985),
and genitalia (e.g. Restrepo-Mejia 1980).
Other observations include Wood and Mor-
ris’ (1974) survey of 100 species of Mem-
bracidae for the presence of articulated hairs
(sensilla trichodea) on the pronotum,
Wood’s (1975) observations on the ultra-
structure of the membracid pronotal integ-
ument, and Kitching and Filshie’s (1974)
study of the anal apparatus of membracid
nymphs. Smith and Littau (1960) and Giin-
thart (1977) documented the presence of
minute spherical excretory granules called
brochosomes covering the bodies of some
leafhoppers. Presently, such observations
Detail of abdominal integument of a cicadellid, Draeculacephala sp. showing microtrichia (m) and
have not been applied to systematic studies
and the integument of body parts other than
those mentioned above has been virtually
ignored.
MATERIALS AND METHODS
Most of the materials examined were dried
museum specimens (see Table | for a list
of taxa examined). All are deposited in the
North Carolina State University Insect Col-
lection (Department of Entomology, North
Carolina State University, Raleigh). The ab-
domen of each was removed, glued to an
aluminum stub with conductive graphite
paint, coated with gold or gold/palladium
in a sputter coater and observed and pho-
tographed in a JEOL T200 or Philips SOST
scanning electron microscope at 10-20 kV.
To determine whether air-drying signifi-
cantly distorted the fine-structural features
being examined, some specimens preserved
in 70% ethanol were dehydrated in a graded
VOLUME 91, NUMBER 2
145
Figs. 2-10. Abdominal sensilla of auchenorrhynchous Homoptera. 2-6, Sensilla coeloconica. 2, Enchenopa
sp. 3, Cymbomorpha amazona. 4, Acutalis tartarea. 5, Oncometopia orbona. 6, Philaenus sp. 7-9, Sensilla
papillosa. 7, Cymbomorpha amazona. 8, Aconophora cultellata Walker. 9, Hypsoprora coronata. 10, Sensillum
placodeum, Centrodontus atlas.
series of ethanol (80, 95, and 100%), dried
in a Tousimis Samdri-PVT-3B critical point
drier and coated and observed as described
above. In this comparison, air-drying did
not appear to significantly distort the fine-
structural features observed.
Many of the cicadellids examined had the
integument covered with brochosomes (Fig.
1; for a description, see Smith and Littau
1960) which obscured other features of the
integument. According to Giinthart (1977),
these can be removed by soaking the spec-
imens in hexane, diethyl ether, or methanol.
(Interestingly, the presence of brochosomes
may itself be of taxonomic significance. They
were found on several cicadellids, an eu-
rymelid, and an aetalionid, but not on any
of the other specimens examined—see Ta-
blew)
To determine the cellular basis for the
individual elements of surface sculpturing,
the abdomens of two specimens preserved
in 70% ethanol, one pharate and one fully-
sclerotized adult, were embedded in Spurr
medium (Hayat 1986), sectioned with an
ultramicrotome, slide mounted, stained with
146 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Summary of observations on the abdom-
inal integument of selected auchenorrynchous Ho-
moptera. (Arrangement of higher taxa follows that of
Evans (1977) except where otherwise noted.)
Other
Taxon Sensilla Features*
Cicadelloidea
Eurymelidae
Eurymela sp. tac m, b
Cicadellidae
Ledra sp. t;c d
Evacanthus sp. lie m, b
Idiocerus sp. m, b
Oncometopia sp. tc m
Draeculacephala sp. m, b
Membracoidea
Membracidae (sensu
Deitz 1975).
Centrotinae
Platycentrini
Tylocentrus sp. t a
Nessorhininae
Nessorhinini
Goniolomus sp. tac d
Membracinae
Aconophorini
Aconophora spp. ticepay sd
Talipedini
Talipes appendiculata tc d
(da Fonseca)
Hoplophorionini
Alchisme sp. t,c,pa m
Hoplophorion sp. t,c,pa m
Ochropepla sp. t,c,pa a
Platycotis vittata
(Fabricius) t,c,pa =m
Potnia sp. t,c,pa d
Stalotypa sp. t,c, pa m
Umbonia crassicornis
(Amyot & Serville) t,c,pa m
Membracin1
Campylenchia latipes
(Say) tc d
Enchenopa sp. me d
Hypsoprorini
Hypsoprora coronata
(Fabricius) t, pa a
Centrodontini
Centrodontus atlas Goding t,¢,pl =m
Darninae
Cymbomorphini
Cymbomorpha amazona
Stal t'¢;;pay ‘mm
Procyrtini
Procyrta sp. t,.c m
Table 1. Continued.
Other
Taxon Sensilla Features*
Darnini
Stictopelta sp. tC m
Hyphinoini
Hyphinoe sp. tc m
Hemikypthini
Proterpia sp. t,c m
Smiliinae
Acutalini
Acutalis tartarea (Say) in d
Ceresini
Spissistilus festinus (Say) tic m
Cyphonia sp. tC m
Amastrini
Vanduzeea arquata (Say) tic m
Smiliini
Atymna querci (Fitch) tc m
Archasia auriculata
(Fitch) tC m
Tragopini
Tragopa sp. tic m
Polyglyptini
Entylia carinata
(Forster) t,c,pa m
Polyrhyssa sp. t,c,pa m
Stegaspidinae
Microcentrini
Microcentrus caryae
(Fitch) tc m
Heteronotinae
Heteronotini
Heteronotus sp. LEXe m
Biturritiidae (= Lampropteridae
sensu Evans 1948)
Tropidaspis sp. tic d
Biturritia sp. tc m
Aetalionidae
Aetalion reticulatum
(Linnaeus) tie d,b
Darthula hardwickii
(Gray) t d
Cercopoidea
Cercopidae
Prosapia bicincta (Say) - =
Aphrophoridae
Philaenus sp. tac m
Cicadoidea
Tibicinidae
Tibicen sp. t,¢; pl’ “m
Fulgoroidea
Cixiidae
Pintalia sp.
VOLUME 91, NUMBER 2
Table 1. Continued.
Other
Taxon Sensilla Features*
Delphacidae
Liburniella ornata (Stal) — _
Fulgoridae
Alphina glauca (Metcalf) ~~ -
Flatidae
Metcalfa pruinosa (Say) - a
* KEY: a = simple acanthae; b = brochosomes; ¢ =
sensilla coeloconica; d = toothed acanthae; m = mi-
crotrichia; pa = sensilla papillosa; pl = sensilla pla-
codea; t = sensilla trichodea; — = absence of above
features.
methylene blue, and examined under a
compound microscope.
RESULTS
Sensilla and non-sensory protuberances
were found on the non-genital abdominal
terga of adult representatives of all the au-
chenorrhynchous superfamilies except Ful-
goroidea (i.e. Membracoidea, Cicadello-
idea, Cercopoidea, and Cicadoidea: see
Table 1).
Richards and Richards (1979) classified
the cuticular protuberances of insects into
four major groups based on their cellular
structure: 1) multicellular undifferentiated
(spines); 2) multicellular differentiated (sen-
silla); 3) unicellular (acanthae); and 4) sub-
cellular (microtrichia). Three of these (2, 3,
and 4) were found on the abdominal integ-
ument of membracids. They are described
as follows: A. Sensilla. Four kinds of sen-
silla-like structures were found and are
named (with one addition) according to the
classifications of Snodgrass (1935) and Za-
charuk (1985) based on external morphol-
ogy. The following designations are neces-
sarily tentative until the functions of these
features are elucidated through ultrastruc-
tural studies.
Sensilla trichodea are socketed, hair-like
structures— setae — present in varying num-
bers and sizes on the abdomen of many Au-
chenorrhyncha (e.g. Fig. 24).
Sensilla coeloconica are pegs or cones set
147
in pits approximately 1-20 um in diameter.
They vary in size, abundance, and fine-
structure. When present they appear as
punctations or foveae or are not visible at
low magnifications (Figs. 2-6).
Sensilla placodea consist of a plate of sen-
sory cuticle surrounded by a membranous
ring 1-5 wm in diameter (Fig. 10). They were
found on only two of the specimens ex-
amined (see Table |) but eventually may
be found to be common in some groups (e.g.
Cicadoidea).
Sensilla papillosa are previously unde-
scribed structures, presumed to be sensilla,
consisting of groups of papillae 2—5 um in
diameter that may be associated with a pore
or membrane. They are less abundant than
sensilla coeloconica and are generally not
visible at low magnifications (Figs. 7-9).
Further study of the ultrastructure of these
features is needed to determine whether they
can be assigned to any of the other classes
of sensilla.
B. Acanthae, microtrichia, and deriva-
tives. The unicellular structures of Mem-
bracidae vary from single tooth-like or pec-
tinate processes (acanthae) to divided groups
of subcellular projections (microtrichia).
They also vary in size and relative abun-
dance, and give the integument a grainy to
finely pubescent appearance at low magni-
fication. Differentiation among acanthae and
microtrichia presents some difficulty as it
requires knowledge of the developmental
origins of these structures (Richards and
Richards 1979). Thin-sections of a pharate
adult Archasia auriculata (Fitch) (Fig. 12,
cf. Fig. 11) indicate that the small hair-like
projections on the integument surface are
relatively numerous compared to the un-
derlying epidermal cells, demonstrating their
subcellular nature. It seems likely that each
of the arcuate groups of these microtrichia
seen in Fig. | 1 corresponds to an individual
epidermal cell and is therefore derived from
an acanthus (sensu Richards and Richards
1979).
There is often a wide range of variation
in the relative amount of development of
Figs. 11-15.
acanthae and microtrichia on individual
specimens. The general trend in Membrac-
idae is from maximal to minimal devel-
opment from anterior to posterior ends of
the individual terga (Fig. 13). Thus, when
comparing taxa, it may be desirable to con-
sider homologous sites on the specimens, or
to compare only the maximal or minimal
conditions of these structures.
Patterns of subcellular sculpturing, other
than microtrichia, that are uniformly dis-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
11-12. Cellular structure of the cuticular protuberances of a membracid, Archasia auriculata.
11, Distribution of microtrichia (m) and sensilla trichodea (t) on abdominal tergum IV. 12, Thin-section through
a portion of abdominal tergum integument showing positions of epidermal cells (e) relative to microtrichia. 13.
A portion of abdominal tergum IV of Enchenopa sp. (Membracidae) illustrating variation in the integument
sculpturing from anterior (left) to posterior (right). 14-15. Microtextures of membracid integument. 14, Micro-
granulate, Acutalis tartarea. 15, Microrugose, Atymna guerci.
tributed over the integument surface may
be referred to as “microtexture.”’ Thus far,
three conditions have been observed and
are named according to the convention of
Harris (1979): microglabrous (smooth, e.g.
Fig. 27); microgranulate (grainy, Fig. 14);
and microrugose (wrinkled, Fig. 15).
DISCUSSION
The observations documented here dem-
onstrate the need to study the fine-structure
VOLUME 91, NUMBER 2
Figs. 16-21. Comparisons of foveae on the abdominal terga of three membracids (lateral views). 16-17,
Goniolomus tricorniger. 16, Terga II] and IV. 17, Detail, tergum IV. 18-19, Talipes appendiculata. 18, Terga
II-V. 19, Detail, tergum IV. 20-21, Hypsoprora coronata. 20. Terga II and IV. 21, Detail, tergum III.
of integumental sculpturing if such features
are to be used in systematics. Simply char-
acterizing integument features as textures
may be misleading with regards to homol-
ogies among such characters at the level of
the individual fine-structural elements. Fur-
thermore, examination of the integument
fine-structure yields a wealth of morpho-
logical information that is potentially ap-
plicable to phylogenetic studies. These
150 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 22-27. Variation of non-sensory protuberances on abdominal tergum IV in two membracid tribes. 22-
24, Hoplophorionini. 22, Ochropepla sp. 23, Potnia sp. 24, Alchisme sp. 25-27, Aconophorini. 25, Aconophora
cultellata Walker (t, sensillum trichodeum). 26, Aconophora grisescens. 27, Aconophora sp.
points are best illustrated with examples Hypsoprora coronata (Fabricius) all have
from the membracid taxa surveyed. large pits on the abdominal terga bearing a
The species Goniolomus tricorniger Stal, superficial resemblance to one another (Figs.
Talipes appendiculata (da Fonseca), and 16, 18, and 20). However, the fine-structure
VOLUME 91, NUMBER 2
of these pits differs greatly: G. tricorniger
(Fig. 17) has simple inornate pits, 7. ap-
pendiculata (Fig. 19) has a single coeloconic
sensillum in each pit, and H. coronata (Fig.
21) has a single papillose sensillum and three
to five non-sensillar depressions per pit.
A trend in the evolution of subcellular
protuberances is evident when representa-
tives of the tribe Hoplophorionini (sensu
Deitz 1975) are compared. In the genus
Ochropepla (Fig. 22), for example, the pro-
tuberances are maximally developed as
small acanthae, each bearing one or two
acute projections. In the genus Potnia (Fig.
23), many of these acanthae are further di-
vided into groups of three or more projec-
tions. The groups of microtrichia found in
the genus Alchisme (Fig. 24) are each ap-
parently homologous to a single acanthus
of Ochropepla.
A similar trend occurs among the species
of the genus 4conophora. The non-sensory
protuberances of Aconophora cultellata
Walker (Fig. 25) are maximally developed
as single to tridentate acanthae. The acan-
thae of Aconophora grisescens (Germar) (Fig.
26) are multidentate. In a third (unde-
scribed) species (Fig. 27), the acanthae each
have several blunt crenulations, suggesting
a secondary reduction of the pointed den-
ticulae found in other species of the genus.
A more extensive survey of the integu-
mental fine-structure of Auchenorrhyncha
is needed to determine whether this previ-
ously overlooked complex of characters
supports current taxonomic classifications
of the group. Furthermore, elucidation of
the functions of the sensilla described herein
may provide valuable information concern-
ing the evolution and ecology of the taxa in
which they occur.
ACKNOWLEDGMENTS
I am indebted to N. A. Leidy for provid-
ing the initial micrographs that inspired this
work, to C. B. Moore for invaluable tech-
nical assistance, and to L. L. Deitz, M. H.
Farrier, S. H. McKamey, H. H. Neunzig,
151
and an anonymous reviewer for many use-
ful comments on the manuscript. This work
was funded, in part, through a mini-grant
from the Center for Electron Microscopy,
North Carolina State University. Paper No.
11529 of the Journal Series of the North
Carolina Agricultural Research Service, Ra-
leigh, NC 27695.
LITERATURE CITED
Backus, E. A. and D. L. McLean. 1983. Sensory sys-
tems and feeding behavior of leafhoppers. II. A
comparison of sensillar morphologies of several
species (Homoptera: Cicadellidae). J. Morphology
176: 3-14.
Bourgoin, T. 1985. Morphologie antennaire des Tet-
tigometridae (Hemiptera, Fulgoromorpha). Nouv.
Revue Entomol. (N. S.) 2(1): 11-20.
Deitz, L. L. 1975. Classification of the higher cate-
gories of the New World treehoppers (Homoptera:
Membracidae) North Carolina Agric. Exp. Sta.
Tech. Bull. 225. 177 pp.
Evans, J. W. 1948. Some observations on the clas-
sification of the Membracidae and on the ancestry,
phylogeny, and distribution of the Jassoidea. Trans.
Royal Entomol. Soc. London 99(15): 497-515.
1977. The leafhoppers and froghoppers of
Australia and New Zealand (Homoptera: Cica-
delloidea and Cercopoidea). Part 2. Rec. Austra-
lian Mus. 31(3): 83-129.
Giinthart, H. 1977. Einfluss des insektenalters auf
bestimmungsmerkmale. Biotaxonomische und
rasterelektronmicroscopische untersuchungen bei
kKleinzikaden (Hom. Auchenorrhyncha, Cicadel-
lidae). Mitt. Schweiz. Entomol. Ges. 50: 189-201.
Harris, R. 1979. A glossary of surface sculpturing.
Occas. Pap. Bur. Entomol. California Dep. Agric.
28: 1-31.
Hayat, M.A. 1986. Basic techniques for transmission
electron microscopy. Academic Press, Orlando,
Florida, 411 pp.
Kitching, R. L. and B. K. Filshie. 1974. The mor-
phology and mode of action of the anal apparatus
of membracid nymphs with special reference to
Sextius virescens (Fairmaire) (Homoptera). J.
Entomol. (sec. A)49(1): 81-88.
Lewis, C. T. and A. T. Marshall. 1970. The ultra-
structure of the sensory plaque organs of the an-
tennae of the Chinese lantern fly, Pyrops cande-
laria L. (Homoptera, Fulgoridae). Tissue & Cell
2(3): 375-385.
Restrepo-Mejia, R. 1980. Membracidos de Colom-
bia-I. Revision parcial de las especies del género
Alchisme Kirkaldy (Homoptera: Membracidae:
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Hoplophorioninae). Caldasia (Bogota) 13(61): 103-
164.
Richards, A. G. and P. A. Richards. 1979. The cu-
ticular protuberances of insects. Int. J. Insect Mor-
phol. Embryol. 8: 143-157.
Smith, D. S. and V. C. Littau. 1960. Cellular spe-
cialization in the excretory epithelia of an insect,
Macrosteles fascifrons Stal (Homoptera). J. Bio-
phys. Biochem. Cytol. 8: 103-133.
Snodgrass, R. E. 1935. Principles of Insect Mor-
phology. McGraw-Hill, New York, 667 pp.
Wood, T. K. 1975, Studies on the function of the
membracid pronotum (Homoptera). II. Histology.
Proc. Entomol. Soc. Washington 77(1): 78-82.
Wood, T. K. and G. K. Morris. 1974. Studies on the
function of the membracid pronotum (Homop-
tera). I. Occurrence and distribution of articulated
hairs. Can. Entomol. 106: 143-148.
Zacharuk, R. Y. 1985. Antennae and sensilla, pp. 1-
69. In Kerkut, G. A. and L. I. Gilbert, eds., Com-
prehensive Insect Physiology, Biochemistry, and
Pharmacology. Vol. 6. Nervous System: Sensory.
Pergamon Press, Oxford.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 153-157
WATER AND TEMPERATURE RELATIONS OF THE PRIMITIVE
XYLOPHAGOUS COCKROACH CR YPTOCERCUS PUNCTULATUS
SCUDDER (DICTYOPTERA: CRYPTOCERCIDAE)
A. G. APPEL AND R. C. SPONSLER
Department of Entomology and Alabama Agricultural Experiment Station, Auburn
University, Alabama 36849-5413.
Abstract. — Percent total body water content, cuticular permeability (CP), rate of water
loss, and critical thermal maxima and minima were determined for the xylophagous
cockroach Cryptocercus punctulatus. There was no difference between female and male
C. punctulatus for any of the measured variables. Mean percent total body water was 75%
and cuticular permeability was 44.1 weg cm~* h~' mm Hg '!. Critical thermal temperatures
were 40.3°C and —7.8°C for maxima and minima, respectively. The CP of C. punctulatus
was similar to that of closely related blattid cockroaches and two sympatric termite species.
Critical thermal temperatures were not similar to those of related taxa.
Key Words:
lations, desiccation
The primitive xylophagous cockroach,
Cryptocercus punctulatus Scudder (Dic-
tyoptera: Cryptocercidae), inhabits moist
decaying hard and soft wood in the southern
and northwestern United States. This species
is considered an example of the close phy-
logenetic relationship between the cock-
roaches and termites (Cornwell 1968). Both
groups have cellulose digesting gut protozoa
and similar proventricular and genital mor-
phology (McKittrick 1964). Within the
Blattaria, C. punctulatus is most closely re-
lated to the Blattidae, the family that con-
tains the pest genera Blatta and Periplaneta.
Not only do C. punctulatus consume
wood, but they live in small social groups
within wood. Colony galleries consist of
networks of horizontal shafts, smaller trans-
verse shafts, and enlarged arena-like areas
(Nalepa 1984). The logs containing colonies
Dictyoptera, Cryptocercus punctulatus, temperature sensitivity, water re-
are quite damp, containing up to 80% mois-
ture by weight (Appel pers. comm.). These
cockroaches are primarily found in their
galleries, but occasionally occur in the soil
and leaf litter near decaying wood. Thus, C.
punctulatus lives in a moist environment,
buffered from rapid humidity and temper-
ature changes.
The purpose of this study was to examine
aspects of the water relations and temper-
ature tolerance of C. punctulatus and to
compare these parameters with those of
closely related cockroaches and termites.
MATERIALS AND METHODS
Specimens of C. punctulatus were ob-
tained 7 March 1987 and 12 January 1988
from decaying logs in Cleburne Co., Ala-
bama, off U.S. 431, 2.8 km SE of the Cle-
burne-Calhoun Co. line (G. W. Folkerts,
154
Department of Zoology and Wildlife Sci-
ence, Auburn University, pers. comm.).
Adults, nymphs, and wood were returned
to the laboratory for study.
Water relations experiments. — The water
content, cuticular permeability (CP), and
percent of total body water (% TBW) lost
over time was determined gravimetrically
with a digital balance (0.01 mg sensitivity)
(Appel et al. 1983, Mack and Appel 1986).
Percent TBW was calculated as the differ-
ence between the initial mass of live spec-
imens and their mass after drying, divided
by the initial live mass multiplied by 100.
Dry mass was determined after the speci-
mens were dried at ca. 50°C for at least three
days or until two successive weighings did
not differ >0.01 mg.
CP was calculated as the ug of water lost
per unit surface area (cm”’) per unit time (h)
per unit saturation deficit (mm Hg) ina des-
iccator chamber. Individual cockroaches
were placed in uncovered 30-ml plastic cups
that were put into an 1|1-liter desiccator jar
maintained at 30 + 0.1°C and 0-2% RH.
The conditions in the desiccator were mon-
itored with a Cole-Parmer LCD Digital Hy-
grometer (Model 3309-50). The hygrometer
sensor was positioned at the same height in
the desiccator as the specimens. Surface area
was estimated for each specimen by Meeh’s
formula: S = 12-M”, where S = surface area
(cm?) and M = initial mass (g) (Edney and
McFarlane 1974). Mass losses were mea-
sured at 2, 4, 6, 8, 10, and 24 h, but cuticular
permeability was only calculated from 2
h-data. Preliminary experiments showed no
significant difference in the amount of water
lost by live and HCN-killed cockroaches at
2 h. Therefore, 2-h respiratory water was
less than the sensitivity of the balance (0.01
mg) and cuticular permeability could be cal-
culated from live cockroaches without com-
plications of respiratory mass losses. Hourly
mass loss and % TBW was used to calculate
% TBW lost over time.
Temperature relations experiments. —
Critical thermal maxima (CTMax) and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
minima (CTMin) were defined as the upper
or lower temperatures, respectively, that in-
duced reversible knock-down when tem-
peratures were rapidly increased (CTMax)
or decreased (CTMin) at 1°C/min and 100%
RH. Adult female and male C. punctulatus
were used for CTMax and CTMin experi-
ments. To determine CTMax, a 130-ml glass
jar containing about 120 ml of water was
sealed with a metal lid and placed into a
0.45-liter glass jar. A 5.5-cm-diameter plas-
tic petri dish bottom containing a mois-
tened disk of Whatman No. 2 filter paper
was placed on top of the inner jar and 65
ml of water added to the space between the
two jars. One cockroach was placed in the
petri dish and the 0.45-liter jar was sealed
with a metal lid and band. A small hole was
cut in the center of the lid and a copper-
constantan thermocouple (0.74-mm diam-
eter) connected to a digital thermometer
(Sensortek BAT-12) was threaded through
the hole to contact the bottom of the petri
dish. The thermocouple wire was attached
to a 15-cm wooden applicator to ease ma-
nipulation. Body temperature and response
to probing were determined by gently
touching the thermocouple to the mem-
brane between the cockroaches’ meso- and
metathoracic coxae. The jar-probe appa-
ratus was placed on a hot plate that was
adjusted to increase the temperature inside
the apparatus at 1°C/min. Cockroaches were
judged as knocked down when they lost co-
ordination and the ability to right them-
selves. Recovery was assessed after 1 min
at 22.8°C.
CTMin were measured in a covered 5.5-
cm-diameter plastic petri dish containing a
moistened disk of filter paper. A small hole
was cut in the petri dish cover to allow ac-
cess of the same thermocouple and probe
as used for the CTMax determinations. A
chill table was used as the cooling device
and a constant rate of 1°C/min decline in
temperature was maintained by varying the
height of the petri dish above the chill table.
Temperatures were measured as above, but
VOLUME 91, NUMBER 2
knock down was defined as the lack of leg
movement when probed. Recovery was as-
sessed after 15 min at 22.3°C. Leg move-
ment was used in CTMin experiments be-
cause chilled cockroaches were not always
immobilized on their dorsum as with
CTMax.
Statistical analysis.—A completely ran-
domized design and a Student’s f-test (SAS
Institute 1982) were used to determine sig-
nificant differences in initial mass, % TBW,
and cuticular permeability between sexes.
Regression was used to determine if cuti-
cular permeability was related to initial live
mass or % TBW, and how % TBW loss was
related to time. Data are expressed as means
+ SE. Completely randomized designs with
sex as the factors were used for CTMax and
CTMin. Student’s t-tests (P < 0.05) were
used to determine significance. Data are ex-
pressed as means + SE.
RESULTS AND DISCUSSION
Masses of C. punctulatus nymphs and
adults ranged from 133.9 to 876.31 mg for
second instars and adults, respectively.
There was no difference between the masses
of adult females (829.76 + 15.78 mg) and
males (781.83 + 17.93 mg). All stages con-
tained about 75% TBW and there was no
difference between the % TBW of adult fe-
males and males (Table 1). Edney (1977)
summarized the % TBW ofa variety of ar-
thropods (45-92% TBW) and Appel et al.
(1983) reported a range of 62.7 to 71.8%
TBW for ten cockroach species. The more
primitive blattid species generally had
greater % TBW than blattellid or blaberid
species. Although the significance of % TBW
is unknown, the close phylogenetic rela-
tionship of C. punctulatus with the blattids
may relate to the similarity of % TBW
among these species. In addition, the % TBW
of two sympatric termite species, the For-
mosan subterranean termite, Coptotermes
formosanus Shiraki and the eastern subter-
ranean termite, Reticulitermes flavipes Kol-
ler (both Rhinotermitidae), ranged from 62.3
155
50 —
@
@ y = 1.78x + 2.73, 2 = 0.99 =e
40 }
O Y = 1.85X + 1.51, r2 = 0.99
Percent Total Body Water Lost
Time (h)
Fig. 1. Cumulative % TBW lost by female (O) and
male (@) Cryptocercus punctulatus during desiccation
at 30°C and 0-2% RH.
to 75.9 (Sponsler and Appel unpublished
data).
Percent TBW loss of female (F = 32,502.1,
P = 0.0001) and male (F = 2340.1, P =
0.0001) C. punctulatus increased linearly
with desiccation time (Fig. 1). There was a
significant difference between the rate of %
TBW loss (slope +2 SE) of female and male
C. punctulatus. This difference in rate prob-
ably resulted in the significant difference in
24-h mortality (Table 1). The greater mean
mass of females, approximately 120 mg or
90 mg of water, accounts for the observed
difference in desiccation-induced mortality.
Linear rates of % TBW loss have been re-
ported for cockroaches (Appel et al. 1983),
termites (Sponsler and Appel unpublished
data), and other insects (Mack and Appel
1986, Mack et al. 1988).
Cuticular permeability determines the rate
of water loss. There was no relationship be-
tween the CP of female or male C. punc-
tulatus and initial mass (F = 0.12; df = 1;
P = 0.74 and F = 0.00; df = 1; P = 0.99,
respectively). Similarly, there was no rela-
tionship between CP and % TBW for males
(F = 1-92; df= 1; P = 0:20), The CP of
females, however, declined linearly (F =
6.47; df = 1; P = 0.03) with % TBW (CP =
—0.91 % TBW + 111.69; r? = 0.45). Appel
et al. (1986) found that water loss of adult
male Periplaneta fuliginosa (Serville) was
156
Table 1.
and male C. punctulatus [x + SE; n = 20].
Sex Initial Mass (mg)
Female 596.69 + 78.20
Male 476.11 + 67.70
«yg cm? h"' mm Hg".
negatively related to body lipid content. The
negative relationship observed with female
C. punctulatus may also be related to lipid
content and factors such as age and repro-
ductive status that affect lipid content.
There was no difference between the CP
of female and male C. punctulatus (Table
1). The CP of C. punctulatus (44.1 ug cm~?
h~' mm Hg _') was similar to those of blat-
tid cockroaches such as Periplaneta amer-
icana (L.), Periplaneta australasiae (Fab.),
Blatta orientalis (L.), and Periplaneta brun-
nea Burmeister with CP values of 53.7, 43.1,
43.4, and 41.7 ug cm-* h-'! mm Hg "|, re-
spectively (Appel et al. 1983). The CPs of
worker C. formosanus and R. flavipes were
also similar to C. punctulatus (37.5 and 27.8
ugcm~*h~'mm Hg |, respectively) (Spon-
sler and Appel unpublished data). Arthro-
pods with CPs of approximately 40 gener-
ally inhabit hygric environments such as
tropical forests, moist wood, and sewers
(Edney 1977).
Even though the CPs of female and male
C. punctulatus were not different, the rates
of % TBW loss (see above) were signifi-
cantly different. Differences in initial mass
explain this apparent discrepancy. Al-
though both sexes had the same CP (44.1
ug cm~* h~' mm Hg~!), there was signifi-
cantly more water in females than in males.
Therefore, females lose a lower percentage
of their % TBW than males at any given
time. —
Table 2.
Sex CTMax
Female 39.6 + 0.28 (39.0-40.5)
Male 40.9 + 0.21 (39.7-40.9)
% TBW
74.87
76.01
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Initial masses, percent total body water (% TBW), cuticula permeability, and mortality of female
Cuticular Permeability* Mortality at 24 h
1.39
0.66
43.52 + 1.89
44.73 + 3.25
30%
90%
tame
There was no difference between the crit-
ical thermal temperatures of female and
male C. punctulatus. Critical thermal tem-
peratures ranged from 40.9 to —8.5°C for
CTMax and CTMin, respectively (Table 2).
The CTMax of C. punctulatus is the lowest
recorded for cockroaches. Previously re-
ported cockroach CT Max ranged from 47.6
to 51.4°C (Appel et al. 1983); the CTMax
of C. punctulatus was nearly 7.5°C lower
than that of the next lowest cockroach
species. Positive correlations between
CTMax and habitat temperatures have been
established for desert cockroaches (Cohen
and Pinto 1977, Cohen and Cohen 1981),
desert termites (Rust et al. 1979), and mel-
oid beetles (Cohen and Pinto 1977). Appel
et al. (1983) found that the CTMax of ten
cockroach species, representing three fam-
ilies, could be divided into four statistically
distinct groups: <48.1°C, 48.7-49.1°C,
49.3-49.5°C, and >49.6°C. There was no
apparent phylogenetic relationship in the
ability to tolerate high temperatures, and
our results with C. punctulatus support this
conclusion.
Few studies have reported the CTMin of
insects. Sponsler and Appel (unpublished
data) determined the CTMin for soldier and
worker C. formosanus and R. flavipes. The
mean CTMin for these termites was 13.2°C
and there was no difference between stages
or species. Although collected at the same
time of year as these termites, the CTMin
Critical thermal temperatures (°C) of adult C. punctulatus [x + SE (minimum—maximum); n = 10].
CTMin
—§8.5 + 0.52 (—10.0——7.2)
=7al 0:55; (—9:0-——7,.1)
VOLUME 91, NUMBER 2
of C. punctulatus ranged from —10.0 to
—7.1°C. In addition, C. punctulatus remain
active at —5.0 to 0°C (Appel unpublished
data). Preliminary studies with P. ameri-
cana and B. orientalis indicated that the
CTMin of these cockroaches was much
higher (ca. 10°C) than that of C. punctulatus.
Differences in temperature tolerance be-
tween C. punctulatus and sympatric ter-
mites are likely due to differences in micro-
habitats. Cryptocercus punctulatus live in
moist, fallen, partially buried logs in dense
forests (Seelinger and Seelinger 1983, Na-
lepa 1984). These logs provide a habitat in-
sulated by moisture. With leaf littered soil
as a heat-sink, these logs maintain lower
than ambient temperatures (Appel unpub-
lished data). During hot spells, evaporation
cools the logs. It is also possible that the
cockroaches move to preferred cooler lo-
cations within their extensive galleries.
Rhinotermitid termites, however, usually
maintain soil contact even when they forage
on surface debris. Since most colonies are
located well below the soil surface (and frost
line) and are composed of thousands of met-
abolically active individuals, these termites
are probably exposed to more constant,
warm temperatures than are log inhabiting
C. punctulatus.
Similar CPs among C. punctulatus, blat-
tid cockroaches, and termites may reflect
their similar high-humidity habitats or close
phylogenetic relationships. The interstitial
spaces in wood or soil containing >17%
moisture are at saturated humidity and do
not contribute to significant body water loss.
Since all of these groups inhabit moist areas,
it 1s not surprising that they all have similar
CPs. Our results concur with the conclu-
sions of Appel et al. (1983) that, in cock-
roaches, CP is related to habitat moisture
and temperature sensitivity is related to
habitat temperature.
ACKNOWLEDGMENTS
We thank George W. Folkerts, Depart-
ment of Zoology and Wildlife Science, Au-
157
burn University, for collecting the C. punc-
tulatus and for reviewing the manuscript.
Alabama Agricultural Experiment Station
Journal Series No. 17-881834P.
LITERATURE CITED
Appel, A. G., D. A. Reierson, and M. K. Rust. 1983.
Comparative water relations and temperature sen-
sitivity of cockroaches. Comp. Biochem. Physiol.
74A: 357-361.
1986. Water relations of the smokybrown
cockroach, Periplaneta fuliginosa. J. Insect. Phys-
iol. 32: 623-628.
Cohen, A. C. and J. L. Cohen. 1981. Microclimate
temperature and water relations of two species of
desert cockroach. Comp. Biochem. Physiol. 69A:
165-167.
Cohen, A. C. and J. D. Pinto. 1977. An evaluation
of xeric adaptiveness of several species of blister
beetles (Meloidae). Ann. Entomol. Soc. Am. 70:
741-749.
Cornwell, P. B. 1968. The Cockroach, Volume I.
Hutchinson & Co. LTD, London. 391 pp.
Edney, E. B. 1977. Water Balance in Land Arthro-
pods. Springer, New York. 282 pp.
Edney, E. B. and J. McFarlane. 1974. The effects of
temperature on transpiration in the desert cock-
roach, Arenivaga investigata and in Periplaneta
americana. Physiol. Zool. 47: 1-12.
Mack, T. P. and A. G. Appel. 1986. Water relations
of immature and adult lesser cornstalk borers,
Elasmopalpus lignosellus (Lepidoptera: Pyrali-
dae). Ann. Entomol. Soc. Am. 79: 579-582.
Mack, T. P., A. G. Appel, C. B. Backman, and P. J.
Trichilo. 1988. Water relations of several ar-
thropod predators in the peanut agroecosystem.
Environ. Entomol. 17: 778-781.
McKittrick, F.A. 1964. Evolutionary studies of cock-
roaches. Cornell Univ. Agric. Exp. Sta. Mem. 389:
1-197.
Nalepa, C. A. 1984. Colony composition, protozoan
transfer and some life history characteristics of the
woodroach Cryptocercus punctulatus Scudder
(Dictyoptera: Cryptocercidae). Behav. Ecol. So-
ciobiol. 14: 273-279.
Rust, M. K., D. A. Reierson, and R. H. Scheffrahn.
1979. Comparative habits, host utilization and
xeric adaptations of the southwest drywood ter-
mites, Jncisitermes fruticavus Rust and Jncisi-
termes minor (Hagen) (Isoptera: Kalotermitidae).
Sociobiology 4: 239-255.
SAS Institute. 1982. SAS user’s guide: Statistics. SAS
Institute, Cary, N.C.
Seelinger, G. and U. Seelinger. 1983. On the social
organisation, alarm and fighting in the primitive
cockroach Cryptocercus punctulatus Scudder. Z.
Tierpsychol. 61: 315-333.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 158-163
FIRST NEARCTIC RECORD OF THE GENUS NORDLANDERIA
(HYMENOPTERA: EUCOILIDAE), WITH DESCRIPTIONS
OF TWO NEW SPECIES
TERRY D. MILLER
Department of Entomology, Washington State University, Pullman, Washington 99164.
Abstract. —The genus Nordlanderia (Hymenoptera: Cynipoidea, Eucoilidae) is reported
from the Nearctic region for the first time and two new species are described. They differ
from their African counterparts by the absence of coriaceous sculpturing below the meso-
pleural carina and the presence of apical punctulations on the abdominal tergites. This
genus is apparently restricted in distribution to arid areas of western North America and
southern Africa.
Key Words:
coilidae
Quinlan (1986) described the genus Nord-
landeria and three new species from ma-
terial collected in southern Africa. The ge-
nus is distinguished by the presence of
triangular projections on the supraclypeal
area and the anterior region of the face. This
is consistent with the general tendency to-
ward facial projections in the Gronotoma
group of genera. This genus, previously
known only from Africa, can be added to
the known Nearctic fauna with the discov-
ery of the two new species described.
METHODS AND TERMS
Descriptions and measurements are
largely based on scanning electron micro-
graphs due to the small size of the specimens
(about | mm total length) and the predom-
inately reflective integument. Terms for sur-
face sculpture follow Harms (1979) and
morphological terms are after Richards
(1977), Quinlan (1978) and Nordlander
(1982).
Type material will be deposited in the
following institutions: British Museum
Distribution, southern Africa, western North America, Cynipoidea, Eu-
(Natural History); California Academy of
Sciences; and the University of Idaho.
Nordlanderia merickeli Miller,
NEw SPECIES
Figs. 1-6
Female.—1.15 mm long. Head and tho-
rax black, antennae dark brown, gaster dark
red-brown, legs yellow-brown and wings
hyaline.
Head subcircular in frontal view. Com-
pound eyes normal size, weakly converging
ventrally and with several very short scat-
tered hairs between facets. Vertex smooth,
with few hairs; ocelli of moderate size; pos-
terior ocellar line longer than lateral ocellar
line which is longer than ocular ocellar line.
Occiput smooth, with few setiferous punc-
tures. Face smooth, with scattered hairs be-
low toruli and line of hairs along the inner
orbital margins. Frons slightly raised. Sub-
ocular sulcus distinct, with a few poorly de-
fined striae above and below (Fig. 1). Cheeks
below sulcus and supraclypeal area strongly
VOLUME 91, NUMBER 2
protruding, forming three spine-like keels
(Figs. 1, 2, 4). Mandibles bidentate; inner
tooth two-thirds length of outer tooth (Fig.
1). Antennae 13-segmented, clavate, with
very weakly defined 8-segmented club, seg-
ment 3 equal in length to segment 4.
Thorax convex in lateral view. Pronotal
plate not protruding; median bridge width
to plate width ratio, 13:35; lateral margins
parallel; foveae on lateral margins open (Fig.
3). Propleuron smooth, with scattered long
hairs except on anterior margin where ves-
titure is both dense and long (Fig. 2). Meso-
scutum smooth, with line of hairs replacing
the notali. Scutellum with lateral bars
smooth; scutellar plate longer than wide,
with large posterior fovea and margins of
plate bordered by setiferous punctures (Fig.
5); dorsal surface of scutellum above margin
carinate-rugose, with circle of inwardly bent
hairs around plate; scutellum below margin
with widely spaced and regularly radiating
striae (Fig. 6). Mesopleural carina complete;
pleuron above and below suture smooth;
area anterior to subalar pit depressed (Fig.
2). Metanotal plate indented posteriorly, as
wide as distance between the propodeal ca-
rinae (Fig. 5). Metapleuron smooth; meta-
pleural groove absent; pleuron depressed
dorsal to obsolete ridge | (Fig. 2); lobe over
propodeal spiracle wing-like, nearly 2x as
long as wide (Fig. 6). Propodeal carinae na-
ked, parallel anteriorly and converging to-
wards nucha posteriorly; lateral carinal ex-
tensions continuing to near spiracle (Figs.
5, 6); propodeal surface smooth, with few
159
hairs between the carinae, and densely pu-
bescent laterally (Fig. 5). Legs normal and
moderately pubescent.
Gaster with segment | obscured by seg-
ment 2 when viewed laterally; segment |
forming narrow ring; tergite 2 with a basal
ring of hairs, lacking any felt-like pubes-
cence; tergite 2 the largest, occupying whole
lateral surface of gaster; posterior margins
of segment 2—4 punctulate; punctulations of
tergite 2 faint.
Wing surface pubescent, with marginal
ciliation of moderate length; second radial
abscissa noticeably longer than first radial
abscissa; radial cell closed on wing margin;
submarginal vein distinct; median, discoi-
dal and subdiscoidal veins barely discern-
ible in some specimens.
Male.—1.18 mm long. Antennae 15-seg-
mented, filiform; segment 3 equal in length
to segment 4, very slightly curved, not swol-
len distally. Habitus similar to female, dif-
fering in that metapleural lobe over pro-
podeal spiracle nearly as wide as long; medial
area of propodeum densely pubescent; fa-
cial protuberances not generally as well de-
veloped.
Material examined.— U.S.A. Idaho: Nez
Perce Co., Hells Gate State Park, T. D. Mil-
ler collector; 13-V-83 (1 male), 5-VI-83 (1
male, | female), 20-VI-83 (1 male), 15-VII-
83 (1 male), 20-VII-83 (3 females), 23-VII-
83 (1 male, | female). Washington: Asotin
Co., 9 mi. W. Clarkston, T. D. Miller col-
lector; 20-V-84 (1 male).
Types.— Holotype female, allotype male,
—
Figs. 1-6. Nordlanderia merickeli. 1, Frontal view of female head (scale line 75 um). 2, Lateral view of female
head and thorax (scale line 80 um). 3, Pronotal plate of male (scale line 30 um). 4, Dorsal view of female facial
projections (scale line 30 um). 5, Posterior view of female thorax (scale line 150 um). 6, Lateral view of female
scutellum (scale line 35 um). (mb = median bridge; Ic = lateral carinae; mpl = metapleural lobe.)
Figs. 7-12. Nordlanderia navajoae. 7, Frontal view of male head (scale line 75 um). 8, Anterior view of male
head and thorax (scale line 100 um). 9, Dorsal view of female head (scale line 105 um). 10, Pronotal plate of
male (scale line 87 um). 11, Lateral view of female thorax (scale line 135 wm). 12, Posterior view of female
thorax (scale line 150 um). (mb = median bridge; rl = metapleural ridge 1; st = scutellar tubercle; mnp =
metanotal plate.)
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
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162 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and 2 female paratypes to be deposited in
the California Academy of Sciences; | male
and | female paratype to be deposited in
the British Museum (Natural History); 1
male and | female paratype gold coated on
scanning electron microscope (S.E.M.) stubs
and 2 uncoated males and | female paratype
deposited in the University of Idaho.
Etymology.—This species is named in
honor of Frank W. Merickel of the Univer-
sity of Idaho for his valued friendship and
support over the years.
Remarks.—This species is distinguished
from the other new Nearctic species, na-
vajoae, by the broader median bridge of the
pronotal plate, size of the metanotal plate,
complete lateral extensions of the propodeal
carinae, lack of sculpture on the mesopleu-
ron, ridge | absent on the metapleuron, and
the well developed lobe over the propodeal
spiracle.
Nordlanderia nayajoae Miller
New SPECIES
Figs. 7-12
Female.—1.1 mm long. Antennae, head,
thorax and gaster black, legs yellow-brown
and wings hyaline.
Head subcircular in frontal view. Com-
pound eyes normal size, not protruding,
nearly parallel. Vertex smooth, with scat-
tered setiferous punctures, ocelli of mod-
erate size; posterior ocellar line longer than
lateral ocellar line which is longer than ocu-
lar ocellar line (Fig. 9). Occiput smooth, with
few hairs. Face smooth, with scant vestiture
below toruli and irregular line of hairs along
inner ocular margin. Frons slightly protrud-
ing. Subocular sulcus distinct, with well de-
fined striae above and below, cheeks com-
pletely striate (Fig. 7). Cheeks below
subocular sulcus and supraclypeal area
moderately protruding to form 3 tooth-like
projections (Figs. 7, 8, 9). Mandibles biden-
tate; inner tooth three-fourths length of out-
er tooth (Fig. 7). Antennae 13-segmented,
clavate, with weakly defined 10-segmented
club; segment 4 greater in length than seg-
ment 3.
Thorax convex in lateral view. Pronotal
plate protruding slightly; median bridge
width to plate width ratio, 9:55; lateral mar-
gins of plate curved, not parallel; foveae on
lateral margins open (Fig. 10). Propleuron
smooth, with scattered long hairs except an-
terior margin which is covered with short,
dense hairs (Figs. 8, 11). Mesoscutum
smooth; notali obsolete, replaced by line of
hairs in some specimens. Scutellum with
lateral bars smooth; scutellar plate longer
than wide, with large posterior fovea and
lateral margins with single puncture at mid-
length; anterior of lateral puncture is small,
setiferous tubercle (Fig. 12), setae visible
only in perfect specimens; dorsal surface of
scutellum above margin areolate-rugose,
with few hairs, scutellum below margin car-
inate-rugose (Figs. 11, 12). Mesopleural ca-
rina complete; area below suture smooth,
light striations present above suture ante-
riorly and just below subalar pit (Figs. 8,
11). Metanotal plate indented posteriorly,
not as wide as distance between the pro-
podeal carinae (Fig. 12). Metapleuron
smooth; metapleural groove absent, ridge 1
distinct (Fig. 11); lobe over propodeal spi-
racle short, wider than long. Propodeal ca-
rinae parallel anteriorly and converging to-
wards nucha posteriorly; lateral carinal
extensions poorly developed, not reaching
spiracles (Fig. 12); propodeal surface
smooth, moderately pubescent, except for
naked carinae. Legs normal, moderately pu-
bescent.
Gaster with segment | obscured by seg-
ment 2 when viewed laterally; segment |
forming narrow ring; tergite 2 with basal
ring of hairs, lacking any felt-like pubes-
cence; tergite 2 the largest, occupying whole
lateral surface of gaster; posterior margins
of tergites 3 and 4 punctulate.
Wing surface pubescent, with short mar-
ginal ciliation; forewing broad, apically
rounded; second radial abscissa slightly
VOLUME 91, NUMBER 2
longer than first radial abscissa; radial cell
closed on wing margin.
Male.—1.1 mm long. Antennae 15-seg-
mented, filiform; segment 3 curved and
swollen distally, greater in length than seg-
ment 4. Habitus similar to female, differing
in that sculpturing of dorsal scutellar surface
is less well defined.
Material examined.—U.S.A. Arizona.
Apache Co., | mi. south of Ganado, T. D.
Miller and F. W. Merickel collectors; 20-
VI/ 1-VII-85 (2 females, 1 male).
Types. — Female holotype to be deposited
in the California Academy of Sciences; |
male and | female paratype gold coated on
S.E.M. stubs deposited in the University of
Idaho.
Etymology.— This species was named for
the Navajo Indians upon whose tribal lands
this species was discovered.
Remarks.—This species is distinguished
from merickeli by the distinct striations on
the cheeks and mesopleuron, the setiferous
tubercles on the scutellar plate, metapleural
ridge | present, the narrow median bridge
of the pronotal plate, absence of lateral pro-
podeal carinae, and the differences in an-
tennal characters.
DISCUSSION
The distribution of Nordlanderia in North
America is apparently restricted to arid areas
of the western United States. In addition to
the two new species described in this paper
several undescribed species await further
study. All Nearctic Nordlanderia species ex-
amined so far, including the undescribed
163
ones, differ from their African counterparts
due to the absence of coriaceous sculpturing
below the mesopleural carina and the pres-
ence of apical punctulations on the abdom-
inal tergites.
ACKNOWLEDGMENTS
I am grateful for the help of Mr. John
Quinlan, British Museum (Natural Histo-
ry), for helpful comments and comparisons
with type material in their collection and to
Dr. Goran Nordlander, Swedish University
of Agricultural Sciences, for the many very
helpful comments on the manuscript. Ap-
preciation 1s also extended to Dr. William
J. Turner, Washington State University and
to Dr. James B. Johnson and Mr. Frank W.
Merickel of the University of Idaho for their
review of the manuscript.
LITERATURE CITED
Harris, R. A. 1979. A glossary of surface sculpturing.
Occas. Pap. Ent., State Calif., Dep. Food Agric.
28. 31 pp.
Nordlander, G. 1982. Systematics and phylogeny of
an interrelated group of genera within the family
Eucoilidae (Insecta: Hymenoptera, Cynipoidea).
Dissertation, Univ. of Stockholm, Dept. of Zo-
ology, Stockholm. 32 pp.
Quinlan, J. 1978. Hymenoptera: Cynipoidea; Eu-
coilidae. Handbk. Ident. Br. Insects, Royal Ent.
Soc. London, 8(1b): 1-58.
1986. A key to the Afrotropical genera of
Eucoilidae (Hymenoptera), with a revision of cer-
tain genera. Bull. Br. Mus. (Nat. Hist.) Ent. Ser.
52(4): 243-366.
Richards, O. W. 1977. Hymenoptera. Introduction
and Keys to Families, 2nd ed. Handbk. Ident. Br.
Insects, Royal Ent. Soc. London, London, 6(1): 1-
100.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 164-168
HOST PLANTS OF NEASPILOTA IN CALIFORNIA
(DIPTERA: TEPHRITIDAE)
RICHARD D. GOEDEN
Department of Entomology, University of California, Riverside, California 92521.
Abstract. —New host-plant records from California are reported for the tephritids, Nea-
spilota achilleae Johnson, N. aenigma Freidberg and Mathis, N. albiseta Freidberg and
Mathis, N. appendiculata Freidberg and Mathis, N. caillistigma Freidberg and Mathis, N.
stecki Freidberg and Mathis, and N. viridescens Quisenberry. Neaspilota stecki is initially
reported from California. The host plants of these tephritids are mainly in the tribe
Astereae of Asteraceae, although N. achilleae was reared for the first time from the western
United States and from capitula of Stephanomeria virgata Bentham in the tribe Cichoreae.
Besides new records from 13 species in six genera of Astereae for N. viridescens, another
record for this tephritid from Lepidospartum squamatum (Gray) Gray is the first from
the tribe Senecioneae for any Neaspilota.
Key Words:
Recent revision of the genus Neaspilota
(Diptera: Tephritidae) by Freidberg and
Mathis (1986) facilitated determination of
specimens that I had reared from capitula
of California Asteraceae (= Compositae) and
enabled this report on the host-plant rela-
tions of this taxon. I hereby respond to their
concern that “... Confirmed rearing rec-
ords are especially scarce from western
United States, where half of the known
species of Neaspilota occur.”
MATERIALS AND METHODS
Host-plant information on Neaspilota was
obtained since 1980 mainly in southern
California, an ecologically diverse region
that is defined and treated botanically by
Munz (1974) and encompasses roughly the
southern third of the State. The materials
and methods used in sampling mature ca-
pitula and rearing Tephritidae from sam-
ples were described by Goeden (1985).
Sweep-net collections supplemented rear-
ings. Flies were identified with keys by
Diptera, Tephritidae, host plants, Neaspilota, subgenus Neorellia, Asteraceae
Freidberg and Mathis (1986). Selected spec-
imens were confirmed by F. L. Blanc, Cal-
ifornia Department of Food and Agriculture
Sacramento, and by A. Freidberg, Tel-Aviv
University, Israel. All host-plant identifi-
cations were confirmed by A. C. Sanders,
Curator of the Herbarium of the University
of California, Riverside. Plant names follow
Munz and Keck (1959) and Munz (1974).
RESULTS AND DISCUSSION
Freidberg and Mathis (1986) reviewed the
sparse information available on the natural
history and host plants of Neaspilota. All
known species develop in flower or seed
heads of Asteraceae without forming galls.
I offer the following information on the host
plants of Neaspilota species in California.
Neaspilota achilleae Johnson.—This
species was reared from three separate sam-
ples of capitula of Stephanomeria virgata
Bentham collected in southern San Diego
County as follows: 2 é and 2 2, SE of Barrett
Junction, 8-X-1987; 7 46 and 6 9, Bratton
VOLUME 91, NUMBER 2
Valley, 20-X-1987; 2 6, Deerhorn Valley,
20-X-1987. This is a new host-plant genus
for N. achilleae and the first rearing record
for this fly from the western United States.
Freidberg and Mathis (1986) list four
species of Aster, three of Chrysopsis, three
of Erigeron, four of Hieraceum, and one
species each of Prenanthes, Sericocarpus,
and Trilisa as hosts of N. achilleae from
eastern United States. I have reared other
genera and species, but not this tephritid,
from one to five samples each of 10 species
of Aster, three of Chrysopsis, nine of Erig-
eron, and two of Hieraceum from Califor-
nia. In addition, I have not reared N. achil-
leae from one sample of Stephanomeria
cichoriacea Gray, three samples of Steph-
anomeria exigua Nuttall, nor six samples
of Stephanomeria pauciflora (Torrey) Nut-
tall. Hieraceum, Stephanomeria, and Trili-
sa belong to the tribe Cichorieae (= Lac-
tuceae); Aster, Chrysopsis, and Erigeron, to
the tribe Astereae. Lack of rearing records
and host-plant tribal affinities suggest that
the “peculiar,” apparently discontinuous,
eastern and western distributions of N.
achilleae by Freidberg and Mathis (1986)
may represent allopatric populations with
different host-plant specificities. They also
reported that specimens examined from
California and Arizona were smaller on the
average than their eastern counterparts,
which may be added evidence of allopatry.
The record for N. achilleae “‘taken ... on
Heracleum sp.” (Umbelliferae) in Phillips
(1946) probably represents adults collected
from a non-host. Sweep records from As-
teraceae may be poor indicators of the host-
plant affinities of nonfrugivorous, nongal-
licolous Tephritidae (Goeden 1987, and
unpublished data). The western populations
of N. achilleae appear more stenophagous
than the eastern populations, possibly mo-
nophagous, and, therefore, could have been
derived from eastern ancestral stock via Ci-
choreae, and could be a separate species.
Neaspilota aenigma Freidberg and Math-
is.—Only five males have previously been
165
found in California, including the holotype
and three allotypes swept in Inyo County
(Freidberg and Mathis 1986). I have since
reared the following specimens from capit-
ula samples: Acamptopappus shockleyi Gray,
7 6 and 6 2, SE end of Kingston (Mountain)
Range, NE San Bernardino Co., 27-V-1982;
Lessingia lemmonii Gray, 1 °, Lockwood
Valley, Los Padres Nat. Forest, Ventura Co.,
at ca. 1700 m elev., 31-VIII-1986; 1 8, Mo-
jave River Forks, San Bernardino Nat. For-
est, SW San Bernardino Co., 13-VIII-1987;
Machaeranthera tortifolia (Gray) Cronquist
and Keck, 2 4, along Death Valley Road,
Last Chance Mountain Range, at 1130 m
elev., Inyo Co., 9-VI-1987.
Freidberg and Mathis (1986) reported its
host plants as Haplopappus gooddingii (A.
Nelson) Munz and Johnston, and possibly
H. hartwegi (Gray) Blake. I have not yet
sampled H. gooddingii or reared N. aenigma
from 17 other California species of Hap-
lopappus sampled. Haplopappus hartwegi
does not occur in California (Munz and Keck
1959, Munz 1974), and may be an invalid
species name (A. C. Sanders in litt. 1988).
Still, all four host genera mentioned above
belong to the tribe Astereae (Munz and Keck
1959), and, thus, constitute a definable
grouping of host Asteraceae as reported for
other oligophagous Tephritidae, e.g. certain
California species of Trupanea (Goeden
1985) and Urophora (Goeden 1987). Freid-
berg and Mathis (1986) noted that the ma-
jority of known hosts of Neaspilota spp. in
the subgenus Neore/lia, with which the pres-
ent report is concerned, belong to the As-
tereae.
Neaspilota albiseta Freidberg and Math-
is.—Freidberg and Mathis (1986) listed the
host plants of this southwestern species as
unknown. The following is the first host-
plant rearing record for N. albiseta: 8 males
and 2 females reared from the same sample
of mature heads of Machaeranthera torti-
folia from along Death Valley Road in the
Last Chance Mountains of Inyo County in
1987 that also yielded N. aenigma. Freid-
166 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
berg and Mathis (1986) noted that some
plant species are hosts to two or three species
of Neaspilota. My rearing record confirms
this co-occurrence in capitula from one lo-
cal host-plant species population. Synphagy
also is acommon mode of resource sharing
in other genera of capitula-feeding Tephriti-
dae, e.g. Trupanea and Urophora spp. (Goe-
den 1985, 1987).
Neaspilota appendiculata Freidberg and
Mathis. —Freidberg and Mathis (1986) also
listed the host plants of this species as un-
known. The following are the first rearing
records for N. appendiculata: Corethrogyne
filaginifolia (Hooker and Arnott) Nuttall, 1
2, N of Thomas Mountain, San Bernardino
Nat. Forest, Riverside Co., 29-IX-1982, 3
é and 5 9, Big Sandy Bluff, Sierra Nat. For-
est, Fresno Co., 235000 23-VIII-1988: Ma-
caeranthera canescens (Pursh) Gray, 2 6 and
10 2, Onyx Peak, San Gorgonio Mountains,
San Bernardino Nat. Forest, SE San Ber-
nardino Co., 22-V-1987. Both hosts are As-
tereae.
Neaspilota brunneostigma Doane.—
Neaspilota brunneostigma, as revised by
Freidberg and Mathis (1986), has not been
found in California. Therefore, all of the
host-plant records for this species listed in
Wasbauer (1972) remain unconfirmed.
Neaspilota callistigma Freidberg and
Mathis.—The only host recorded for this
species is Haplopappus venetus (Humboldt,
Bonpland, Kunth) Blake var. vernonioides
(Nuttall) Haller (Foote and Blanc 1963,
Freidberg and Mathis 1986), which I con-
firmed when I reared one female from ca-
pitula collected near the Stanton Ranch Air-
field on Santa Cruz Island, Santa Barbara
Co., 13-X-1983. This female was identified
as Neaspilota sp. in Goeden (1986). I sub-
sequently reared two males and 16 females
from capitula of H. venetus collected in
Marrow Valley, SE of Dulzura, San Diego
Co., 20-X-1987. My other rearing records
are from H. acradenius (Greene) Blake: 1 4
and | 9, Coyote Wells, E of Ocotillo,
100-m elev., SW Imperial Co., 6-XI-1986;
3 4, Sentenac Canyon, San Diego Co.,
7-X-1987; 10 6 and 5 2, Mountain Springs
Pass, SE San Diego Co., 7-X-1987. I have
reared other tephritid genera and species,
but not N. callistigma from one to five sam-
ples each of 15 other species of Haplopap-
pus, therefore, this tephritid may be nearly
monophagous.
Neaspilota stecki Freidberg and Math-
is.—Heretofore known only from 6 males
swept in New Mexico (Freidberg and Math-
is 1986). One male and 2 females were reared
by me from capitula of Acamptopappus
sphaerocephalus (Harvey and Gray) Gray
collected at Snow Creek, Riverside Co.,
7-V-1981. This is a new state record and
the first host record for N. stecki.
Neaspilota viridescens Quisenberry.—As
revised by Freidberg and Mathis (1986), N.
viridescens is the most common and wide-
spread species in this genus in California.
Formerly reported only from Aster spinosus
Bentham (Freidberg and Mathis 1986), this
tephritid was additionally reared from ca-
pitula of: 4. eatonii (Gray) Howell, 6 4 and
13 2, Perazo Meadow, NW of Truckee, Ta-
hoe Nat. Forest, 2200-m elev., Plumas Nat.
Forest, Plumas Co., 10-IX-1986; A. scro-
pulorum Gray, 46 é and 42 2, Cowhorn Val-
ley, Inyo Nat. Forest, NE Inyo Co., 30-VI-
1982; Chrysothamnus teretifolius Durand
and Hilgard, 53 6 and 53 9, Mountain
Springs, SW Imperial Co., 6-XI-1986:; Evrig-
eron argentatus Gray, 3 6 and | 2, White
Mountain, 1630-m elev., Inyo Nat. Forest,
Inyo Co., 18-VI-1986; E. divergens Torrey
and Gray, 7 é and 11 2, Seven Oaks, along
Santa Ana River, San Bernardino Nat. For-
est, SW San Bernardino Co., 5-VII-1983;
Gutierrezia sarothrae (Pursh) Britton and
Rusby, | ¢, Mountain Springs, SW Imperial
Co., 6-XI-1986; Haplopappus cooperi (Gray)
Hall, 284 46 and 280 2, Walker Well, Walker
Pass, Kern Co., 21-V-1986; H. ericoides
(Lessing) Hooker and Arnott subsp. blakei
C. B. Wolf, 1 ¢ and 3 2, Orcutt, Santa Bar-
bara Co., 12-XI-1980; H. /aricifolius Gray,
13 d6and 12 2, NW of Kessler Peak at S end
VOLUME 91, NUMBER 2
of Ivanpah Mountains, NE San Bernardino
Co., 21-X-1982; H. linearifolius de-
Candolle, 33 6 and 20 2, N Meadow Creek,
Sequoia Nat. Forest, Tulare Co., 21-V-1986;
H. palmeri Gray, 3 4 and 4 2, N of Barrett
Junction, San Diego Co., 20-X-1987; H.
propinquus Blake, 27 4 and 35 2, along
Kitchen Creek, Cleveland Nat. Forest, San
Diego Co., 14-X-1981; Lepidospartum
squamatum (Gray) Gray, 11 éand 10°, NW
of Stepladder Mountains, SE San Bernar-
dino Co., 22-X-1982; Machaeranthera ca-
nescens (Pursh) Gray, | 6 and 4 2, Keystone
Canyon, Inyo Nat. Forest, Inyo Co., 1-IX-
1982; M. gracilis (Nuttall) Gray, 2 6 and 2
9, Cedar Canyon, New York Mountains, NE
San Bernardino Co., 20-IX-1983.
In addition, I confirmed the host record
for Aster spinosus with flies reared as fol-
lows: 9 6 and 13 2, Obsidian Butte, Imperial
Co., 7-XII-1983; 4 3, Calipatria, Imperial
Co., 28-XI-1984; 17 6 and 28 2, NW of
Blythe, Riverside Co., 2-XTI-1987.
The hosts of N. viridescens include plant
species from two tribes of Asteraceae: As-
tereae (Aster, Chrysothamnus, Erigeron,
Gutierrezia, Haplopappus, Macaeranthera)
and Senecioneae (Lepidospartum). Appar-
ently, this is the first rearing record of a
Neaspilota from the Senecioneae (Freidberg
and Mathis 1986). I have confirmed Lepi-
dospartum squamatum as a host as follows:
7 6 and 7 2, SE of Essex at Danby Cross-
roads, SE San Bernardino Co., 20-X-1982.
The coexistence in southern California of
an apparently generalist feeder like N. vir-
idescens with oligophagous and monopha-
gous congeners has been shown for several
generalist 7rupanea species, including T. jo-
nesi Curran, T. nigricornis (Coquillett), and
T. radifera (Goeden 1985). However,
another interpretation of these rearing rec-
ords is that N. viridescens, as presently de-
fined, is composed of several oligophagous
and monophagous species not yet separable
morphologically.
Neaspilota wilsoni Blanc and Foote.—In
stating that host plants of N. wi/soni are un-
167
known, Freidberg and Mathis (1986) missed
my rearing record for this tephritid from
Haplopappus squarrosus Hooker and Ar-
nott subsp. grindeloides (deCandolle) Keck
(Goeden 1983). They listed Coreopsis cal-
liosidea deCandolle in the tribe Heliantheae
as an unconfirmed, but suspected host. My
rearing record from a member of the As-
tereae casts doubt on the validity of this
unconfirmed record for this possible mono-
phage. Also, I sampled, but was unable to
obtain specimens from up to five samples
of four other species of Coreopsis or 16 other
species of Haplopappus, many of which
yielded other Tephritidae.
CONCLUSION
The genus Neaspilota, as now known in
California, is composed of species with a
full range of host-plant specificities, includ-
ing apparently strictly monophagous N. al-
biseta, N. stecki and possibly N. wilsoni;
nearly monophagous N. callistigma; oligo-
phagous N. aenigma and N. appendiculata
as well as general feeders, N. viridescens and
possibly N. achilleae. Other species record-
ed from California (Freidberg and Mathis
1986), but as yet not reared by me, for which
host data are unknown or too scant to as-
sess, include N. footei Freidberg and Mathis
and N. pubescens Freidberg and Mathis.
My host-plant records for California
species of Neaspilota generally support
Freidberg and Mathis’ (1986) ideas about
the monophyletic lineages of the two sub-
genera, Neaspilota sensu stricto, with hosts
mostly in the tribe Vernoniaeae, and Neo-
rellia, with hosts mostly in the Astereae. My
records associate N. aenigma, N. albiseta,
N. appendiculata, N. callistigma, N. stecki,
N. viridescens, and N. wilsoni with hosts in
the tribe Astereae. Hosts for Neaspilota
(Neorellia) species other than Astereae, were
confirmed by my rearing records from Ci-
choreae and Senecioneae. These departures
may be evidence of active speciation in the
genus Neaspilota involving changes in host
plants (Bush 1974, 1975).
168 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ACKNOWLEDGMENTS
My thanks to F. L. Blanc, A. Freidberg,
D. H. Headrick, and W. N. Mathis for their
helpful comments on earlier drafts of this
manuscript. The technical support of D. H.
Headrick and D. W. Ricker is gratefully ac-
knowledged.
LITERATURE CITED
Bush, G. L. 1974. Mechanisms of sympatric host race
formation of the true fruit flies (Tephritidae), pp.
3-23. In White, M. J. D., ed., Genetic Mecha-
nisms of Speciation in Insects. Australia and New
Zealand Book Co., Sydney, Australia.
1975. Modes of animal speciation. Annu.
Rev. Ecol. Syst. 6: 339-394.
Foote, R. H. and F. L. Blanc. 1963. The fruit flies or
Tephritidae of California. Bull. Calif. Insect Surv.
Te St pp:
Freidberg, A. and W. N. Mathis. 1986. Studies of
Terelliinae (Diptera: Tephritidae): A revision of
the genus Neaspilota Osten Sacken. Smithson.
Contrib. Zool. 434: 1-75.
Goeden, R. D. 1983. Initial host-plant records for
five species of fruit flies from southern California
(Diptera: Tephritidae). Proc. Entomol. Soc. Wash.
85: 399-400.
1985. Host-plant relations of Trupanea spp.
(Diptera: Tephritidae) in southern California. Proc.
Entomol. Soc. Wash. 87: 564-571.
1986. New records of Tephritidae (Diptera)
from Santa Cruz Island, California. Pan-Pac.
Entomol. 62: 326-328.
1987. Host-plant relations of native Uro-
phora spp. (Diptera: Tephritidae) in southern Cal-
ifornia. Proc. Entomol. Soc. Wash. 89: 269-274.
Munz, P. A. 1974. A Flora of Southern California.
Univ. Calif. Press, Berkeley. 1086 pp.
Munz, P. A. and D. D. Keck. 1959. A California
Flora. Univ. Calif. Press, Berkeley. 1681 pp.
Phillips, V. T. 1946. The biology and identification
of trypetid larvae (Diptera: Trypetidae). Mem.
Amer. Entomol. Soc. 12: 1-161.
Wasbauer, M. W. 1972. An annotated host catalog
of the fruit flies of America north of Mexico (Dip-
tera: Tephritidae). Calif. Dep. Agric. Bur. Ento-
mol. Occas. Pap. 19. 172 pp.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 169-174
A NEW SPECIES OF NEOLASIOPTERA (DIPTERA: CECIDOMYIIDAE)
FROM BACCHARIS (ASTERACEAE) IN SOUTHERN
UNITED STATES AND THE DOMINICAN REPUBLIC
RAYMOND J. GAGNE AND PAUL E. BOLDT
(RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA,
% U.S. National Museum NHB 168, Washington, D.C. 20560; (PEB) Grassland, Soil and
Water Research Laboratory, Agricultural Research Service, USDA, 808 East Blackland
Road, Temple, Texas 76502.
Abstract.— A new species, Neolasioptera rostrata Gagné (Diptera: Cecidomyiidae), which
galls flower receptacles of several species of Baccharis (Asteraceae), is described and
illustrated. It is known from Maryland, Florida, Texas, and New Mexico in the USA, and
from the Dominican Republic. A field study in Texas showed that N. rostrata has two
generations per year and that overwintering larvae diapause from November to the fol-
lowing September.
Key Words:
A new species of gall midge, Neolasiop-
tera rostrata, 1s described that forms a gall
in the receptacle of male and female flowers
of several species of Baccharis in Maryland,
Florida, Texas, and New Mexico, and the
Dominican Republic. The life history of N.
rostrata was studied on saltwillow, Bac-
charis halimifolia L. (Asteraceae: Astereae),
in Texas. Saltwillow is a woody, perennial,
dioecious shrub (2 to 3 m ht), that grows
from Texas to Florida and north to New
York (Tarver et al. 1979). This shrub in-
vades pastures, rangeland, and fallow fields
(Hardin 1959), but may be toxic for cattle
(Kingsbury 1964, Manley et al. 1982). It is
currently being studied in Texas as a poten-
tial target for biological control (DeLoach
et al. 1986).
MATERIALS AND METHODS
Anatomical terminology in the species
description follows the Manual of Nearctic
Diptera (McAlpine et al. 1981). Some spec-
imens were permanently mounted in Can-
gall midges, Baccharis, saltwillow
ada balsam on slides. The specimens used
for the description have been deposited in
the National Museum of Natural History
(USNM), Washington, D.C.
The field study of N. rostrata in Texas
was part of a general survey of the phy-
tophagous insect fauna of B. halimifolia.
Observations were made on three estab-
lished plots, one to two ha each, in aban-
doned pastures at Waller, Waller Co., and
Port Lavaca, Calhoun Co., and along three
kilometers of roadside ditch at Indianola,
Calhoun Co., Texas. Each plot contained 50
to 100 plants of B. halimifolia. Flowers were
sampled at about two week intervals from
September 14 to November 10, 1987. At
each plot, two to four stems were removed
from 10 male and !0 female plants and
bagged. Sample stems grew one to two m
above the ground and bore 50 to 100 flow-
ers. In the laboratory, each field sample was
divided into four groups, and 25 flowers
from each group were removed without bias
for dissection. The number of galled flowers
170
and the presence of larvae, pupae, and par-
asites were recorded. Galls collected on No-
vember 10 at Port Lavaca were measured
in cross-section at 100 with a calibrated
ocular micrometer in a stereo microscope.
The plots were inspected monthly from De-
cember, 1981 to February, 1988, but few
flowers were collected because they did not
remain on the plants.
In addition to the foregoing collections,
large numbers of infested flowers of B. hal-
imifolia were collected between October and
December, 1986 at Conroe, Montgomery
Co.; Liberty, Liberty Co.; and Monroe City,
Chambers Co., Texas. Some flowers were
dissected; others were held for emergence
of adults. On July 10, 1986, a collection was
made of infested flowers of Baccharis ne-
glecta Britt. at Fort Leaton, Presidio Co.,
Texas.
Neolasioptera rostrata Gagne,
New SPECIES
Adult.—Scale color pattern: frons white;
posterior surface of head and all of scutum
brown; legs white ventrally, brown dorsally;
leading edge of wing brown except for white
spot at juncture of RS and costa; anterior
half of tergites 1-7 brown, posterior half
silvery-white. Antenna with 11 flagello-
meres in 6 (n = 10) (Fig. 5), 12 in 2 (n =
10). Mouthparts (Figs. 4, 5): labrum long-
attenuate; labellum elongate, broadly
rounded at apex, second segment more than
3 times as long as first; palpus 4-segmented.
Thoracic vestiture: scutum with dorsocen-
tral and lateral rows of setae, covered else-
where with scales; anepisternum with scales
on dorsal half to two-thirds; katepisternum
bare; anepimeron with 9-12 setae and 0-2
scales. Wing length: 4, 1.2-1.6 mm (n = 10);
Figs. 1-10.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
9, 1.4-1.6 mm (n = 10). Length RS to re-
mainder of wing: 6, 0.55—0.60; 2, 0.54—-0.57.
Male abdomen (Figs. 7-10): Tergites 1-7
short, with single row of setae along entire
posterior margin, covered elsewhere with
scales; tergite 8 short, unsclerotized and
without scales on mesal third, sclerotized
and covered with scales on lateral third.
Sternites 2-6 with setae inside periphery,
scales elsewhere; sternites 7-8 short, with
setae posteriorly only, scales elsewhere.
Genitalia as in Figs. 8-10, setulae on gono-
stylus extending to midlength on venter.
Female abdomen (Fig. 6): Tergites 1-6
longer than in 4, vestiture as in 4; tergite 7
less than half width of 6, double row of setae
present along posterior margin, scales pres-
ent on posterior half; tergite 8 approxi-
mately twice length of tergite 7, longitudi-
nally divided except near anterior end.
Sternites 2-7 similar to 1-6 of 4, sternite 8
slightly shorter than the preceding, its setae
scattered across sclerite; tergite 6 is 0.21-
0.23 length of distal half of ovipositor (n =
10).
Last instar.—Length, 1.5-1.7 mm (n =
10). Integument pebbled. Spatula (Fig. 2)
anteriorly with 2 triangular lobes. Papillae:
three laterals on each side of spatula, two
with short setae, one without; inner pleurals
without setae on prothorax, with setae on
remaining segments; four terminals on anal
segment (Fig. 1), each with short seta; re-
mainder characteristic of genus but setae
short.
Specimens examined.—Holotype: 4,
FLORIDA, Dodge I., Miami, emerged XI-
1970 from seedheads of Baccharis glome-
ruliflora, collected XI-1970, C. E. Steg-
maier, Jr. Paratypes: FLORIDA: 1 4, same
data as holotype. MARYLAND: 6 4, 6 2
Neolasioptera rostrata (1-2, 4-10) and N. lathami (3). 1, Eighth and anal larval segments. 2,
Spatula and associated papillae. 3-5, Heads, 3, 5 in frontal view, 4 in side view. 6, Female postabdomen. 7,
Male abdominal segments 5-8. 8, Male genitalia (one gonopod shown). 9, Male genitalia, lateral view. 10,
Gonostylus. Scale line for Figs. 1-5, 8, 9 = 0.10 mm; 6, 7 = 0.05 mm; 10 = 0.01 mm.
171
VOLUME 91, NUMBER 2
172 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs JH.
Baccharis halimifolia. Left, flowering branchlet and leaf, 1 x. Center, flower heads, one in sagittal
section to show receptacle swelling, 2. Right, flower in sagittal section to show swollen receptacle and curled
larva, 6x.
Talbot Co., emerged VIII 1985 from flower
heads of Baccharis halimifolia, collected
X-1984, V. Krischik. NEW MEXICO: 2
last-instar larvae, 4 mi. e. Loving, Eddy Co.,
from flower heads of Baccharis salicina, T.
O. Robbins. TEXAS: 3 2, 3 2, Ft. Leaton
St. Pk., Presidio Co., reared from flower re-
ceptacle galls on Baccharis neglecta, col-
lected VI-10-1986, T. O. Robbins; 10 last-
instar larvae, Indianola, Calhoun Co., from
flower receptacle galls on Baccharis hal-
imifolia, X1-10-1987, P. E. Boldt; 1 4, Lib-
erty, Liberty Co., Hwy U.S. 90, from flower
receptacle galls on Baccharis halimifolia,
X-10-1986, P. E. Boldt; 1 2°, Port Lavaca,
Calhoun Co., from flower receptacle galls
on Baccharis halimifolia, X-28-1987, P. E.
Table 1.
halimifolia at three locations in Texas, 1987.
Boldt; 1 4, 1 9, Waller, Waller Co., reared
X-11-1986 from flower receptacle galls on
Baccharis halimifolia, collected XI-1987, P.
E. Boldt. DOMINICAN REPUBLIC: 5 4,
4 °, road from Constanza to San Jose de
Ocoa, Prov. La Vega, emerged VII-1987
from flowers of Baccharis myrsinites, col-
lected VII-17-1987, A. L. Norrbom.
Distribution.— Maryland on Baccharis
halimifolia, Florida on B. glomeruliflora;
Texas on B. halimifolia and B. neglecta; New
Mexico on B. salicina; and Dominican Re-
public on B. myrsinites.
Remarks. — Adults of N. rostrata are sim-
ilar to those of Neolasioptera lathami Gagné,
which causes stem galls on Baccharis spp.
from New York to Texas (Gagné 1971). The
Mean percentage (+SD) of galls of Neo/asioptera rostrata in male and female flowers of Baccharis
Waller Port Lavaca Indianola
Date Male Female Male Female Male Female
September 24 0 0 8.0 + 3.2 0 8.0 + 11.2 0
October 14 1:0. .2:0 0 5.0+3.6 16.0 + 8.6 3/0¥a) 5.2 2.0 + 2.4
October 28 35:0 = 11.4 3010 78:3 20.0 + 7.3 6.0 + 7.6 51.0 + 8.4 12.0 + 8.9
November 10* a 11.0 + 6.0 _ 10.0 + 9.4 - 32.0 + 10.8
* No male flowers remaining on plant.
VOLUME 91, NUMBER 2
principal difference between adults of these
species 1s the length of the mouthparts. The
labrum and labella of N. rostrata are very
long, reaching almost to the distal end of
the outstretched third palpal segment (Figs.
4, 5). The labrum and labella of N. lathami
are similar to those of other neolasiopteras
and reach only to the distal end of the sec-
ond palpal segment (Fig. 3).
Larvae of both N. rostrata and N. lathami
have three lateral thoracic papillae, two with
setae and one without on each side of the
spatula. The inner pleural prothoracic pa-
pilla, slightly lateral to the group of lateral
papillae, has no seta in N. rostrata (Fig. 2).
Further, N. rostrata has four terminal pa-
pillae (Fig. 1) instead of the six found in N.
lathami.
BIOLOGICAL NOTES
Neolasioptera rostrata produces an en-
largement of the receptacle of either male
or female flowers of B. halimifolia (Fig. 11).
The gall is usually single-celled with one
single larva. Two of the 227 galled flowers
dissected during this work contained two
larvae, each in its own cell and separated
from the other by a thin wall. The full-grown
larva is curled in a circle and fills the cham-
ber. The gall in the receptacle is not appar-
ent externally until the mature larva pushes
the top of the gall into a conical shape and
forms an exit hole at the apex. The hole is
then covered with a silky membrane pre-
sumably produced by the larva. The larva
then pupates or diapauses. Fifteen galls
measured 0.69 + 0.12 mm wide by 1.06 +
0.20 mm high (< + SD).
The development of the midge is closely
related to the bud and flower development
of its host. In Maryland, adults emerged in
August and September, 1984 from flowers
collected in October of the previous year.
Emergence coincided with normal flowering
of B. halimifolia in Maryland. In 1987, galls
were first observed in Texas on September
24, in male flower buds at two of three lo-
cations (Table 1). Male flowers were prob-
3
ably attacked first because they developed
sooner and were larger than female flowers.
Pupae were present in both male and female
flowers on October 14 at Port Lavaca. Emp-
ty galls, indicating previous adult emer-
gence, were found on October 28, the only
sample date when N. rostrata was present
in both male and female flowers at all plots.
Of 600 senescent or blooming flowers
sampled, 151 contained galls (25.2% infes-
tation). The actual infestation rate was
probably smaller because most of the se-
nescent flowers had already dropped from
the plant by that date and were uninfested.
Of the 151 galls, adult midges had emerged
from 37. Eight others contained pupae, nine
contained small larvae, and 97 contained
medium or large larvae. The presence of
galls in the receptacles did not appear to
affect pollen or seed production. Adult
emergence in late October indicated the
probability ofa second generation. The small
larvae present at that time were probably
the progeny of the last adults of the first
generation to oviposit.
On November 10, all male flowers had
dropped, but some senescent female flowers
remained on the shrubs. Of the 300 flowers
sampled, 53 were galled (17.7% infestation)
and contained medium to large quiescent
larvae. Only a few empty galls or galls con-
taining larvae were found in each of the
following months through February, 1988
when sampling was discontinued.
One species of Tetrastichus sp. (Eulophi-
dae: Hymenoptera) was found feeding ex-
ternally on a larva collected October 28,
1987 at Port Lavaca. Another was reared
from a pupa and from galls collected Oc-
tober 10, 1986 at Liberty. Platygaster sp.
(Platygasteridae) was also reared from galls
collected October 10, 1986, at Liberty, Tex-
as.
ACKNOWLEDGMENTS
We thank the following individuals for
their help with this publication: Deborah
Leather Roney drew Fig. 11 and inked Figs.
174
1-10; V. Krischik, A. L. Norrbom, T. O.
Robbins, and C. E. Stegmaier, Jr. collected
specimens of the new species; M. E. Schauff
identified the Hymenoptera; and D. R. Fer-
guson, K. M. Harris, V. Krischik, W. A.
Palmer, and R. V. Peterson reviewed a draft
of the ms.
LITERATURE CITED
DeLoach, C., P. E. Boldt, H. Cordo, H. Johnson, and
J. Cuda. 1986. Weeds common to Mexican and
U.S. rangelands: Proposals for biological control
and ecological studies, pp. 49-68. Jn Patton, D.,
V. Gonzales, C. Medina, L. Segura, and R. Hamre,
eds., Management and Utilization of And Land
Plants: Symposium Proceedings. Saltillo, Mexico.
USDA For. Serv. Gen. Tech. Rep. RM-135.
Gagné, R. J. 1971. Two new species of North Amer-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ican Neolasioptera from Baccharis (Diptera: Ceci-
domyiidae—Compositae). Proc. Entomol. Soc.
Wash. 73: 153-157.
Hardin, J. 1959. Some notes on weeds in North Car-
olina. J. South. Appalachian Bot. Club 24: 22-23.
Kingsbury, J. M. 1964. Poisonous Plants of the United
States and Canada. Prentice-Hall Inc., Englewood
Cliffs, New Jersey.
Manley, G. D., G. T. Edds, and S. F. Sundlof. 1982.
Cattle deaths from poisonous plants. Fla. Vet. J.
P1220:
McAlpine, J. F., B. V. Peterson, G. E. Shewell, H. J.
Teskey, J. R. Vockeroth, and D. M. Wood, eds.
1981. Manual of Nearctic Diptera. Vol. 1. Re-
search Branch, Agriculture Canada. Monograph
No. 27. vi + 674 pp.
Tarver, D., J. Rodgers, M. Mahler, and L. Lazor. 1979.
Aquatic and wetland plants of Florida. Fla. Dept.
Nat. Resources. 72 pp.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 175-178
TWO NEW ANTS OF THE GENUS SOLENOPSIS (DIPLORHOPTRUM)
FROM EASTERN TEXAS (HYMENOPTERA: FORMICIDAE)
WILLIAM P. MAcKAy AND S. BRADLEIGH VINSON
Department of Entomology, Texas A&M University, College Station, Texas 77843.
Abstract. — Solenopsis subterranea, n. sp. and S. puncticeps, n. sp. are described from 10
K N of Kurten, Brazos Co., Texas, USA. Solenopsis subterranea also occurs in Louisiana.
Both species are members of the subgenus Dip/orhoptrum as it 1s presently defined. These
two species have potential importance as natural enemies of founding queens of the
imported fire ant, So/enopsis invicta Buren.
Key Words:
Diplorhoptrum
Ants of Solenopsis (Diplorhoptrum) are
difficult to identify as the workers are very
small (often less that 2 mm long) and the
species demonstrate considerable conver-
gence in the worker caste (Creighton 1950).
They are primarily hypogaeic or “‘geo-
bionts” (Kempf 1961), and most species are
seldom collected unless special techniques
are used. Their nests are often found in close
proximity to the nests of other ant species
from which they presumably steal brood or
food.
Ants of this subgenus are important as
predators of founding queens of the im-
ported fire ant, Solenopsis invicta Buren
(Lammers 1987). There are undoubtedly
many undescribed species in the subgenus,
and it is in great need of revision. However,
we are describing these two species at this
time because of the need to place names on
natural enemies which may be important in
controlling populations of the imported fire
ant.
Solenopsis (Diplorhoptrum) subterranea
MacKay and Vinson,
New SPECIES
Figs. 1-4
Description (worker).—Head length
(HL—anterior median border of clypeus to
Formicidae, red imported fire ant, Solenopsis invicta predation, biocontrol,
median occipital margin) 0.34-0.38 mm;
head width (HW—maximum, at eye level)
0.24-0.25 mm; maximum eye length (EL)
0.02-0.03 mm: scape length (SL—excluding
basal condyle) 0.19-0.21 mm; Weber’s
length (WL—anterior pronotal declivity to
metasternal lobe) 0.30-0.35 mm; maxi-
mum petiolar width (PW) 0.08-0.09 mm;
maximum postpetiolar width (PPW) 0.09-
0.10 mm; cephalic index (CI = HW x 100/
HL) 67-74; ocular index (OI = EL x 100/
HL) 6-7; scape index (SI = SL x 100/HL)
52-63. Mandible with 4 well defined teeth
(Fig. 1), smooth and shining with scattered
punctures; clypeus with 2 well developed
carinae which end in long, well developed
teeth on both sides of median lobe, area
between carinae depressed, smooth, shin-
ing; dorsum of head smooth, polished, with
numerous scattered coarse punctures which
are much greater in diameter than the hairs
which arise from them (Fig. 1); occiput
somewhat concave in full face view; anten-
nal scape smooth and shining with punc-
tures which are much smaller in diameter
than those on dorsum of head, with nu-
merous erect and suberect hairs: eye small
with a total of about 3 ommatidia.
Mesosoma smooth and shining, with
176 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4 Holotype of S. subterranea: |. Full face view showing representative section of punctures; 2. Lateral
view of mesosoma; 3. Lateral view of petiole and postpetiole; 4. Dorsal view of petiole and postpetiole.
Figs. 5-8. Holotype of S. puncticeps: 5. Full face view showing representative section of punctures; 6. Lateral
view of mesosoma; 7. Lateral view of petiole and postpetiole; 8. Dorsal view of petiole and postpetiole. All
illustrations are drawn to same scale.
scattered punctures not much greater in di- _ petiole (Fig. 4); gaster smooth and shining
ameter than the hairs which arise from them _ with scattered punctures not much greater
(Fig. 2); petiole and postpetiole rounded (Fig. in diameter than hairs which arise from
3), width of postpetiole greater than that of them.
VOLUME 91, NUMBER 2
Entire dorsa of head and mesosoma cov-
ered by short (0.01-0.04 mm), erect and
suberect hairs (Fig. 2). Light yellowish-
brown with gaster and legs slightly darker,
tips of mandibular teeth and eyes dark brown
or black.
Female and male: Unknown.
Etymology.—From Latin, subter—be-
neath, and terra—soil, as we have found this
species only below the soil surface.
Type material.—Holotype and 20 para-
types to be deposited in the Museum of
Comparative Zoology, Harvard University;
additional paratypes (180) to be deposited
in the United States National Museum,
American Museum of Natural History, Cal-
ifornia Academy of Sciences, Los Angeles
County Museum of Natural History, British
Museum of Natural History, Museo de His-
toria Natural (México), the Field Museum
of Natural History, the Florida State Col-
lection of Arthropods, Universidade de Sao
Paulo, Museo Argentino de Ciéncias Nat-
urales, Universidad Nacional de Colombia,
the Insect Collections of Texas A&M Uni-
versity and Mississippi State University, and
the collection of WPM.
Distribution.— Known from the type lo-
cality, 10 K N Kurten, Brazos Co. TX (east-
ern Texas, on N side of Ferrill Creek Rd.,
3.94 K E of turnoff from Farm Road 2038
North) and southwestern Louisiana. A sin-
gle worker was captured in a subterranean
trap baited with a mealworm in Louisiana
(not designated as a paratype). It was col-
lected in Calcasieu Parish, Sam Houston
Jones State Park on 17 Aug. 1987, collection
number 9724-1. We have sampled inten-
sively throughout the southeastern region of
the United States with subterranean baits
and have not collected this species in any
other locality.
Biology.—Captured in subterranean pit-
fall traps at 10 cm depth, baited with a
cooked mixture of eggs, hamburger and
honey, as well as a trap baited with live
mealworms. We have not captured this
species on the soil surface at the type lo-
177
cality, despite intensive searches of the area
by five different individuals.
Solenopsis (Diplorhoptrum) puncticeps
MacKay and Vinson,
New SPECIES
Figs. 5-8
Description (worker).—HL 0.50-0.51
mm; HW 0.48-0.49 mm; EL 0.04 mm; SL
0.35-0.36 mm; WL 0.59-0.61 mm; PW
0.14-0.15 mm; PPW 0.16-0.18 mm; CI 95;
OI 7-8: SI 68-73. Mandible with 4 well
defined teeth (Fig. 5), smooth and shining
with scattered punctures; clypeus with 2 well
defined teeth, depressed, smooth and shin-
ing between the teeth; dorsum of head
smooth and shining, but with numerous
large, coarse punctures (Fig. 5); occipital
border slightly concave; scape smooth and
shining with much smaller punctures than
on the surface of head; mesosoma smooth
and shining with a few scattered punctures;
spiracle on propodeum very large and round
in shape (Fig. 6); ventral peduncular tooth
of petiole well developed, flattened and
moderately pointed (Fig. 7); petiolar node
not as wide as postpetiolar node (Fig. 8).
Entire body surface with erect hairs (Fig.
6): those on head short (0.02—0.05 mm)
and subequal in length, those on dorsum of
mesosoma longer (up to 0.8 mm) and un-
even in length (Fig. 6), hairs on petiole, post-
petiole and gaster similar to those on me-
sosoma. Color medium yellowish-brown
with mandibular teeth and eyes somewhat
darker.
Female and male: Unknown.
Etymology. — From Latin, puncta—punc-
ture and ceps—derivation of caput for head.
Type material.—Holotype and 10 para-
types will be deposited in the Museum of
Comparative Zoology, 110 paratypes will
be distributed to the institutions mentioned
previously.
Distribution. — Known only from the type
locality, 10 K N Kurten, Brazos Co. TX
(same type locality as S. suwbterranea).
Biology. — Captured in a subterranean trap
178 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
baited with the cooked mixture of eggs,
hamburger and honey.
Discussion.—Both of these species are
members of the group of thief ants in which
the dorsum of the head is covered with
coarse punctures which are much larger in
diameter than the hairs which arise from
them (couplets 10-13 of Creighton’s 1950
key to species). Solenopsis subterranea is ap-
parently most closely related to, and could
be confused with S. tennesseensis Smith (S.
longiceps in Creighton’s key). It differs in
that the postpetiole is not circular as seen
from above, and the head and mesosoma
are covered by short erect and suberect hairs
(0.01-0.04 mm long and essentially equal
in length in S. subterranea, uneven in length
with a range of 0.04-0.08 mm in S. fen-
nesseensis). Solenopsis tennesseensis 1S
somewhat larger (HL 0.41, HW 0.30, WL
0.44), but the CI(73), OI(7) and SI(59) are
within the range of S. subterranea. This new
species differs from S. krockowi Wheeler as
it is much smaller (8S. Arockowi: HL 0.51-
0.54, WL 0.53-0.57), and has much smaller
eyes (diameter in S. krockowi = 0.05, OI 9-
10). The clypeal teeth are very small in S.
krockowi and the erect hairs on the head
and mesosoma of S. krockowi are uneven
in length and range from 0.03-0.13 mm. It
differs from S. salina Wheeler in that the
tooth on the anterior peduncle of the petiole
of the new species is blunt and compressed
(it is usually sharply pointed in S. salina).
Hairs on the body surface are longer in S.
salina (0.04-0.08 mm) and are greatly un-
equal in length.
Solenopsis puncticeps is distinctive in that
the punctures on the dorsum of the head
are very large and coarse. It could be con-
fused with the closely related S. pergandei
Forel which has similar punctures, but the
postpetiole from above is not round in shape,
as it is in S. pergandei. It can be distin-
guished from S. krockowi by the coarser
punctures on the dorsum of the head, and
the diarneter of the propodeal spiracle which
is about twice that of the spiracle of S.
krockowi. It is easily separated from S. sub-
terranea by the hairs on the propodeum
which are much longer and unequal in
length.
We suspect these species are important
natural enemies of the founding queens of
the imported fire ant, as are others in the
subgenus Dip/orhoptrum (Lammers, 1987).
The fire ant density at the type locality of
the two new species 1s much lower than in
the surrounding area, possibly due in part
to the presence of these species as well as
an abundance of other species of the same
subgenus.
ACKNOWLEDGMENTS
Dr. David Smith of the Systematic Ento-
mology Laboratory, Agriculture Research
Service, Washington, DC loaned us para-
types of S. tennesseensis Smith. Dr. Alex
Mintzer suggested the study area, assisted
in collecting ants in the area and critically
read the manuscript. Dr. Awinash Bhatkar
and an anonymous reviewer critically read
the manuscript. Mark Strain, Shelley Stone-
cipher and Cecil Pinder helped search the
area for colonies. The research was partially
supported by Texas State Funds #2000-
1925. Approved as TA-23930 by the Di-
rector of the Texas Agriculture Experiment
Station, College Station.
LITERATURE CITED
Creighton, W. S. 1950. The ants of North America.
Bull. Mus. Comp. Zool. 104: 1-585 + 57 plates.
Kempf, W. W. 1961. A survey of the ants of the soil
fauna in Surinam (Hymenoptera: Formicidae).
Studia Entomol. 4: 481-524.
Lammers, J. 1987. Mortality factors associated with
the founding queens of Solenopsis invicta Buren,
the red imported fire ant: A study of the native
ant community in Central Texas. Unpublished MS
Thesis, Texas A&M University, 206 pp.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 179-184
BEHAVIOR AND DEVELOPMENT OF THE WASP
PTEROMBRUS RUFIVENTRIS HYALINATUS KROMBEIN
(HYMENOPTERA: TIPHIIDAE), A PARASITE OF LARVAL
TIGER BEETLES (COLEOPTERA: CICINDELIDAE)
C. BARRY KNISLEY, DARREN L. REEVES, AND GREGORY T. STEPHENS
Department of Biology, Randolph-Macon College, Ashland, Virginia 23005.
Abstract. —Pterombrus rufiventris hyalinatus was found attacking third instar larvae
of three Cicindela species in Arizona. Its parasitic behavior is generally similar to that
described for tropical species of Prerombrus, but different in having a more elaborate
burrow plugging behavior and one generation per year. Female wasps enter open burrows
or occasionally dig into plugged burrows, sting the host larvae several times under the
head or thorax, and oviposit on the second abdominal sternum. Burrows are closed with
a primary plug of compacted soil immediately above the larvae and a secondary plug
loosely packing the remainder of the burrow. Mean development time is 2.93 days for
the egg stage and 8.70 days for larvae. When mature, wasp larvae detach from the host,
spin a cocoon in the larval burrow, and emerge the following July with the onset of the
““monsoon”’ rains.
Key Words:
Tiphudae, wasp
Two genera of tiphiid wasps of the
subfamily Methochinae, Methocha and
Pterombrus, are parasites of tiger beetle lar-
vae. Some of the natural history and be-
havior of several species of Methocha has
been described (Adlerz 1906, Williams
1916, 1919, Iwata 1934, 1936, Burdick and
Wasbauer 1959, Wilson and Farish 1973).
Pterombrus is primarily tropical in distri-
bution and the limited knowledge about it
comes from studies of P. cicindelicidus and
P. theringi in Brazil (Williams 1928) and P.
piceus in Costa Rica (Palmer 1976). These
studies included field observations and notes
of parasitic behavior and development
times.
Pterombrus rufiventris is the only known
Cicindela, larvae, parasite, parasitic behavior, Pterombrus, tiger beetle,
United States species of the genus. Krom-
bein (1949) distinguished two subspecies, P.
r. rufiventris from eastern Texas, Louisiana,
North Carolina, and Virginia, and P. r. hy-
alinatus from western Texas, Arizona, and
California. Nothing has been published on
the biology of this species. During ecological
studies of tiger beetles in southeastern Ar-
izona, one of us (CBK) discovered P. r. hy-
alinatus parasitizing larvae of Cicindela ob-
soleta, C. pulchra, and C. marutha. Because
of its possible importance in limiting these
tiger beetle populations, field and labora-
tory studies of this wasp were initiated. We
present here observations on parasitic be-
havior and development. Other studies
present rates of parasitism of Cicindela
180
species (Knisley 1987, Knisley and Juliano,
1988) and factors that influence rates of
parasitism (Knisley, in prep.).
METHODS
Field studies. — Field studies were done in
the Sulphur Springs Valley, near Willcox,
Cochise Co., AZ, during July and August
from 1983-87. Eight sites representing four
habitat types (playa, saltbush flat, grassland,
sand ridge), each with one or more species
of adult or larval Cicindela (Knisley 1987)
were checked for wasps one to two times
per week. During these checks all Prerom-
brus were counted to determine relative
abundance and seasonal activity. Wasp be-
havior was observed and recorded.
Laboratory studies.—The behavior of
Pterombrus was studied under laboratory
conditions of summer photoperiod (13L:
11D) at 24—28°C in an observation cham-
ber. The chamber was made from a trans-
parent plastic “shoe” box (17 x 32 x 9 cm)
and supported by 20 cm wooden legs placed
under each corner. An acrylic tube (2.5 x
18 cm) filled with soil and containing an
active third instar of C. pulchra (Knisley
and Pearson 1984) was inserted through a
hole in the bottom of the box so that the
top of the tube was flush with the surface
of a 5—6 cm layer of soil in the box. The top
of the tube was wrapped with tape to ensure
a tight fit of the tube into the hole. For clear
observation of wasp behavior, only tubes in
which the larva had dug a burrow along the
side wall of the tube were used for the trials.
To begin an observation trial a female wasp
was placed into the covered plastic box. We
observed the complete sequence of parasitic
behavior for three wasps and partial se-
quences for three others.
Development time of Pterombrus was de-
termined from host preference studies
(Knisley, in prep.) in trials in which a tiger
beetle larva in a rearing tube was confined
with a wasp. Each parasitized larva was
carefully dug from the tubes and transferred
onto a disc of moistened paper towel in a
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
plastic petri dish (10 cm dia). Develop-
mental progress and body length of larval
stages were recorded daily.
RESULTS AND DISCUSSION
Field studies.— Distribution and behav-
ior of wasps in the field were based on over
70 sightings of adult wasps. Only four males
were observed and all were flying near fe-
males and appeared to be attempting to
mate. No mating was seen. Adult wasps were
found at six sites. Only one wasp was ob-
served each at two of these sites, a ditch
edge and a pond edge. No wasps were ob-
served in the playa or playa edge habitats.
Wasps were relatively common (two to five
observed during each survey) at two sand
ridge sites where there were large popula-
tions (over 500) of C. marutha larvae and
ata nearby (400 m) saltbush flat where there
were several hundred C. pulchra. Pterom-
brus was also common at a higher elevation
grassland site near Chiricahua National
Monument which had a population of over
300 C. obsoleta larvae.
Adult wasps were first observed at field
sites within one or two days of the first sig-
nificant July rainfall (>8-10 mm). The ac-
tual dates of first sightings were July 6 in
1986, July 10 in 1985 and July 16 in 1987.
Wasp emergence may be triggered by the
onset of the summer “monsoon” rains, as
is tiger beetle adult emergence and larval
activity. Numbers of wasp sightings de-
creased from July through late August at all
sites, but the pattern of abundance at the
grassland site was about two weeks later than
at the other sites. In the field, we observed
P. r. hyalinatus parasitizing only third in-
stars of C. pulchra, C. obsoleta, and C. ma-
rutha, but in the laboratory some second
instars of these species were attacked. Palm-
er (1976) found that P. piceus parasitizes
primarily second instars of Pseudoxychila
tarsalis Bates.
Searching behavior.—We observed the
searching behavior of 15 wasps in the field.
Female wasps walked rapidly over the
VOLUME 91, NUMBER 2
ground, frequently turning and darting, in
what appears to be a random pattern. The
wasps stopped to examine or probe with
their antennae holes, cracks or surface
depressions. Often they passed within sev-
eral centimeters of active larval burrows then
moved away. Wasps initiated an attack only
when they came in contact with the burrow
edge. Pterombrus piceus searches in a sim-
ilar manner and seems to prefer to attack
larvae that move down the burrow when
they approach (Palmer 1976).
We observed four instances of wasps dig-
ging out plugged burrows of third instar C.
marutha, a species which typically plugs its
burrows during most of the day (Knisley
1987). In each case, the wasp used its man-
dibles to grasp bits of sand from the plugged
area, moved back 4 to 6 centimeters and
dropped the sand, eventually forming a small
arc or circle of sand around the burrow
mouth. Removal of the 2 to 4 cm plug took
19 to 33 minutes, after which the wasp en-
tered the burrow to attack the larva. Meth-
ocha (Padgen 1926) and P. itheringi (Wil-
liams 1928) have also been observed digging
out plugged burrows. In addition, Methocha
reportedly digs in soil where tiger beetle lar-
vae had previously occurred (Williams
1919). These observations suggest that wasps
may detect the odor of host larvae or their
burrows once they come in close proximity
to them. In the laboratory chambers, Pre-
rombrus initiated attacks on larvae in 4 to
38 minutes (x = 16.2). In the field, wasps
were observed to search for 15-30 minutes
without finding a larva.
Pterombrus did not seem to fly when
searching for burrows, and seldom flew un-
less disturbed. Short flights (7-20 m), ob-
served when wasps searched unsuccessfully
in an area for 15 to 20 minutes, could in-
dicate dispersal to new areas of potential
larval habitat.
Attack, stinging and oviposition.—At-
tacks of host larvae in the laboratory were
initiated by female P. r. hyalinatus moving
quickly into the burrow head first, causing
181
the larva to retreat. The wasp continued
down the burrow until her head touched the
larva’s head. Sometimes the wasp used its
mandibles to grasp the head of the larva
before quickly curving her abdomen for-
ward and stinging the larva under the head
or thorax. This caused the larva to move
further down the burrow. The wasp re-
mained inactive for 2-5 minutes, then cau-
tiously approached the larva and probed the
larva’s head with its antenna. In two of the
trials with large third instars of C. obsoleta,
the larvae responded with slight movement
and were again stung once or twice.
The wasp used one or more of three meth-
ods to slightly raise the larva in the burrow:
grasping the larva’s head with its mandibles
and pulling from above, pushing up on the
caudal portion of the larva’s abdomen, or
grasping the larva’s forelegs with its man-
dibles and pushing while positioned beside
the larva. These movements often wedged
the larva’s cephalothoracic shield in the
burrow. In two cases another sting was then
applied to the larva’s venter. We never ob-
served larvae flipping out of the burrow.
This may be a defense reaction of the larva
and is reported in studies of Pterombrus
(Palmer 1976) and Methocha (Mury Meyer
1983). We also did not observe wasps being
seized by tiger beetle larvae. Methocha has
been observed to allow the larva to seize it
with its mandibles before quickly stinging
the larva. Palmer (1976) reported one in-
stance of this behavior in P. piceus.
After positioning the larva or sometimes
after oviposition the wasp used the tip of
its abdomen to pack the bottom of the bur-
row. Pterombrus theringi shows similar
packing behavior after using its mandibles
to loosen the soil (Williams 1928). We ob-
served one wasp grasp at the larva’s second
abdominal sternum. The wasp may have
been chewing, feeding or preparing an ovi-
position site. Burdick and Wasbauer (1959)
reported that M. californica commonly feeds
on fluids exuding from sting punctures and
suggested that this may provide nutrients
182
that are necessary for successful oviposi-
tion.
To oviposit, the wasp positioned itself
venter to venter with the larva, moved for-
ward using the tip of its abdomen to sweep
back and forth over the larva’s abdominal
sternites, then stopped and deposited an egg
on the midline of the second abdominal
sternum. In two cases the wasp next stung
the beetle larva one or more times near the
egg. These stings or the chewings near the
Oviposition site may prepare an access for
the newly hatched wasp larva to enter the
host to feed.
The number of stings given the beetle lar-
va by both Pterombrus and Methocha is
variable. Any movement by the larva seems
to be sufficient to trigger a stinging response
in Methocha (Burdick and Wasbauer 1959).
Our observations suggested this may be true
for Pterombrus. Stings also seemed to be
given after certain behavioral events, such
as positioning the larva and oviposition.
Host larvae were apparently completely
paralyzed by the stings because they did not
move in their burrows, and exhibited only
slight movement when probed. The effect
of the stings of Methocha species has been
correlated with the site of oviposition. Host
larvae stung by species that oviposit on the
host’s abdomen are permanently paralyzed
by the host; host larvae stung by species that
Oviposit on the venter of the metathorax
recover and move actively in their burrows
(Williams 1919, Iwata 1936).
Burrow preparation and plugging. — After
Oviposition the wasp moved above the larva
and constructed two plugs, a compact pri-
mary plug immediately above the larva and
a loosely packed secondary plug which filled
the rest of the burrow. To form the primary
plug, the wasp positioned itself 24 cm above
the paralyzed larva, curved its abdomen
forward across the burrow shaft and used
its mandibles to dislodge soil particles from
the sides of the burrow. Soil fell onto the
underside of the abdomen, was compacted
there with the hind legs and then pressed
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
onto the sides of the burrow with the ab-
domen. As the wasp moved in a circle, the
activity eventually formed a circular shelf
of soil. To complete the primary plug, the
wasp moved to near the top of the burrow,
used its mandibles to dislodge soil onto the
shelf, then moved down and appeared to
use its hind legs to cover the hole in the
shelf left by its abdomen. The primary plug
was 1.5-3.0 cm thick and required 7-15
minutes to complete.
Next, the wasp left the burrow and
searched the area within 10-20 cm of the
burrow entrance for bits of soil, stones, twigs
and other materials for the secondary plug.
The wasp carried each piece in its mandibles
and usually dropped it into the burrow from
without. Filling of the burrow and comple-
tion of the secondary plug required from 21
to over 140 trips and 13-41 minutes (x =
18, n = 12). Types of materials available
around the burrow and depth of the burrow
seemed to account for the variation in num-
ber of trips and time. For example, filling
of burrows at the saltbush flat site required
more time because plugging materials were
scarce. At a sand ridge site and in two lab-
oratory chambers where the soil was nearly
pure sand, wasps formed most of the sec-
ondary plug by pushing sand into the bur-
row with the hind legs. Plugging was then
completed by filling in with larger materials.
In general, wasps seemed to be more selec-
tive of materials during the final phase of
burrow plugging.
The surface appearance of the plugged
burrows was distinctive at each of the sites.
The secondary plugs at the grassland site
consisted of small pebbles. Those at the salt-
bush flat had small twigs, bits of plant ma-
terial or flakes of soil. Sand ridge burrows
were filled with sand and often eroded
around the edge. Methocha plugged bur-
rows with a variety of materials (Bouwman
1909, Champion and Champion 1914),
possibly selecting them on the basis of avail-
ability and ease of transport (Burdick and
Wasbauer 1959). Complete closure of the
VOLUME 91, NUMBER 2
Table 1.
at 24-28°C.
183
Duration and size (length in mm) of developmental stages of P. r. hyalinatus reared in the laboratory
Egg length (mm)
Duration egg stage (days)
Larval length (mm) on day after hatch:
iB
ANDO Hh WwW
8
Duration of larval stage (to detachment from host) (days)
Total development time (days)
burrow may be necessary to protect the de-
veloping wasp against natural enemies. We
noticed that several incompletely plugged
burrows were dug into by ants and the host
larvae were eaten.
Plugging burrows with two plugs by P. r.
hyalinatus involves more elaborate behav-
ior than the single plugging by other species
of Pterombrus. This may be an adaptation
to the dry environment of P. r. hyalinatus
where infrequent rains are less likely to nat-
urally close the burrow. The single plug con-
structed by P. piceus (Palmer 1976) resem-
bles the secondary plug described here. The
single plug of P. iheringi (Williams 1928) is
similar to the primary plug of P. r. hyali-
natus. Observations of Methocha indicate
plugging involves a complete filling of the
burrow (Alderz 1906, Bouwman 1909, Bur-
dick and Wasbauer 1959).
Development.— Upon hatching the wasp
larva typically remained at the site of ovi-
position on the second abdominal sternum
and began extracting the liquid contents of
the host larva. Growth was rapid with lar-
vae increasing by about 1-1.5 mm per day,
from a mean length of 2.8 mm at hatching
to 14.8 mm when fully grown after eight
days. Detachment from the host and initi-
ation of cocoon spinning began when the
host was completely consumed and shriv-
eled. The cephalothoracic shield of the host
N Mean SD Range
15 2.39 0.14 2.10-2.52
14 2:93 0.59 2.30-3.90
9 2.86 0.38 2.52-3.57
8 3.61 0.34 2.45-4.34
9 4.70 0.58 3.90-6.09
7 6.56 0.95 5.18-7.42
11 5.76 1.01 4.64-6.24
6 9.12 1.35 5.76-10.88
8 9.04 2.26 8.00-11.20
8 14.88 1.29 13.12-16.16
[5 8.70 1.35 7.80-10.50
25: 12.33 2.65 10.70-15.30
was typically attached to the top of the pupal
cocoon and apparently served as a starting
point for cocoon spinning. The pupal co-
coon was similar to P. cicindelidicus (Wil-
liams 1928). Adult Pterombrus collected at
the saltbush flat and grassland sites which
had larger host tiger beetle larvae (C. pul-
chra and C. obsoleta, respectively) had a
mean length of 13.5 mm and those collected
at the sand ridge sites with the smaller C.
marutha had a mean length of 10.5 mm. In
laboratory host preference studies pupal size
was highly correlated with host larva size
(Knisley, in prep.).
Mean time from oviposition to comple-
tion of the pupal cocoon was 13.2 days (SD
= 1.6, range 10.2-17.3) (Table 1). Mean du-
ration of stages was 2.93 days for the egg,
8.70 days for the larva, and 1.60 days from
larval detachment to completion of the co-
coon. Larval development is about 12 days
for P. piceus (Palmer 1976) and 10-12 days
for several Methocha species.
The behavior we observed for P. r. hy-
alinatus was similar in most aspects to that
of other species of the genus. The more elab-
orate plugging behavior and the one gen-
eration per year may be adaptations to a
different environment. Our observations
also indicate similarity between the behav-
ior of Pterombrus and Methocha and sug-
gest that these behaviors have evolved in
184
response to the highly specialized life of lar-
val tiger beetles. The apparent consistent
difference between these genera is the initial
attack behavior in which Methocha allows
itself to be seized by the larva. Studies on
searching, host selection, and abundance of
these two genera of parasitic wasps could
provide important additional information
for determining how they limit tiger beetle
populations.
ACKNOWLEDGMENTS
Mark R. Carter, David R. Beatty, and
Frank W. Ellis assisted in the field work.
Karl V. Krombein identified P. r. hyalina-
tus. This research was supported in part by
grants from the National Science Founda-
tion (DEB8119256 and BSR8415949).
LITERATURE CITED
Adlerz, G. 1906. Methoca ichneumonides Latr., dess
lefnadsattoch utvecklingsstadier. Arkiv for Zoo-
logie, Bd. 3, No. 4, p. 1-48, 1 pl. Uppsala and
Stockholm.
Bouwman, B. E. 1909. Uber die Lebenweise von
Methoca ichneumonides Latr. Tijdschr. voor Ent.
52: 284-294.
Burdick, D. J. and M. S. Wasbauer. 1959. Biology
of Methocha californica Westwood (Hymenop-
tera: Tiphiidae). Wasmann Jour. Biol. 17: 75-88.
Champion, H. G. and R. J. Champion. 1914. Ob-
servations on the life-history of Methoca ichneu-
monides Latr. Ent. Monthly Mag., Ser. 2, 25: 266-
270.
Iwata, Kunio. 1934. Comparative studies on the hab-
its of solitary wasps. Tenthredo 4(1-2): 1-146.
1936. Biology of two Japanese species of
Methoca with the description ofa new species (Hy-
menoptera, Thynnidae). Kontyu 10: 67-89.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Knisley, C. B. 1987. Habitats, food resources, and
natural enemies of a community of larval Cicin-
dela in southeastern Arizona (Coleoptera: Cicin-
delidae). Can. J. Zool. 65: 1191-2000.
Knisley, C. B. and S. A. Juliano. 1988. Survival,
development, and size of larval tiger beetles: Ef-
fects of food and water. Ecology 69: 1983-1992.
Knisley, C. B. and D. L. Pearson. 1984. Biosystem-
atics of larval tiger beetles of the Sulphur Springs
Valley, Arizona. Descriptions of new species and
a review of characters for Cicindela (Coleoptera:
Cicindelidae). Trans. Amer. Entomol. Soc. 110:
465-551.
Krombein, K. V. 1949. A new subspecies of Prerom-
brus rufiventris (Cresson). Pan-Pacific Entomol. 25:
88-89.
Mury Meyer, E. J. 1983. An analysis of survivorship
and foraging methods in larvae of three sympatric
species of tiger beetles occurring in central Penn-
sylvania. Ph.D. dissertation. Pennsylvania State
Univ., University Park, PA.
Padgen, H. T. 1926. Observations on the habits and
parthenogenesis of Methoca ichneumonides Latr.
Trans. Entomol. Soc. London Ser. 4, 26: 591-597.
Palmer, M. K. 1976. Notes on the biology of Pterom-
brus piceus Krombein (Hymenoptera: Tiphiidae).
Proc. Entomol. Soc. Wash. 78: 369-375.
Williams, F. X. 1916. Notes on the life-history of
Methoca stygia Say. Psyche 23: 121-125.
. 1919. Phillipine wasp studies II. Descriptions
of new species and life history studies. Bull. Exp.
Stn. Hawaiian Sugar Planters Assoc., Entomol. Ser.
14: 19-180.
. 1928. Pterombrus, a wasp-enemy of the larva
of tiger beetles. /n Studies in Tropical Wasps—
Their Hosts and Associates (with Descriptions of
New Species). Bull. Exp. Stn. Hawaiian Sugar
Planters Assoc., Entomol. Ser. 19: 144-151.
Wilson, E. O. and D. J. Farish. 1973. Predatory be-
haviour in the ant-like wasp Methocha stygia (Say)
(Hymenoptera: Tiphiidae). Anim. Behav. 21: 292-
295.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 185-189
DESCRIPTION OF THE PREDACEOUS LARVA OF
PSEUDOGAURAX SIGNATUS (LOEW) (DIPTERA: CHLOROPIDAE)
ADAM ASQUITH AND DAviID ADAMSKI
(AA) Systematic Entomology Laboratory, Entomology Department, Oregon State Uni-
versity, Corvallis, Oregon 97331; (DA) Department of Entomology, Drawer EM, Missis-
sippi State University, Mississippi State, Mississippi 39762.
Abstract.—A description of the predaceous larva of Pseudogaurax signatus (Loew)
(Diptera: Chloropidae) includes an illustration of the cephalopharyngeal skeleton, spinule
pattern and posterior spiracles and scanning electron micrographs of the cuticular ar-
mature, facial mask and the anterior and posterior spiracles.
Key Words:
Pseudogaurax Malloch is a large, mostly
tropical group (Sabrosky 1966) whose lar-
vae are predators of the immature stages of
several species of Araneae and Insecta.
Pseudogaurax signatus (Loew) was origi-
nally placed in the genus Gaurax Loew. Sa-
brosky (1945, 1966) has provided the only
taxonomic treatment on this group since
Hall (1937) transferred the species to the
genus Pseudogaurax. P. signatus has been
reared from egg sacs of the spiders Latro-
dectus mactans (Fabr.), Argiope riparia (Lu-
cas) and Araneus sp., oothecae of Mantidae
(Davidson 1896, Hall 1937, Breland 1941),
and the pupae of Hyalophora cecropia (L.)
(Saturniidae) and Euproctis chrysorrhoea
(L.) (Lymantriidae), (Sabrosky 1945).
Chloropid larvae display diverse feeding
habits, including phytophagy, scatophagy,
necrophagy, and predation. The larval stages
of most chloropid species are unknown, and
most available descriptions are for phy-
tophagous species. The only known larval
descriptions of predaceous chloropids are
for Steleocerellus latiseta (Lamb) (Kirk-
Spriggs 1986) and Pseudogaurax anchora
(Loew) (Howard 1916); in the latter how-
ever, only the anterior spiracles and pupar-
larval morphology, ultrastructure, spider-parasite
ium were described. Roberts (1971) and
Teskey (1981b) have emphasized that de-
tailed larval descriptions of all immature
stages together with descriptions of adults
will undoubtedly be useful in elucidating
phylogenetic relationships and the evolu-
tion of trophic diversity in this group of
flies. We thus describe the third-instar larva
of Pseudogaurax signatus (Loew).
MATERIALS AND METHODS
One Argiope sp. egg sac was collected in
the field and taken to the laboratory for dis-
section. A portion of the total number of
third-instar larvae of P. signatus found
within the egg sac were extracted, killed in
KAAD, and preserved in 70% ethanol. The
incision made in the egg sac was closed and
the egg sac placed in a rearing chamber at
room temperature until adult flies emerged.
For SEM examination, the larvae were
dehydrated in ethanol and degreased in pen-
tane. The specimens were then rehydrated,
washed in 0.1 M phosphate buffer (pH 7.3),
and postfixed in 2% osmium tetroxide in
the same buffer. After dehydration in eth-
anol, specimens were critical point dried,
186 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vs vuwuyvyy Yee YEE Yuriy vy On
vuwy
OV UUY YY
Pseudogaurax signatus, larva. 1, Lateral view. Line scale = 0.25 mm. 2, Ventral view. Line scale
Figs. 1-3a.
= 0.25 mm. 3, Spinules on anteroventral portion of Irst abdominal segment (enlargement of area indicated by
arrow in Fig. 1). Line scale = 10 um. 3a, Spinule pattern on 4th abdominal segment (ventral view). Line scale
= 0.05 mm.
mounted on stubs with silver paint and S-2R scanning electron microscope at an
coated with gold-palladium in a Polaron accelerating voltage of 20 kV.
e5100 sputter coater. The ultrastructure of For study of the cephalopharyngeal skel-
the larva was studied with an Hitachi HH-_ eton, larvae were cleared in 20% hot potas-
PastmB Hyphar
4 — ———
cephalopharyngeal skeleton (lateral view). D Corn = Dorsal Cornu,
Hyphar = Hypopharyngeal Sclerite, Md = Mandible, Pastm B = Parastomal Bar, V Corn = Ventral Cornu.
Line scale = 0.12 mm.
Fig. 4. Pseudogaurax signatus, larva,
VOLUME 91, NUMBER 2
Figs. 5-11. Pseudogaurax signatus, larva. 5, Thorax, and Irst abdominal segment (dorsal view). 6, Head,
facial mask (anterolateral view). 7, Head and prothorax (lateral view). 8, Head (ventrolateral view). 9, Maxillary
palpus (ventrolateral view). 10, Anal plate (posterior view). Line scale for Figs. 5-7 = 100 um. Line scale for
Figs. 8-10 = 10 um. A = Antenna, ASp = Anterior Spiracle, FR = Frontal Rami, MxP = Maxillary Palpus,
MxR = Maxillary Ring, PSp = Posterior Spiracle. 11, Posterior spiracle (posterior view). Line scale = 0.05 mm.
188
11
sium hydroxide, mounted on microscope
slides in glycerine, and examined with an
Olympus compound microscope.
DESCRIPTION
Larva vermiform, tapering anterior of
third abdominal segment, length 4.0-5.0
mm, greatest width 0.96 mm, creamy white;
anterior margin of thoracic and first abdom-
inal segments with several irregular rows of
spinules on creeping welts (Figs. 1-3, 5).
Spinules arranged in 5 regular but broken
rows on ventral surface of abdominal seg-
ments 2-10; spinules coarse, oriented pos-
teriorly, variable in size with those on 4th
row larger (Fig. 3a); all abdominal segments
with two annulations, thoracic segments
without annulations.
Facial mask on cephalic segment bilobed,
hooded by first thoracic segment (Figs. 6-
7); frontal rami consisting of two oral ridges,
serrate and overlapping (Fig. 8); antenna
prominent, one-segmented, directed an-
teroventrally (Figs. 6-8); maxillary ring in-
complete, forming a semicircle, opening
dorsally; maxillary palpus in two sensillar
groups (Figs. 8-9); a ventral triad is delim-
ited by fleshy folds above the maxillary ring,
abutting a dorsally located pair.
Mandibles of cephalopharyngeal skeleton
separate, strongly sclerotized, basal portion
deepest, narrowing distally, curved apically
(Fig. 4); hypopharyngeal sclerite nearly same
length as mandibles, similar in degree of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
melanization and sclerotization; parasto-
mal sclerites slender, appearing as narrow
bands above hypopharyngeal sclerite in lat-
eral view, fused basally with tentorophar-
yngeal sclerite, but much less sclerotized;
labial sclerite present, small, located be-
tween anterior ends of hypopharyngeal
sclerites, not visible in lateral view; ventral
cornu of tentoropharyngeal sclerite broader
and longer than dorsal cornu, both pro-
cesses heavily sclerotized along inner mar-
gins. Anterior spiracles palmate, with seven
short papillae (Figs. 6-7); posterior spiracles
0.06 mm wide, convergent with protuber-
ant bases 0.18 mm apart, three spiracular
openings, oval, longitudinal axis of middle
opening directed dorsally, lateral openings
angled, longitudinal axes converging to-
wards middle opening dorsally (Fig. 1 1); anal
plate oval and fleshy (Fig. 10).
Specimens examined.—Larvae (n = 28):
Label data Miss(issippi), Oktibbeha Co.,
Dorman Lake; 21 Jan(uary) 1986; Coll.
Adam Asquith. Collected from Argiope sp.
egg sac. Adults: (6 males, 7 females): Label
data same as above except for second label:
Reared from Argiope sp. egg sac. All vouch-
er specimens are deposited in the Missis-
sippi Entomological Museum at Mississippi
State University.
DISCUSSION
The larva of Pseudogaurax signatus 1s
comparable to that of P. anchora. Both lack
the frond-like scales found on the posterior
spiracles of other chloropid species such as
Meromyza laeta Meigen (Fedoseyeva 1966),
Polyodaspis ruficornis Macquart (Kiauka
1974), and Ectocephala capillata (Coquil-
let), (Deeming 1977, 1985). The antennae
of P. signatus appear to be longer and more
prominent than those of P. anchora, but
being a contractile organ (Yamada et al.
1981) the variation in length may be a result
of differential extension during fixation. We
note that the terms antenna and maxillary
palpi are adopted here because of their use
in the taxonomic literature (Teskey 198 1a),
VOLUME 91, NUMBER 2
but these structures also represent the dorsal
and terminal organs respectively of Bolwig
(1946), that are commonly used in descrip-
tive morphology (Chu-Wang and Axtell
1971, 1972, Yamada et al. 1981).
The spinules found on the body of P. sig-
natus are typical for many described chlo-
ropid larvae. However, in Meromyza they
are absent (Fedoseyeva 1966) and in Poly-
odaspis ruficornis Macquart the spinules are
restricted to the ventral surface of the ab-
domen (Kiauka 1974). The mandibles of P.
signatus are similar to those of the necroph-
agous chloropid Conioscinella hinkleyi
(Malloch) (Norrbom 1983), and a preda-
ceous species Steleocerellus latiseta (Lamb)
(Kirk-Spriggs 1986), in being slightly more
elongate and pointed than those of phy-
tophagous larvae. Unfortunately, no other
characters seem to indicate the predatory
habits of this larva.
ACKNOWLEDGMENTS
We thank Curt Sabrosky, Systematic
Entomology Laboratory, USDA, U.S. Na-
tional Museum, for providing us with larvae
of P. anchora for examination. We thank
Anna Asquith for assistance with the illus-
trations. We also thank John D. Lattin, De-
partment of Entomology, Oregon State Uni-
versity, Gerald T. Baker and Larry D.
Corpus, Department of Entomology, Mis-
sissippi State University, for review of the
manuscript; and Greta E. Tyson, Head,
Electron Microscope Center, Mississippi
State University, for providing us with the
use of the electron microscope and for the
preparation of photographic plates.
LITERATURE CITED
Bolwig, N. 1946. Senses and sense organs of the an-
terior end of the house fly larva. Vid. Medd. dansk
nat.-hist. Foren. 109: 81-217.
Breland, O. P. 1941. Podagrion mantis Ashmead and
other parasites of praying mantid egg cases (Hym.:
Chalcidoidea; Dipt.: Chloropidae). Ann. Ent. Soc.
Am. 34: 99-113.
Chu-Wang, I. W. and R. C. Axtell. 1971. Fine struc-
ture of the dorsal organ of the house fly larva,
Musca domestica L. Z. Zellforsch. 117: 17-34.
189
. 1972. Fine structure of the terminal organ of
the house fly larva, Musca domestica L. Z. Zell-
forsch. 127: 287-305.
Davidson, A. 1896. Parasites of spider eggs. Entomol.
News 7: 319-320.
Deeming, J. C. 1977. The immature stages of Ela-
chiptereicus abessynicus Becker and Pachylophus
lugens Loew (Diptera: Chloropidae). Bull. Ento-
mol. Res. 67: 325-328.
1985. A description of the adult and im-
mature stages of Ectecephala capillata (Coquillet)
(Diptera: Chloropidae). Proc. Entomol. Soc. Wash.
87: 402-407.
Fedoseyeva, L. I. 1966. Larval diagnostics of grain-
grass flies of the genus Meromyza Mg. (Diptera,
Chloropidae). Entomol. Rev. 45: 108-113.
Hall, D. G. 1937. The North and Central American
spider parasites of the genus Pseudogaurax (Dip-
tera: Chloropidae). J. Wash. Acad. Sci. 27: 255-
261.
Howard, L. O. 1916. The Insect Book. Doubleday,
Page and Company, New York. 429 pp.
Kiauka, G. F. 1974. The larvae of Polyodaspis rufi-
cornis Mcq. (Diptera, Chloropidae), living in wal-
nuts. Entomol. Rev. 53: 154-156.
Kirk-Spriggs, A. H. 1986. The puparia of some West
African species of Mepachymerus and Steleoce-
rellus (Diptera: Chloropidae). J. Nat. Hist. 20: 767—
776.
Norrbom, A. L. 1983. Four acalyptrate Diptera reared
from dead horseshoe crabs. Entomol. News 9: 1 17—
PAL
Roberts, M. J. 1971. The structure of the mouthparts
of some calypterate dipteran larvae in relation to
their feeding habits. Acta. zool. Stockh. 52: 171-
188.
Sabrosky, C. W. 1945. Three new African Chloropi-
dae with a discussion of the status of Pseudogaurax
(Diptera). Proc. Zool. Soc. London 114: 456-461.
1966. Three new Brazilian species of Pseu-
dogaurax with a synopsis of the genus in the west-
ern hemisphere (Diptera, Chloropidae). Papeis
Avulsos de Dept. Zool. 19: 117-127.
Teskey, H. J. 1981a. Morphology and terminology—
Larvae, pp. 65-88. Jn McAlpine, J. F., et al., eds.,
Manual of Nearctic Diptera. Vol. 1, Monograph
27, Research Branch Agriculture Canada, vi + 674
pp.
1981b. Key to families—Larvae, pp. 125-
147. In McAlpine, J. F., et al., eds., Manual of
Nearctic Diptera. Vol. 1, Monograph 27, Research
Branch Agriculture Canada, vi + 674 pp.
Yamada, Y., Y. Ishikawa, T. Ikeshoji, and Y. Mat-
sumoto. 1981. Cephalic sensory organs of the
onion fly larva, Hylemya antiqua Meigen (Dip-
tera: Anthomyiidae) responsible for host-plant
finding. Appl. Entomol. Zool. 16: 121-128.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 190-202
NEW SPECIES OF MICRO-CADDISFLIES
(TRICHOPTERA: HYDROPTILIDAE) FROM NEW CALEDONIA,
VANUATU AND FIJI
ROBERT W. KELLEY
Biologist, Enwright Laboratories, 25 Woods Lake Rd., Greenville, South Carolina 29607.
Abstract. —New species of micro-caddisfly genera Oxyethira Eaton, Paroxyethira Mose-
ly, Acritoptila Wells and Caledonotrichia Sykora are described from the South Pacific
islands of New Caledonia, Vanuatu (Efate) and Fiji. Range extensions of genera Parox-
yethira and Acritoptila are included and a new subgenus, endemic to New Caledonia, is
added to the genus Oxyethira.
Key Words:
Numerous species of micro-caddisflies
have been reported from the Australasian
biogeographic region. Wells has noted a to-
tal of 12 micro-caddisfly genera from Aus-
tralia: Hydroptila Dalman, Oxyethira
Eaton, Orthotrichia Eaton, Orphninotrichia
Mosely, Maydenoptila Neboiss, Xuthotri-
chia Mosely, Hellyethira Neboiss, Austra-
trichia Wells, and Tricholeiochiton (Kloet
& Hincks). Only two hydroptilid genera,
Oxyethira and Paroxyethira, are known
from New Zealand and one genus, Cale-
donotrichia Sykora, from New Caledonia.
Genera of the tribe Hydroptilini Stephens
seem to be the predominant micro-caddis-
flies in the region. An examination of ma-
terial from the Bishop Museum in Hawaii
revealed previously unpublished species of
four established hydroptilid genera on the
Melanesian islands of New Caledonia, Efate
(Vanuatu) and Fiji.
Described from New Caledonia are five
new species of Oxvethira, two new species
of Caledonotrichia and six new species of
Acritoptila. This brings to 17 the number of
species, distributed among four genera, of
micro-caddisflies known from the island.
caddisfly, new species, New Caledonia, Hydroptilidae
One new species of Oxyethira is described
from Vanuatu and one from Fiji. These dis-
coveries extend the range of Oxyethira to
include Melanesia, of Paroxyethira to in-
clude New Caledonia as well as New Zea-
land, and of Acritoptila which was previ-
ously considered endemic to Australia. New
species of Oxyethira from Melanesia 1n-
clude species in the subgenus Trichoglene
Neboiss and a new subgenus, Pacifica. The
collections also showed a high incidence of
endemism at the species level in New Cal-
edonia. The length and number of antennal
segments were unavailable for several
species. All type specimens are deposited in
the Bishop Museum in Hawaii.
Genus Acritoptila Wells, 1982
Type species: Acritoptila globosa Wells,
1982, by original designation.
Six New Caledonian species of the genus
Acritoptila are described below. This genus
in the Hellyethira complex of genera, was
previously known only from Australia, with
two species in Western Australia and one
species in Queensland. The males of these
VOLUME 91, NUMBER 2
crinita
7 glossocercus
Figs. 1-8.
191
disjuncta
4 ;
ap planichela
Male terminalia of Acritoptila Wells, lateral and dorsal views. A. amphapsis: 1, lateral. A. chiasma:
2, lateral; 3, tergum X (dorsal). 4. crinita: 4, lateral. A. disjuncta: 5, lateral; 6, tergum X (dorsal). 4. glossocercus:
7, lateral. A. planichela: 8, lateral. ap, apico-mesal process; bp, bilobed process; 1a, inferior appendages; sg,
subgenital processes; tX, tergum X; vp, ventrolateral process.
new species share with the Australian species
of the genus, fused inferior appendages (Fig.
17); a complex tergum X with spines and
projections (Fig. 3) and long rod-like pro-
cesses (Fig. 7), described as parameres by
Wells (1982), projecting from the lateral
margin of tergum X. In most of the new
species the fusion of the inferior appendages
is more complete than in the Australian
species. Females, although not described
here, share with the Hellyethira generic
complex shortened apodemes on segments
VIII and IX. The spur formula, wing ve-
nation and other characters of the new
species are consistent with the generic de-
scription by Wells (1982).
Acritoptila amphapsis,
NEw SPECIES
Figs. 1, 9, 10
Male.—Length 2.7 mm. Segment VII: ap-
ico-mesal process of venter elongate. Seg-
ment IX: pleuron produced into blunt pos-
terior lobe; venter excised anteriorly and
posteriorly. Segment X: dorsum split near
base into sclerotized processes which con-
verge at apices; ventrolateral processes
reaching base of subgenital processes. In-
ferior appendages: truncate, joined by nar-
row bridge. Subgenital processes: conver-
gent, arch-shaped in lateral view. Aedeagus
(Fig. 10): elongate, bifurcate apically.
192 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
WwWwS'0
ap q
15. disjuncta ali
glossocercus
at
1 3 crinita
11° chiasma
20
ap
1 9 planichela
Figs. 9-20. Male terminalia of Acritoptila Wells, aedeagus and ventral views. 4. amphapsis: 9, ventral; 10,
aedeagus. 4. chiasma: 11, ventral; 12, aedeagus. A. crinita: 13, ventral; 14, aedeagus. 4. disjuncta: 15, ventral
with inset of caudal view of inferior appendages; 16, aedeagus. 4. glossocercus: 17, ventral; 18, aedeagus. A.
planichela: 19, ventral; 20, aedeagus. ap, apico-mesal process; bp, bilobed process; ej, ejaculatory duct; ia, inferior
appendages; IAe, inferior appendage extensor muscle; sg, subgenital processes; vp, ventrolateral processes.
Female.— Unknown.
Etymology.— Named for the arch-shaped
subgenital processes.
Holotype.—Male. New Caledonia: Ho-
nailu River, 26 Oct. 1958, C. R. Joyce.
Paratypes.— None.
Diagnosis.—The semi-fused inferior ap-
pendages (Fig. 9) and relatively short ven-
trolateral lobes of tergum X (Fig. 1) are more
similar to male genitalia of the Australian
Acritoptila than to the genitalia of other New
Caledonian species.
Acritoptila chiasma,
New SPECIES
Figss2,.3 sll 2
Male.—Antennae 34 segmented. Seg-
ment VIII: ventral apico-mesal process
elongate. Segment IX: pleuron with narrow
sclerotized process on lateral margin. Ter-
gum X: elaborately sculptured with mesal,
blackened tips and elongate ventrolateral
rod-like processes (Fig. 3). Inferior append-
ages: completely fused; membranous caudal
lobe with 2 short setae. Subgenital process-
es: mesal margin bilobed, with inner lobe
auriculate; single lateral seta; bilobed pro-
cess with long lobes. Aedeagus: apex largely
membranous with small apical point (Fig.
12)s
Female. — Unknown.
Etymology.—Named for the membra-
nous mesal area dividing tergum X into right
and left halves.
VOLUME 91, NUMBER 2
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same as holotype, 26 4. Pla-
teau du Dogny, 20 Nov. 1958, | 2.
Diagnosis.—This species is most closely
related to the New Caledonian 4. glosso-
cercus and A. crinita. Like those species, it
bears a lateral process on pleuron IX. But,
the tenth tergum is quite distinctive.
Acritoptila crinita,
NEw SPECIES
Figs. 4, 13, 14
Male.—Segment VII: ventral apico-mesal
process elongate (AP). Segment VIII: dor-
sum with lateral fringe of elongate setae (not
in illustration). Segment IX: lateral process
short, broad, setose. Tergum X: largely
membranous except for sclerotized lateral
margin and caudolateral process. Inferior
appendages: completely fused into rounded
structure (ventral view); membranous cau-
dal lobe with two short setae. Subgenital
processes: mesally bifid and blackened; two
lateral setae; rod-like sclerite connecting lat-
eral lobes of subgenital processes to base of
tergum X; bilobed process with short lobes
covered with minute setae. Inferior ap-
pendages: extensor muscle inserted on lat-
eral lobe of subgenital processes. Aedeagus:
apex with several sclerotized processes (Fig.
14).
Female.— Unknown.
Etymology.— Named for long fringe of se-
tae on dorsum VIII.
Holotype.— Male. New Caledonia: head-
waters of Honailu River, 26 Oct. 1958, C.
R. Joyce.
Paratypes.— None.
Diagnosis. — This species is a sister species
of A. glossocercus, with nearly identical ven-
trolateral rod-shaped processes on tergum
X (Figs. 4, 7). It may be distinguished from
A. glossocercus by the rounded fused infe-
rior appendages (Fig. 13), the setose bilobed
process (Fig. 13) and the non-dilated cau-
dolateral processes of tergum X (Fig. 4).
193
Acritoptila disjuncta,
NEw SPECIES
Figs. 5, 6, 15, 16
Male.—Antennae 35 segmented. Seg-
ment IX: pleuron acutely pointed; venter
broadly excised. Tergum X: ventral pro-
cesses broadened distally with narrow
pointed lobe, asymmetrical; remainder of
tergum membranous. Inferior appendages:
fused and broadly oval in ventral view;
membranous mesal lobe with 3 setae at tip
and one short seta laterad of base of lobe
on each side. Subgenital processes: mesally
connected dorsad of inferior appendages and
with convergent distal processes; paired
caudolateral setae; apices directed ventrally;
bilobed process short. Aedeagus: simple,
with distal sinuate process.
Female.— Unknown.
Etymology.—Named for the appearance
that tergum X 1s not joined to segment IX.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, 3 Nov.
1958, C. R. Joyce.
Paratypes.—Same data as holotype, 5 2.
New Caledonia, Plaine des Lacs area, 3 Nov.
1958, C. R. Joyce.
Diagnosis.— This species is grouped with
chiasma, crinita and glossocercus because
the males bear an elongate ventrolateral
process on tergum X. It differs because its
processes are thicker and asymmetrical, and
the inferior appendages are broad.
Acritoptila glossocercus,
NEw SPECIES
Figs. 7, 17, 18
Male.—Segment VII: ventral apico-mesal
process elongate. Segment IX: lateral pro-
cess of pleuron attenuate, moderate in
length. Tergum X: largely membranous,
with curved lateral process; elongate ven-
trolateral rod-shaped processes widened
subapically. Inferior appendages: complete-
ly fused; tongue-shaped in caudal view;
membranous caudal lobe with two short se-
tae. Subgenital processes: mesally heavily
194
sclerotized and bilobed; one lateral seta; bi-
lobed process with short lobes. Aedeagus:
single apical sclerotized process (Fig. 18).
Female. — Unknown.
Etymology.—Named for the tongue-
shaped inferior appendages.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes. — None.
Diagnosis. —A sister species of 4. crinita,
with similar ventrolateral rod-shaped pro-
cesses on tergum X (Figs. 4, 7), this species
may be most easily recognized by the di-
lated condition of these processes (Fig. 7)
and the tongue-shaped inferior appendages
(caudal view—Fig. 17)
Acritoptila planichela,
New SPECIES
Figs. 8, 19, 20
Male.—Segment VII. Apico-mesal pro-
cess of venter short, acutely pointed. Seg-
ment IX: pleuron truncate; venter anterior-
ly truncate. Tergum X: distally flared with
ventral flat, heavily sclerotized, claw-shaped
processes. Inferior appendages: fused into
broad plate with pair of ventral protuber-
ances and single spine at each lateral mar-
gin. Sclerotized subgenital structure ventrad
of aedeagus, with narrow sclerotized band
connecting it to tergum X. Aedeagus: sim-
ple, elongate, lacking titillator (Fig. 20).
Female.— Unknown.
Etymology.—Named for the flat claw of
tergum X.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, 3 Nov.
1958, C. R. Joyce.
Paratypes.—Same data as holotype, 2 é.
Diagnosis. — This 1s the most divergent of
the New Caledonian species of Acritoptila.
The subgenital structures are difficult to
homologize with the subgenital processes of
other species of Acritoptila. It is the only
species in the genus with a short apico-me-
sal process of venter VII. Relationships with
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
other New Caledonian Acritoptila are un-
clear.
Genus Caledonotrichia Sykora, 1967
Type species: Caledonotrichia iiliesi Sykora,
1967.
As noted by Wells (1983), males of Ca/e-
donotrichia closely resemble those of May-
denoptila Neboiss. Indeed, if only genitalic
characters are analyzed, the two genera could
be synonymous. Adult males of Caledono-
trichia may be distinguished by the paired
subgenital processes. In Maydenoptila, the
subgenital processes are fused or absent.
Caledonotrichia species have a transverse
suture on the mesoscutellum, as do
Maydenoptila species. In the generic de-
scription given by Marshall (1979), the su-
ture was described as absent in Caledono-
trichia.
Caledonotrichia charadra,
New SPECIES
Figs. 21-23
Male.—Segment IX: anterior and caudal
margins of venter truncate; pleuron sinuate;
dorsum a broad band. Subgenital processes:
parallel, contiguous, caudally spatulate in
ventral view and anteriorly divergent.
Membranous sac within segment IX. Infe-
rior appendages: bilobed and attached to
dorsolateral margin of segment IX; upper
lobe sickle-shaped with blackened, ventral-
ly directed tooth on mesal margin; ventral
lobe triangular in shape. Aedeagus: largely
membranous with elongate sclerotized pro-
cess originating at base of aedeagus (Fig. 23).
Female.— Unknown.
Etymology.— Named habitat of holotype
in mountain stream.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, 31 4.
Diagnosis.—This species is closely relat-
ed to C. minor Sykora, particularly in the
shape of the inferior appendages and ae-
VOLUME 91, NUMBER 2
Figs. 21-32.
22, ventral; 23, aedeagus. C. extensa: 24, lateral; 25, ventral; 26, aedeagus. P. dumagnes: 27, lateral; 28, ventral;
29, aedeagus. P nigrispina: 30, lateral; 31, ventral; 32, aedeagus. ap, apio-mesal process; as, aedeagal sheath; ej,
ejaculatory duct; ia, inferior appendages; iae, inferior appendage extensor muscle; ial, inferior appendage lower
lobe; iau, inferior appendage upper lobe; ms, membranous sac; sg, subgenital process; ti, titillator; vl, ventral
lobe.
deagus. It may be distinguished by the shape
of the subgenital processes.
Caledonotrichia extensa,
NEw SPECIES
Figs. 24-26
Male.—Antennae 27 segmented. Seg-
ment IX: venter protruded anteriorly within
segment VIII; caudal margin broadly ex-
cised. Subgenital processes: elongate and
narrow. Tergum X: elongate, tongue-shaped.
Inferior appendages: bilobed; attached to
dorsolateral margin of segment IX; dorsal
lobe long, with ventrally directed tooth and
+ re]
P. nigrispina
Male terminalia of Caledonotrichia Sykora and Paroxyethira Mosely. C. charadra: 21, lateral;
peg-like setae at base; ventral lobe broadly
oval. Aedeagus: elongate; dorsal crease along
most of length; membranous lobe extends
distally beyond tip of ejaculatory duct.
Female.— Unknown.
Etymology.—Named for divergent con-
dition of inferior appendages.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, | 4.
Diagnosis. —C. extensa is distinctive, with
the lobes of the inferior appendages longer
than wide. It is likely a sister to the distinct
196 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
group composed of the other species in the
genus.
Genus Oxyethira Eaton, 1873
Subgenus Trichoglene (Neboiss, 1977)
Type species: Oxyethira albiceps (Mac-
Lachlan, 1862).
Two species of Oxyethira from New Cal-
edonia are placed in subgenus 7richoglene
Neboiss along with the New Zealand O. al-
biceps and the Australian brevis Wells, co-
lumba (Neboiss), mienica Wells, triangu-
lata Wells and retracta Wells. Males of the
two new species share with these other
species a plesiomorphic character; a com-
plete, non-excised segment VIII, and two
apomorphic characters; aedeagus with re-
curved sub-distal spinous process and
subgenital processes widely separated and
partly fused with each pleuron of segment
IX. Males of O. caledoniensis n. sp. are sim-
ilar to O. brevis in having a short titillator,
but seem to be more primitive than all Aus-
tralian and New Zealand species in the re-
tention of a sclerotized mesal connection
between the subgenital processes. O. insu-
laris n. sp. males are aberrant, with an en-
larged forked structure, apparently the
modified inferior appendages, on elongate
venter IX. The forked structure is like that
found in males of the subgenus Dactylotri-
chia Kelley but is probably not homologous.
The type species of 7richoglene was 1n-
correctly identified as O. columba (Neboiss)
in Kelley (1984). It should be O. albiceps
(MacLachlan). A redescription of the sub-
genus was given in Kelley (1984).
Oxyethira caledoniensis,
New SPECIES
Figs. 33, 42, 56
Male.—Segment VIII: cylindrical, with-
out excisions. Segment IX: dorsum narrow
in lateral view; venter reaching anterior end
of segment VIII. Inferior appendages: re-
duced to small, widely separated bilobed
areas of sclerotization. Subgenital process-
es: widely separated, narrow, connected by
thin mesal sclerotized strip; dorsolaterally
fused to pleuron IX; bilobed processes short.
Aedeagus: titillator short; recurved subdis-
tal process.
Female.— Unknown.
Etymology.—Named for island where
holotype was collected.
Holotype. — Male. New Caledonia: Plum,
20-60 m, malaise trap, 23-25 Mar. 1968,
J. L. Gressitt & T. C. Maa.
Paratypes.— None.
Oxyethira insularis,
New SPECIES
Figs. 34, 43, 57
Male.—Length 1.7 mm. Segment VIII:
cylindrical, without excisions. Segment IX:
dorsum broad; venter elongate, reaching an-
terior end of segment VII; venter with large
forked process, fused inferior appendages,
subtending membranous, serrately tipped
plate. Subgenital processes: acutely pointed,
connected mesally by curved sclerotized
band and connected dorsolaterally to pleu-
ron IX; bilobed processes short. Aedeagus:
ttillator lacking; sclerotized, recurved pro-
cesses at apex.
Female.— Unknown.
Etymology.— Named for its island habi-
tat.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, | é.
Genus Oxyethira Eaton, 1873
Subgenus Pacificotrichia, n. subgen.
Seven new species of Oxyethira from New
Caledonia, Vanuatu and Fiji are arranged
here in a new subgenus endemic to the South
Pacific islands. Relationships with other
subgenera are unclear, although Pacificotri-
chia is most similar to Dampfitrichia Mose-
ly. Similarities between the males include
the shallowly excised venter VIII (Fig. 62),
the deeply excised dorsum VIII (Fig. 37),
fused R, and R, forewing veins, configura-
VOLUME 91, NUMBER 2
eye) Vill
caledoniensis
36
indorsennus
39
scutica
ap
Figs. 33-41.
197
a7 Se 35
insularis dorsennus
38
oropedion
41
fijiensis
Male terminalia of Oxyethira Eaton, lateral views. 33, O. caledoniensis. 34, O. insularis. 35, O.
dorsennus. 36, O. indorsennus. 37, O. melasma. 38, O. oropedion. 39, O. scutica. 40, O. efatensis. 41, O. fijiensis.
ap, apico-mesal process; bp, bilobed process; ia, inferior appendages; sg, subgenital processes; sl, tapered seta.
tion of the subgenital processes, and reduc-
tion or loss of the pre-apical spur on the
meso-tibia. The relatively simple male gen-
italia, with a reduced ninth segment, are like
the genitalia of the minima group of sub-
genus Dampfitrichia, the only other known
Australasian group of Oxyethira besides the
endemic subgenus Trichoglene. But, there
are important differences: unlike males of
the minima group, the subgenital processes
are distally fused (Fig. 56), do not bear distal
setae (Fig. 63) and retain the bilobed pro-
cess. The lack of significant shared apo-
morphies prohibit the placement of these
species in subgenus Dampfitrichia.
Two distinct groups can be identified in
subgenus Pacificotrichia. The oropedion
group includes species O. dorsennus n. sp.,
indorsennus n. sp., melasma n. sp., orope-
dion n. sp. and scutica n. sp. In males of
these species, venter IX does not reach the
anterior end of segment VIII (Fig. 35) and
the fused subgenital processes bear a pro-
nounced mesodistal projection (Fig. 58).
Females are characterized by elongate apo-
demes (Fig. 50) and cerci (Fig. 53), as well
as by a sclerotized lateroventral projection
on tergum VIII. The efatensis group in-
cludes O. efatensis n. sp. and fijiensis n. sp.
Males of this group are distinguished by
venter IX which extends into segment VII
(Fig. 40), subgenital processes which are not
connected anteromesally (Fig. 63) and an
aedeagus with a tapered dorsal sclerotized
198 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
42 43 44 45 46 47 48
®
dorsennus
SCcutica
22
scutica
scutica
Figs. 42-55.
as
o q
S \ 50
T= 2 , oropedion
Si = r
3 x c)
a
=) | 3 :
° eh 2 ®
Ss a
c
o ® |
= i=)
o
E 3
® =}
oO
®
c
= 8
al
oO
S {\
2 \
L \
BY
sacl oil
oropedion
sp
efatensis
Aedeagi and female terminalia of Oxyethira Eaton. Figs. 42-49, aedeagi. 42, O. caledoniensis.
43, O. insularis. 44, O. dorsennus. 45, O. melasma. 46, O. oropedion. 47, O. scutica. 48, O efatensis. 49, O
fyiensis. Figs. 50-53, female terminalia. O. oropedion: 50, lateral; 51, ventral. O. scutica: 52, lateral; 53, ventral.
O. efaensis: 54, lateral; 55, ventral. ap, apodeme IX; e}, ejaculatory duct; hl, horizontal lamella; sp, spermathecal
process; ss, spermathecal sclerite; ti, titullator; tX, tergum X; vl, ventral lobe.
process and a membranous ventral tube (Fig.
49). Females bear short apodemes and cerci
(Fig. 54).
Subgenus Pacificotrichia, n. subgen.
Type species: Oxyethira oropedion, n. sp.
Spur formula: 0-2-4 or 0-3-4.
Forewings: R, and R, fused.
Male.—Length 2.1-2.4 mm. Antennae
24-30 segmented; sparse placoid sensilla.
Segment VIII: venter shallowly excised (Fig.
63); dorsum deeply excised. Segment IX:
dorsum narrow (Fig. 40); posterolateral pro-
cesses lacking; venter reaching to anterior
end of segment VIII or posterior end of seg-
ment VII (Figs. 37, 40). Inferior append-
ages: small or lacking (Fig. 40); setal lobes
usually indistinct. Subgenital processes:
fused distally (Fig. 63); bilobed process short
(Fig. 62). Aedeagus: titillator absent (Fig.
48).
Female.—There is too much variability
among females to describe subgeneric char-
acters.
Range. — New Caledonia; Vanuatu; Fi1.
VOLUME 91, NUMBER 2
caledoniensis
t 0.5mm 4
61
scutica
oropedion
Figs. 56-63.
57
insularis
199
Vill- J =; —bp
58
dorsennus
59
melasma
62
efatensis
fijiensis
Male terminalia of Oxyethira Eaton, ventral views. 56, O. caledoniensis. 57, O. insularis. 58,
O. dorsennus. 59, O. melasma. 60, O. oropedion. 61, O. scutica. 62, O. efatensis. 63, O. fyiensis. bp, bilobed
process; ia, inferior appendages: mr, mesal ridge; sg, subgenital processes; sl, setal lobe.
oropedion Group
Oxyethira dorsennus,
New SPECIES
Figs. 35, 44, 58
Spur formula: 0-2-4.
Male.—Length 2.1 mm. Antennae 25 seg-
mented. Segmented VII: ventral apico-me-
sal process present. Segment VIII: dorsum
with deep excision in lateral view. Inferior
appendages: darkened, nearly contiguous
mesally. Subgenital processes: triangular
fused apex. Aedeagus: lacking distal pro-
cesses.
Female.— Unknown.
Etymology.—Named for the hump on
dorsum VIII.
Holotype.— Male.
New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, | 2.
New Caledonia, Plum, 20-60 m, 23-25 Mar.
1958, 1 3.
Oxyethira indorsennus,
NEw SPECIES
Figs. 36, 44, 58
Spur formula: 0-3-4.
Male.—Length 2.1 mm. Antennae: 25
segmented. Segment VII: ventral apico-
mesal process present. Segment VIII: dor-
sum gradually excised in lateral view. In-
ferior appendages: darkened, nearly contig-
uous mesally. Subgenital processes:
triangular fused apex. Aedeagus: lacking
distal processes.
200
Female. — Unknown.
Etymology.— Named for absence of dor-
sal hump on dorsum VIII.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, 5 2.
Diagnosis.—This species differs from O.
dorsennus, n. sp. only in the shape of dor-
sum VIII (Figs. 35, 36) and the spur for-
mula.
Oxyethira melasma,
NEw SPECIES
Figs. 37, 45, 59
Spur formula: 0-2-4.
Male.—Antennae: 23-30 segmented.
Segment VII: ventral apico-mesal process
large. Segment VIII: venter with shallow,
acutely pointed excision; setae stout with
tapered curved apices. Segment IX: venter
with mesal ridge, caudal end tapered to
acutely pointed tip; dorsum membranous.
Inferior appendages: indistinct; setal lobes
lacking. Subgenital processes: with ventral
black spot at fused apex; elongate bilobed
process with mesally contiguous lobes. Ae-
deagus: apical sclerotized process.
Female.— Unknown.
Etymology.—Named for the black spot
on the subgenital processes.
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, | 4.
New Caledonia, Honailu River headwaters,
26 Oct. 1958, 1 6.
Oxyethira oropedion,
New SPECIES
Figs. 38, 46, 50, 51, 60
Spur formula: 0-3-4.
Male.—Antennae: 26 segmented. Seg-
ment VII: ventral apico-mesal process pres-
ent. Segment VIII: venter not excised; dor-
sum deeply excised. Inferior appendages:
widely separated; setal lobes forming setose
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
membranous bridge between inferior ap-
pendages. Aedeagus: without apical pro-
cesses.
Female.—Segment VIII: tergum and apo-
demes elongate; sternum with ventral black
spot. Spermathecal sclerite indistinct.
Etymology.—Greek for plateau, the ho-
lotype habitat.
Holotype.— Male. New Caledonia: Pla-
teau de Dogny, light trap, 20 Nov. 1958, C.
R. Joyce.
Paratypes.—Same data as holotype, | é.
Diagnosis.—In ventral aspect, the males
are distinguished by the combination of
widely separated inferior appendages and
broad subgenital processes. The females are
easily distinguished by the black spot on
sternum VIII.
Oxyethira scutica,
New SPECIES
Figs. 39, 47, 52, 53, 61
Spur formula; 0-2-4.
Male.—Antennae: 26 segmented. Seg-
ment VII: ventral apico-mesal process pres-
ent. Segment VIII: pleuron roundly trun-
cate; venter moderately excised. Inferior
appendages: minute contiguous lobes at
caudal tip of venter IX; setal lobes lacking.
Subgenital processes: with paied black spots
bordering caudomesal protrusion; bilobed
process with elongate lobes. Aedeagus: nar-
row elongate process extending from apex.
Female.— Antennae: 20 segmented. Seg-
ment VIII: tergum and apodemes elongate.
Spermathecal process distinct. Sclerotized
sac present caudad of spermathecal process.
Etymology.—Named for the elongate
whip-like process of the aedeagus (Fig. 47).
Holotype.—Male. New Caledonia:
mountain stream up Boulari River, light
trap, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, 3 2.
Same locality as holotype, 17 Nov. 1958,
17 6.
Diagnosis. — Males of this species may be
recognized by the minute inferior append-
VOLUME 91, NUMBER 2
ages and triangular (ventral view) subgenital
processes, in addition to the shape of the
aedeagal process. Oxyethira scutica is the
only species of subgenus Pacificotrichia with
paired black teeth along the posterior fused
margin of the subgenital processes. This
character 1s common among species of other
neotropical subgenera of Oxyethira.
efatensis Group
Oxyethira efatensis,
NEw SPECIES
Figs. 40, 48, 54, 55, 62
Spur formula: 0-3-4.
Male. —Length 2.4 mm. Antennae 26 seg-
mented. Segment VII: ventral apico-mesal
process present. Segment VIII: venter and
pleuron slightly excised. Inferior append-
ages: triangular in shape, discrete. Aede-
agus: with dorsal, sclerotized, attenuate band
subtended by membranous tube (Fig. 48).
Female.—Length 2.4 mm. Antennae 21
segmented. Segment VIII: tergum short with
anterior margin heavily sclerotized. Seg-
ment IX: lacking. Spermathecal process
teardrop-shaped; horizontal lamella dis-
tinctly sclerotized.
Etymology.—Named for locality of col-
lection.
Holotype.— Male. Vanuatu: Efate (NW),
Maat, Ambryn Village, 3M., 18 Aug. 1957,
light trap, J. L. Cressitt.
Paratypes.—Same data as holotype, 4 4.
Same locality as holotype, 19 Aug. 1957,
5 6.
Diagnosis.—This species may be distin-
guished from O. fijiensis n. sp. by the excised
pleuron VIII and the median ventral pro-
jection of the fused subgenital processes.
Oxyethira fijiensis,
New SPECIES
Figs. 41, 49, 63
Spur formula: 0-3-4.
Male. —Segment VII: ventral apico-mesal
process lacking. Segment VIII: venter with
small mesal excision; pleuron truncate. Seg-
201
ment IX: pleuron with dorsolateral point on
anterior margin; venter pointed anteriorly.
Inferior appendages: small, indistinct, widely
separated. Subgenital processes: fused cau-
dal margin with mesal tooth. Aedeagus: one
sclerotized blade-like process and one
membranous process enclosing ejaculatory
duct (Fig. 49).
Female.— Unknown.
Etymology.— Named for collection local-
ity of holotype.
Holotype.— Male. Fiji: Levu, Nandari-
vatu, Jan. 1955, N. L. H. Krauss.
Paratypes.— None.
Diagnosis.—This species may be distin-
guished by the distomesal tooth on the
subgenital processes.
Genus Paroxyethira Mosely, 1924
The two new species of Paroxyethira
Mosely described here are the first known
from this genus outside New Zealand. Both
species are similar to previously described
species of Paroxyethira in the morphology
of the terminal abdominal segments as well
as wing venation, spur formula and other
non-genitalic characters. The genus is char-
acterized by the elongate ventral process of
segment VIII (Fig. 30) and the asymmetrical
aedeagal sheath in the males (Fig. 30). A
description of the genus was given by Mar-
shall (1979).
Paroxyethira dumagnes
NEw SPECIES
Figs. 27, 29, 31
Male.—Antennae 25 segmented. Seg-
ment VII: ventral apico-mesal process pres-
ent. Segment VIII: length short; venter with
elongate spatulate process; pleuron with
elongate setae. Segment IX: dorsum asym-
metrical and tilted anteriorly; pleuron pro-
truded caudally. Inferior appendages: elon-
gate with numerous black setae on inner
margin; extensor muscles originate on an-
terior pleuron IX and insert on anterolateral
corners of inferior appendages. Aedeagus:
202
sheath short, asymmetrical, with several
short setae and a long anterior apodeme;
one muscle originates caudally on venter IX
and inserts on apex of apodeme, another
inserts on caudal end of sheath; aedeagus
elongate, with titillator and two distal lobes.
Female. — Unknown.
Etymology.— Named for the inferior ap-
pendages, which have black setae which ap-
pear like magnetic filings.
Holotype.— Male. New Caledonia: Bou-
lari River, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, 2 2.
Same locality as holotype, 17 Nov. 1958,
165 3.
Diagnosis.—Paroxyethira dumagnes
males have a short aedeagal sheath with an
anterior apodeme (Fig. 27) as do males of
P. tillyardi Mosely. However the apex of the
aedeagal sheath is different from all other
known species of the genus.
Paroxyethira nigrispina,
New SPECIES
Figs. 30-32
Male.—Antennae 22 segmented. Seg-
ment VII: short ventral apico-mesal process
present. Segment VIII: length short; venter
with elongate, flattened, gradually tapered
process. Segment IX: dorsum excised to an-
terior margin, asymmetrical; pleuron pro-
truded caudally. Inferior appendages: short-
er than segment IX. Aedeagus: sheath
elongate, with small spine at midlength and
thick spine at base of apical process; apical
process sinuate and black at tip: aedeagus
elongate with titillator and two distal lobes.
Female. — Unknown.
Etymology.— Named for black tip of ae-
deagal sheath.
Holotype.— Male. New Caledonia: Bou-
lari River, 3 Nov. 1958, C. R. Joyce.
Paratypes.—Same data as holotype, 86 ¢.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Same locality as holotype, 17 Nov. 1958,
9 8.
Diagnosis.—Paroxyethira nigrispina
males have an elongate aedeagal sheath sim-
ilar to males of P. kimminsi Leader. The
apex of the sheath however is distinct.
ACKNOWLEDGMENTS
I am grateful to the Bishop Museum in
Hawaii and Alice Wells of the University
of Adelaide for the loan of specimens. John
Morse of Clemson University and Enwright
Laboratories of Greenville, South Carolina
were also supportive. This publication is
Technical Contribution #2806 of the Clem-
son University Agricultural Experiment
Station.
LITERATURE CITED
Eaton, A. E. 1873. On the Hydroptilidae, a family of
Trichoptera. Trans. Entomol. Soc. London 1873:
125-150.
Kelley, R. W. 1984. Phylogeny, morphology and clas-
sification of the micro-caddisfly genus Oxyethira
Eaton (Trichoptera: Hydroptilidae). Trans. Amer.
Entomol. Soc. 110: 435-463.
Mac Lachlan, R. 1862. Characters of new species of
exotic Trichoptera. Trans. Entomol. Soc. London
(3)1: 301-311.
Marshall, J. E. 1979. A review of the genera of the
Hydroptilidae (Trichoptera). Bull. Br. Mus. (Nat.
Hist.) Entomol. 39: 135-239.
Mosely, M.E. 1924. New Zealand Hydroptilidae (Or-
der Trichoptera). Trans. R. Soc. N.Z. 55: 670-673.
Neboiss, A. 1977. A taxonomic and zoogeographic
study of Tasmanian caddis-flies (Insecta: Trichop-
tera). Mem. Natl. Mus. Victoria 38: 1-208.
Sykora, J. 1967. Trichoptera collected by Prof. J.
Illies in New Guinea and New Caledonia. Pac.
Insects 9: 585-595.
Wells, A. 1982. Tricholeiochiton Kloet & Hincks and
new genera in the Australian Hydroptilidae (Tr-
choptera). Aust. J. Zool. 30: 251-270.
1983. New species in the Australian Hydrop-
tilidae (Trichoptera), with observations on rela-
tionships and distributions. Aust. J. Zool. 31: 629-
649.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 203-205
WASP PARASITES OF THE BURDOCK SEED MOTH,
METZNERIA LAPPELLA ZELLER (GELECHIIDAE):
NEW HOST RECORD FOR BRACON MELLITOR SAY
(HYMENOPTERA: BRACONIDAE)
FOSTER FORBES PURRINGTON AND DEBORAH H. STINNER
Department of Entomology, The Ohio State University, The Ohio Agricultural Research
and Development Center, Wooster, Ohio 44691.
Abstract. —Bracon mellitor parasitizes the burdock seed moth, Metzneria lappella. This
is anew host record for this wasp, best known as a parasite of the boll weevil, Anthonomus
grandis. Agathis malvacearum, another wasp parasite of this moth, is newly reported from
the Pacific coast of Washington, a westward range extension of 2000 km. We also report
a transcontinental North American distribution for the host—parasite association of M.
lappella with Hyssopus thymus, the paramount Nearctic wasp parasite of the European
pine shoot moth, Rhyacionia buoliana.
Key Words:
The burdock seed moth, Metzneria lap-
pella Zeller (Lepidoptera: Gelechiidae) eats
the seeds of burdock (the Palearctic genus,
Arctium L. (Compositae)) by excavating in
the indehiscent flower head, or bur. Natu-
ralized in North America, this univoltine
moth is found wherever common burdock,
A. minus (Hill) Bernhardi, is established
(Purrington 1970). Field collections of bur-
dock burs made in the United States have
yielded several wasp species that parasitize
M. lappella \arvae (Juhala 1967, Purrington
1970, 1979, Purrington and Uleman 1972).
In this paper, we report a new primary wasp
parasite of this moth and record new dis-
tributions for two other parasites.
We obtained about 100 burs in Matta-
poisett, MA (12-II-86), which yielded 250
M. lappella larvae and 7 overwintering pre-
pupal larvae of Bracon mellitor Say (Hy-
menoptera: Braconidae). This was the first
record of B. mellitor as a parasite of M.
lappella. Wasp larvae were enclosed in pa-
Braconidae, burdock, distribution, Gelechiidae
pery silken cocoons inside host excavations
within the burs. Bracon mellitor is a solitary
univoltine ectoparasite of several larval
weevils and moths in the Nearctic region
(Adams etal. 1969, Cross and Chesnut 1971,
Krombein et al. 1979, Cuda and Burke
1983). Itis an important parasite of the boll
weevil, Anthonomus grandis Boheman (Co-
leoptera: Curculionidae) (Adams et al. 1969,
Sturm and Sterling 1986, Pencoe and Phil-
lips 1987).
Agathis malvacearum Latreille (= A.
metzneriae Muesebeck) (Hymenoptera:
Braconidae) is an Old World solitary uni-
voltine endoparasitic wasp introduced into
the Western Hemisphere, probably with
burdock containing M. lappella. In the Pale-
arctic, 4. malvacearum is also found on the
moths, /. carlinella Stainton (Gelechiidae),
Evetria resinella L. (Gelechiidae), and Co-
leophora otitae Zeller (Coleophoridae)
(Shenefelt 1970). North American distri-
bution records are Quebec, New England,
204
Long Island, and west to North Dakota (Ju-
hala 1967, Krombein et al. 1979). We found
A. malvacearum on M. lappella in burdock
collected at Portland, OR (Reed College, 25-
XII-85, col. C. B. Purrington), Bellfountain,
OR (10-I-86), and Seattle, WA (Pike Place,
6-1-86). The new records extend the wasp’s
known range by 2000 km westward to the
Pacific coast.
Evidently burdock has been present in
western Washington since at least the 1930's
(Gunther 1973). It arrived in the New World
with European settlers in the same way that
it has become ubiquitous: its burs are armed
with hooked spiny bracts that engage fur
and cloth. Although Fyles (1899) speculates
that M. lappella arrived in Quebec from Eu-
rope only late in the 19th century, burdock
infested with M. lappella and its braconid
parasite, A. malvacearum, likely became es-
tablished in North America soon after the
first settlements were made along the At-
lantic coast. Gross et al. (1980) review the
introduction and early distribution of bur-
dock in North America.
We recovered Hyssopus thymus Girault
(Hymenoptera: Eulophidae), a multivol-
tine, gregarious, larval ectoparasite from M.
lappella in burrs obtained at Mattapoisett,
MA (12-II-86), Wooster, OH (10-IV-86),
and Kent, WA (2-I-86). These new collec-
tion records establish a coast-to-coast North
American distribution for the M. /appella—
H. thymus host-parasite association, a re-
lationship first reported from North Dakota
(Purrington 1970). According to Syme
(1974), H. thymus is the most common,
widespread, and effective Nearctic parasite
of the European pine shoot moth, Rhy-
acionia buoliana (Schiffermiiller) (Tortrici-
dae).
We anticipate that wherever burdock be-
comes established it will have arrived with
some members of a small integrated insect
community. We predict this community will
include the Old World co-immigrants, !.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lappella and A. malvacearum, as well as
eclectic elements from native fauna.
ACKNOWLEDGMENTS
We thank P. M. Marsh (USDA-Beltsville)
for identifying A. malvacearum and B. mel-
litor, and M. E. Schauff (USDA-Beltsville)
for identifying H. thymus. C. B. Purrington
supplied burdock burs from Portland, OR;
G. Smith and S. Blue provided support fa-
cilities in Seattle, WA. Salaries and research
support provided by State and Federal
Funds appropriated to The Ohio Agricul-
tural Research and Development Center,
The Ohio State University. Manuscript
number / 10-88.
LITERATURE CITED
Adams, C. H., W. H. Cross, and H.C. Mitchell. 1969.
Biology of Bracon mellitor, a parasite of the boll
weevil. J. Econ. Entomol. 62: 889-896.
Cross, W. H. and T. L. Chesnut. 1971. Arthropod
parasites of the boll weevil, Anthonomus grandis:
1. An annotated list. Ann. Entomol. Soc. Am. 64:
516-526.
Cuda, J. P. and H. R. Burke. 1983. Trichobaris brid-
welli, a new host for Bracon mellitor. Southwest.
Entomol. 8: 65-66.
Fyles, T. W. 1899. Notes on the season of 1899. Ann.
Rept. Entomol. Soc. Ontario 19: 105-106.
Gross, R. S., P. A. Werner, and W. R. Hawthorn. 1980.
The biology of Canadian weeds. 38. Arctium mi-
nus (Hill) Bernh. and A. /appa L. Can. J. Plant Sci.
60: 621-634.
Gunther, E. 1973. Ethnobotany of western Washing-
ton. Univ. Washington Press, Seattle, 71 pp.
Juhala, C. 1967. Notes on parasitic Hymenoptera
associated with a gelechiid moth, Metzneria lap-
pella, in the common burdock, and description of
a new species of Agathis (Braconidae). Ann. Ento-
mol. Soc. Am. 60: 95-97.
Krombein, K. V., P. D. Hurd, Jr., D. R. Smith, and
B. D. Burks. 1979. Catalog of Hymenoptera in
America north of Mexico. Smithsonian Institution
Press, Washington, DC, 2209 pp.
Pencoe. N. L. and J. R. Phillips. 1987. The cotton
boll weevil: Legend, myth, reality. J. Entomol. Sci.
Suppl. 1: 30-51.
Purrington, F. F. 1970 Ecology of Metzneria lappella
(Lepidoptera: Gelechiidae) and its hymenopterous
VOLUME 91, NUMBER 2
parasites in eastern North Dakota. Ann. Entomol.
Soc. Am. 63: 942-945.
1979. Biology of the hyperparasitic wasp
Perilampus similis (Hymenoptera: Perilampidae).
Great Lakes Entomol. 12: 63-66.
Purrington, F. F.and J.S. Uleman. 1972. Brood size
of the parasitic wasp Hyssopus thymus (Hyme-
noptera: Eulophidae): Functional correlation with
the mass of a cryptic host. Ann. Entomol. Soc.
Am. 65: 280-281.
205
Shenefelt, R. D. 1970. Braconidae 3: Agathidinae,
pp. 342-343. /n Ferriere, C. and J. van der Vecht,
eds., Hymenopterorum Catalogus, W. Junk, Dor-
drecht.
Sturm, M. M. and W. L. Sterling. 1986. Boll weevil
mortality factors within flower buds of cotton. Bull.
Entomol. Soc. Am. 32: 239-247.
Syme, P. D. 1974. Observations of the fecundity of
Hyssopus thymus (Hymenoptera: Eulophidae).
Can. Entomol. 106: 1327-1332.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 206-229
THE COCKROACH GENUS MARGATTEA SHELFORD, WITH A NEW
SPECIES FROM THE KRAKATAU ISLANDS, AND
REDESCRIPTIONS OF SEVERAL SPECIES
FROM THE INDO-PACIFIC REGION
(DICTYOPTERA: BLATTARIA: BLATTELLIDAE)
Louis M. RoTH
Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts
02138. (Correspondence: Box 540, Sherborn, Massachusetts 01770.)
Abstract. — Diagnostic characters for the cockroach genus Margattea Shelford are given.
Theganosilpha Kumar and Princis 1s synonymized with Margattea. Margattea paracey-
lanica, n. sp. from the Krakatau Islands is described. The following Wargattea species are
redescribed: ceylanica (Saussure), nimbata (Shelford) (previously considered a synonym
of ceylanica), perspicillaris (Karny), anceps (Krauss), contingens (Walker) [= humeralis
(Walker)], and /ongealata (Brunner). A key to the above males is provided. Symploce
obtusifrons (Walker) (sp. incertae sedis) is transferred to Margattea. The following species
of Margattea are transferred to the genus Ba/ta Tepper: aurea Hanitsch, setifera (Hanitsch),
parvula (Bolivar), crassivenosa (Bolivar), and /ongicercata (Bolivar). The position of the
male’s genital hook and rotation of the ootheca in subfamily placement (Pseudophyllo-
dromiinae or Blattellinae) is discussed.
Key Words:
Princis (1969: 862, 1971: 1162) listed 44
species of Margattea Shelford. It 1s a widely
distributed genus with species occurring in
Asia, Indonesia, Malaysia, South Pacific,
and Africa. This paper is a result of my
attempt to identify a few specimens of Mar-
gattea collected in the Krakatau Islands and
Australia. After reading the many species
descriptions it was evident that the genus
needed revision and that the various taxa
should be redescribed with particular em-
phasis on male genitalia. In the original de-
scriptions of known species, 12 were based
on both sexes, 17 on males, and 15 on fe-
males only. Even when males were de-
scribed, a number of characters which are
known to be important today (e.g. type of
front femur, whether or not the tarsal claws
are toothed, whether or not the male has a
Cockroaches, Blattellidae, taxonomy, Krakatau Islands
tergal gland, and the shapes of the male gen-
ital phallomeres were not mentioned). Iden-
tification of females, when only that sex is
known, is difficult unless the specimens are
topotypic and the species have distinctive
color patterns. Based on descriptions a
number of species of Margattea belong in
another genus. For example, I have exam-
ined the types of the following species and
they all belong in the genus Ba/ta Tepper:
Margattea aurea Hanitsch (HDEO), M. se-
tifera Hanitsch (NRSS), M. parvula Bolivar
(BMNH), 7. crassivenosa Bolivar (BMNH),
and M. longicercata Bolivar (BMNH).
The Krakatau specimens were collected
by personnel from institutions in Australia,
Indonesia, and the United Kingdom during
1984 and 1985 expeditions (Thornton and
Rosengren, in press). Thirteen species of
VOLUME 91, NUMBER 2
cockroaches were taken and I will report on
these elsewhere. Only one species of Mar-
gattea was collected and it did not agree
with the four species of the genus previously
reported from these islands; I am describing
it as new. I will present the diagnostic char-
acters of the genus and redescribe the fol-
lowing taxa that have been recorded from
the Krakataus by others: M. ceylanica
(Saussure), Mf. contingens (Walker) [= M.
humeralis (Walker)], M. anceps (Krauss),
and M. nimbata (Shelford). Kuchinga He-
bard has been considered a junior synonym
of Margattea, and I have concluded that
Theganosilpha Kumar and Princis also is a
junior synonym and therefore will rede-
scribe their type species [Kuchinga
longealata (Brunner) and Theganosilpha
perspicillaris (Karny)].
The following museums and their cura-
tors or collection managers loaned me spec-
imens:
(ANIC)—Australian National Insect
Collection, CSIRO, Canberra,
A:G@-r.. Australia: Dr. D.C.F.
Rentz.
(ANSP)—Academy of Natural Sciences
of Philadelphia, PA, U.S.A.;
Mr. Donald Azuma.
(BMNH)-—British Museum (Natural His-
tory), London, England; Mrs.
Judith Marshall.
(BPBM)—Bernice P. Bishop Museum,
Honolulu, HI, U.S.A.; Mr.
Gordon M. Nishida.
(DEIG)—Deutsches Entomologisches
Institut, Eberswald, Germany;
Dr. G. Petersen.
(HDEO)—Hope Department of Ento-
mology, University Museum,
Oxford, England; Dr. George
C. McGavin.
(MCZH)— Museum of Comparative Zo-
ology, Harvard University,
Cambridge, MA, U.S.A.
(MNHG)— Museum d’ Histoire Naturelle,
207
Geneva, Switzerland; Dr.
Bernd Hauser.
(NRSS)—Naturhistoriska Riksmuseet,
Stockholm, Sweden; Per Inge
Persson.
(RNHL)—Riksmuseum van Natuurlijke
Historie, Leiden, The Neth-
erlands; Dr. Jan van Tol.
(TUVA)—La Trobe University, Bun-
doora, Victoria, Australia; Mr.
Patrick J. Vaughan and Prof.
I.W.B. Thornton.
(ZILS)—Zoological Institute, Lund,
Sweden; Dr. R. Danielsson.
Genus Margattea Shelford
Margattea Shelford 1911:155. Type species,
by monotypy: Blatta ceylanica Saussure,
Rehn 1931: 302; Bey-Bienko 1938: 121;
1950: 145; Princis 1969: 862.
Kuchinga Hebard 1929:39, 41. Type species,
by selection: Phyllodromia longealata
Brunner; Hanitsch 1931b: 392 (synony-
mized with Margattea), Bey-Bienko 1938:
121 (synonymized with Margattea).
Theganosilpha Kumar and Princis 1978: 33:
Asahina 1979: 114, 119. Type species by
monotypy: Theganopteryx perspicillaris
Karny. New Synonymy.
Diagnosis.—The following characteriza-
tion is based on the species mentioned ear-
lier as well as a study of more than 25 de-
scribed and undescribed species which I plan
to present elsewhere: Third and fourth max-
illary palpal segments usually longer than
the fifth (Fig. 40). Tegmina and wings fully
developed or variably reduced, hind wings
rarely absent; if reduced, hind wings are gen-
erally shorter than the tegmina and may be
vestigial: if fully developed, hind wings have
the discoidal and median veins simple, sub-
costa and most costal veins clubbed or
thickened distad, cubitus vein essentially
straight or weakly curved, with 1-4 com-
plete (rarely O branches in reduced wings)
and no incomplete branches, apical triangle
small (Figs. 6, 13). Abdominal terga unspe-
208
cialized, or with a group of setae near the
posteromedial margin of eighth tergum
(Figs. 24, 28); this specialization may be
hidden under the seventh tergum. Antero-
ventral margin of front femur Type B, or B,
(with | to 4 proximal spines) (Fig. 34) (both
types may occur in the same species, dif-
fering between the sexes), rarely Type C,;
all 4 proximal tarsomeres with pulvilli, tar-
sal claws symmetrical with inner margins
generally denticulate (Figs. 7, 20, 32; the
serrations often subobsolete and sometimes
only seen at high magnification of a com-
pound microscope), arolia present. Supra-
anal plate usually relatively short, trans-
verse, right and left male paraprocts similar,
relatively simple plates (Figs. 8, 16). Male
subgenital plate symmetrical or subsym-
metrical styles simple, cylindrical, generally
similar (Figs. 1, 29, 44); some species with
a stylelike process at the base of each style
giving the appearance of having 4 rather
than 2 styles (Figs. 51, 69, 72). Male geni-
talia with hooklike phallomere on the right
side; median sclerite may have an associ-
ated sclerite sometimes with setal brushes
(Figs. 1, 26, 29, 69).
Comments.—The principal diagnostic
characters used by Kumar and Princis to
distinguish Theganosilpha from Thegan-
opteryx were hind wings reduced in both
sexes, eighth tergum of male weakly spe-
cialized, anteroventral margin of front fe-
mur Type B, (some large proximal spines
followed by a row of piliform spinules and
terminating in 3 distal spines), inner mar-
gins of tarsal claws finely denticulate. Asa-
hina (1979: 119) accepted Theganosilpha as
a valid genus and correctly pointed out that
it was closer to Wargattea than to Thegan-
opteryx.
Rehn (1931) stated that the front femur
of M. cevlanica was Type B, and that the
tarsal claws were equal with internal mar-
gins unspecialized (i.e. not denticulate). Bey-
Bienko (1938: 121) believed that the tarsal
claws of MW. cevlanica and M. longealata are
equal and unspecialized, but later (Bey-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Bienko 1958: 587) correctly stated that the
former has barely visible notched tarsal
claws. The claws of !. /ongealata also have
subobsolete serrations. Asahina (1979: 119)
pointed out that since some species of Var-
gattea have toothed tarsal claws, this char-
acter cannot be used to distinguish it from
Theganosilpha. The size of the denticles may
be so small that they cannot be seen at the
magnifications usually used in examining
pinned specimens with a dissecting micro-
scope. If slides are prepared and the claws
are examined at high magnification with a
compound microscope, the minute teeth, or
their remnants (Figs. 7, 20) can be seen.
The male abdominal terga of 7. ceylan-
icaand M. longealata are unspecialized. The
male of Theganosilpha perspicillaris has a
setal tuft on the eighth tergum. There are
many examples of genera in which the males
of a genus have or lack abdominal tergal
glands (used in courtship behavior), and in
those species that have them the specializa-
tions may be found on one or more seg-
ments. For example, species of Blattella may
have glands on T7 only, or on T7 and T8,
and the morphology of the glands may differ
considerably between different species (Roth
1985a). In species of Parcoblatta Hebard,
the males have glands on T1 only (6 spp.),
Tl and T2 (4 spp.), or they are absent (2
spp.) (Hebard 1917).
Asahina (1979: 119) separated Thegano-
silpha from Margattea as follows: In The-
ganosilpha the front femur is Type B,, the
tegmina are narrowed apically, somewhat
nail shaped, rather coriaceous, and the hind
wings are rudimentary but sometimes one
half the length of the tegmina. In Vargattea
the front femur is Type B,, the tegmina are
not narrowed apically or nail shaped, and
it and the hind wing are fully developed.
Hebard (1929: 4) pointed out that wing re-
duction often is an unreliable generic char-
acter (unless correlated with other more sta-
ble features), and the armament on the front
femur also is subject to individual variation.
Kumar and Princis (1978: 33) described the
VOLUME 91, NUMBER 2
hind wings in both sexes of 7. perspicillaris
as reduced to small narrow lappets about
half the length of the tegmina. The male
lectotype and two male paralectotypes of
this species actually have vestigial wings
distinctly less than half the length of the
tegmina. The hind wings of a male 7. per-
spicillaris (fig. 25 in Asahina 1979) are more
than half the length of the tegmina and their
veins are well developed.
It is possible for a genus to have more
than one type of front femur. Species of
Balta may have Type B or C (Hebard 1943:
39). Bey-Bienko (1938: 121) claimed that
Margattea inermis Bey-Bienko is aberrant
in having a Type C, rather than Type B
femur. Margattea hemiptera Bey-Bienko
(Bey-Bienko 1958) and Margattea elongata
Kumar (Kumar 1975: 105) have reduced
tegmina and wings and Type B, front fe-
murs. I have examined specimens of The-
ganosilpha ogatai Asahina and Thegano-
silpha satsumana Asahina (identified by
Asahina) and in both species the males have
Type B, and the females Type B, femurs.
There is no correlation between Type B,
femurs and wing reduction, and Type B,
femurs with fully developed organs of flight.
The difference of 1 distal spine on the
anteroventral margin of the front femur
(Type B, vs. Type B,), the variable reduc-
tion of tegmina and wings, and the presence
or absence of a male tergal gland cannot be
used to distinguish between Margattea and
Theganosilpha, and Iam synonymizing the
latter genus. The diagnostic characters for
the genus Margattina Bey-Bienko are so
similar to those of Margattea (Bey-Bienko
1958: 587) that I believe the former genus
(known only from the type species Margat-
tina trispina Bey-Bienko, a single male from
China) should be reevaluated.
SUBFAMILY PLACEMENT OF MARGATTEA
The genitalia of male cockroaches are
strongly asymmetrical and one of the more
conspicuous structures is a hooklike phal-
lomere (Fig. 29) although in some species
209
(Blaberidae) it may be greatly reduced. Ac-
cording to McKittrick (1964) in the Blat-
tellidae the hook is on the left side in the
Blattellinae, Ectobiinae, and Nyctiborinae,
or on the right in the Plectopterinae (= Pseu-
dophyllodromuinae in Vickery and Kevan’s
1983, classification). McKittrick also
claimed that the females of Blattellinae, Ec-
tobiinae, and Nyctiborinae rotate their
oothecae during oviposition, whereas the
Plectopterinae do not. This concept holds
for a large number of species but Brown
(1975) found that Ectobius panzeri Ste-
phens has the male genital hook on the right
side although the female rotates its ootheca
(Brown 1973). Bohn (1987) found that the
male of Phyllodromica (Lobolampra) mon-
tana Chopard and several other species of
the ““montana-group” of Phyllodromica
(Ectobiinae) show a similar reversal of the
right-left asymmetry in male genitalia and
have the hook on the right whereas some
other species of the genus have the hook on
the left. Although Bohn did not mention
oviposition behavior of P. montana in his
paper, he (personal communication) was
“convinced” that it rotates its ootheca and
that it and Ectobius panzeri are true Ecto-
biinae, all of which rotate their oothecae.
In Margattea the male’s genital hook is
on the right side which in McKittrick’s sys-
tem places it in the Plectopterinae. How-
ever, 2 females of Margattea nimbata were
carrying oothecae in the rotated position
(Fig. 25) a behavior which is characteristic
of the Blattellinae. As Bohn (1987: 303)
pointed out, it is not sufficient to use the
position of the genital hook in placing blat-
tellids in their respective subfamilies. He
also claimed that symmetry reversal has oc-
curred several times independently during
the evolution of the Blattaria and that it is
possible there are species of Blattellinae with
reversed symmetry as those found in the
Ectobiinae. Bohn concluded that other
characteristics such as oothecal rotation, and
especially structure of the phallomeric scle-
rites be considered.
210
Since the position of the male’s genital
hook cannot always be used to distinguish
Plectopterinae from Blattellinae, oothecal
rotation or its absence appears to be the best
discriminating character, and I used it to
place a number of Blattellidae into these 2
subfamilies (Roth 1968a, 1971: 134). Thus
M. nimbata, which rotates its ootheca, be-
longs in the Blattellinae even though the
genital hook is on the right side. But the
problem here is more complex. The ovi-
position behavior of the species other than
nimbata is not known. Three females of
Margattea longealata (Brunner) were car-
rying oothecae whose keels were directed
dorsad (i.e. non-rotated) (Fig. 62). It is pos-
sible that these specimens were collected be-
fore they could rotate their oothecae prior
to depositing them. However, the ootheca
is distinctly different in morphology from
that of nimbata. It is particularly unusual
in being wider than high and strongly re-
sembles the oothecae of Lophoblatta brevis
Rehn and Lophoblatta arlei Albuquerque
from South America (see figs. 1-6 77 Roth
1968b). These are the only known members
of the Plectopterinae that carry their oothe-
cae with their keels dorsad (i.e. non-rotated)
until the eggs hatch; other species in this
subfamily drop the non-rotated oothecae
shortly after it is formed. One ootheca of
M. longealata (KOH treated) lacked cal-
cium oxalate crystals. The oothecae of L.
brevis and L. arlei have relatively few cal-
cium oxalate crystals, but egg cases 1n other
species in the Plectopterinae have large
amounts of calcium oxalate crystals.
The amount of calcium oxalate in cock-
roach oothecae decreased during the evo-
lution of ovoviviparity and viviparity in the
Blaberoidea. Oothecae of the Blattellinae
usually have large amounts of calclum ox-
alate crystals but the amount of this com-
pound varies from dense to sparse to absent
in species of Blattella, all of which carry
their oothecae externally in the rotated po-
sition until the eggs hatch (Roth 1968a).
Species of the African blattellid genus Stay-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ella Roth (Roth 1984) have an ootheca that
is typical of species of Blattella, but it is
retracted and carried internally until the eggs
hatch (Roth 1982). The egg cases of Stayvella
lack calcium oxalate as do species of Bla-
beridae, all of which incubate their eggs in-
ternally.
The oothecae ot /. /ongealata are some-
what transparent in the specimens studied
and their eggs are undeveloped (the eggs
from one of the oothecae were removed and
cleared and showed no embryonic devel-
opment). It is possible that (7. longealata
carries its ootheca in the vertical position
for the entire gestation period, as in Lo-
phoblatta, but the 3 females reported here
were collected shortly after their oothecae
were formed and therefore their eggs show
no development. It is probable that Mar-
gattea contingens (Walker) has an ootheca
similar to that of /ongealata because the male
and female genitalia of these 2 taxa are so
much alike (Figs. 51, 57-59, 61, 63, 68-70).
The male genitalia (particularly the left and
right phallomeres) of M. /ongealata and M.
contingens differ considerably from those of
other species in the genus. Perhaps these
two species should be placed in a separate
genus; if this is done Hebard’s Kuchinga
(type species, /ongealata) would be resur-
rected. For the present I am placing them
in a species group of Margattea.
SPECIES GROUPS OF .WARGATTEA
I have examined about 25 described
species of Margattea and about 10 taxa that
apparently are new. Thus far my study in-
dicates that the genus can be arranged in
seven species groups. The seven species dis-
cussed in this paper belong to 3 groups as
follows (other species in the groups will be
presented elsewhere):
Ceylanica species group.—Eighth ab-
dominal tergum unspecialized. Accessory
stylelike structures near the base of the styles
absent (Fig. 1). Tegmina and wings fully de-
veloped. Front femur Type B;. Species: cey-
lanica.
VOLUME 91, NUMBER 2
Perspicillaris species group.—Eighth ab-
dominal tergum specialized (Figs. 9, 24, 28,
43). Interstylar region may be simply a con-
tinuation of the lateral margins of the plate
(Figs. 23, 44), or there is a small plate some-
times bearing small dark spines (Fig. 29).
Accessory stylelike structures absent (Figs.
18, 29). Tegmina and wings fully developed
or variably reduced, the wings sometimes
vestigial. Front femur Type B, or B,. Species:
perspicillaris, nimbata, paraceylanica, an-
ceps. This is the largest group and contains
at least an additional twelve species.
Contingens species group.—Eighth ab-
dominal tergum unspecialized. Lateral cor-
ners of subgenital plate produced having
the appearance of styles (accessory stylelike
structures; Figs. 55, 56). Male genital phal-
lomeres: hook portion of right phallomere,
slender, elongate, strongly curved; median
phallomere needlelike, accessory median
phallomere absent; left phallomere roundly
bulbous basally, relatively simple (Figs. 51,
57-59) (cp. the differences in the taxa
belonging to the other 2 species groups).
Tegmina and wings fully developed. Front
femur Type B,. Species: contingens, lon-
gealata.
Key TO SOME MALES OF
INDo-PAcIFIC Ma4RGATTEA*
|. Eighth abdominal segment without a tergal
gland 2
— Eighth abdominal segment with a tergal gland
2. Subgenital plate with lateral corners produced
giving the appearance of accessory stylelike
structures (Figs. 51, 55, 56, 69) 3}
— Lateral corners of subgenital plate rounded, not
stylelike (Fig. 1) ceylanica
3. Head brown with a yellowish orange band be-
tween antennal sockets (Fig. 65). Pronotal disk
with a broad dark brown macula (Fig. 64)
longealata
* Includes only males discussed in this paper. I have
seen specimens only of nimbata and paraceylanica from
the Krakataus. The earlier records of ceylanica, con-
tingens, and anceps from these islands are question-
able.
— Head with an orangish macula on vertex be-
tween the antennal sockets blending into the
rest of the lighter face (Fig. 47). Pronotal disk
light brown without distinct markings (Fig. 53)
Fsthcky <parepeene ie tpertiort. 2 contingens
4. falcata region a aiororeccnen lan plate with
short robust spines on posterior margin (Fig.
29). Front femur Type B, (Fig. 34). Wings
shorter than the tegmina, sometimes vestigial
nee perspicillaris
- initerstylar 1 region not as above. Front femur
Type B,. Tegmina and wings fully and equally
developed : 5
5. Tegmina with a dark fon macula on basal
region (Fig. 41), sometimes extending the full
length of the wing cover : anceps
— Tegmina without dark etkies 6
6. Apex of median genital phallomere terminat-
ing in a pair of heavy spinelike structures (Figs.
18, 23, 26) nimbata
— Apex of median beni analloniene slender,
acute (Fig. 10) paraceylanica
DESCRIPTIONS OF SPECIES OF
MARGATTEA
Margattea ceylanica (Saussure)
Figs. 1-7
Blatta ceylanica Saussure 1868: 355 (9);
1869: 247; Walker 1871: 20.
Allacta ceylanica (Saussure): Kirby 1904:
100.
Phyllodromia ceylanica (Saussure): Shel-
ford 1908b: 12; Dammerman 1948: 483.
Margattea ceylanica (Saussure): Shelford,
1911: 155; Rehn 1931: 302, fig. 1; Han-
itsch 1933a: 232; 1933b: 310; 1934: 118;
1936: 392; Bruning 1947: 221; 1948: 62;
Bey-Bienko 1957: 899; Princis 1969: 862
(references to generic combinations and
records; incorrectly included /. nimbata
as a synonym); Asahina 1979: figs. 69-
Tile
Material examined.—Sri Lanka.
(MNHG): holotype female of Blatta ceylan-
ica, Peradenia, ““Ceylan.” (ANSP): The fol-
lowing specimens were determined by He-
bard as M. ceylanica (Sauss.) and many were
labelled topotypes: Belihuloya, | @ (termi-
nalia slide 398), 1.ix.1928; Bibile Estate, 1
212 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-7.
PY SS
gs Ne,
pom } =f
oe
OC ,
7 + = 7 Sh
Cae a PAB Css es)
Margattea ceylanica (Saussure) from Sri Lanka. 1, 3-6, males from Udahamulla, and 7, from
Battaramulla, 2, female from Labugama: 1, subgenital plate and genitalia (dorsal); 2, supraanal plate and genitalia
(ventral); 3, distal ends of median, accessory median, and hooklike right genital phallomeres; 4, pronotum; 5,
supraanal plate and paraprocts; 6, hind wing; 7, tarsal claws. Abbreviations: a, left phallomere; b, median
phallomere; c, accessory median phallomere; d, hooklike right phallomere; e, style; f, supraanal plate; g, paraproct;
h, intercalary sclerite; 1, first valvifer. Scales (mm): 1, 2, 0.5; 3, 0.25; 4, 1.0; 5, 0.5; 6, 2.0; 7, 0.15.
4, 26.vii.1929; Wellaway, 1 4, 28.iv.1931;
Rakwana, | 9, 3.v.1929; Horawupotana, |
6, 1 2, 13.x.1924; Labugama, | 6, 2 2(1 with
genitalia slide 404), 15-18. viii.1932;
Woodside, Urugalla, 2 4, 22.1v.1924; Bat-
taramulla, W.P., 1 46, 20.v.1931; Udaha-
mulla, W.P., 1 ¢ (terminalia slide 399),
ix. 1926.
Male.—Interocular space decidedly less
than distance between antennal sockets.
Tegmina and wings fully developed; hind
wing with costal veins clubbed, cubitus vein
with 2 complete and 0 incomplete branches,
apical triangle small (Fig. 6). Pronotum sub-
oval (Fig. 4). Abdominal terga unspecial-
ized. Anteroventral margin of front femur
Type B, (with 3 or 4 large spines on prox-
imal half); tarsal claws weakly serrated (Fig.
7; serrations sometimes difficult to see un-
der binocular microscope). Supraanal plate
VOLUME 91, NUMBER 2
transverse, hind margin trigonal, apex
sometimes weakly indented; paraprocts dis-
similar (Fig. 5). Subgenital plate almost
symmetrical, styles similar, cylindrical, in-
terstylar margin weakly convex (Fig. 1).
Genitalia as in Figs. | and 3; hooklike right
phallomere with a small apical spine; apex
of median phallomere enlarged, the swollen
portion terminating in a transparent fila-
ment; apex of accessory median sclerite nar-
row and slightly curved; left phallomere with
spinelike processes.
Coloration.—Light brown. Vertex with
dark brown band connecting eyes at top of
head, occiput pale, whitish band along eye
margins between tops of antennal sockets
followed by a light transverse band, below
this a pair of light brown spots. Pronotal
disk with brown pattern as in Fig. 4, the
remainder of pronotum hyaline. Tegmina
hyaline without markings. Abdominal terga
infuscated, with light brown medial areas.
Abdominal sterna light brown with small
dark maculae laterally, and some infusca-
tion on lateral borders. Legs pale without
dark markings. The markings are variable;
the pronotal pattern may be intense or sub-
obsolete, with gradations in between; also
the amount of tergal infuscation and the size
of the sternal maculae varies.
Female.—Supraanal plate transverse,
hind margin weakly convex, medially weak-
ly concave (Fig. 2). Genitalia as in Fig. 2:
intercalary sclerites greatly reduced.
Measurements (mm) (2 1n parentheses).
Length, 8.0-9.2 (7.8-8.9); pronotum length
x width, 2.1-2.2 x 2.8-3.2 (2.1-2.3 x 3.0-
3.2); tegmen length, 9.1-10.5 (8.7-10.1).
Comments.—This species was not col-
lected on the 1984/85 Krakatau expedi-
tions. Dammerman (1948: 483), and
Bruijning (1948: 62) recorded it from Kra-
katau, the latter from a single female col-
lected in May, 1908. Princis (1969: 862)
listed the species from Ceylon, Java, Bor-
neo, and Krakatau (probably based on the
above two authors). Most likely the Kra-
katau records were misidentifications and
213
referred to M. nimbata (see remarks under
that species), or MM. paraceylanica.
Asahina (1979) illustrated the terminal
abdominal segments and right hooklike
genital phallomere (which he incorrectly la-
belled the left phallosome) (his figs. 69, 70)
of a Sri Lankan specimen which he deter-
mined as M. ceylanica with a query. His
drawings of these structures agree with the
specimens of ceylanica which I have illus-
trated here.
Margattea paraceylanica Roth
New SPECIES
Figs. 8-14
Holotype.— Male, Rakata, Krakatau Is-
lands, 200 m, s. face, 24.vili.1985, Zool.
Exp. Krakataus.
Paratypes.—Krakatau Islands: Rakata,
Zwarte Hoek, 850 ft., ridge, water trap, | 4
(terminalia slide no. 3), 1984; 6.09S
105.25E, under rocks, 1 (abdomen missing),
| 2 (genitalia slide no. 4), 6.1x.1984; Rakata,
s. face, 200 m, 1 2, 400 m, | nymph,
24.vi1.1985; Panjang, 6.05S 105.28E, ex
litter, | 9, 14.1x.1984, 1 9, beating,
20.1x.1984; Sertung, 6.04S 105.24-25E, spit,
transition zone, sweep, | 4, 18.vili.1985. [All
specimens were collected on the 1984 and
1985 expeditions to the Krakatau Islands.
The holotype and some of the paratypes are
deposited in the Zoological Museum, Bo-
gor, W. Java. Some specimens will be kept
at (TUVA).]
Male.—Pronotum subelliptical (Fig. 12).
Tegmina and wings fully developed extend-
ing beyond end of abdomen. Hind wing with
costal veins thickened distad, cubitus with
4 complete and 0 incomplete branches, api-
cal triangle very small (Fig. 13). Anteroven-
tral margin of front femur Type B, (with 3
or 4 large proximal spines); tarsal claws
symmetrical, minutely serrated, pulvilli on
4 proximal tarsomeres, arolia present.
Eighth abdominal tergum with a postero-
medial tuft of setae, hind margin concave
(Fig. 9). Supraanal plate transverse, hind
214
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Wy ae |
ce
oe
Figs. 8-14.
Margattea paraceylanica Roth, paratypes from Rakata. 8-13, male: 8, supraanal plate and
paraprocts (ventral); 9, abdominal terga 8 and 9; 10, subgenital plate and genitalia (dorsal); 11, hooklike mght
phallomere; 12, pronotum; 13, hind wing; 14, female, supraanal plate and genitalia (ventral). Abbreviations: a,
supraanal plate; b, accessory median phallomere; c, median phallomere; d, right phallomere; e, left phallomere;
f, subgenital plate; g, style; h, paraproct; i, intercalary sclerite; }, ovipositor valve; k, paratergite; 1, first valvifer.
Scales (mm): 8-10, 0.5; 11, 0.15; 12, 1.0; 13, 2.0; 14, 0.5.
margin weakly concave medially; right and
left paraprocts weakly defined, similar (Fig.
8). Subgenital plate weakly asymmetrical,
styles similar, cylindrical, interstylar mar-
gin weakly convex (Fig. 10). Genitalia as in
Fig. 10; apex of hooklike right phallomere
acute, without an apical or preapical spine;
the tip looks like a spine but it is not artic-
ulated (Fig. 11); median phallomere slender
throughout; associated median phallomere
with a small preapical branch; left phallo-
mere with spinelike processes.
Coloration (specimens originally pre-
served in alcohol, then pinned and dried). —
VOLUME 91, NUMBER 2
Light brown. Head with dark band on ver-
tex extending as faint stripes on occiput.
Pronotal disk with light symmetrical spots
and lines (variable in intensity), surround-
ing regions mostly hyaline (Fig. 12). Teg-
mina hyaline, humeral region colorless, re-
mainder light brown; hind wings appear
colorless but when mounted on a white card
the apical region of the anterior field has a
faint yellowish tinge.
Female.—Supraanal plate (Fig. 14) sim-
ilar to that of the male (Fig. 8). Subgenital
plate extends well beyond hind margin of
supraanal plate. Genitalia as in Fig. 14; in-
tercalary sclerites very large dark plates that
extend laterally well beyond ovipositor
valves, first valvifer slender rods without
swellings.
Measurements (mm) (2 in parentheses).
Length, 6.8-8.0 (7.3-8.5); pronotum length
x width, 2.0-2.1 x 2.7-3.0 (2.2 x 2.9-3.0);
tegmen length, 7.2-8.6 (7.3-7.9).
Comments.—The male genital phallo-
meres and female genitalia clearly show a
close relationship to cey/anica. Although the
subgenital plate and styles are similar in both
paraceylanica and ceylanica, the former has
a tergal gland on T8 which is lacking in the
latter. The female genitalia of both taxa are
distinctly different. It is likely that paracey-
lanica occurs on other Indonesian islands.
Margattea nimbata (Shelford)
Figs. 15-26
Phyllodromia nimbata Shelford 1907b: 31
(6 2); 1908b: 13; Hanitsch 1915: 57;
1923b: 410.
Margattea nimbata (Shelford): Hanitsch
1928223; 1929a7 13; 1931b: 392; 1932a:
5; 1933b: 310 (incorrectly synonymized
with ceylanica), Bruijning 1947: 221;
1948: 62; Princis 1969: 863 [Bruijning
and Princis accepted Hanitsch’s synony-
my].
Kuchinga nimbata (Shelford): Hebard 1929:
42.
215
Material examined.—Lectotype (here
designated). Male (terminalia slide 200),
Kuching, N.W. Borneo [Sarawak], pres.
1905 by Sarawak Museum, 15.x11.1898.
Type Orth. 89 1/4 in (HDEO). Paralecto-
types. Sarawak. (HDEO): 2 ¢ (genitalia slides
201 and 203), same locality as lectotype,
Dyak coll. pres. 1900 by R. Shelford,
25.1.1900 and 27.vui.1900, Type Orth.
89 3/4 and 89 4/4.
Additional material.—Krakatau. (RNHL):
1 6 (terminalia slide 139) | 2 (with rotated
ootheca, keel to right, Fig. 25) (det. as Mar-
gattea ceylanica by Hanitsch); 1 2, v.1908,
E. Jacobson (labelled MM. cevlanica).
Australia (Northern Territory). (BPBM):
Holmes Jungle, Palm Cr., 15 km NE of Dar-
win, 5 m, 2 4 (1 with terminalia slide 468),
| (abdomen missing), light trap, 11.11.1961,
1 6, 14.111.1961, J. L. and M. Gressitt.
Kei Island. (RNHL): 3 2, 1922, H. C. Sie-
bers (labelled Phyllodromia nimbata Shelf.,
by Hanitsch).
Christmas Island (Indian Ocean). (ANIC):
National Park, | 4 (terminalia slide 210), 2
2, 3 nymphs, x.1983, L. Hill [ANPWS].
Thailand. (ZILS): 30 km south of Pak
Thong Chai, 2 2, 1 2 (reared from 2 taken
in a dry stream bed) (det. as M. ceylanica
by Princis), 13.iv.1967, 1 4, 14.iv.1967, 2 2
(on shrubbery at night), 26.1v.1967 (det. as
M. ceylanica by Princis), | 4 (reared from 2
taken on shrubbery at night) [det. as Mar-
gattea punctulata (Brunner) by Princis],
26.iv.1967, L. M. Roth.
Sarawak. (ANSP): Kapit, | 2 (terminalia
slide 405), 9.vu1.1910, Wm. Beebe [reported
as Kuchinga nimbata (Shelford), by Hebard,
1929:42]. (NRSS): Medan, 5 4, | (abdomen
missing), Mjoeberg (labelled Phyllodromia
or Margattea nimbata Shelf., by Hanitsch):
Tjinta Radja, | 2, Mjoeberg (labelled Phy/-
lodromia nimbata Shelf., by Hanitsch).
Borneo (Kalimantan). (NRSS): Tand-
jong, Redeb, O. Borneo, 3 4 (1 with ter-
minalia slide 67) (labelled Margattea cey-
lanica Sauss., by Hanitsch), 2 2 [1 with
rotated ootheca and labelled argattea cey-
216
Figs. 15-22
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
/
Margattea nimbata (Shelford). 15, 16, 18, 20, 21, male lectotype, 17, 19, female paralectotype,
22, female from Kapit, Sarawak: 15, pronotum; 16, supraanal plate and paraprocts (ventral); 17, pronotum; 18,
subgenital plate and genitalia (dorsal); 19, supraanal plate and genitalia (ventral); 20, tarsal claws and arolium;
21, right phallomere; 22
. hind wing. Abbreviations: a, supraanal plate; b, paraproct; c, left phallomere; d, median
phallomere; e, accessory median phallomere; f, right phallomere; g, style; h, intercalary sclerite; 1, paratergite; j,
ovipositor valve. Scales (mm): 15, 1.0; 16, 0.5; 17, 1.0; 18, 19, 0.5; 20, 21, 0.15; 22, 2.0.
lanica Sauss. (= M. nimbata Shelf.), by
Hanitsch]. (ZILS): Pelawan besar, | ? (det.
as M. ceylanica by Princis), vi.1937, Mrs.
M. E. Walsh.
Java. (RNHL): Ardja Sari, Preanger, | 2
(det. as V/. nimbata by Bruijning 1949).
Male.—Interocular space less than dis-
tance between antennal sockets. Pronotum
subelliptical (Fig. 15). Tegmina and wings
fully developed, extending beyond end of
abdomen. Hind wing with subcosta and cos-
tal veins clubbed distad, discoidal and me-
dian veins straight, unbranched: cubitus vein
straight with 2-5 complete and 0 incom-
plete branches, apical triangle small (Fig.
22). Front femur Type B, (with 3-5 large
proximal spines); pulvilli present on 4 prox-
imal tarsomeres, tarsal claws symmetrical
with subobsolete denticles on ventral mar-
gins (Fig. 20), arolia present. Eighth abdom-
inal tergum medially concavely arched and
bearing posteromedially a group of setae
(tergum 8 in the lectotype is damaged, but
the specialization is visible in other males)
(Fig. 24). Supraanal plate transverse, con-
vexly rounded, apex of hind margin weakly
VOLUME 91, NUMBER 2
217
Figs. 23-26. Margattea nimbata (Shelford). 23-25, from Krakatau: 23, male subgenital plate and genitalia
(dorsal); 24, male, setal modification on eighth abdominal tergum; 25, female with rotated ootheca attached to
terminal segments (ventral); 26, male from Northern Territory of Australia, genital phallomeres (dorsal). Scales
(mm): 23, 0.5; 24, 0.25; 25, 1.0; 26, 0.25.
excavated, reaching to about hind margin
of subgenital plate; right and left paraprocts
similar (Fig. 16). Subgenital plate subsym-
metrical, styles similar, cylindrical, inter-
stylar margin weakly convex, accessory
stylelike structures absent (Figs. 18, 23).
Genitalia as in Figs. 18, 21, 23, 26; left phal-
lomere large and complex with spinelike
processes; apex of median phallomere with
a pair of spinelike structures (visible in
pinned specimens), accessory median scler-
ite present; hooklike right phallomere with-
out a subapical or apical spine.
Coloration.— Yellowish or light brown.
Head with pale occiput, a reddish transverse
band on vertex, sometimes with a lighter
brown interocellar band. Pronotal disk yel-
lowish brown with symmetrical reddish or
light brown dots and lines, surrounding area
mostly hyaline (Fig. 15). Tegmina hyaline,
pale brown; hind wing with clubbed region
of costal veins dark, remainder essentially
not infuscated (Fig. 22). Abdominal terga
pale with narrow dark brown transverse
bands along anterior margins of segments
that do not reach lateral bands, narrow pos-
terior segmental bands connect with broad
lateral infuscation; supraanal plate with a
pair of dark brown spots on posterior half
separated by a narrow whitish spot behind
the concave apex, basal half of segment yel-
lowish brown (Fig. 16). Abdominal sterna
pale with broad lateral dark brown borders
that become narrower on posterior seg-
ments; subgenital plate pale except for nar-
row dark interstylar margin. Cerci with ba-
solateral area of segment 8 and most of
segment 9 dark brown, remainder (includ-
218
ing 3 terminal segments) pale (cercal color
varies and the dark areas may be practically
absent).
Female.—Supraanal plate transverse,
hind margin with a distinct narrow, concave
excavation (Fig. 19). Genitalia as in Fig. 19;
sclerites very darkly pigmented, paratergites
very wide. Pronotal disk markings similar
to, or more pronounced than in male (Fig.
17). Ootheca as in Fig. 25.
Measurements (mm) (2 in parentheses).
Length, 7.2-9.0 (8.0-9.6); pronotum length
3.4); tegmen length, 7.7-10.3 (7.3-10.1).
Comments.— Hebard (1929: 42) suggest-
ed that nimbata may be a pale form of ob-
tusifrons Walker, but Hanitsch (1931b: 392)
was correct in stating that these 2 species
are distinct. The female genitalia of nimbata
(Fig. 19) and obtusifrons (fig. 14L in Roth
1985b) differ. I have reexamined the type
female of Blatta obtusifrons which has been
placed in ?Kuchinga (Hebard 1929), Sym-
ploce (Princis 1969: 9) and sp. incertae sedis
(Roth 1985b: 156). The female’s front legs
are missing but the tarsal claws on 2 other
intact legs are minutely toothed. The male’s
front femur is Type B, [with 5 large proxi-
mal spines, then 2 short heavy spines the
same length as the following piliform spi-
nules, terminating in 3 large spines; this type
of armament is intermediate between Type
A and Type B, and I incorrectly stated it
was Type A, (Roth 1985b)]. The tarsal claws
of the male are minutely but distinctly
toothed so I am placing obtusifrons in Mar-
gattea.
Hanitsch (1931b: 392) suspected that
nimbata was a synonym of ceylanica, but
he retained the former name because he had
not seen the type of ceylanica. Later, he
(Hanitsch 1933b: 310) claimed to have con-
firmed this synonymy. Bruijning (1947,
1948) and Princis (1969) listed nimbata as
a synonym of ceylanica. Superficially the
two species resemble one another, but the
male and female genitalia of ceylanica and
nimbata are distinctly different (cp. Fig. |
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
with 18, 23, 26, and Fig. 2 with 19). Mar-
gattea nimbata 1s very widely distributed.
Margattea ceylanica is only known from Sri
Lanka, and all records of this species outside
of that island probably refer to nimbata.
One female paralectotype (Type Orth.
89 2/4) in (HDEO) of nimbata (supraanal
plate and part of genitalia on slide 202) with
the same data as the other paralectotypes
probably is another species. Unfortunately
most of the genitalic structures were lost
during slide preparation. However, the hind
margin of the supraanal plate lacks the dis-
tinctive medial indentation, and the inter-
calary sclerites and remnant of the parater-
gites differs from those of nimbata.
Margattea perspicillaris (Karny)
NEw ComBINATION
Figs. 27-35
Theganopteryx perspicillaris Karny 1915:
103 (42); Hanitsch 1927: 33; Shiraki 1931:
209, Princis; 1969: 1012 (sp. incertae se-
dis).
Theganosilpha perspicillaris (Karny): Ku-
mar and Princis 1978: 33, figs. 27, 28;
Asahina 1979: 114, figs. 11-13, 25, 43-
50, 60.
Material examined.— Lectotype (labelled
by Kumar 1977). Male, Hoozan, Formosa
[Taiwan], 1910, H. Sauter; in (DEIG).
Paralectotypes. Taiwan. (DEIG): same
data as lectotype, 2 4(1 with terminalia slide
6), 1 8g.
Male. — Pronotum subparabolic (Figs. 33,
35). Tegmina reaching slightly beyond end
of abdomen. Hind wings vestigial, narrow,
lateral, reaching between T1 and T2, some-
times longer with venation. Front femur
Type B,, tarsal claws minutely serrated, the
teeth visible under dissecting microscope
(Figs. 32, 34). Eighth abdominal tergum with
hind margin concavely indented medially
with a posteromedial tuft of setae (Fig. 28),
hidden under T7. Supraanal plate trans-
verse, hind margin convex entire, or with a
weak indication of a medial indentation;
VOLUME 91, NUMBER 2
219
Figs. 27-35. Margattea perspicillaris (Krauss), male types. 27-34, paralectotype: 27, supraanal plate and
paraprocts (ventral); 28, abdominal terga 8 and 9; 29, subgenital plate and genitalia (dorsal); 30, left phallomere;
31, hooklike right phallomere; 32, tarsal claws and arolium; 33, pronotum; 34, front femur (anterolateral); 35,
lectotype, pronotum. Abbreviations: a, paraproct; b, median phallomere; c, accessory median phallomere; d,
hooklike right phallomere; e, left phallomere; f, style. Scales (mm): 27-29, 0.5; 30-31, 0.25; 32, 0.15; 33, 1.0:
34, 0.5; 35, 1.0.
paraprocts similar with a straight spinelike
process (Fig. 27). Subgenital plate essen-
tially symmetrical, convex, exposed sides
and interstylar regions reflexed dorsad;: styles
similar, cylindrical, widely separated, be-
tween them a rectangular plate, corners
rounded, with short robust spines on its hind
margin (Fig. 29). Genitalia as in Fig. 29:
hooklike right phallomere relatively small,
without a preapical spine, apex acute, curved
220
(Fig. 31); left phallomere with a large spine-
like process (Fig. 30); accessory median
phallomere with a large setal brush.
Coloration.—Light brown. Head with a
weak indication ofan interocular band. Pro-
notal disk with distinct (Fig. 33) or faint
(Fig. 34) markings, remaining portion hya-
line.
Female.—Tegmina not reaching beyond
end of abdomen, extending to about T9;
hind wings vestigial. Supraanal plate short,
transverse, hind margin broadly convex.
Measurements (mm) (2 in parentheses).
Length, 9.3-9.5 (11.5); pronotum length x
width, 2.7-3.1 = 4.0-4.3 (3.0 x 4.5); teg-
men length, 7.2-7.9 (7.7).
Comments.—The spined interstylar plate
distinguishes the male of perspicillaris from
the other species described in this paper.
Margattea anceps (Krauss)
Figs. 36-46
Blatta (Phyllodromia) anceps Krauss 1903:
749 (9).
Blatta anceps Krauss: Kirby 1910: 563.
Phyllodromia anceps Krauss: Shelford
1908b: 14; Hanitsch 1915: 50; 1923b: 463.
Margattea anceps (Krauss): Caudell 1927:
12; Hanitsch 1928: 23 (incorrectly syn-
onymized nigrovittata Hanitsch with an-
ceps, see remarks below); 1929b: 276:
1932b: 61; 1933b: 310; 1933a: 232; Dam-
merman 1948: 483, 555; Bruyning 1948:
63: Princis 1969: 864.
Kuchinga anceps (Krauss): Hebard, 1929:
42 (8).
Holotype (not examined). Female, Tji-
bodas, Java. (According to Krauss, the spec-
imens he described in his paper were pre-
served “in Spiritus’ in the Zoology
Department of the “Universitats-Institute
zu Jena.” Dr. D. v. Knorre of the Friedrich-
Schiller-Universitat zu Jena wrote me 28
May 1987 that the type of anceps is not in
their museum. He also wrote me on 14 Oc-
tober 1987 that the type is not at the Zoo-
logical Institute in Tubingen where Krauss
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
worked at that time and that the specimen
probably no longer exists.)
Material examined.—Neotype (here des-
ignated). Male, Tjibodas, Java, 1500 m,
vili.1921, Karny; in (RNHL).
Additional material. Java. (RNHL): same
data as neotype, 2 4 (one with terminalia
slide 141), 1400 m, 1 ¢ (terminalia slide
143), 3 2, vili.1921, 2 2, vi.1929, Karny;
Panggerango, West Java, 1000 m, 1 9,
7.v1.1932, M. A. Lieftinck.
Male.—Interocular space less than dis-
tance between antennal sockets and ocelli.
Pronotum flat, anterior and posterior edges
almost straight, lateral edges convex (Fig.
37). Tegmina and wings fully developed.
Hind wings with costal veins thickened on
distal halves or clubbed apically, discoidal
vein straight; median and cubitus veins
straight, the former simple, the latter with
3 complete (1 may be forked) and 0 incom-
plete branches, apical triangle small (Fig.
39). Front femur Type B, (with 4 large prox-
imal spines); pulvilli present on 4 proximal
tarsomeres, tarsal claws symmetrical, ven-
tral margins minutely toothed, arolia pres-
ent. Eighth abdominal tergum with hind
margin concavely indented and arched,
bearing a tuft of setae (Fig. 43). Supraanal
plate transverse, hind margin narrowly
truncate medially, reaching to hind margin
of subgenital plate; right and left paraprocts
similar, without spinelike processes (Fig. 42).
Subgenital plate subsymmetrical, exposed
margin rounded, styles cylindrical, similar,
separated by about their lengths, interstylar
margin straight, accessory stylelike process-
es absent (Fig. 44). Genitalia as in Figs. 44—
46; apex of hooklike right phallomere con-
cavely excavated; median phallomere with
a preapical branch, distal region broadened
and terminating in a spine.
Coloration. — Head yellowish brown with
a broad transverse dark brown band on ver-
tex, occiput pale, sometimes with pale spots
between antennal sockets and on face (Fig.
36). Pronotal disk with reddish brown sym-
metrical pattern (Fig. 37). Tegmina with
VOLUME 91, NUMBER 2
@) \e *
|
=e 2 |
=
ts See
| Oy aot
| 1 ol t
OMe null i
|
a f |
R
Figs. 36-41.
i
in)
—
=——<—LUeze
Margattea anceps (Krauss), female from Panggerango, West Java: 36, head; 37, pronotum; 38,
terminal abdominal segments (dorsal); 39, hind wing; 40, maxillary palp; 41, tegmen. Scales (mm): 36-38, 1.0;
39, 4.0; 40, 0.5; 41, 4.0.
dark brown on basal portion; that part of
the right tegmen covered by the left may be
darkened whereas the similar area on the
left tegmen is pale; the extent of dark areas
on the tegmina varies from a small region,
e.g. Fig. 41, to large areas extending the
length of the wing cover. Abdominal terga
light to dark brown, if the former with dark
infuscation along posterior borders of the
segments; tergal gland area on T8 and pos-
terior part of T7 pale, supraanal plate with
a small pale dot on posteromedial region
near the hind margin. Abdominal sterna
light or dark brown; pale specimens may
have a small dot and some infuscation lat-
erally. Cerci pale dorsally. Legs pale.
Female.—Interocular space about the
same as interocellar distance, less than the
space between antennal sockets (Fig. 36).
Supraanal plate transverse, hind margin
weakly convex, not reaching hind margin of
subgenital plate which is large and laterally
overlaps several of the abdominal terga (Fig.
38). Pronotum as in Fig. 37. Tegmina and
wings (Figs. 39, 41), and maxillary palps
(Fig. 40) as in male. Abdominal terga and
sterna very dark brown, subgenital plate with
a large mediobasal yellowish spot.
Measurements (mm) (2 in parentheses).
Length, 8.5-10.0 (9.0-10.1); pronotum
3.1-3.2); tegmen length, 11.0-12.0 (11.0-
11.8).
Comments.— Although the holotype of A.
anceps apparently is lost, the present ma-
terial is from the same locality (Tjibodas)
and the females agree closely with Krauss’s
description of that sex. Hanitsch synony-
mized M. nigrovittata (Hanitsch) with an-
ceps, probably because of the tegminal
markings and interocular band in both taxa.
Nevertheless, tegmina with brownish mark-
ings and interocular bands occur in species
other than anceps. I have seen the types of
222 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 42-46. Margattea anceps (Krauss), male from Tjibodas, Java: 42, supraanal plate and paraprocts
(ventral); 43, abdominal tergum 8; 44, subgenital plate and genitalia (dorsal); 45, distal part of right genital
phallomere; 46, left genital phallomere. Scales (mm): 42-44, 0.5; 45, 0.1; 46, 0.15.
nigrovittata and the male’s styles and gen-
italia differ markedly from those of anceps.
Both are valid species.
Margattea contingens (Walker)
Figs. 47-61
Blatta contingens Walker 1868: 229 (2).
Phyllodromia contingens (Walker): Kirby
1904: 92: Shelford (1906) 1907a: 490;
1908a: 27; 1908b: 13; Hanitsch 1915: 45;
1923a: 198; 1923b: 402; Dammerman
1929: 112; 1948: 483.
Blattella contingens (Walker): Dammerman
1922: 107; Karny 1925: 188, fig. 20.
Kuchinga contingens (Walker): Hebard
1929: 45.
Margattea contingens (Walker): Hanitsch
1929a: 13; 1936: 392; Bruning, 1948:
63.
Blatta humeralis Walker 1869:140 (6); Han-
itsch 1915: 45.
Phyllodromia humeralis (Walker): Kirby
1904: 91; Shelford (1906) 1907a: 490, pl.
30, fig. 4 (synonymized with contingens),
1908b: 13; Hanitsch 1923a: 198.
Kuchinga humeralis (Walker): Hebard 1929:
45.
Margattea humeralis (Walker): Hanitsch
1928: 21 (contingens and humeralis are
distinct taxa); Bruijning 1948: 64; Princis
1950: 174.
Phyllodromia abrupta Hanitsch 1923b: 399,
figs. 2, 3 (9).
Margattea abrupta (Hanitsch) 1928: 21
(synonymized with humeralis), Bruyning
1948: 64.
Material examined.— Holotype. Female
(genitalia slide 204) of Blatta contingens,
VOLUME 91, NUMBER 2
+
1@ = @
@ CY
}
a
ee = LLLZZTTTZ :
| SS = = = ———|
Ny]
aa
——=—=——~
— a
@
Figs. 47-52.
and paraprocts (ventral); 49, pronotum; 50, hind wing;
223
(
PE ;
<< h
Se ily
C=’
ao
@ Wy
Margattea contingens (Walker), male from Long Petak, Borneo: 47, head; 48, supraanal plate
51, subgenital plate and genitalia (dorsal); 52, tegmen.
Scales (mm): 47, 1.0; 48, 0.5: 49, 1.0; 50, 3.0; 51, 0.5; 52, 3.0.
Sarawak, Wallace, coll. (1830-73), W. W.
Saunders, purchased and pres. ’73 by Mrs.
F. W. Hope; Type Orth. 77 (HDEO).
Singapore. (HDEO): Male (terminalia
slide 199) holotype of Blatta humeralis,
Wallace, E. coll. (1830-73), W. W. Saun-
ders, purchased and pres. ’73 by Mrs. F. W.
Hope, Type Orth. 78.
Borneo. (RNHL): Long Petak, 450 m, |
4 (terminalia slide 145), 2° (1 with genitalia
slide 146), ix.1925, H. C. Siebers, M. O.
Borneo Exp.
Male.—Interocular space about the same
as distance between the antennal sockets
(Fig. 47). Maxillary palps with third seg-
ment slightly longer than the fourth, each
distinctly longer than the fifth. Pronotum
suboval (Figs. 49, 53). Tegmina and wings
fully developed extending well beyond end
of abdomen. Hind wing with costal veins
weakly thickened distad, median vein sim-
ple, cubitus vein straight with 3 complete
and 0 incomplete branches, apical triangle
small (Fig. 50). Front femur Type B, (with
5 proximal spines). Eighth abdominal ter-
gum unspecialized. Supraanal plate trans-
verse, rectangular, hind margin truncate not
reaching hind margin of subgenital plate;
right and left paraprocts similar (Figs. 48,
54). Subgenital plate weakly asymmetrical,
lateral corners produced, stylelike (not ar-
ticulated), styles similar, cylindrical, each
arising basally on the inner margin of the
lateral projections (appear to be 4 styles),
224
64) =
me —/ ¥
(53) aaa
|
\ Wn i}
G) {N\ 4
|
Figs. 53-59.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Margattea contingens (Walker), male holotype of Blatta humeralis Walker: 53, pronotum; 54,
supraanal plate and paraprocts (ventral); 55, distal region of subgenital plate showing hind margin and styles
(dorsal); 56, right posterior corner of subgenital plate; 57-59, left, right, and median genital phallomeres. Ab-
breviation: a, style. Scales (mm): 53, 1.0; 54, 55, 0.5; 56-58, 0.15; 59, 0.25.
interstylar margin slightly asymmetrical, not
or only slightly produced beyond the tips of
the styles (Figs. 51, 55, 56). Genitalia as in
Figs. 51, 57-59; hook on right side, strongly
curved, apex rounded without an apical
spine or incision; the median phallomere is
a Slender, tapering, apically acute rod lying
on the right side near the hook (generally
the median phallomere lies more or less be-
tween the right and left phallomeres; I was
careful not to move this structure during
slide preparation so its position as shown
in Fig. 51 may be normal); left phallomere
bulbous on basal half.
Coloration.—Head with an orangish
macula on vertex to between the antennal
sockets blending into the lighter facial area
(Fig. 47). Pronotal disk light brown without
distinct markings, surrounding regions hya-
line (Figs. 49, 53). Tegmina very pale, yel-
lowish hyaline with a dark streak along the
radial vein (this mark may be reduced and
only indicated at the base of the vein; Fig.
52). Hind wing infuscated (Fig. 50). Ab-
dominal terga light brown, edges pale. Ab-
dominal sterna lighter, edges darker. Cerci
pale dorsally and ventrally.
Female.—Pronotum as in Fig. 60. Su-
VOLUME 91, NUMBER 2
Figs. 60-63.
225
Margattea spp. 60, 61, M. contingens (Walker), female holotype: 60, pronotum; 61, supraanal
plate and genitalia (ventral); 62, 63, M/. /ongealata (Brunner), females: 62, from 10 miles south of Kuching,
Sarawak, ootheca attached to the end of the female abdomen (dorsal); 63, from Sandakan, Sabah, supraanal
plate and genitalia (ventral). Abbreviations: a, paraproct; b, supraanal plate; c, paratergite; d, first valvifer. Scales
(mm): 60, 1.0; 61, 0.5; 62, 1.0; 63, 0.5.
praanal plate strongly transverse, short,
rectangular, hind margin entire (Fig. 61).
Genitalia as in Fig. 61; paratergites slender,
first valvifer broad, enlarged, connected to
the paratergites by slender sclerotizations.
Dark streak on radial vein of tegmina great-
ly reduced. Hind wings lightly infuscated.
Measurements (mm) (2 1n parentheses).
Length, 7.4-9.6 (7.7-9.5); pronotum length
3.5); tegmen length, 10.4-10.5 (11.0-12.0).
Comments.—Shelford (1907a: 490) syn-
onymized humeralis (8) with contingens (2).
Hanitsch initially agreed with him but later
(Hanitsch 1928: 27) considered both species
distinct, basing his conclusion on the pres-
ence of a brown streak along the radial vein
of the tegmen of humeralis, and absent in
contingens. Hebard (1929: 45) agreed with
Hanitsch. I do not believe that this color
difference is sufficient to separate these two
taxa. In the male contingens from Long Pe-
tak Borneo, the streak on the tegmen is sub-
obsolete, but the important characters (ab-
sence of a tergal gland, genital phallomeres,
subgenital plate, and styles) are so similar
to the holotype of Aumeralis that I am cer-
tain they are the same taxon. Also the fe-
males from Long Petak are very similar to
the 2 holotype of contingens from Singa-
pore.
Although M. /ongealata is strikingly dif-
ferent in coloration from contingens the male
characters, so similar in both taxa, leave no
doubt they are very closely related. The fe-
male genitalia of contingens (Fig. 61) also
226 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 64-72. Margattea longealata (Brunner). 64, 65, female from 10 miles south of Kuching, Sarawak,
pronotum and head. 66-72, male from Sandakan, Sabah: 66, hind wing; 67, supraanal plate and paraprocts;
68, left genital phallomere; 69, subgenital plate and genitalia (dorsal); 70, hooklike right phallomere; 71, tarsal
claws and arolium; 72, right corner of hind margin of subgenital plate (dorsal). Scales (mm): 64, 65, 1.0; 66,
2.0; 67, 0.5; 68, 0.15; 69, 0.5; 70-72, 0.15.
are very similar to that of /ongealata (Fig. Phyllodromia longealata Brunner: Kirby
63). 1904: 92: Shelford 1908b: 13; Hanitsch
1915: 42; 1923b: 463; 1925: 83 (8).
Margattea longealata (Brunner) Kuchinga longealata (Brunner): Hebard
Figs. 62-72 1929: 46.
Margattea longealata (Brunner): Hanitsch
Phyllodromia longe-alata Brunner 1898: 1933a: 232; Princis 1969: 865 (the fol-
205, pl. 16, fig. 9 (2). lowing references erroneously listed by
VOLUME 91, NUMBER 2
Princis under Margattea longealata ac-
tually refer to Balta longealata (Hanitsch)
Hanitsch, 1930: 254; 193la: 43; Bruy-
ning 1947: 214—Princis correctly listed
these references under Ba/ta on page 977
of his catalogue).
Holotype (not examined).— Female, Sa-
rawak; in the Vienna Museum.
Material examined.—Sarawak. (NRSS):
Kuching, N. W. Borneo, 2 2, 25.v.1900,
Dyak coll., pres. 1900 by R. Shelford (det.
as longealata by Shelford). (ANSP): same
data as above, | 2, | 2 (with ootheca),
27.11.1900, 1 4, 30.111.1900; 10 miles south
of Kuching, | 2 (with ootheca), 27.vi.1910,
Beebe.
Sabah. (ANSP): Sandakan, Borneo, | 4,
| (terminalia slide 401), 1 2 (genitalia slide
403), 1 2 (carrying ootheca, wall of the
ootheca on slide 400), Baker. [All (ANSP)
specimens reported as Kuchinga longealata
by Hebard (1929: 46).]
Borneo. (RNHL): Long Petak, M.O., 450
m, 6 6, 10 9, ix.1925 or 1x.-x.1925, H. C.
Siebers, Borneo Exp. (NRSS): Pajau River,
O. Borneo, 2 4, 2 2°, Mjoeberg (det. as Mar-
gattea longe-alata Br., by Hanitsch).
Male.—Interocular space slightly less than
space between antennal sockets. Maxillary
palps with segments 3 and 4 about equal,
each longer than the swollen fifth segment.
Pronotum suboval (Fig. 64). Tegmina and
wings fully developed extending well be-
yond the end of the abdomen. Hind wing
with costal veins thickened on distal halves,
median vein simple, cubitus with 2-3 com-
plete and 0 incomplete branches, apical tri-
angle small (Fig. 66). Front femur Type B,
(with 3 or 4 large proximal spines), tarsal
claws with subobsolete teeth on ventral
margins (Fig. 71; the denticles cannot be
seen under the dissecting microscope but
after the claws are treated with KOH and
examined under a compound microscope,
the weakly defined truncate teeth are seen).
Abdominal terga unspecialized. Supraanal
plate transverse, narrow, subrectangular, not
221
reaching hind margin of subgenital plate;
right and left paraprocts similar, simple
plates without spinelike processes (Fig. 67).
Subgenital plate symmetrical, its hind mar-
gin and styles (Figs. 69, 72) similar to that
of contingens (cp. Figs. 51, 55, 56). Geni-
talia as in Figs. 68-70; phallomeres similar
to those of contingens (cp. Figs. 51, 57-59).
Coloration.— Head brown with a yellow-
ish orange band between antennal sockets
(Fig. 65). Pronotal disk with a broad dark
brown band whose oblique sides may or
may not be margined by narrow yellowish
stripes, lateral border regions semi-hyaline
(Fig. 64). Tegmina with anterior borders
hyaline, remainder dark brown, a contin-
uation of the dark area of the pronotal disk.
Hind wings darkly infuscated, thickened re-
gion of costal veins whitish or yellowish.
Abdominal terga and sterna dark brown.
Legs dark brown. Cerci dark ventrally, light
on dorsal surface.
Female.—Interocular space less than dis-
tance between antennal sockets (Fig. 65).
Cubitus vein of hind wing with 2-3 (rarely
4) complete branches. Supraanal plate nar-
rowly subrectangular; paraprocts weakly
dissimilar (Fig. 63). Genitalia as in Fig. 63,
similar to that of contingens (cp. Fig. 61).
Ootheca as in Fig. 62; length, 5.5 mm, width,
2.5; height, 1.5.
Measurements (mm) (2 in parentheses). —
Length, 7.2-11.8 (9.3-11.2) pronotum
length x width, 2.3-2.7 x 3.0-3.3 (2.7-3.0
x 3.4-3.9): tegmen length, 7.9-11.9 (11.0-
13.8).
Comments.—The color markings of
Maregattea longealata are strikingly differ-
ent from those of WZ. contingens. However,
male structures (subgenital plate, styles,
genitalia) and female genitalia are so similar
in both taxa that one might be tempted to
consider /ongealata a color morph, variant,
or subspecies of contingens. Additional
specimens from other localities should be
studied to see how variable the color mark-
ings are. The unusual ootheca of /ongealata
is discussed in the section on subfamily
228 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
placement. It is likely that contingens has a
similar ootheca.
ACKNOWLEDGMENTS
I thank the curators and collection man-
agers listed earlier who sent me specimens,
Dr. Syoziro Asahina for specimens of 7he-
ganosilpha ogatai and T. satsumana, and
Dr. I. W. B. Thornton for the Margattea
paraceylanica, collected on the 1984 and
1985 Zoological Expeditions to the Kra-
katau Islands. I am grateful to the Bureau
of Flora and Fauna, Australian Biological
Resources Study, for partial support.
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PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 230-236
NOTES ON THE GENUS ZAGRAMMOSOMA
(HYMENOPTERA: EULOPHIDAE) WITH
DESCRIPTION OF A NEW SPECIES
JOHN LASALLE
Department of Entomology, University of California, Riverside, California 92521.
Abstract.—The genus Zagrammosoma Ashmead is characterized. The genus Mirza-
grammosoma Girault is synonymized with Zagrammosoma, and the type species, and
only included species, W/. /ineaticeps Girault, is transferred to Zagrammosoma. New
Nearctic distributional records are given for Z. lineaticeps from Texas and California, and
Z. mirum from Washington. New host records are given for Z. lineaticeps from Kieferia
lycopersicella (Gelechiidae) and Z. mirum from Kieferia lycopersicella and Phthorimaea
operculella (Gelechiidae) and Phyllonorycter elmaella (Gracillaridae). A new species of
Zagrammosoma, Z. hobbesi LaSalle, is described from California. A key is given to
separate the three species of Zagrammosoma which have an entirely black metasoma: Z.
mirum Girault, /Ineaticeps (Girault), and hobbesi LaSalle, n. sp.
Key Words:
During a study involving Mirzagram-
mosoma lineaticeps Girault, the type species
and only included species in the genus M/ir-
zagrammosoma Girault, I realized that this
genus was a synonym of Zagrammosoma
Ashmead. At the same time, I came across
specimens of a new species of Zagrammo-
soma from Southern California. The pur-
pose of this paper 1s to formally synonymize
Mirzagrammosoma with Zagrammosoma,
characterize Zagrammosoma, describe this
new species, and provide new host and dis-
tributional records for Z. /ineaticeps and Z.
mirum Girault.
Morphological terminology follows that
of Graham (1969), except that the term
mesosoma is used for the thorax (including
propodeum), and metasoma used for the
combined petiole and gaster (abdomen mi-
nus propodeum).
Abbreviations for collections are as fol-
lows: AEI, American Entomological Insti-
tute, Gainesville, Florida; BMNH, British
Hymenoptera, Eulophidae, Wirzagrammosoma, Zagrammosoma
Museum (Natural History), London; CNC,
Canadian National Collection, Ottawa; LAS,
personal collection of the author; UCR,
University of California, Riverside; USNM,
United States National Museum, Washing-
tons DC:
Genus Zagrammosoma Ashmead
Hippocephalus Ashmead, 1888: viii. Type
species Hippocephalus multilineatus Ash-
mead (monotypy). Preoccupied by Hip-
pocephalus Swainson 1839, in fishes.
Zagrammosoma Ashmead 1904: 354, 393.
Replacement name for Hippocephalus
Ashmead.
Zagrammatosoma Schulz 1906: 142. Un-
justified emendation.
Atoposoma Masi 1907: 276. Type species
Atoposoma variegatum (monotypy).
Virzagrammosoma Girault 1915: 279.
Type species Mirzagrammosoma linea-
ticeps Girault (monotypy). N. syn.
VOLUME 91, NUMBER 2
Figs. 1-6.
4. C. flavoviridis, 2, mesoscutum. 5. Z. hobbesi, 2,
bars = 0.1 mm.
Diagnosis.—Scutellum with 4 setae (2
pairs); submarginal vein with at least 3 dor-
sal setae; funicle 2-segmented: notauli com-
plete, curving to meet axilla at a distance
well separated from posterior margin of
mesoscutum; head with vertex vaulted be-
tween compound eyes.
1. Zagrammosoma mirum, 2, head. 2. Cirrospilus flavoviridis, 2, head. 3. Z. mirum, 2, mesoscutum.
mesoscutum. 6. Z. hobbesi, °,
dorsum of mesosoma. Scale
Discussion.— Members of the genus Za-
grammosoma are parasitic on leafmining
Lepidoptera and Diptera. This genus is in
the subfamily Eulophinae, tribe Elachertini.
The subfamily Eulophinae is characterized
by: 4 setae on the scutellum:; submarginal
vein not broken before it reaches marginal
232
vein, and having 3 or more setae on its dor-
sal surface; postmarginal vein present and
well developed. Elachertini have complete
notauli, Eulophini have the notauli incom-
plete or absent. Zagrammosoma 1s very
close to Cirrospilus, as both have a 2-seg-
mented funicle, and the postmarginal vein
shorter than or equal in length to the stigmal
vein. Zagrammosoma has been treated as
a subgenus of Cirrospilus by European au-
thors (Peck et al. 1964, Boucek and Askew
1968), but as a distinct genus by American
authors (Peck 1963, Burks 1979). Gordh
(1978) also treated Zagrammosoma as a
distinct genus, and provided a key to Nearc-
tic species. He gave two morphological
characters to support Zagrammosoma as
distinct from Cirrospilus: Zagrammosoma
species have an elongate head which 1s dis-
tinctly vaulted dorsally so that the vertex
extends much higher than the compound
eyes (Fig. 1); and Cirrospilus species have a
well developed median carina on the pro-
podeum, while in Zagrammosoma this ca-
rina is weakly developed or absent. The
vaulted vertex is a solid diagnostic char-
acteristic, as it represents a derived char-
acter which is unique to Zagrammosoma
(within the Eulophinae) and this character
does not appear in Cirrospilus (Fig. 2). How-
ever the propodeal carina does not appear
to be a good character for the separation of
these two genera as it is not constant
throughout all the species.
An additional character which may be
used to distinguish these two genera is that
in Zagrammosoma the notaulus curves to
meet the axilla, and never reaches, or ap-
proaches, the posterior margin of the meso-
scutum (Figs. 3, 5); in Cirrospilus the no-
taulus is straight, and extends to the posterior
margin of the mesoscutum (Fig. 4).
The genus Mirzagrammosoma Girault is
herewith synonymized with Zagrammoso-
ma. The type species, and only included
species, M. lineaticeps Girault displays
characters consistent with the definition of
Zagrammosoma: the elongate head with a
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
vaulted vertex; and the notaulus curving to
meet the axilla.
There are presently eight species of Za-
grammosoma known in North America
(Gordh 1978); the inclusion of Z. /ineaticeps
and Z. hobbesi, n. sp. brings the number up
to ten. Gordh treated only one species of
Zagrammosoma with an entirely black or
dark metasoma: Z. mirum Girault (Fig. 7).
Z. lineaticeps also has an entirely black or
very dark metasoma, as well as the new
species described in this paper, Z. hobbesi
LaSalle. These three species are the only
Nearctic Zagrammosoma without any yel-
low markings on the metasoma, and the fol-
lowing key will serve to separate females of
these species.
1. Forewing (Fig. 8) with a longitudinal dark stripe
which extends from near the base of the wing
to the anterior margin just before the apex, and
connects to transverse stripes which extend to
the anterior margin of the wing at the junction
of marginal and submarginal vein, and at the
base of the stigmal vein; apical margin of wing
with a transverse stripe in posterior half .
b aos Mon tienen ae Z. lineaticeps (Giant)
I Rorewing (Figs. 9, 10) without such a long lin-
ear stripe, at most with a U-shaped pattern
connecting the junction of the submarginal and
marginal veins with the base of the stigmal
vein; apical margin of wing without any mark-
1a: CE eM tr kon BORO OOTS c 2
2. Mesosoma dopenlly aah ibroad: median yel-
low stripe (Fig. 7); venter of mesosoma yellow.
Funicular segments both the same color, dark
dorsally, brown to yellow ventrally
Bi eee ae eee Z. mirum Girault
2" Nesesomea entirely black, or with only slight
yellow markings, not with markings as above.
First funicular segment dark, second funicular
segment yellow (Fig. 11) :
es ety Zi nobbest Pasatles n. sp.
Zagrammosoma lineaticeps (Girault),
New ComMBINATION
Fig. 8
Mirzagrammosoma lineaticeps Girault
1915: 279. Holotype °¢, MEXICO, San
Rafael, Jicoltepec (USNM) [Examined].
Diagnosis.— Z. /ineaticeps can be distin-
guished from the other species of Zagram-
VOLUME 91, NUMBER 2
mosoma with a uniformly black or dark or
metasoma by the following characters.
Forewing (Fig. 8) with a wide longitudinal
dark stripe from the base of the wing which
curves to meet the anterior margin of the
wing before the apex and which connects to
two small transverse stripes which join the
anterior margin of the wing at the junction
of marginal and submarginal vein, and the
junction of marginal and stigmal vein; ad-
ditionally there is a transverse dark stripe
bordering the apex of the wing in the pos-
terior half. Mesosoma entirely black or dark
except longitudinal yellow stripe dorsolat-
erally on pronotum, small longitudinal yel-
low stripe may be present laterally on meso-
scutum; pronotum ventrally yellow. Fore
coxa yellow, middle and hind coxae black;
fore and middle legs yellow, hind femur and
tibia predominantly black, tarsi yellow.
Z. lineaticeps is known in the Neotropical
region from Mexico, Central America and
the Caribbean (De Santis 1979), however it
has only been recorded in the Nearctic re-
gion from Florida (Burks 1979). Known
hosts were in the Agromyzidae and Lyone-
tiidae. The following represent new distri-
butional records from California and Texas,
and a new host record from the tomato pin
worm, Kieferia lycopersicella (Walsingham)
(Gelechiidae).
CALIFORNIA: San Diego Co., Batequi-
tas Lagoon, | mi. E. Leucadia, 8.vili.1979,
C. W. Melton (8 2, UCR).
TEXAS: Cameron Co., Brownsville,
27.vi.1979, E. R. Oatman, ex. Kieferia ly-
copersicella (6 2, UCR).
Zagrammosoma mirum Girault
Figs. 1, 3, 7, 9
Zagrammosoma mira Girault 1916: 119.
Holotype 2, USA, California, mountains
near Claremont (USNM) [Examined].
Diagnosis.—Z. mirum can be distin-
guished from the other species of Zagram-
233
mosoma with a uniformly black or dark or
metasoma by the following characters.
Forewing (Fig. 9) without a longitudinal dark
stripe or transverse stripe along apical mar-
gin; with a transverse stripe at level of base
of stigmal vein, these stripes connected pos-
teriorly to form a U-shaped pattern; an ad-
ditional transverse stripe is present 1n an-
terior half of wing between postmarginal
vein and wing apex. Dorsum of mesosoma
(Fig. 7) black or dark, with a broad longi-
tudinal yellow stripe medially on mesoscu-
tum and scutellum, this stripe becoming two
parallel stripes on pronotum; additional thin
longitudinal stripe laterally on pronotum.
Entire venter of mesosoma yellow. All cox-
ae yellow except hind coxa black basally on
dorsum; fore and middle legs yellow, hind
femur black except extreme base and apex,
remainder of hind leg yellow.
As noted by Gordh (1978), the species of
Z. flavolineatum Crawford (1913) is very
similar to Z. mirum, differing only in col-
oration; flavolineatum 1s slightly lighter col-
ored than mirum. In flavolineatum the
metasoma 1s uniformly dark brown, but
there are small but distinct yellow spots lat-
erally on the metasomal tergites; the pro-
podeum has the callus yellow, as opposed
to the propodeum completely black or dark
in mirum, the hind femur has the entire
basal half yellow, as opposed to only the
extreme base (only about 0.1 x the length
of the femur) in mirum. Z. flavolineatum is
known from a single female specimen from
Colorado, and this specimen may represent
nothing more than a color variant of mirum.
As Gordh points out, additional material
will be necessary to resolve this problem. If
they do prove to be synonymous, the name
flavolineatum would have precedence over
mirum. Using the key in this paper, speci-
mens of flavolineatum would be determined
as Z. mirum.
Zagrammosoma mirum has previously
been recorded only from California. Known
hosts are in the genera Lithocolletis (Gra-
cillaridae) and Liriomyza (Agromyzidae).
234 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-11. 7. Zagrammosoma mirum, °, body (from Gordh, 1978; length 2 mm). 8. Z. /ineaticeps, °,
forewing. 9. Z. mirum, °, forewing. 10. Z. hobbesi, °, forewing. 11. Z. hobbesi, 2, antenna. Scale bars = 0.2 mm.
Dr. Michael Schauff has kindly informed maella (Gracillaridae). Additional new host
me that there are specimens of Z. mirumin records for this species are:
the USNM collection from Wenatchee, CALIFORNIA: Riverside Co., Corona,
Washington, reared from Phyllonorycter el-31.x.1931, A. J. Baringer, ex. Kieferia ly-
VOLUME 91, NUMBER 2
copersicalla (Gelechiidae) (1 2, UCR); Los
Angeles Co., Pasadena, vii.1915, J. E. Gray,
ex. Phthorimaea operculella (Gelechiidae)
(1 2, UCR).
Zagrammosoma hobbesi LaSalle,
NEw SPECIES
Bigs .65 OM el
Diagnosis.—Z. hobbesi can be distin-
guished from the other species of
Zagrammosoma with a uniformly black or
dark metasoma by the following characters.
Forewing (Fig. 10) without a longitudinal
dark stripe or transverse stripe along apical
margin; with a transverse stripe at level of
junction of marginal and submarginal veins,
and a transverse stripe at level of base of
stigmal vein, these stripes usually not con-
nected posteriorly, but may be faintly con-
nected to form a U-shaped pattern; an ad-
ditional transverse stripe is present in
anterior half of wing between postmarginal
vein and wing apex. Entire mesosoma black
or dark, except edges of some of the sclerites
may be brown or yellow. All coxae black;
fore and middle legs yellow; hind femur
black except for extreme apex, hind tibia
dusky to black basally, yellow apically.
Female.—Length 1.5-2.9 mm. Mesoso-
ma and metasoma entirely black except a
small yellow mark laterally on axilla, and
anteriorly on tegula. Fore and middle fem-
ora and tibiae yellow, hind femur black ex-
cept yellow apically, hind tibia yellow ex-
cept usually black basally. Tarsi yellow,
distal segments may be brown. Head black
and yellow; occiput black with median yel-
low stripe which extends to vertex and small
yellow spot bordering eye; face and frons
yellow, with numerous black stripes. An-
tenna (Fig. 11) with scape yellow, dark dor-
soapically; pedicel yellow ventrally, dark
dorsally; first funicular segment dark, sec-
ond funicular segment yellow; basal two club
segments dark, third club segment dark ba-
sally, yellow apically.
235
Head |.0—1.2 times higher than wide, the
vertex extending distinctly higher than the
height of the eyes. Face and frons reticulate.
Toruli situated at level of lower eye margin.
Eyes 1.2—1.4 times longer than malar sulcus.
Antenna (Fig. 11) with scape 4.3—5.2 times
longer than wide. Pedicel 1.3-1.5 times
longer than wide. First funicular segment
1.25-1.5 times longer than second. Club
about equal in length to both funicular seg-
ments taken together. Club and funicle about
equal in width.
Mesosoma (Figs. 5, 6) with pronotum,
mesoscutum and scutellum distinctly retic-
ulate, metanotum and propodeum lightly
sculptured to smooth. Propodeum with me-
dian carina; propodeal callus with 4-7 setae.
Forewing (Fig. 10) with a transverse stripe
at level of junction of marginal and sub-
marginal veins, and a transverse stripe at
level of base of stigmal vein, these stripes
usually not connected posteriorly, but may
be faintly connected to form a U-shaped
pattern; an additional transverse stripe is
present in anterior half of wing between
postmarginal vein and wing apex. Veins dark
except marginal vein yellow. Submarginal
vein with 5-8 setae. Submarginal vein 1.45-
1.65 times longer than marginal vein; mar-
ginal vein 3.0-3.6 times longer than post-
marginal vein, 2.15—2.6 times longer than
stigmal vein; stigmal vein 1.2-1.7 times
longer than postmarginal vein.
Metasoma 3.0-4.0 times longer than wide,
pointed apically.
Male.—Length |.1-1.5 mm. Differs from
female only in genitalia.
Distribution.— Known only from South-
ern California.
Material examined.— Holotype °. CAL-
IFORNIA, San Bernardino Co., Summit
Valley, 14.v.1985, G. Gordh, on Eriodic-
tyon (mounted on point, USNM).
18 2, 10 6 Paratypes. CALIFORNIA: as
holotype (7 2, 2 6, UCR; 3 2, USNM); as
holotype but on Haplopappus (2 2, 1 4,
UCR); San Bernardino Co., Mojave River
236
Forks, ~9 mi. S. Hesperia, 2.v.1985, J. D.
Pinto (3 2, 4 6, LAS; 1 9, 1 6: BMNH, CNC,
AEI).
Etymology.— The coloration of the face,
yellow with numerous black stripes, 1s rem-
iniscent of the face of a tiger. This species
is named for Calvin’s tiger friend, Hobbes.
ACKNOWLEDGMENTS
I thank G. Gordh and J. D. Pinto for
supplying material of Z. hobbesi, M. E.
Schauff for loans of material from the USNM
and for host information on USNM speci-
mens, and G. Gordh for supplying the il-
lustration of Z. mirum.
LITERATURE CITED
Ashmead, W. H. 1888. Descriptions of some un-
known parasitic Hymenoptera in the collection of
the Kansas State Agricultural College, received
from Prof. E. A. Popenoe. Bull. Kansas St. Agric.
Coll. 3: I-VI (Appendix).
1904. Classification of the chalcid flies, or
the superfamily Chalcidoidea, with descriptions of
new species in the Carnegie Museum, collected in
South America by Herbert H. Smith. Mem. Car-
negie Mus. 1(4): i-xi, 225-551, pls. 31-39.
Boucéek, Z. and R. R. Askew. 1968. Palearctic Eu-
lophidae (excl. Tetrastichinae). (Hym. Chalcidoi-
dea.) Index of Entomophagous Insects. Le Fran-
cois, Paris. 260 pp.
Burks, B.D. 1979. Family Eulophidae, pp. 967-1022.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
In Krombein, K. V. et al., eds., Catalog of Hy-
menoptera in America North of Mexico. Vol. I.
Symphyta and Apocrita (Parasitica). Smithsonian
Institution Press, Washington, D.C. 1198 pp.
Crawford, J. C. 1913. Descriptions of new Hyme-
noptera, No. 6. Proc. U.S. Natl. Mus. 45: 241-
260.
De Santis, L. 1979. Catalogo de los himenopteros
calcidoideos de américa al sur de los estados un-
idos. Publicacion especial. Comision de Investi-
gaciones Cientificas de la Provincia de Buenos
Aires. La Plata. 488 pp.
Girault, A. A. 1915. New chalcidoid Hymenoptera.
Ann. Entomol. Soc. Amer. 8: 279-284.
1916. Three new chalcid flies from Califor-
nia. Jour. Entomol. and Zool. 8: 119-122.
Gordh, G. 1978. Taxonomic notes on Zagrammo-
soma, a key to the nearctic species and descriptions
of new species from California (Hymenoptera: Eu-
lophidae). Proc. Entomol. Soc. Wash. 80(3): 344—
359.
Graham, M. W.R. de V. 1969. The Pteromalidae of
Northwestern Europe (Hymenoptera: Chalcidoi-
dea). Bull. Brit. Mus. (Nat. Hist.), Entomol. Supp.
16. 908 pp.
Masi, L. 1907. Contribuzioni all conoscenza dei Cal-
cididi italiani. Boll. Lab. Zool. gen. agr. Portici 1:
231-295.
Peck, O. 1963. A Catalogue of the Nearctic Chalci-
doidea (Insecta: Hymenoptera). Can. Entomol.,
Suppl. 30: 1-1092.
Peck, O., Z. Bouéek, and A. Hoffer. 1964. Keys to
the Chalcidoidea of Czechoslovakia (Insecta: Hy-
menoptera). Mem. Entomol. Soc. Canada 34: l-
120.
Schulz, W.A. 1906. Strandgut. Spolia hymenop. Pad-
erborn: 77-269.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 237-247
TWO NEW PHYCITINE MOTHS OF THE GENUS COENOCHROA
(LEPIDOPTERA: PYRALIDAE) FROM BRAZIL
JAy C. SHAFFER
Department of Biology, George Mason University, Fairfax, Virginia 22030.
Abstract. —Two new sibling species of phycitine moths, Coenochroa dentata n. sp. and
C. prolixa n. sp., are described from Rio Brilhante, Mato Grosso do Sul, Brazil. Reference
is made to North American species of the genus. Adult moths and genitalia of both sexes
are illustrated and scanning electron micrographs of denuded head capsules and of certain
genital structures are included.
Key Words:
The genus Coenochroa Ragonot, 1887 was
previously known from three species, all
North American. C. bipunctella inhabits the
Atlantic and Gulf coastal plains, while i//i-
bella and californiella are mainly western,
the former extending eastward 1n the Great
Lakes region, the latter ranging south into
Panama (Shaffer 1984).
While recently examining a series of
pyralid moths collected by Dr. Vitor Becker
in Brazil I segregated for further study a
small series of specimens from Rio Bril-
hante, Mato Grosso do Sul which externally
resemble californiella in size, wing pattern,
venation, and frons modification. On dis-
section these proved to be two undescribed
sibling species. The male and female geni-
talia of these two species fit well within the
parameters of Coenochroa, are quite dis-
tinct in spite of the great external similarity
of these moths, and exhibit unusual apo-
morphies in the male genitalia of both
species. These two Brazilian species are de-
scribed herein with a view to inclusion in
the pyralid section of the Checklist of Neo-
tropical Lepidoptera (J. B. Heppner, ed., in
prep.).
Although a review of the genus would be
premature, I have included a key to the five
C. dentata, C. prolixa, taxonomy, neotropics
known species. It is useful to know that: a)
only i/libella has a white costal band, b) all
but i/libella have the dark forewing spot, c)
bipunctella is the only species in the south-
eastern U.S.A., d) californiella and illibella
are sympatric and have very similar geni-
talia for both sexes, but are readily distin-
guished externally, e) dentata and prolixa
are sympatric and difficult to separate ex-
ternally, but both sexes have distinctive
genitalia. One should refer to Shaffer (1968,
1984) for illustrations and other informa-
tion pertaining to North American species
of Coenochroa.
KEY TO SPECIES OF COENOCHROA
1. Male genitalia with valve rounded, unmodi-
fied, not dentate; forewing spot present or ab-
sent (i/libella), costal band present (i/libella) or
absent; frons with central beak minute to
prominent; North and Central America 2)
— Male genitalia with valve dentate; forewing with
dark spot at lower outer angle of cell; costal
band absent; frons with central beak minute
(e.g. Fig. 17) or absent (Fig. 20), varying in-
traspecifically; Brazil 4
2. Forewing spot absent; white costal band pres-
ent; frons with central beak large, protruding
well beyond rim of frons modification; western
North America and Great Lakes region
illibella (Hulst)
238 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ati
Figs. 14. Adult moths and head profiles. 1-2, Coenochroa dentata, male paratype (USNM slide no. 57864).
3-4, Coenochroa prolixa, male paratype (USNM slide no. 57867). Scale bar = 2 mm (Figs. 1, 3), | mm (Figs.
2, 4).
— Forewing with dark spot at lower outer angle — Forewing pale yellow, not darker anterior to
of cell; costal band absent; frons with the cen- cell; costa of valve with single large tooth, sac-
tral beak small to absent, not extending beyond cus very long (Fig. 32); ductus bursae with loop
rim 3 (Fig. 41) prolixa Shaffer
3. Central beak of frons extending to rim; aede-
agus with vesica bearing numerous small cor-
nuti and single larger one; Atlantic and Gulf METHODS
coastal plains :
bipunctella (Barnes and McDunnough)
— Central beak of frons minute, not reaching rim;
Macerated and cleared genitalia were
stained with 0.3% aqueous solution of mer-
aedeagus with a single long slender cornutus; curochrome, mounted in Euparal, and pho-
western North America through Central Amer- tographed with a Nikon AFM camera on a
ica californiella Ragonot_ Nikon S-Ke II microscope set for Koehler
4. Forewing light yellowish brown, darker ante- illumination. All scanning electron micro-
rior to cell; costa of valve with serrate flange
(Fig. 24), vinculum with saccus very short (Fig. StODE work was done with a Hitachi S-530
23); ductus bursae straight (Fig. 38) SEM at 5 Kv. One head capsule for each of
dentata Shaffer the two species was macerated in hot 10%
VOLUME 91, NUMBER 2 239
Figs. 5-10. Scanning electron micrographs of denuded head capsules in frontal (top row), anterodorsal (middle
row), and lateral (bottom row) views, all to same scale. 5-7, Coenochroa dentata, paratype (USNM slide no.
57865). 8-10, Coenochroa prolixa, holotype (J. Shaffer slide no. 2239). Scale bar = 0.5 mm.
KOH, cleaned, denuded of scales, mounted Coenochroa dentata Shaffer,
from 95% ethanol using silver paint, air New SPECIES
dried, and sputter coated with gold. The Figs. 1, 2, 5-7, 11-13, 17-19,
dentata valve (Figs. 30-31) was similarly 23-31, 38-40
treated. All original photographs were taken Diagnosis.—Externally very similar to
on Kodak Technical Pan Film 2415. prolixa, but with forewing ground light yel-
Figs. 11-16.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ES,
Denuded head capsules showing ventral aspect (top row), mouthparts (middle row; g = galea,
mp = maxillary palpus, pf = pilifer), and ventral view of frons protuberance. 11-13, Coenochroa dentata,
paratype (USNM slide no. 57865). 14-16, Coenochroa prolixa, holotype. Scale bar = 250 um (Figs. 11, 14), 50
um (Figs. 12, 15), 100 wm (Figs. 13, 16).
lowish brown, darker and orange brown an-
terior to cell. Male genitalia with costa of
valve bearing serrate flange (Fig. 23); saccus
very short; aedeagus with two strong sub-
equal apical spines (Fig. 29). The serrate
flange is unique to this species of the genus.
Female genitalia with ductus bursae straight
(Fig. 38); ostium with lip-like sclerotization.
Description.—Frons light brown, darker
laterally in some specimens; protuberance
completely covered with scales except for
extreme tip, form variable (see below). La-
bial palpus (Fig. 2) deflected, slender, about
3.3 times as long as eye diameter; basal seg-
ment about as long as second, third about
“4 as long as second; light brown with nu-
merous scattered brown-tipped scales,
overall color similar to frons. Maxillary pal-
pus 3-segmented, minute. Proboscis greatly
reduced. Ocellus rudimentary. Vertex, oc-
VOLUME 91, NUMBER 2
Figs. 17-22. Enlargments of frons protuberances in frontal (top row), anterodorsal (middle row), and lateral
(bottom row) views. 17-19, Coenochroa dentata, paratype. 20-22, Coenochroa prolixa, holotype. Scale bar =
100 um (Figs. 17, 20), 200 um (Figs. 18, 19, 21, 22).
ciput, patagium, and tegula light brown.
Forewing radius about 8-9 mm: venation
variable; R, usually free, sometimes stalked
with R,,;; R, always stalked with R,,;, the
common stalk variable in length; R,;,,
stalked with R,, the common stalk variable
in length; M, from upper outer angle of cell;
M,,, fused, stalked with Cu,, the common
stalk variable in length, from lower outer
angle of cell. Ground light yellowish brown,
242 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 23-31. Coenochroa dentata, male genitalia. 23-36, holotype (J. Shaffer slide no. 2254). 27, paratype
(USNM slide no. 57865). 28-31, paratype (USNM slide no. 57864). 23, male genitalia. 24, detail showing tubular
anellus and serrate costa flange. 25, aedeagus, lateral view, same scale as fig. 23. 26, medial process of gnathos.
27, detail showing gnathos teeth. 28, aedeagus, dorsal view, same scale as fig. 23. 29, aedeagus tip, enlarged.
30, SEM, right valve, ventral view of costa flange. 31, same, detail of costa tip. Scale bar = 100 um (Fig. 30),
25 um (Fig. 31).
VOLUME 91, NUMBER 2
set with numerous scattered brown-tipped
scales, with orange brown cast anterior to
cell and posterior to A,, somewhat more
yellowish cast on distal half of cell, between
M, and M,,,;, and in A, fold. Veins indis-
tinctly marked with white, best developed
on cubitus and A,. Distinct brown spot at
lower outer angle of cell.
Hindwing light brown, darker at apex.
M,,,; fused, long stalked with Cu,, from
lower outer angle of cell.
Male genitalia (Figs. 23-31) with medial
process of gnathos U-shaped, arms minute-
ly spinose, spines irregular, variable (Fig.
27), not extending onto midregion of
U-shaped process. Anellus forming sclero-
tized tube around aedeagus, complete ex-
cept at dorsal midline. Vinculum broadly
triangular, hoodlike, midregion membra-
nous except near saccus; saccus short, about
as long as broad. Valve with costa carinate,
projecting beyond midregion of valve, dor-
sal surface a horizontal somewhat concave
flange bearing row of strong, irregular teeth
on its inner margin (Figs. 24, 30, 31); valve
unmodified elsewhere, tip rather broadly
rounded. Aedeagus slender, about 10 times
as long as maximum width, most slender
just anterior to middle, broadest at posterior
end; posterior end with a pair of strong par-
allel spines fused to aedeagus tube, left spine
longer and curved at tip, right spine straight,
about %4 as long as left; vesica unarmed.
Female genitalia (Figs. 38-40) with ovi-
positor triangular, about 7 as wide at base
as long, moderately setose; typical of genus.
Apophyses straight, moderately robust;
posterior about 1.25-1.5 times as long as
anterior. Eighth segment with scattered
moderate setae, except bare anterior third
of dorsal surface, ventrally forming pair of
triangular lobes which approach each other
most closely at posterior of segment, here
separated by roughly one-fifth of segment
width; ventrally these lobes joined by mem-
brane roughened with numerous minute,
closely-set cusps; this membrane extending
anterior to ostium and posteriorly to ovi-
243
positor lobes as 8-9 intersegmental mem-
brane; 7-8 intersegmental membrane sim-
ilarly roughened (Fig. 40). Ostium heavily
sclerotized, posteriorly lip-like, broadly
lobed, lateral margins curving anteriorly.
Ductus bursae with anterior half membra-
nous, longitudinally rugose; posterior por-
tion smooth, flat, heavily sclerotized, grad-
ually broadening posteriorly. Corpus bursae
round to elongate, membranous, lacking
signum; its surface set with minute, rather
widely separated scale-like structures rather
difficult to discern optically. Ductus sem-
inalis from middle of ductus bursae.
Holotype. —4, labelled: “Rio Brilhante M
Grosso, Brasil 22. I. 1971 Becker leg.”’; “*d
genitalia on slide 2254 J. C. Shaffer”:
“Holotype Coenochroa dentata Shaffer”
[NMRJ].
Paratypes.—3 6, same locality as holo-
type, dates: 23-I-1971 (USNM slide 57865),
[USNM], 25-I-1971 (USNM slide 57864),
[USNM}]; 25-I-1971, Becker col. no. 13800,
undissected [NMRJ]. 2 2, same locality as
holotype, dates: 22-I-1971 (J. Shaffer slide
2251), [NMRJ]; 23-I-1971 (USNM slide
57866), [USNM]. All labelled: ‘‘Paratype
Coenochroa dentata Shaffer.” Specimen de-
position given in brackets.
Distribution. — Known only from the type
locality.
Etymology.—The specific epithet is an
adjective derived from the Latin dentata
(toothed) in reference to the toothed flange
of the valve.
Coenochroa prolixa Shaffer,
NEw SPECIES
Figs. 3, 4, 8-10, 14-16, 20-22,
32-37, 41-43
Diagnosis.—Externally very similar to
dentata, but with forewing ground pale yel-
low and not darker anterior to cell. Male
genitalia (Fig. 32) with valve costa lacking
serrate flange, terminating in blunt tooth:
saccus extremely elongated; aedeagus with
single apical spine (Fig. 35). Female geni-
talia with midregion of ductus bursae form-
244
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 32-37.
no. 57867). 32, male genitalia. 33, aedeagus, same scale as Fig. 32. 34, medial process of gnathos. 35, aedeagus
tip, enlarged. 36, detail showing gnathos teeth. 37, labial palpus.
ing sclerotized loop (Fig. 41). The single large
costa tooth, elongate saccus, and ductus bur-
sa loop are each unique to this species of
Coenochroa.
Description. — Forewing radius about 8-
9 mm: venation similar to C. dentata.
Ground pale yellow throughout, set with
Coenochroa prolixa. 32-35, 37, Holotype (J. Shaffer slide no. 2239). 36, Paratype (USNM slide
numerous darker brown and rust colored
scales. Veins marked with white, most no-
tably cubitus and A,. Distinct brown spot
at lower outer angle of cell.
Hindwing nearly uniformly grayish white,
somewhat darker at apex in some speci-
mens. Venation as in dentata.
VOLUME 91, NUMBER 2 245
Figs. 38-43. Female genitalia. 38-40, Coenochroa dentata. 41-43, Coenochroa prolixa, paratype (USNM
slide no. 57868). 38, paratype, dorsal view (USNM slide no. 57866). 39, paratype, lateral view (J. Shaffer slide
no. 2251). 40, detail of 7-8 intersegmental membrane (slide 2251). 41, dorsal view. 42, detail of corpus bursae,
inner surface. 43, detail of 7-8 intersegmental membrane. Scale bar = 25 um (Fig. 40), 10 um (Fig. 42), 25 um
(Fig. 43).
246
Male genitalia (Figs. 32-36) with medial
process of gnathos U-shaped, covered with
numerous, minute, rather regular recurved
teeth (Fig. 36), these extending onto midline
of gnathos, though there much diminished.
Juxta platelike, dorsally membranous with
sclerotized portion emarginate, ventrally
with a pair of short strong protuberances.
Vinculum broadly triangular, hoodlike,
membranous along midline; saccus ex-
tremely elongate, very slender, anterior end
somewhat bulbous. Valve with blunt costal
projection on distal 74; straight single row
of about 8-10 setae extending from base of
projection toward base of valve, row par-
allel to and about '4 distance across valve
from costal margin; valve unmodified else-
where, tip rounded. Aedeagus very slender,
about 20 times as long as wide, distal % with
numerous parallel lateral diagonal folds (Fig.
35), on distal “4 these folds meeting mid-
ventrally to form pattern of chevrons; distal
end of aedeagus with a single short stout
medially angled spine; vesica unarmed.
Female genitalia (Figs. 41-43) similar to
those of dentata, differing as follows: broad
sclerotized ostium lip absent; sclerotized
posterior portion of ductus bursae longer,
extremely flattened, curved, not broadened
posteriorly; midregion of ductus bursae
forming a prominent, heavily sclerotized,
thickened loop (Fig. 41).
Holotype.—4, labelled: “Rio Brilhante,
Mato Grosso, Brasil 23-27. X. 1970 V. O.
Becker col.”’; “*é genitalia on slide 2239 J.C.
Shaffer’; ‘‘Holotype Coenochroa_ prolixa
Shaffer”; [NMRJ].
Paratypes.—4, same locality as holotype,
date: 25-1-197(1), (USNM slide 57867),
[USNM]. 6 2, same locality as holotype,
dates: 25-X-1970 (USNM< slide 57863),
[USNM]; 27-X-1970, Becker col. no. 13304
(J. Shaffer slide 2256) [NMRJ]; 21-I-1971
(USNM slide 57868), [USNM]; 23-I-1971
(USNM slide 57869), [USNM], 25-I-1971,
Becker col. no. 13925, (J. Shaffer slide 2241),
[NMRJ]; 25-I-1971 (J. Shaffer slide 2253),
[NMRJ]. All labelled: “‘Paratype Coeno-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
chroa prolixa Shaffer.” Specimen deposi-
tion given in brackets.
Distribution.— Known only from the type
locality.
Etymology.—The specific epithet is an
adjective derived from the Latin prolixa
(stretched out) in reference to the elongated
saccus.
Apomorphies.—In dentata the costal
flange and tubular anellus of the male gen-
italia, and in prolixa the elongate slender
saccus and loop of the ductus bursae are
derived features not shared with any other
known Coenochroa species.
Immature stages and hosts.— Unknown.
The habitus of Coenochroa species suggests
that the larvae may be associated with grass-
es.
Discussion.— Dr. Becker (pers. com.)
notes that these moths were collected by
mercury vapor light in savanna habitat bor-
dering a gallery forest.
These two species are extremely similar
externally, and while the forewing color dif-
ferences noted are useful, I have not found
external structural differences that will re-
liably distinguish all specimens. The frons
modification clearly separates the North
American C. californiella and illibella, and
obvious differences in the frons structure are
seen in the scanning electron micrographs
of the two specimens illustrated herein (Figs.
17-22). Unfortunately, optical examination
of the remaining specimens suggests that
these differences are possibly more a matter
of individual than interspecific variability.
It will be necessary to examine a larger series
of specimens to determine if any frons char-
acters useful in separating these two species
exist.
ACKNOWLEDGMENTS
I am grateful to Dr. Vitor Becker for the
opportunity to examine a portion of his col-
lections, to Mr. Jan Endlich for assistance
with photoprocessing and SEM work, and
to Mr. Vichai Malikul for his comments on
preparing the illustrations.
VOLUME 91, NUMBER 2
The holotype and selected paratypes are
deposited in the National Museum, Rio de
Janeiro, Brazil [NMRJ]. The remaining
paratypes are in the National Museum of
National History, Smithsonian Institution,
Washington, D.C., USA [USNM]. The
scanning electron microscope used in this
study was supported in part by NSF Grant
No. BSR-8511148.
LITERATURE CITED
Heppner, John B., ed. Checklist of Neotropical Lepi-
doptera. Part 2. E. J. Brill, Leiden. In prep.
Shaffer, J.C. 1968. A Revision of the Peoriinae and
Anerastiinae (Auctorum) of America North of
Mexico (Lepidoptera: Pyralidae). U.S. Natl. Mus.
Bull. 280: 1-124.
. 1984. Neotropical Pyralid moths transferred
from Anerastiinae (Auctorum) to Phycitinae. Proc.
Entomol. Soc. Wash. 86(2): 383-395.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 248-256
TYPE MATERIAL OF FOUR AFRICAN SPECIES OF
NOTARCHA MEYRICK, WITH DESIGNATIONS OF
LECTOTYPES AND CHANGES IN SYNONYMY
(LEPIDOPTERA: CRAMBIDAE: PYRAUSTINAE)
JAY C. SHAFFER AND EUGENE MUNROE
(JCS) Department of Biology, George Mason University, Fairfax, Virginia 22030; (EM)
Granite Hill Farm, R.R. #2, Dunrobin, Ontario KOA 1T0, Canada
Abstract. —Lectotypes are designated for three Zeller species: Notarcha quaternalis, N.
temeratalis, and N. muscerdalis. These and the holotype of N. cassusalis Walker, the type
of Notarcha, are redescribed and the wings, head profiles, and female genitalia illustrated.
Key Words:
types, African Pyraustinae
In his revision of the Pyralidae Hampson
(1898: 728) synonymized seven names un-
der Lygropia quaternalis (Zeller). In re-
searching our paper (in prep.) on the Cram-
bidae of Aldabra Atoll, we found it necessary
to reexamine this synonymy to determine
the identity of an Aldabran species erro-
neously identified in the literature as qua-
ternalis. On examining type specimens we
discovered that most, perhaps all, of these
seven names represent distinct species. The
Aldabra species matches none of them and
will be described as new.
The purpose of this paper primarily is to
designate lectotypes for guaternalis and two
related African species to provide stability
for the names, and secondarily to redescribe
and illustrate the three lectotypes and the
holotype ofan additional species to separate
previously confused forms and to facilitate
identification.
The African species that Hampson syn-
onymized under quaternalis are temeratalis
Zeller, and cassusalis Walker. All three
species are properly referred to Notarcha
Meyrick, 1884, for which cassusalis is the
Notarcha quaternalis, N. temeratalis, N. cassusalis, N. muscerdalis, \ecto-
type species. In this paper we include the
related N. muscerdalis, not part of Hamp-
son’s synonymy, but of which we studied
Zeller’s type. Notarcha 1s a large genus with
many undescribed species and deserving of
extensive study. The scope of this paper is
limited to delineating described African
species.
Type material referred to herein is in the
collections of the British Museum (Natural
History) [BMNH], and the Naturhistoriska
riksmuseet, Stockholm [NHRM].
Key TO INCLUDED SPECIES OF NOTARCHA
1. Forewing uniformly yellow, with single dark
discal spot (Fig. 4) . muscerdalis
— Forewing yellow with transverse lines or dif-
fuse bands of darker yellow; with dark discal
spot and 3 similar spots along costal margin 2
2. Labial palpus with third segment mostly dark
brown and first segment with brown medial
spot (Fig. 5, arrows) .... : . quaternalis
— Labial palpus without dark markings ........ 3
3. Forewing with anterior half of transverse pos-
terior band in form of diagonal line (Fig. 3) .
(Fig. 2) . cassusalis
VOLUME 91, NUMBER 2
Notarcha quaternalis (Zeller)
Figs. 1, 5, 9, 13-16
Botys quaternalis Zeller, 1852, pp. 44-45.
Diagnosis.—Among the described Afri-
can species of the Notarcha quaternalis
complex the dark spot on the basal segment
of the labial palpus and the dark third seg-
ment (Fig. 5, arrows) are each unique to this
species, as 1s the spiny knob near the en-
trance to the corpus bursae.
Description (female).—Frons smooth,
covered with appressed yellow scales. La-
bial palpus obliquely ascending; first seg-
ment yellow with prominent dark brown
medial spot adjacent to eye; second segment
yellow; third segment short, subcylindrical,
dark brown with yellow apex. Maxillary pal-
pus slender, cylindrical, yellow with sub-
apical dark-brown band. Proboscis scales
yellow. Antenna filiform, finely ciliate and
with single long cilium near base of each
segment; scales light yellow. Eye diameter
about 0.75 mm, black. Ocellus prominent,
with clear lens surmounted on black ellip-
tical base. Vertex yellow. Occiput and tuft
of scales just posterior to ocellus straw col-
ored. Patagium, tegula, and thorax yellow.
Forecoxa brownish yellow; forefemur
brownish yellow, yellow approaching apex,
dark brown at apex; foretibia yellow, dark
brown at apex; foretarsus yellow, black
markings on basal half and on distal third.
[Meso- and metathoracic legs lost on type.]
Forewing (Fig. 1) radius 11 mm; with four
subequal dark brown spots; first (most bas-
al) spot elliptical, on costa near wing base
and separated from it by its own width: sec-
ond spot elliptical, on costa at one-fifth dis-
tance to wing apex; third (discal) spot nearly
circular, very slightly larger than first two,
on closing vein of cell; fourth spot slightly
smaller and more narrowly elliptical than
others, very near to costal margin of wing,
but separated from margin by about one-
third its length. Ground yellow, marked with
diffuse transverse lines of darker yellow;
three short lines on basal half of forewing,
249
the first descending from the first spot and
separated from wing base by its own width;
the second from between the first and sec-
ond spots; the third from just distal to sec-
ond spot; a fourth line (transverse posterior)
descending from fourth spot to just beyond
center of wing, angling sharply basad to just
beyond posterior outer angle of cell, then
angling sharply posteriorly to posterior wing
margin; a sixth runs very near to outer mar-
gin of wing, broad near wing apex, narrow-
ing posteriorly, and absent from posterior
third of wing.
Hindwing with first line indistinct, de-
scending from second of forewing; second
line better developed and descending from
third of forewing; third line not matched
with any forewing line, descending from
lower outer angle of cell; fourth line de-
scending from fourth of forewing, some-
what sinuate, its two most distal portions
between M, and M, and on Ist A; fifth line
relatively broad, narrowing posteiorly and
approaching posterior end of fourth line;
outer margin of wing with distinct dark yel-
low terminal line; fringe light yellow, darker
on basal half.
Lines developed on undersides of both
sets of wings; discal spot prominent on un-
derside of forewing, other spots not devel-
oped on undersides.
Female genitalia (Figs. 13-16) with ovi-
positor compressed, with one zone of setae
along its inner margin (seen extended in Figs.
13, 14), and a second zone along its outer
margin; setae of outer margin densely set
and three to four times as long as setae of
inner margin. Anterior apophysis nearly
twice as long as posterior. Ostial chamber
small; immediately adjacent to a flat scler-
otized trough: posterior part of inner surface
of trough studded with numerous minute,
sharp, posteriorly directed spines; anterior
part spineless, somewhat granular in ap-
pearance. Ductus bursae membranous, with
round expanded pouch, studded with nu-
merous minute cusps. Corpus bursae slight-
ly over twice as long as wide; nearly uni-
250 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4. Wing patterns. 1, Notarcha quaternalis, lectotype; 2, N. cassusalis, holotype; 3, N. temeratalis,
lectotype; 4, N. muscerdalis, lectotype. Scale bar = 2 mm.
Figs. 5-8. Head profiles of above specimens. 5, N. quaternalis, 6, N. cassusalis, 7, N. temeratalis, 8, N.
muscerdalis. Scale bar = 1 mm (Figs. 5-8).
formly finely scobinate, each scobination set its posterior end folded into a short knob-
in center of irregular plate, many plates hex- shaped pouch with spines facing outward,
agonal or nearly so; posterior part of bursa these spines numerous, slender, sharp
with irregular sclerite, spinose on both sides, pointed; signum absent. Ductus seminalis
VOLUME 91, NUMBER 2
[@genitalia on |
slide /870
J.C. Shaffer
Riksmuseum
| Stockholm
genitalia on
slide /8 i |
J.C. Shaffer
Figs. 9-12.
10
+
Olid
musceroaus
Types with labels. 9, Notarcha quaternalis, lectotype (1.6 x); 10, N. cassusalis, holotype, insert
shows reverse side of “Pt Natal” label (1.4 =); 11, N. temeratalis, lectotype (1.6 x); 12, N. muscerdalis, lectotype
(1.4 x).
from membranous posterior part of corpus
bursae.
Type locality.— Natal, South Africa.
Lectotype, hereby designated, labelled:
““Caffraria.”; “197”; ““Riksmuseum Stock-
holm”; “g genitalia on slide 1870 J. C. Shaf-
fer’; ““Botys quaternalis Lectotype by J.
Shaffer & E. Munroe, 1989” [NHRM].
Notarcha cassusalis (Walker)
Figs. 2, 6, 10, 17-21
Zebronia cassusalis Walker, 1859, p. 477.
Diagnosis.—Among the described Afri-
can species of the Notarcha quaternalis
complex this species 1s externally similar to
quaternalis, but lacks dark markings on the
labial palpus, and has broader more diffuse
transverse bands on the wings. The spinose
triangular plate at the entrance to the corpus
bursae and the internally spinose ductus
seminalis are each characteristic of this
species.
Description (female).—Frons smooth,
covered with appressed yellow scales. La-
bial palpus obliquely ascending, third seg-
ment short, subcylindrical; all segments
uniformly yellow on outer side, lacking dark
spots of guaternalis. Maxillary palpus light
yellow. Antenna as in quaternalis. Eye di-
ameter about 0.5 mm. Ocellus as in qua-
ternalis. Vertex yellow; patagium vivid yel-
low centrally, lighter peripherally; tegula
extending nearly to abdomen, vivid yellow.
Outer side of forecoxa yellow basally,
brown elsewhere; forefemur brown on inner
side, light yellow on outer side, small dark
brown spot on apex; foretibia yellow on bas-
al half, dark brown on distal half; foretarsus
252 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 13-21. Female genitalia. 13, Notarcha quaternalis, lectotype (20 x); 14, ovipositor, enlarged (45 =); 15-
16, armature at entrance to corpus bursae, two focal levels (50x); 17, N. cassusalis, corpus bursae surface,
enlarged to show surface texture (220 x); 18, female genitalia (28 x); 19, armature at entrance to corpus bursae
(110); 20, rotated specimen showing origin of ductus seminalis (28 x ); 21, ductus seminalis, enlarged to show
internal spines (220 x).
yellow, brown spot at apex of first segment, midtibia nearly uniformly light yellow,
smaller spot at apex of second segment, third lacking dark markings; midtarsus light
segment yellow, fourth segment brown, fifth brownish yellow. Metathoracic leg color-
segment yellow. Midfemurrather uniformly ation similar to that of mesothoracic leg.
light yellow, small dark brown spot at apex; Forewing (Fig. 2) radius about 9 mm.
VOLUME 91, NUMBER 2
Figs. 22-27.
27
Female genitalia. 22, Notarcha temeratalis, lectotype (28 x ); 23, corpus bursae surface, enlarged
to show surface texture (200). 24, Notarcha muscerdalis, \lectotype (23 x); 25, signum, enlarged (55 =); 26,
signum and adjacent bursa surface, enlarged (220 =); 27, rotated specimen showing origin of ductus seminalis
(28 x).
Ground light yellow with broad diffuse viv-
id yellow bands. Costa with three promi-
nent dark brown spots; spot at basal band
reniform, spot on antemedial band the larg-
est of the three, circular, distal spot just be-
yond midregion of wing and smallest of the
three spots. Discal spot dark brown, large
and prominent, somewhat triangular with
longest side transverse and distal.
Hindwing light yellow with broad diffuse
vivid yellow bands.
Female genitalia with ovipositor com-
pressed, moderately setose. Anterior
apophysis 1.5 times as long as posterior,
slightly decurved; posterior apophysis slen-
der, angled at anterior third and at posterior
third. Ostial chamber well sclerotized,
broadly expanded posteriorly; anterior one-
fifth devoid of spines and somewhat gran-
ular in appearance; posterior four-fifths with
numerous minute, hairlike spines, these di-
rected inward or posteriorly and shortest
254
along lateral regions of ostial chamber. Duc-
tus bursae membranous on posterior one-
fourth; anterior three-fourths with irregular
sheetlike sclerotization, this folded longi-
tudinally about one and one-half times, the
nearly closed fold with patch of strong in-
wardly directed spines near its anterior end;
other side of sclerotized sheet extending into
corpus bursae as strongly setose triangular
plate (Fig. 19); anterior part of ductus bur-
sae with small membranous pouch. Corpus
bursae nearly twice as long as broad, lacking
signum, nearly uniformly finely scobinate
(Fig. 17), each scobination set in center of
minute plate, plate hexagonal or approxi-
mately so. Ductus seminalis from near pos-
terior end of corpus bursae (Fig. 20), mem-
branous, its inner surface set with numerous
slender setae (Fig. 21).
Type locality.— Natal, South Africa.
Holotype, labelled: ““Holotype’’; “Type”;
“Pt Natal [& on reverse side] 57 3°; “‘Ze-
bronia Cassusalis”’; “@ Pyralidae Brit. Mus.
Slide No. 18060” [BMNH].
Notarcha temeratalis (Zeller)
New COMBINATION
Figs. 3, 7, 11, 22-23
Botys temeratalis Zeller, 1852, pp. 45-46.
Diagnosis.—Among the described Afri-
can species of Notarcha only temeratalis has
a portion (anterior half) of the transverse
posterior line of the forewing developed as
a narrow diagonal line. The species is also
unique in that the corpus bursae is unmod-
ified.
Description (female).—Frons smooth,
covered with appressed yellow scales. La-
bial palpus obliquely ascending, third seg-
ment short, subcylindrical; all segments
white to straw yellow on outer sides, first
with indistinct light-brown medial spot ad-
jacent to eye, first and second somewhat
darker apically. Maxillary palpus cylindri-
cal, straw yellow. Base of proboscis clothed
with straw-yellow scales. Antenna as de-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
scribed for N. quaternalis. Eye diameter 0.6
mm, black. Ocellus as described for N. gua-
ternalis. Occiput white; patagium yellow an-
teriorly, white posteriorly; tegula white with
yellow medial band. Thoracic vesture of
broad white scales beneath head. Forecoxa
light brown on inner side, white on outer
side; forefemur similar, but with dark-brown
apical spot; foretibia yellow basally, distally
with tuft of dark-brown scales; foretarsus
with dark-brown subapical spot. Midtho-
racic leg rather uniformly straw yellow, with
small dark-brown spot at apex of femur.
Forewing (Fig. 3) radius 9 mm. Ground
white, bearing four subequal dark-brown
spots; first (most basal) spot nearly round,
on costa near wing base and separated from
it by its own width, second spot slightly larg-
er, nearly round, somewhat pointed poste-
riorly, on costa at one-fourth distance to
wing apex; third (discal) spot slightly small-
er than first two, nearly circular, at outer
margin of cell; fourth spot less well devel-
oped than other three, in form of oblique
dash near costa, not quite reaching wing
margin. Ground white with yellow mark-
ings in form of line, diffuse bands, and
broader diffuse patches; a broad band de-
scending obliquely from second spot to pos-
terior wing margin near base; a second par-
allel band descending from point midway
between second and third (discal) spots to
posterior margin, gradually broadening pos-
teriorly; a small yellow patch immediately
distal to discal spot; a narrow line descend-
ing obliquely distad from fourth spot to Ist
A, angled basad between Cu, and Cu,, then
obliquely and basad to posterior margin as
broad diffuse band; a broad yellow patch in
anterior preterminal area, bordered entirely
by white ground; a smaller patch from angle
of narrow line to terminus; a well developed
thin terminal line of yellow on outer margin
of wing.
Hindwing ground white; a diffuse yellow
band running basally from cubitus in cell,
then angled posteriorly to anal margin; a
broad yellow band runs obliquely from pos-
VOLUME 91, NUMBER 2
terior outer angle of cell to posterior wing
margin; a narrow yellow line descending
obliquely from fork of Sc and Rs to Cul,
there broadening to an oval yellow patch
just anterior to anal angle, oval patch sep-
arated from wing margin by narrow white
ground; a large yellow patch near apex bor-
dered entirely by white ground; outer mar-
gin with a distinct narrow yellow terminal
line, and a similar line on fringe near its
base, best developed on posterior half of
outer margin.
Undersides of both wings with lines and
patches showing, but indistinctly so; discal
spot poorly developed, other three spots ab-
sent.
Female genitalia (Figs. 22, 23) with ovi-
positor compressed. Anterior apophysis
about 1.7 times as long as posterior. Ostial
chamber moderately well sclerotized, some-
what flattened, with margins turned dorsad
and rolled inward, posterior half with nu-
merous, sharp, posteriorly directed spines.
Ductus bursae short, membranous, with
small lateral pouch, expanded toward cor-
pus bursae, bearing fine granulations, dis-
tinct spines absent. Corpus bursae about 1.5
times as long as wide; signum absent; sur-
face finely scobinate (Fig. 23), each minute
scobination borne on an irregular somewhat
hexagonal plate; scobinations best devel-
oped on dorsal surface. Ductus seminalis
arising dorsally from junction of corpus
bursae and ductus bursae.
Type locality.—South Africa, roughly the
region of the Transvaal and Orange Free
State. Zeller (p. 46) cites the type locality as:
“Patria ad fluvios Limpoponem et Garie-
pem.” Gariep (Gareep) 1s an obsolete name
which according to Skead (1973: 61, 171,
239) applied to both the lower and upper
(above its junction with the Vaal) Orange
River and to the lower Vaal River. Zeller’s
material came from Wahlberg, whose col-
lecting localities in South Africa are impre-
cisely known. Horn and Kahle (1936: 293)
record Wahlberg’s first journey (1838-1845)
there as to southern Africa, only later (1853)
255
traveling to southwestern Africa. Therefore,
it is probable that Zeller’s reference to the
Gariep applies to the upper (eastern) Orange
or possibly to the lower Vaal, but not to the
lower (western) Orange.
Lectotype, hereby designated, labelled:
“Caffraria.”; “436; ““Riksmuseum Stock-
holm”’; ‘“*? genitalia on slide 1871 J. C. Shaf-
fer’; ‘‘“Botys temeratalis Lectotype by J.
Shaffer & E. Munroe, 1989” [NHRM].
Notarcha muscerdalis (Zeller)
New ComMBINATION
Figs. 4, 8, 12, 24-27
Botys muscerdalis Zeller, 1852, pp. 43-44.
Diagnosis. — This species 1s distinguished
externally from other African species of No-
tarcha by having a dark discal spot on an
otherwise uniformly yellow forewing and
internally by the presence of a signum on
the corpus bursae.
Description (female).—Frons smooth,
covered with appressed yellow scales. La-
bial palpus obliquely ascending, third seg-
ment short, subcylindrical; second and third
segments uniformly yellow on outer sides,
first segment similar distally, lighter basally;
all segments devoid of dark spots. Maxillary
palpus cylindrical, slender, extending to base
of third segment of labial palpus; yellow.
Antenna as in N. quaternalis. Ocellus well
developed, with clear round lens on black
elliptical base. Vertex yellow; occiput light
yellow; patagium and tegula vivid yellow.
Forewing (Fig. 4) radius 12 mm; ground
nearly uniformly yellow with a single large
dark-brown elliptical, obliquely set discal
spot.
Hindwing uniformly yellow.
Forecoxa brownish yellow; forefemur
brown on inner side, light yellow on outer
side; foretibia brownish yellow on basal half,
brown on distal half; foretarsus with first
segment yellow, second similar but with
suggestion of brown at apex, third yellow
on basal half, brown on distal half, fourth
256
and fifth brown. Meso- and metathoracic
legs nearly uniformly yellow.
Female genitalia (Figs. 24-27) with ovi-
positor compressed; lobes narrow, moder-
ately setose. Anterior apophysis about 1.4
times as long as posterior, curved upward
slightly, foliate at base; posterior apophysis
with distal two-thirds straight. Ostial cham-
ber well sclerotized, long and narrow, four
times as long as central width, expanded
slightly at posterior end; smooth, devoid of
spines or setae. Ductus bursae with poste-
rior one-third membranous, unarmed but
for exceedingly minute widely spaced cusps;
anterior two-thirds sclerotized, set with nu-
merous minute cusps. Corpus bursae nearly
round, slightly longer than wide, posterior
end tapering to ductus bursae; surface finely
scobinate (Fig. 25), each scobination in cen-
ter of minute plate, roughly hexagonal to
foliate; a single small longitudinal signum
(Fig. 25) in center of corpus bursae, formed
of several irregular longitudinal rows of small
sclerotized papillae (Fig. 26). Ductus sem-
inalis (Fig. 27) from extreme posterior end
of corpus bursae, inner surface set with
widely separated minute triangular cusps,
devoid of setae.
Type locality.—South Africa, roughly the
region of the Transvaal and Orange Free
State. (Comment under temeratalis above
applies here also.)
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Lectotype, hereby designated, labelled:
“Type”; “Botys muscerdalis Z. Caffr. 43.
Led 85. Caffraria”’; “Zell. Coll. 1884’; ‘“‘¢
Pyralidae Brit. Mus. Slide No. 18061”; “Bo-
tys muscerdalis Lectotype by J. Shaffer & E.
Munroe, 1989” [BMNH].
ACKNOWLEDGMENTS
We thank Michael Shaffer of the Depart-
ment of Entomology, British Museum (Nat-
ural History) and Bert Gustafsson of the
Section for Entomology, Naturhistoriska
riksmuseet, Stockholm for their faithful cor-
respondence and for generously making
types available for study.
LITERATURE CITED
Hampson, G. F. 1898. A Revision of the Moths of
the Subfamily Pyraustinae and the Family Py-
ralidae. Part 1. Proc. Zool. Soc. Lond. 1898: 590-
761, figs. 1-87, pl. 49, 50.
Horn, Walther and Ilse Kahle. 1936. Uber entomo-
logische Sammlungen. Ent. Beih. Berl.-Dahlem 3:
161-296, pl. 17-26.
Skead, C. J. 1973. Zoo-Historical Gazetteer. Ann.
Cape Prov. Mus. 10: i-v, 1-259.
Walker, Francis. 1859. List of the Specimens of Lep-
idopterous Insects in the Collection of the British
Museum, 1859. 17: 255-508.
Zeller, P. C. 1852. Lepidoptera Microptera, quae J.
A. Wahlberg in Caffrorum Terra Collegit. Kongl.
Vetenskaps-Akademiens Handlingar for Ar 1852:
1-120.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 257-268
DESCRIPTION OF MALE OSTROCERCA RICKER
(PLECOPTERA: NEMOURIDAE) USING THE
SCANNING ELECTRON MICROSCOPE
DERON C. YOUNG, Boris C. KONDRATIEFF, AND RALPH F. KIRCHNER
(DCY, BCK) Colorado State University, Department of Entomology, Fort Collins,
Colorado 80523; (RFK) Department of the Army, Huntington District Corps of Engineers,
Water Quality Section, 502 8th Street, Huntington, West Virginia 25701.
Abstract. —Scanning electron micrographs of the male terminalia are given for all six
species of the stonefly genus Ostrocerca Ricker: O. albidipennis (Walker), O. complexa
(Claassen), O. dimicki (Frison), O. foersteri (Ricker), O. prolongata (Claassen), and O.
truncata (Claassen). Structures of the male genitalia previously not illustrated are de-
scribed. Distribution records for each species are also reported.
Key Words:
graphs
Scanning electron microscopy (SEM) has
been used recently to elucidate fine struc-
tures of the male genitalia of stoneflies
(Kondratieffand Kirchner 1984, Nelson and
Baumann 1987). Newly recognized struc-
tures have been used in making determi-
nations and preparing relationships.
The Nearctic genus Ostrocerca Ricker
contains six species and is known for its
small size (4-8 mm) and complex male gen-
italia (Ricker 1952). This genus occurs in
eastern (four species) and northwestern (two
species) North America (Stark et al. 1986).
Nymphs usually inhabit crenon habitats, and
adults are rare in collections, but can be
collected by sweeping or beating.
Keys to males and females are provided
by Ricker (1952) in his review of the genus
and an updated key to females is given in
his 1965 paper. Hitchcock’s (1974) study
enables the separation of the four eastern
species. Baumann’s (1975) generic descrip-
tion is based primarily on O. truncata
Claassen. Our previous studies of speci-
mens from Virginia (Kondratieffand Kirch-
Stonefly, Plecoptera, Nemouridae, Ostrocerca, scanning electron micro-
ner 1987) and West Virginia (Tarter and
Kirchner 1980) showed that it is difficult to
see the arrangement of structures of the
complex epiproct with standard light mi-
croscopes. Using published illustrations, es-
pecially of the eastern species (Claassen
1923, Hitchcock 1974), it often appeared
that additional taxa could be involved.
Therefore, we used SEM to study the male
genitalia of Ostrocerca. The higher magni-
fication and greater depth of field in SEM
made it possible to illustrate structures not
previously recognized.
METHODS AND MATERIALS
Specimens were prepared for study by re-
moving them from 70-80% ethanol, sub-
mersing them in liquid Freon 12 at — 154°C
for several minutes, and then placing them
into stored liquid nitrogen (—110°C to
— 80°C) and into a Balzers 301 Freeze-Frac-
ture Unit at 10-° my. Specimens were then
mounted and sputter coated with gold in a
Hummer V Sputter Coater. Specimens were
258 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4.
viewed using a Phillips 505 SEM at 30,000
accelerating voltage.
The following institutions and individu-
als provided additional material for this
study: R. W. Baumann, Brigham Young
University, Monte L. Bean Life Science
Museum (RWB); O. S. Flint, Jr., Smithson-
Ostrocerca albidipennis. 1, Epiproct, dorsal view, 312 x. 2, Hypoproct, apex, 845 x. 3, Terminalia,
ventral view, 120. 4, Terminalia, ventral view, 241 x. Terms: hp, hypoproct; vs, ventral sclerite; la, lateral
arms; il, inner lobe; c, cerci; v, vesicle.
ian Institution, National Museum of Nat-
ural History (USNM): R. Foottit, Biosys-
tematics Research Centre, Canadian
National Collection (CNC); J. K. Liebherr,
Cornell University (Cornell), Kathryn C.
McGiffen, Illinois State Natural History
Survey Division (INHS); C. Vogt, Museum
VOLUME 91, NUMBER 2
of Comparative Zoology, Harvard Univer-
sity (MCZ); and J. R. Voshell, Jr., Virginia
Tech.
Morphological terms of the following de-
scriptions follow Baumann (1975). Bau-
mann (1975) provides excellent illustra-
tions of the male terminalia of O. truncata,
clearly indicating the orientation of the
structures of the male epiproct. Abbrevia-
tions used on the SEM photographs include
hp, hypoproct; ds, dorsal sclerite; vs, ventral
sclerite; la, lateral arms; ol, outer lobe; ml,
middle lobe; il, inner lobe; c, cerci; and v,
vesicle.
Ostrocerca albidipennis (Walker)
Nemoura albidipennis Walker, 1852: 191.
Type locality: Nova Scotia, Canada.
Nemoura serrata Claassen, 1923: 289. Type
locality, Bar Harbor, (Hancock Co.)
Maine; Ricker, 1952: 39 (syn.).
Ostrocerca albidipennis, Mlies, 1966: 217.
The distinctive epiproct has a hammer-
head shaped ventral sclerite, which is over-
lapped by the lateral arms of the dorsal
sclerite (Fig. 1). The hypoproct is long and
slender, and terminates in a complex serrate
structure (Fig. 2). The paraprocts have inner
lobes that are hooked medially and beak-
like apically (Figs. 3 and 4). The figures pre-
sented by Claassen (1923) as N. serrata and
by Hitchcock (1974) do not accurately de-
pict these complicated structures.
The female is accurately illustrated by
Hitchcock (1974) and by Ricker (1965).
Material examined.—Holotype M (N.
serrata), Bar Harbor, Maine, 8 VI 1921, C.
W. Johnson (Cornell #1199). Paratype: same
data as holotype, | M (Cornell #1199).
Other specimens: CANADA: Ontario,
Algonquin Park, Costello Lake, 26 V 1938,
W. M. Sprules, | M (INHS); same locality,
29 V 1939, 2 M (INHS); same locality, 26
V 1941, 1 M (INHS); Maynooth, Spring
Creeks, 22 VI 1953, J. F. McAlpine, 2 M
(CNC); Petawawa, Meridian Road, Forest
Station, 28 V 1959, J. R. Vockeroth, | M,
259
1 F(CNC). Quebec, Harrington Lake, Gat-
ineau Park, 27 V 1954, H. J. Huckel, 1 M,
6 F (CNC); Harrington Lake, 30 V 1954, E.
E. Sterns, 3 M, 3 F(CNC); Wakefield, Lind-
say’s Creek, 31 V 1930, J. McDunnough, |
M (CNC). CONNECTICUT: Naugatuck
State Forest, Beacon Falls, 17 V 1961, S.
W. Hitchcock, 5 M, 1 N (USNM). MAINE:
on dropnet, (no locality) 4-14 VI 1970, D.
E. Leonard, 4 M (USNM). MASSACHU-
SETTS: Whately, 18 V 1939, W. B. Nutting,
1 M, 4 F (USNM). NEW HAMPSHIRE:
Warren, 21 VI 1941, T. H. Frison & H. H.
Ross, | M (INHS). TENNESSEE: Cumber-
land Co., small stream, 2 mi. N of Adams
Bridge, Obed River, Rt. 298, 13 V 1988, R.
W. Baumann, C. R. Nelson, RFK & BCK,
1M, 2 F(CSU). VERMONT: Mt. Hay Stack,
2400’, 28 V 1938, W. B. Nutting, 1 M
(USNM). VIRGINIA: Giles Co., Mountain
Lake, 2 VI 1962, J. R. Vockeroth, 2 F (CNC);
Grayson Co., springs into Lewis Fork, Co.
Rt. 603, 4 mi. W of Troutdale, 9-30 VI
1981, 16 M, 5 F, 3 N, RFK & BCK (RFK),
11 M, 9 F, 6 N BCK (VPI); Smyth Co.,
springs to Big Laurel Creek, Trail 166, Jef-
ferson National Forest, 30 VI 1981, RFK
& BCK, 2 F(RFK); Washington Co., spring
to Brumley Creek, Clinch Mountain at Low
Gap, Co. Rt. 690, 3600’, 23 VI 1981, RFK
& BCK, 9 M, 6 F (RFK): Wise Co., Phillips
Creek Rec. Area, N Fork Pound Reservoir,
off Co. Rt. 671, 29 IV 1987, RFK & BCK,
1 M(BCK). WEST VIRGINIA: Pocahontas
Co., Island Lick Run of Greenbriar R., Wa-
toga State Park, Rt. 27, 16 VI 1981, RFK,
1 M (RFK); springs to Hills Creek, Rt. 39,
Monongahela National Forest, 16 VI 1981,
RFK, | M (RFK).
Ostrocerca complexa (Claassen)
Nemouracomplexa Claassen, 1937:43.Type
locality: Artists Brook, Essex County, New
York.
Ostrocerca complexa, Mlies, 1966: 217.
The elaborate epiproct (Figs. 5, 6) of this
species has never been completely illustrat-
260
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 5-8.
ed (see Claassen’s 1937, Fig. 2 and Hitch-
cock’s 1974, Fig. 107). The ventral sclerite
is fork-like with a lower process that is elon-
gate and apically recurved. The membra-
nous basal cushion of the dorsal sclerite has
five to six cuticular spine-like projections
Ostrocerca complexa. 5, Terminalia, dorsolateral view, 194.6, Epiproct, dorsal view, 212 x. 7,
Basal cushion of the dorsal sclerite, 965 «. 8, Terminalia, ventral view, 163 x. Terms: hp, hypoproct; vs, ventral
sclerite; il, inner lobe: c, cerci; v, vesicle.
proximally and four to five distally (Fig. 7).
These projections often appear membra-
nous in SEM photomicrographs. The inner
lobes of each paraproct are triangularly pro-
duced. The hypoproct is bifurcate apically
(Fig. 8). The cerci are notched (Fig. 8).
VOLUME 91, NUMBER 2
The terminalia of the female are illus-
trated by Hitchcock (1974) and Ricker
(1952, 1965).
Material examined.—Holotype M: Art-
ists Brook, Essex Co., New York, 11 VI
1933, C. R. Crosby & H. Deitrich (Cornell
#1692). Paratypes: same locality as holo-
type, 28 June 1923, P. W. Claassen, 1 M
(Cornell #1692).
Other specimens: VERMONT: brooklet
top of Brandon Gap, 2170’, 25 VI 1966, S.
W. Hitchcock, 1 M (USNM). VIRGINIA:
Nelson Co., tributary to South Fork Tye
River, Montebello Fish Culture Station, jct.
Rt. 690 & Rt. 56, 3 V 1980, BCK, | M, 2
F (VPI). WEST VIRGINIA: Pendleton Co.,
Spruce Knob, 4 mi. below summit, 16 V
1963, Field & O. S. Flint, 1 M (USNM);
Pocahontas Co., springs to Hills Creek, 15
mi. E Richwood, Hills Creek Scenic Area,
Rt. 39, Monongahela National Forest, 16 V
1981, RFK, 14 M, 6 F(RFK), same locality
15 V 1986, RFK & BCK, 2 M, | F (BCK);
Tea Creek, Tea Creek Campground, Mo-
nongahela National Forest, 21 V 1978, RFK
& R. M. Meyer, | M (RFK).
Ostrocerca dimicki (Frison)
Nemoura dimicki Frison, 1936: 256. Type
locality: Corvallis, Oregon.
Ostrocerca dimicki, Mlies, 1966: 217.
This species is easily recognized by the
hook-shaped ventral sclerite with erect se-
tose processes (Figs. 9, 10, 11). The inner
lobes of the hypoproct are robust and elon-
gate, apically acute, and setose (Fig. 12). The
hypoproct is broadly truncate apically and
bears a medial sclerotized slender process
(Fig. 12).
The female terminalia is illustrated by
Ricker (1965).
Material examined.— Holotype M, Cor-
vallis, Oregon, branch Dixon Creek,
emerged 28 IIT 1935, R. W. Prentiss (INHS).
Paratypes: Corvallis, drainage stream, 16 III
1934, E. E. Ball, 5 M (USNM); same data,
2 F (INHS); Corvallis, Brooklane Creek, 28
IT 1935, R. W. Prentiss, 1 M (USNM).
261
Other specimens: CANADA: British Co-
lumbia, 30 V-18 VI 1955, R. Coyles, 1 M
(CNC). OREGON: Washington Co., North
Plains, 7 IV 1968, K. Goeden, 1 M, 2 F
(USNM).
Ostrocerca foersteri (Ricker)
Nemoura foersteri Ricker, 1943: 70. Type
locality: Reservoir Creek, Cultus Lake,
British Columbia.
Ostrocerca foersteri, Mlies, 1966: 218.
The epiproct is simple (Figs. 13, 14), with
the ventral sclerite knob-shaped and sur-
rounded by the hook-like lateral lobes of
the dorsal sclerite. The inner lobe of the
paraprocts is slender and slightly hook-like
(Fig. 14).
Material examined.— Paratype: Benton
Co., Oregon, Rock Creek, Corvallis, 9 IV
1935, K. Grey, 1 M (INHS).
Other specimens: CANADA: British Co-
lumbia, Qualicum Bay, 15 V 1955, G. E.
Shewell, 1 M, 11 F(CNC). CALIFORNIA:
Humboldt Co., Willow Cr., Hwy 299, below
Berry summit, 24 TV 1987, R. W. Baumann,
C. R. Nelson, B. P. Stark & S. A. Wells, 6
M, 2 F (BYU). OREGON: Clatsop, Vic.,
Gronnel Road, ca. 2 mi. E Elsie, 1 IV 1967,
S. G. Jewett, Jr., 7 M, 3 F (INHS); Douglas
Co., 10 mi. E Gardner, 23 IV 1964, Vertrees
& Schuh, 3 M, 13 F (USNM); Multnomah
Co., Johnson Cr., trib. near SE 82 Ave., 8
III 1987, G. R. Fiala, 7 M (BYU).
Ostrocerca prolongata (Claassen)
Nemoura prolongata Claassen, 1923: 289.
Type locality: Bretton Woods, New
Hampshire.
Ostrocerca prolongata, Mlies, 1966: 218.
The complex ventral sclerite is charac-
terized by two recurved erect processes and
distally terminating in an elongate process
(Figs. 15-18). The dorsal sclerite terminates
into diverging lobes (Fig. 17). The inner
lobes of the paraprocts are long and apically
hooked (Figs. 19-21). The cerci are long and
curved.
262 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 9-12. Ostrocerca dimicki. 9, Terminalia, dorsal view, 163. 10, Epiproct, dorsal view, 462. 11,
Ventral sclerite, lateral view, 356. 12, Terminalia, ventral view, 120x. Terms: hp, hypoproct; vs, ventral
sclerite; c, cerci; v, vesicle
The allotype female is badly damaged and — subgenital plate (as illustrated), and vena-
the terminalia are missing. As Hitchcock tion is very similar to the sympatric Soye-
(1974) points out, the female illustrated by — dina washingtoni (Claassen). Females found
Claassen (1923) may be misidentified. The in collections with male O. prolongata ex-
VOLUME 91, NUMBER 2
263
Figs. 13-14. Ostrocerca foersteri. 13, Terminalia, dorsal view, 163. 14, Terminalia, dorsal view, 462.
Terms: vs, ventral sclerite; il, inner lobe; c, cerci.
amined during this study were similar to
females of O. complexa and O. albidipennis.
And, because Ostrocerca prolongata was
usually collected with the other two species,
made positive association impossible. This
species appears to be rare throughout its
range.
Material examined.— Holotype M: Bret-
ton Woods, New Hampshire, C. W. John-
son, 26 VI 1913 (MCZ). Paratype: Orono,
Maine, | M (Cornell #1200). Allotype F:
Orono, Maine (Cornell #1200).
Other specimens; NEW HAMPSHIRE:
Huntington Ravine, 4000’, 15 VIII 1935,
C. P. Alexander, 1 M (USNM). NEW
YORK: Heart Lake, Adirondacks, 2100’,
10 VII 1938, C. P. Alexander, 1 M(USNM).
MAINE: on dropnet, no locality, 4-14 VI
1970, D. E. Leonard, 5 M (USNM). VIR-
GINIA: Nelson Co., small tributary of South
Fork Tye River, Rt., 687, 3 V 1980, BCK,
2M (VPI). WEST VIRGINIA: Tucker Co.,
Red Creek at Laneville, 26, 27 May 1973,
O. S. Flint, 2 M (USNM).
Ostrocerca truncata (Claassen)
Nemoura truncata Claassen, 1923: 290.
Type locality, Walden, Massachusetts.
Ostrocerca truncata, Mllies, 1966: 218.
This species is easily recognized by the
bell-shaped ventral sclerite of the epiproct
and thickened, short cerci (Figs. 22 and 23).
The slender diverging inner lobes of the
paraprocts are also distinctive (Figs. 24-25).
The middorsal groove terminating in two
sharp projections was not illustrated by
Claassen (1923) (same illustrations used by
Needham and Claassen, 1925) and Hitch-
cock (1974). Baumann (1975) illustration
(Fig. 76) does not indicate the extent of these
projections.
The female has been illustrated by Bau-
mann (1975), Hitchcock (1974) and Ricker
264
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 15-18.
17, Terminalia, lateral view, 156 =
ventral sclerite; c, cerci.
(1965). This species was once considered
the rarest of the Ostrocerca. But, we found
it to be common and widespread. The AI-
abama collection represents the southern-
most extension of its range and is a new
state record. The holotype of Nemoura trun-
cata could not be located at the Museum of
Comparative Zoology (C. Vogt, personal
communication).
Material examined.—ALABAMA:
DeKalb Co., spring seep, DeSoto State Park,
River Trail, cabins 8 & 9, 17 V 1988, RFK
& BCK, 1 F(BCK). CONNECTICUT: Bea-
con Falls, Naugatuck St. Forest, 17 VI 1961,
S. W. Hitchcock, 1 M (USNM); Portland,
20 VI 1963, S. W. Hitchcock, 1 M (USNM);
Ostrocerca prolongata. 15, Terminalia, dorsal view, 170 x. 16, Process of ventral sclerite, 655 x.
. 18, Ventral sclerite, bifurcate process, 356 x. Terms: ds, dorsal sclerite; vs,
Newtown, 10 V 1962, S. W. Hitchcock, 3
M, 2 F (USNM). KENTUCKY: Bell Co.,
Stream at Pinnacle Rd. entrance, Cumber-
land Gap National Park, 1OTV 1971, R.A.
Haick & C. R. Haick, 4 M, 8 N (USNM);
Lawrence Co., Bridge Hollow of Hood Cr.,
1.5 mi. S of Blaine Rt. 201, 20 IV 1984,
RFK, 10 M (RFK). MASSACHUSETTS:
Whately, Whately Glen, 18 V 1938, J. F.
Hanson, | M (USNM); OHIO: Athens Co.,
Canaan Township, 25 IV 1941, J. Walker,
1 M(INHS). PENNSYLVANIA: York Co.,
spring seepage, Davidsburg, 5 mi. NW
Woodland, 20 V 1962, P. J. Spangler, 12
M, 22 F, 12 N(USNM). VIRGINIA: Dick-
enson Co., spring seep into Hunts Creek,
VOLUME 91, NUMBER 2 265
Figs. 19-21. Ostrocerca prolongata. 19, Terminalia, lateral view, 137 x. 20, Terminalia, ventral view, 106 x.
21, Inner lobe of paraproct, 573 x. Terms: hp, hypoproct; vs, ventral sclerite; il, inner lobe: c, cerci: v, vesicle.
Rt. 80, Breaks Interstate Park, 29 April M, 14 F (RFK); Washington Co., spring to
1987, RFK, BCK, 6 F (BCK); Tazwell Co., Brumley Cr., Clinch Mtn. at Low Gap,
Station Springs Cr., Burkes Garden 3840’, 3600’, Rt. 690, 23 V 1981, RFK & BCK,
off Rt. 666, 20 V 1982, RFK & BCK, 48 15 M, 8 F (RFK); Wythe Co., Stony Fk. of
266 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 22-25. Ostrocerca truncata. 22, Terminalia, dorsal view, 178 x. 23, Epiproct, dorsal view, 263 x. 24,
Terminalia, ventral view, 120. 25, Terminalia, ventral view, 356. Terms: hp, hypoproct; ds, dorsal sclerite;
vs, ventral sclerite: il, inner lobe; ol, outer lobe; c, cerci; v, vesicle.
VOLUME 91, NUMBER 2
Reed Cr., Dark Horse Hollow Rec. Area,
Jefferson Nat. For., 29 IV 1978, RFK, 1 M
(RFK). WEST VIRGINIA: Cabell Co.,
Lusher Hollow of Mud River, near Bar-
boursville, US-60, 29-IV-1979, RFK, 4 M,
2 F (RFK); same locality 28 IV 1978, 4 M,
1 N (RFK); Greenbrier Co., Coats Run, Rt.
39, 26 IV 1987, RFK, BCK, 2 M (RFK);
Logan Co., Trace Fork of Copperas Mine
Fork, Holden, 15 TV 1973, RFK, | F(RFK);
Pendleton Co., Spruce Knob, 4 mi. below
summit, 16 V 1963, W D. Field & O. S.
Flint, | M, | F (USNM): Pocahontas Co.,
Hills Cr., 15 mi. E of Richwood, Rt. 39,
Monongahela NF, 20 V 1978, RFK, 2 F
(RFK); East Fork Greenbrier River, Forest
Rd., off Rt. 28, 26 IV 1987, RFK & BCK,
2 M (BCK); Tucker Co., Red Creek near
Laneville, 26, 27 VI 1973, O. S. Flint, 1 M,
3 F (USNM).
DISCUSSION
Examination of the males of Ostrocerca
with SEM indicated that no taxonomic
changes were necessary. However, the higher
magnification and greater depth of field al-
lowed for the first time a complete study of
the male terminalia of all six species. The
SEM micrographs elucidated structures of
the epiproct previously not illustrated. For
example, the distinctive fork-like ventral
sclerite of O. complexa easily distinguishes
this species; the serrate apex of the hypo-
proct is only found in O. alhidipennis; and
the ventral sclerite terminating in an elon-
gate process is characteristic of O. prolon-
gata. Consulting these micrographs will
make it easier to identify male specimens
using a dissecting microscope by confirming
characters not illustrated or mentioned in
original descriptions or redescriptions.
ACKNOWLEDGMENTS
We thank Dr. Robert E. Lee, Department
of Anatomy and Neurobiology, Colorado
State University, for his kind assistance with
the preparation of specimens and SEM, and
Dr. David Walter, United States Depart-
267
ment of Agriculture, Agriculture Research
Service, Orlando, Florida for his sugges-
tions and help. Dr. Richard W. Baumann,
Brigham Young University provided valu-
able comments on an earlier draft of this
manuscript. A Colorado State University
Electron Microscopy Seed Grant supported
this study. The views of the authors do not
purport to reflect the position of the De-
partment of the Army or the Department
of Defense.
LITERATURE CITED
Baumann, R. W. 1975. Revision of the stonefly fam-
ily Nemouridae (Plecoptera): A study of the world
fauna at the generic level. Smithsonian Contrib.
Zool. 211. 74 pp.
Claassen, P. W. 1923. New species of North Amer-
ican Plecoptera. Can. Entomol. 55: 257-263, 281-
292.
1937. New species of stoneflies (Plecoptera).
J. Kansas Entomol. Soc. 10: 42-51.
Frison, T. H. 1936. Some new species of stoneflies
from Oregon (Plecoptera). Ann. Entomol. Soc.
Amer. 29: 256-265.
Hitchcock, S. W. 1974. Guide to the insects of Con-
necticut: Part VII. The Plecoptera or stoneflies of
Connecticut. Bull. State Geol. Nat. Hist. Surv.
Conn. Bull. 107. 262 pp.
Illes, J. 1966. Katalog der Rezenten Plecoptera. Das
Tierreich, 82. Walter de Gruyter and Co., Berlin.
632 pp.
Kondratieff, B. C. and R. F. Kirchner. 1984. A new
species of Nemouridae (Plecoptera) from the Great
Dismal Swamp, Virginia, USA. Proc, Entomol.
Soc. Wash. 86: 578-581.
. 1987. Additions, taxonomic corrections, and
faunal affinities of the stoneflies (Plecoptera) of
Virginia, USA. Proc. Entomol. Soc. Wash. 89: 24—
30.
Needham, J. G. and P. W. Claassen. 1925. A mono-
graph of the Plecoptera or stoneflies of America
north of Mexico. Thomas Say Found. Entomol.
Soc. Amer. 2. 397 pp.
Nelson, C. R. and R. W. Baumann. 1987. Scanning
electron microscopy for the study of the winter
stonefly genus Capnia (Plecoptera: Capniidae).
Proc. Entomol. Soc. Wash. 89: 51-56.
Ricker, W.E. 1943. Stoneflies of southwestern British
Columbia. Indiana Univ. Publ., Sci. Ser. 12. 145
pp.
1952. Systematic studies in Plecoptera. In-
diana Univ. Publ., Sci. Ser. 18. 200 pp.
1965. New records and descriptions of Ple-
268 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
coptera (Class: Insecta). J. Fish. Res. Bd. Can. 22: stoneflies (Plecoptera) of West Virginia. Entomol.
475-S01. News 91: 49-53.
Stark, B. P., S. W. Szczytko, and R. W. Baumann. Walker, F. 1852. Catalogue of the specimens of neu-
1986. North American stoneflies (Plecoptera): ropterous insects in the collection of the British
systematics, distribution, and taxonomic refer- Museum. Part I. Phryganides-Perlides. London:
ences. Great Basin Natur. 46: 383-397. British Museum. 192 pp.
Tarter, D. C. and R. F. Kirchner. 1980. List of the
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 269-285
KEY TO THE KALOTERMITIDAE OF EASTERN UNITED STATES
WITH A NEW NEOTERMES FROM FLORIDA (ISOPTERA)
Davip A. NICKLE AND MARGARET S. COLLINS
(DAN) Systematic Entomology Laboratory, BBII, Agricultural Research Service, %
National Museum of Natural History, Washington, D.C. 20560; (MSC) Research Asso-
ciate, Smithsonian Institution, National Museum of Natural History, Washington, D.C.
20560.
Abstract. —A new species of drywood termite of the genus Neotermes is described and
figured, and an identification key is included to separate it from nine other kalotermitids
from Florida. This species was first recognized as distinct from another morphologically
very similar species, Neotermes jouteli (Banks), on the basis of karyotypic and isozymatic
differences.
Key Words:
Termites resembling Neotermes jouteli
(Banks and Snyder) 1920 but differing from
that species in chromosome number and
isozyme patterns were recently noted by Dr.
Peter Luykx during the course of a cytogenic
survey of species of the family Kalotermi-
tidae. We were requested to seek, and here-
with report, morphometric features for
characterizing this new taxon to make the
name available for his research. Although
we are confident that most of the termite
species of the United States have been de-
scribed, we find that refinements in taxo-
nomic techniques are beginning to uncover
complexes among already described species
(Haverty and Thorne 1989, in press). When
chromosomal, isozymatic, or behavioral
studies are used in termite investigations,
more species may be discovered in North
America.
Samples were preserved in 85% ethanol;
morphological features were measured with
the device described in detail by Grant
(1965). Measurements consisted of the fol-
lowing values in millimeters: head length,
in lateral view, the distance from the left
ventral mandibular condyle to the occiput;
Neotermes, Kalotermitidae, termites
head width, in dorsal view, the greatest
breadth at the genae; head depth, in lateral
view, the distance between the vertex and
the ventral margin of the head capsule; eve
diameter, the width of the eye of the imago
measured from its anterior to posterior
margin; postocular distance, in dorsal view
on the imago, the shortest distance from the
midpoint of the occipital margin of the head
to the imaginary line connecting the pos-
terior margins of the compound eyes; sub-
ocular distance, in lateral view, the shortest
distance connecting the ventral margin of a
compound eye with the ventral margin of
the head capsule; ocellus length and width,
the greatest and shortest diameters, respec-
tively, of the ocellus; postmentum length,
the distance from the midpoint of the labial
suture to the midpoint of the inner posterior
margin of the postmentum; postmentum
width, the minimum width of the postmen-
tum at its constriction; /ength and width of
pronotum, in dorsal view, the medial length
and greatest width of the pronotum; /ength
and width of wing, the total length of the
forewing, including the scale, and the great-
est width of the forewing; total length, in
270
lateral view, the length of the body from the
tip of the face to the apex of the abdomen;
length of the mandible, in dorsal view, the
length of the soldier mandible measured
from the notch at the base of the outer man-
dibular condyle to the tip of the mandible
(Tables 2, 4, 5).
Morphometric ratios found to have some
value in identifying Florida kalotermitids
included the following: Head L/W, the ratio
of head length to head width; Head L/D,
the ratio of head length to head depth; Head
WD, the ratio of head width to head depth;
Postocular distance/Eye diameter and Sub-
ocular distance/Eye diameter; Pronotum
W/L, the ratio of the greatest width to medial
length of the pronotum; Wing L/W, the ra-
tio of the wing length to wing width; Wing
L/Wingscale L, and Postmentum L/W (Ta-
bles 3, 6, 7).
Except for a previously unidentified sol-
dier and nymph in the National Museum
Collection that had been collected by E. M.
Miller in 1930, all of the material upon
which this description 1s based was collected
in south Florida by Peter Luykx. Specimens
used in this study have been deposited in
the following institutions: National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington, D.C. [NMNH], and
the American Museum of Natural History,
New York, NY [AMNH].
DESCRIPTION
Neotermes luykxi Nickle and Collins
New SPECIES
Holotype.—Short-headed soldier. Flori-
da: Broward Co. Dania. PL 389C. 30 June
1984. (Coll. P. Luykx). [NMNH]. Morpho-
type. Imago. Florida: Broward Co. Holly-
wood. PL 558. 21 August 1986. (Coll. P.
Luykx). [NMNH].
Paratype <evies.—26 soldiers, 15 ima-
goes, 20 nymphs. Florida: Broward Co.,
Hollywood, PL 439, (P. Luykx), 6 soldiers,
5 imagoes, 4 nymphs [NMNH]; Same lo-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. A list of chromosome numbers (2n) for
Florida species of Kalotermitidae (Luykx and Syren
1979, Luykx pers. comm.).
Species Chromosome Number
29 (6)
37 (8), 36 (2)
Calcaritermes nearcticus
Cryptotermes brevis
Cryptotermes cavifrons 40
Incisitermes milleri 28
Incisitermes schwarzi 32
Incisitermes snyderi 44
Kalotermes approximatus 32
Neotermes castaneus 38
Neotermes luykxi 45 (6), 44 (2)
Neotermes jouteli 56
cality, PL 558 [morphotype sample], VIII-
21-1986, 7 soldiers, 10 imagoes, 3 nymphs
[NMNH, AMNH]; Florida: Broward Co.,
Dania, PL 389, VI-20-1984, (P. Luykx), 3
soldiers, 3 nymphs [NMNH]; PL 389C,
same locality as PL 389, VI1984, 3 soldiers,
3 nymphs [NMNH]; Same locality, PL 400,
VII-21-1984, (P. Luykx), 6 soldiers, 6
nymphs [NMNH, AMNH]; Florida: Dade
Co., Key Largo, XII-6-1930, (Coll. E. M.
Miller), | soldier, | nymph [NMNH].
Short-Headed soldier.—Head: 1.21 x
longer than wide, 1.46 longer than deep
(Figs. 1E, 2E). From above, sides of head
weakly convex, with greatest width at mid-
dle of head capsule. Labrum about as long
as wide, broadly rounded. Antennal fossa
oval; dorsal ridge well developed, extending
over first article of antenna; ventral ridge
weakly developed, lying flush with head
capsule; from above, antennal fossa extend-
ing as a ridge to dorsal margin of clypeus.
Antennae with 13 articles (rarely with 14),
beadlike; article | broad, barrel shaped,
equal in length to articles 2+3; article 2
smaller than 4; article 3 enlarged, subequal
in length to 4+ 5; each successive article 4
through !3 similar in size, but becoming
increasingly more elongated toward apex of
antenna. Eyes comprised of a cluster of cells
with central core darkly pigmented; eye
271
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VOLUME 91, NUMBER 2
273
Table 4. TERMINTE LONG-HEADED SOLDIERS: means (x) and ranges of measurements (in mm) of
morphological features of Neotermes and Incisitermes species from Florida. Numbers in parentheses after value
indicate size of sample. NA = not available for measurement.
peered Sa Nii Ga + achue es Gane
# antennal articles Range 16 (2) 15-16 (6) NA 12 (2) 13-15 (8) 11 (1)
Head length x 3.75 (10) 3.24 (6) 3.14 (4) 2.41 (2) 3.57 (8) 2.00 (2)
Range 3.50-4.25 3.04-3.42 3.00-3.42 2.37-2.46 3.21-3.92 2.00
Head width x 2.85 (10) 2.30 (6) 2.00 (4) 1.48 (2) 2.13 (8) 1.12 (2)
Range 2.67-3.04 2.04-2.46 1.96-2.04 1.46-1.50 1.96-2.21 iN 2
Head depth ae 1.96 (10) 1.67 (6) 1.49 (4) 1.23 (2) 1.56 (8) 1.02 (2)
Range 1.87-2.21 1.62-1.79 1.46-1.54 1.21-1.25 1.42-1.67 —_1.00-1.04
Mandible length Xx 2.25 (10) 1.84 (6) 1.62 (4) 1.37 (2) 1.71 (8) 1.06 (2)
Range 272537 1.58-2.21 1.58-1.67 1.37 1.67-1.75 1.04-1.08
Postmentum length X 2.62 (10) 2.38 (6) 2.34 (4) 1.75 (2) 2.75 (8) 1.54 (1)
Range 2.12-3.08 2.04-2.58 2.12-2.50 1.75 2.42-3.12 1.54
Postmentum width {x 0.41 (10) 0.48 (6) 0.33 (4) 0.29 (2) 0.28 (8) 0.19 (2)
Range 0.37-0.46 0.25-0.58 0.29-0.37 0.29 0.25-0.37. 0.17-0.21
Width pronotum x 2.59 (10) 2.50 (6) 2.16 (4) 1.44 (2) 2.33 (8) 1.08 (1)
Range 2.46-2.87 2.29-2.71 2.12-2.25 1.42-1.46 2.17-2.50 1.08
Length pronotum x 1.14 (10) 1.21 (6) 1.08 (4) 0.89 (2) 1.23 (8) 0.79 (1)
Range 1.04-1.25 1.17-1.29 1.04-1.17 0.83-0.90 — 1.12-1.33 0.79
Total length x 11.09 (8) 9.68 (6) NA NA 10.72 (8) 6.83 (1)
Range 10.67-11.58 8.75-10.75 10.08-11.25
cluster situated a distance of less than di-
ameter of cluster from antennal fossa. Man-
dibles 74 as long as head capsule; in lateral
view, curving upward anteriorly; mandible
dentition and shape of postmentum as in
Figs VE:
Thorax: Pronotum broader than head
capsule. Anterior margin broadly concave
but not incised; posterior margin weakly
notched medially. Range of greatest width/
medial length 2.04-2.37.
Legs: Femora inflated, 2.3-2.5 = longer
than wide. Tibial spurs 3:3:3. Arolia absent.
Color: Mandibles glossy black. Head cap-
sule reddish-brown anteriorly, becoming
more yellowish-brown along posterior mar-
gin. Antennae light reddish-brown, darker
basally. Labrum reddish-brown; clypeus
translucent or whitish. Eyes purplish-gray.
Body and legs whitish-yellow.
Imago morphotype.—Head: In dorsal
view, head length from base of mandibles
0.78-0.83 x as long as head width just be-
hind compound eyes. Compound eyes 0).35—
0.39 mm in diameter; subocular distance
ca. 0.62 x diameter of compound eye; post-
ocular distance ca. 1.78 x diameter of com-
pound eye. Compound eye separated from
antennal socket by less than 0.1 mm. Ocelli
oval, with greatest diameter 0.15 mm: sep-
arated from compound eye by less than
ocellar diameter. Antenna with 18 articles,
2.1-2.3 mm in length, ca. 1.7 x longer than
head length to base of mandibles.
Thorax: Pronotum 1.87 broader than
long; anterior margin broadly concave; pos-
terior margin medially notched.
Legs: Femora slightly enlarged or inflat-
ed, 2.6-2.8 x longer than wide. Tibial spurs
3:3:3. Arolia present.
Wings: Hyaline, yellowish with brown
anterior veins. Venation as in all Neotermes,
with M nearly as sclerotized as Rs, running
closer to Rs than to Cu. Length of forewing
9.9-10.8 mm (xX = 10.3 mm), 3.77 x longer
than its greatest width (Fig. 3B).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
274
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VOLUME 91, NUMBER 2
275
Table 6. Ratios of morphological features of long-headed soldier termites from Florida. Numbers in paren-
theses after value indicate size of sample. NA = not available for measurement.
Neotermes Neotermes Neotermes Incisitermes Incisitermes Incisitermes
castaneus jouteli luykxi snyder schwarz millert
Head L/W Xx 1.32 (10) 1.42 (6) 1.57 (4) 1.63 (2) 1.78 (8) 1.78 (2)
Range 1.22-1.41 = -1.27-1.65 -1.50-1.67 —1.63-1.64 —_1.60-1.84 1.78
Head L/D Xs 1.91 (10) 1.94 (6) 2.12 (4) 1.96 (2) 2.29 (8) 1.89 (2)
Range 1.83-2.09 1.82-2.08 2.06-2.21 1.96-1.97 2.22-2.46 —1.78-2.00
Head W/D x 1.45 (10) 1.37 (6) 1.35 (4) 1.20 (2) 1.37 (8) 1.10 (2)
Range 1.38-1.51 = -1.26-1.45) -1.32-1.37)) -1.20-1.21 = -1.27-1.43—-1.08-1.12
Head L/Mandible L 38 1.67 (10) 1.77 (6) 1.93 (4) 1.76 (2) 2.08 (8) 1.88 (2)
Range 1.53-1.89 1.55-2.13 1.89-2.05 1.73-1.79 = 1.88-2.35. 1.85-1.92
Pronotum W/L xX 2.28 (10) 2.07 (6) 2.00 (4) 1.61 (2) 1.89 (8) 137\(2)
Range 2.03-2.44 = 1.83-2.21 = 1.93-2.04 -1.48-1.75 —-1.74-2.00 E37,
Postment. L/min. W xX 6.41 (10) 4.62 (5) 7.09 (4) 6.00 (2) 8.04 (8) 8.32 (2)
Range 5.18-7.11 3.77-6.02 6.37-8.57 6.00 6.44-9.37 7.40-9.25
Color: Head, pronotum, legs, and tergites
of abdomen uniformly yellowish; clypeus
whitish-yellow; labrum same color as head
capsule. Sternites lighter yellow than ter-
gites.
Variation: Twenty-two soldier paratypes
are short-headed forms and four are long-
headed forms (Figs. 1B, E and 2D, E). Al-
though all have pigmented eyespots, the de-
gree of pigmentation varies from light to
dark grey, and in some specimens they ap-
pear purplish. Both soldier forms vary in
size (Tables 3, 4). Although there are no
overlaps in most measurements between
soldier forms, the largest short-headed and
smallest long-headed soldiers share the same
values for head depth, postmentum width,
and pronotum width. Among morphomet-
ric ratios for identification of soldier forms,
only Head W/D overlaps in the two forms
(Tables 6, 7). Imago paratypes also vary in
size (Tables 2, 5).
Diagnosis. — Soldier: We recognize luykxi
as a species of Neotermes on the basis of
the shape of the pronotum, which is not
deeply incised as in species of /ncisitermes.
Of the Florida species of Neotermes, luykxi
and jouteli are more closely related and are
easily distinguished from castaneus (Bur-
meister 1839), a species with only one form
of soldier. Neotermes castaneus 1s larger, has
a slightly broader head as seen in dorsal
view (Fig. 2A—C), and has unpigmented eye-
spots. Mandible dentition may also help to
separate these species (Fig. 4B). Neotermes
luykxi and N. jouteli both have pigmented
eyespots and resemble one another in the
head shapes of long- and short-headed sol-
diers. The L/W ratio for short-headed sol-
diers of both N. jouteli and N. luykxiis 1.21;
for long-headed soldiers, the ratio is 1.42
for N. jouteli and 1.57 for N. luykxi. The
L/W ratio for N. castaneus 1s 1.32. The long-
headed soldier of N. /iykx7 usually has both
a narrower head and a longer, narrower
postmentum than N. jouteli. Although N.
luykxi is generally smaller than N. jouteli,
there is overlap for all measurements in sol-
diers of the same head form, and the mean
values reflect only trends for each species
(See Tables). The most significant difference
between the two species is chromosome
number (45[4] 44[2] for N. /uyAxi, 56 [both
sexes] for N. jouteli) (Table 1). The mor-
phometric difference that seems to be most
useful in differentiating these two species 1s
the ratio of the postmentum length to its
minimum width. For short-headed soldiers
this ratio is 4.41 for N. /uwykxi and 4.98 for
N. jouteli. For long-headed soldiers it 1s 7.09
276
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 7. Ratios of morphological features of short-headed soldier termites from Florida. Numbers in pa-
rentheses after value indicate size of sample available for measurement. NA = not available for measurement.
Neotermes Neotermes Incisitermes Incisitermes Incisitermes Kalotermes
youteli luykxi snyderi schwarz milleri approximatus
Head L/W x W211 (13) 1.21 (17) 1.46 (8) 1.23 (5) 1.80 (4) 1.62 (3)
Range 1.16-1.39 1.10-1.31 = 1.42-1.57 1.19-1.32 1.72-1.87 1.58-1.66
Head L/D x 1.77 (13) 1.72 (17) 1.78 (8) 1.68 (5) 2.11 (4) 1.91 (3)
Range =1.59-2.43—-1.60-1.84 =1.71-1.86 1.59-1.80 2.00-2.14 — 1.87-1.97
Head W/D x 1.42 (13) 1.42 (17) 1.21 (8) 1.37 (5) NA 1.16 (3)
Range =1.36-1.54 =1.31-1.52. 1.18-1.26 — 1.28-1.43 1.15-1.19
Head L/Mandible L x 1.53 (13) 1.51 (17) 1.53 (8) 1.44 (5) 1.94 (4) 1.73 (2)
Range =1.39-1.84 =-1.38-1.73 —-1.42-1.62.—_1.30-1.64 =-1.87-2.00 —_1.71-1.75
Pronotum W/L x 2.1'5' (13) 2.20 (17) 1.75 (8) 2.26 (5) 1.36 (4) 1.95 (2)
Range =-1.95-2.54 2.04-2.37. -1.55-1.88 2.05-2.44 = 1.25-1.47 —_1.91-2.00
Postmentum L/min. W ¥ 4.98 (13) 4.41 (17) 5.57 (8) 4.33 (5) 7.17 (4) 6.70 (3)
Range 4.22-5.83 3.71-6.14 4.71-6.80 4.25-4.62 5.67-8.00 5.85-7.67
for N. luvkxi, 4.62 for N. jouteli, and 6.41
for N. castaneus. Neotermes luykxi and N.
jouteli also differ in isozyme composition
(Luykx et al., in prep.).
Imago: Assignment of the imago of N.
luykxi to Neotermes was made on the basis
of wing venation and length of the anterior
margin of the second marginal tooth of the
left mandible. Differentiating the three Flor-
ida species of Neotermes depends primarily
upon a comparison ofall morphometric fea-
tures listed in Tables 2 and 5, none of which
by itself is useful in identifying an individ-
ual specimen. Neotermes luykxi is the
smallest of the three species, however, and
the diameter of its compound eye is con-
sistently smaller. The presence of long setae
on the head and pronotum is a diagnostic
character; N. castaneus generally has long
setae on the head and pronotum, while N.
jouteli and N. luykxi have only scattered
short setae (Fig. 6A-I). Wing length, wing
scale length, and head width are generally
largest for N. castaneus and smallest for N.
luykxi (Tables 2, 5).
Discussion.— Morphologically, Neoter-
mes jouteli is most similar to this new
species, but Luykx et al. (in prep.) found
that the chromosome number (2n) of N. jou-
teli was 56 [both sexes] and only 45 [é] and
44 [9] for N. /uykxi. Luykx also found in-
terspecific differences in a series of isozymes
among five selected kalotermitids (includ-
ing N. jouteli and N. luykxi).
It has become increasingly common to
identify sibling species of animals by using
chromosomal differences. Once phena have
been segregated on the basis of such differ-
ences, it is usually possible to identify mor-
phological characters which are helpful in
distinguishing the forms. Such was the case
with a European mole cricket, Grvllotalpa
septemdecimchromosomica Ortiz (1958),
and more recently with a complex of other
sibling species of Grylletalpa: quindecim,
sedecim, octodecim, and viginti (Baccetti and
—
Fig. |. Termite soldiers, head and pronotum, dorsal view. Also figured beside each soldier head: postmentum,
ventral view. A, Neotermes jouteli, long-head (LH); B, N. luykxi, LH; C, N. castaneus; D, N. jouteli, short-head
(SH); E, NV. /uykxi, SH; F, Kalotermes approximatus, G, Calcaritermes nearcticus, H, Incisitermes snydert; I, I.
schwarzi, LH; J, I. schwarzi, SH; K, I. milleri, LH; L, I. milleri, SH.
VOLUME 91, NUMBER 2
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277
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
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= a, —_——_x,
( . . ee ane oe
( SE oh— ( on aN
I Nets r | Sey 2 K Se | Pees ie
I. snyderi 1. milleri LH milleri SH C. nearcticus
Fig. 2. Termite soldiers, head profile. A, Neotermes castaneus; B, N. jouteli, long-head (LH); C, N. jouteli,
short-head (SH); D, N. /uvkxi, LH; E, N. luvkxi, SH; F, Incisitermes schwarzi, LH; G, I. schwarzi, SH; H,
Kalotermes approximatus, 1, I. snyderi; J, I. milleri, LH; K, I. milleri, SH; L, Calcaritermes nearcticus.
Capra 1978). These species, otherwise mor-
phologically very similar to Gryllotalpa
gryllotalpa (L.), have chromosome comple-
ments of 17, 15, 16, 18, and 14, respec-
tively, compared with 12 for G. gryllotalpa.
Once it was determined that different chro-
mosomal numbers occurred within the
complex, it was possible to establish mor-
phometric parameters of the variable char-
acters for each species.
As in Incisitermes milleri (Emerson), J.
snyderi (Light), I. schwarzi (Banks), and
Neotermes jouteli, N. luykxi soldiers occur
as long-headed or as short-headed forms.
Short-headed forms are usually more com-
mon than long-headed ones within most
samples we have seen. The different head
forms probably result from differences in
the stage of nymphal development when
soldier differentiation occurs. Grassé and
Noirot (1958) demonstrated that young
Kalotermes flavicollis (F.) colonies devel-
oped soldiers from 2nd and 3rd instar lar-
vae, while older colonies developed soldiers
from 4th and Sth instar larvae.
There are five genera of drywood termites
in the eastern United States: Calcaritermes
(nearcticus (Snyder 1933)), Cryptotermes
(cavifrons Banks 1906 and brevis (Walker
1853)), Incisitermes (snyderi, schwarzi, and
milleri), Kalotermes (approximatus Banks
and Snyder 1920), and Neotermes (casta-
neus, jouteli, and the new species /uykxi).
All of these species are known to occur in
Florida.
Kalotermes approximatus 1s not usually
economically important, except in rare cases
when it becomes locally abundant. It is gen-
erally associated with dead wood in sand
dunes in northern Florida, Alabama, Mis-
sissippi, and Louisiana, and northward along
the coastal plain into southern Virginia.
Weesner (1965) reported this species in
homes in Waco, Texas, in 1957 and 1958
VOLUME 91, NUMBER 2
N. jouteli
N. castaneus Si We =
I. schwarzl —~ S —
Fig. 3.
279
B
N. luykxi
C. cavifrons
ee N
RK 3 )
: i: caiderl
AR a
NS
\
le
Wings of alates of kalotermitids. A, Neotermes jouteli, B, N. luykxi, C, N. castaneus; D, Incisitermes
schwarz; E, I. snydert, F, Cryptotermes cavifrons, G, Calcaritermes nearcticus. (M = medius vein.)
according to National Pest Control Asso-
ciation Records, but we have no specimens
to confirm its presence in that state.
The two genera of kalotermitids most
likely to be encountered in southern Florida
are Neotermes and Incisitermes. Neotermes
jouteli occurs in southern Florida, Jamaica,
the West Indies, and Mexico. Neotermes
castaneus occurs in southern Florida, West
Indies, Central and South America. Neo-
termes luykxi 1s presently known only from
Monroe, Dade, ard Broward Counties,
Florida. Neotermes species tend to be more
moisture-dependent than those of J/ncisi-
termes (Collins 1969). Neotermes jouteli may
enter man-made structures, infesting wet
wood or wood that becomes wet from time
to time. Neotermes castaneus also has been
280 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1OKU WO11MM = =©$:88109 P:e0006
16KY WO: 8MM $:00107 P:60001
eA 1@KV WO:16MM 8 $:63629 P: 68886
Fig. 4. Scanning electron micrographs of morphological features of kalotermitid termites. A, Calcaritermes
nearcticus, apex of left foretibia, lateral aspect. B, Neotermes castaneus soldier, left mandible, dorsal aspect. C,
D. Neotermes castaneus, imago mandibles: C, left, D, right. E, F. Cryptotermes brevis, imago mandibles: E, left,
F, right.
recorded as a pest of avocado, guava, and
citrus trees in south Florida and other parts
of its range, attacking dead wood of injuries
and then invading adjacent living parts of
the trees (Miller 1949, Araujo 1970).
Incisitermes snyderi is widespread along
the Coastal Plain from Texas to Florida and
northward to South Carolina and in Central
America and several islands in the Carib-
bean. It occurs commonly in dead cypress
(Taxodium spp.) in the United States, caus-
ing extensive damage to untreated wood and
VOLUME 91, NUMBER 2
Bigs:
Scanning electron micrographs of morphological features of kalotermitid termites. A, C, E. Crypto-
termes brevis, soldier: A, dorsal, C, lateral, E, frontal aspect. B, D, F. Cryptotermes cavifrons, soldier: B, dorsal,
D, lateral, F, frontal aspect
fence posts (Miller 1949). Incisitermes
schwarzi, found commonly in south Flori-
da, also occurs in eastern Mexico (Yucatan)
and throughout the West Indies. Like Neo-
termes jouteli, I. schwarzi is found in hab-
itats with more available moisture and
higher temperatures than J. snyderi. Incis-
itermes milleri is known only from the for-
ested areas of the Florida Keys and Jamaica
and is found in the sound wood of dead
stumps.
The remaining drywood termites in this
region all have soldiers with phragmotic
(stopper-like) heads. Ca/caritermes has nu-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
VOLUME 91, NUMBER 2
merous species in the Neotropics but only
a single species, nearcticus (Banks 1918), in
the United States. Calcaritermes nearcticus
283
Media (M) of forewing curving anteriorly and
joining radial sector at about two-thirds length
of wing from suture (Fig. 3F) [Cryptotermes] 5)
3 3 : : 3. Body length 7-8 mm, body dark brown
is found in natural habitats in northem _ Incisitermes milleri
Florida but has not yet been recorded from - Body length greater than 10 mm, but usually
man-made structures. In contrast, termites not exceeding 16 mm; body yellow to yellowish
A if : BIOWN 25.02. e 4
of the genus Cryptotermes are major pests. Aa Hea gacnete 15-16 dee peices seat eae
Cryptotermes cavifrons has been found iol length (including wing scale) greater than 10
natural habitats throughout peninsular mm (Fig. 3D); head shape as in Fig. 6M-O
Florida, islands of the Caribbean and Cen- _................ see. Incisitermes schwarzi
tral America (Araujo LOT): but Crypto- = Body jengili 10-12 mm; yellowish; wing length
termes brevis, the more important pest less ‘than 10'mmi (Fig. 3E); head shape asin
; ; 1 ON Vi oneuvaereecieveage as cite Incisitermes snyderi
species of the two, is known only from the 5 Back ieneth 212 meme head widih behind
dry wood of man-made structures in the eyes greater than 1.0 mm; antenna usually with
United States and elsewhere in the world. 16-18 articles _ Cryptotermes brevis
It 1S acommon household pest from North = Body length less than 10 mm, head width less
Carolina through Florida to Texas and is than 1.0 mm; antenna usually with fewer than
: 5 ; 16 articles ........ Cryptotermes cavifrons
frequently found in furniture, picture frames, 6. M vein of forewing unsclerotized, located mid-
and other wooden items that have been way between Rs and Cu; compound eye less
transported to northern states from areas of than 0.35 mm in diameter; ocellus less than
infestation in the South and other warm re- 0.12 mm in diameter; head length/width ratio
gions of the world. greater than 1.0; body sooty black; wings sults
: : : : er than body .... _. Kalotermes approximatus
The following identification key resolves — M vein of forewing nearly as sclerotized as Rs,
some of the difficulties in determining ter- located closer to Rs than to Cu; without other
mites from the eastern United States. above combination of characters . . : i
7. Rs and M of forewing close together, without
crossveins; Cu equidistant between M and anal
margin (Fig. 3G) Calcaritermes nearcticus
Key TO THE TEN SPECIES OF — Rsand M of forewing wider apart, with several
KALOTERMITIDAE OCCURRING IN crossveins extending from Rs to costal border;
EASTERN UNITED STATES Cu closer to M than to anal margin (Fig. 3C)
[Neotermes] . 8
IMAGOES 8. Body length 15- 16 mm; wing length preater
1. Anterior edge of second marginal tooth of left than 12 mm; wing dark brown; forewing with
mandible elongated, longer than straight line numerous crossveins between Rs and M (Fig.
transect of posterior edge of first marginal tooth 3C); body darkly pigmented; head and prono-
(Fig. 4E) Se eeaeye ING aves onsets NS oh as 2 tum usually with both long and short setae (Fig.
— Anterior edge of eeeend areal tooth of left 6A-C) Neotermes castaneus
mandible not elongated, subequal in length to - Body length less than 15 mm, wing length 9.0—
straight line transect of posterior edge of first 11.5 mm, wing clear or only slightly pigment-
marginal tooth (Fig. 4C) 6 ed; forewing with a few crossveins between Rs
2. Media (M) of forewing running midway be- and M (Fig. 3A, B); body golden or yellow;
tween radial sector (Rs) and cubitus (Cu) and head and pronotum with short setae only (Fig.
extending to apex of wing (Fig. 3D, E) [/ncis- 6D-I) ..... .. 9
itermes]| . 3. 9. Mean total body length 8. 11 mm (range 7.8-9.0
a2
Fig. 6. Scanning electron micrographs of morphological features of kalotermitid termites. A, B, C, Neotermes
castaneus, dorsal, dorso-lateral, lateral, respectively. D, E, F, Neotermes jouteli. G, H, 1,
K, L, Incisitermes snyderi. M, N, O, Incisitermes schwarzi.
Neotermes luykxi. J,
284
mm); mean head length 1.44 mm, mean head
width 1.66; eye diameter 0.4-0.5 mm
Be A pena eee raha ces ene Neotermes jouteli
Mean total body length usually less than 7.5
mm (range 6.50-—7.92 mm); mean head length
1.31 mm, mean head width 1.46 mm; eye di-
ameter less than 0.4 mm Neotermes luykxi
SOLDIERS
Li
to
Head phragmotic, with front of face rising
sharply at an angle of 50-90° to longitudinal
axis; mandibles small relative to size of head
capsule (Figs. 2L, SA-F)
Head not phragmotic, with front of face son
ing gradually at an angle less than 45° to lon-
gitudinal axis; mandibles well developed . . 4
One anterior tibial spur on foretibia enlarged,
well developed (Fig. 4A) a
NCE SACOM Calcaritermes nearcticus
Tibial spurs on foretibia subequal in length
[Cryptotermes] .
tN
. Dorsum of head capsule wealdy concave; sur-
face of forehead smooth or finely crenulated
(Fig. 5B, D, F) Cryptotermes cavifrons
Dorsum of head capsule distinctly concave;
surface of forehead deeply convoluted (Fig.
SAGE) . Cryptotermes brevis
. Anterior margin of pronotum neither deeply
concave nor incised. 5
Anterior margin of pronotum deeply c concave
or incised (Fig. | H-L) [/ncisitermes] 10
. Third antennal article only slightly longer than
fourth; hindfemur moderately inflated, with
L/W ratio greater than 3.0; head length greater
than 3.4 mm; head depth greater than 1.7 mm
Bid hh tandis, Mateos Gece Neotermes castaneus
Third antennal article subequal to or greater
than articles 4 plus 5; hindfemur strongly in-
flated, with L/W ratio less than 2.6; head length
less than 3.4 mm; head depth less than 1.5
mm
Eyes unpigmented ... . Kalotermes approximatus
Eyes pigmented, purplish, gray, or sooty black
oaseitntlans acted ise tone a. dae a tet ahe ee ene Sent 7
Postmentum length greater than 2.00 mm .— 8
Postmentum length less than 2.00 mm 2.
Mean ratio of length of postmentum to its
minimum width 7.09; head width 1.9-2.1 mm;
width of pronotum usually less than 2.3 mm
: Neotermes luykxi [long-headed]
Mean ratio of length of postmentum to its
minimum width 4.62; head width 2.0-2.5 mm;
width of pronotum usually greater than 2.3
mm . Neotermes jouteli [long-headed]
Head length ca. 2.3 mm (range: 2.0-2.9 mm);
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
head width ca. 1.9 mm (range: 1.7—2.3 mm);
head depth ca. 1.3 mm (range: 1.2—1.7 mm);
ratio of postmentum length to minimum width
usually greater than 4.7
Neotermes jouteli [short-headed]
Head length less than 2.1 mm (range: |.8—2.1
mm); head width less than 1.8 mm (range:
1.6-1.8 mm); head depth 1.2 mm (range: 1.0-
1.2 mm); ratio of postmentum length to min-
imum width usually less than 4.5
Neotermes luykxi [short-headed]
. Body length less than 6.5 mm; restricted to
Florida keys Incisitermes milleri
Body length greater than 6.5 mm; range not
limited to southern tip of Florida
. Head length greater than 3.0 mm; postmen-
tum length greater than 2.4 mm
Aaya eee Incisitermes schwarzi [long-headed]
Head length less than 2.6 mm; postmentum
length less than 1.9 mm
. Head length greater than 2.3 mm, and head
width less than 1.6 mm
Me etotrceei Incisitermes snyderi {long-headed]
Head length less than 2.3 mm, and head width
either less than 1.5 mm or greater than 1.6
mm
13
. Head width greater than 1.6 mm; third an-
tennal article usually longer than 4 and 5 com-
bined; head nearly as broad as long; head L/W
less than 1.40 (Fig. 1J); antenna usually with
16 articles . Incisitermes schwarzi [short-headed]
Head width less than 1.5 mm; third antennal
article no longer than 4 and 5 combined; head
longer than broad; head L/W greater than 1.40
(Fig. 1H); antenna usually with fewer than 16
articles Incisitermes snyderi [short-headed]
ACKNOWLEDGMENTS
We wish to thank the following individ-
uals for providing valuable insights and sug-
gestions for this manuscript: Dr. William B.
Nutting,
retired, University of Arizona,
Tucson, AZ, and Drs. Eric E. Grissell and
Manya B. Stoetzel, Systematic Entomology
Laboratory, BBI, USDA, % National Mu-
seum of Natural History, Smithsonian In-
stitution, Washington, DC. Finally, we are
indebted to Dr. Peter Luykx, University of
Miami, Coral Gables, FL, for the specimens
upon which this paper is based and for shar-
ing information regarding the biology of the
drywood termites of Florida.
VOLUME 91, NUMBER 2
LITERATURE CITED
Araujo, R. 1970. Termites of the neotropical region.
Chapter 12, pp. 527-571. Jn Krishna, K. and
Weesner, F. M., eds., Biology of Termites, vol. 2.
Academic Press, New York and London.
1977. Catalogo dos Isoptera do Novo Mun-
do. Academia Brasileira de Ciencias. Rio de Ja-
neiro. 92 pp.
Baccetti, B. and F. Capra. 1978. Notulae Orthopter-
ologicae. XXXIV. Le Specie Italiane del genere
Gryllotalpa L. Redia 61: 401-464.
Banks, N. 1906. Two new termites. Entomol. News
17: 336-337.
1918. The termites of Panama and British
Guiana. Bull. Am. Mus. Nat. Hist. 38: 659-667.
Banks, N. and T. E. Snyder. 1920. A revision of the
nearctic termites (Banks) with notes on biology
and geographic distribution (Snyder). U.S. Natl.
Mus. Bull. No. 108: 1-128.
Burmeister, H. C. C. 1839. Handbuch der Entomo-
logie, II(1): 757-1050. Berlin.
Collins, M. S. 1969. Water relations in termites.
Chapter 14, pp. 433-458. /n Krishna, K. and
Weesner, F. M., eds., Biology of Termites, vol. 1.
Academic Press, New York and London.
Grant, H. J., Jr. 1965. A measuring device for use in
insect systematics. Entomol. News 76: 249-251.
Grassé, P.-P. and C. Noirot. 1958. La société de Cal-
285
otermes flavicollis de la fondation au premier es-
saimage. Comp. Rev. 246: 1789-1795.
Haverty, M. I. and B. L. Thorne. 1989. (In press.)
Agonistic behaviour correlated with hydrocarbon
phenotypes in dampwood termites, Zootermopsis
(Isoptera: Termopsidae). Animal Behaviour.
Luykx, P., D. A. Nickle, and B. I. Crother. (In prep.)
A morphological, electrophoretic, and karyotypic
assessment of the phylogeny of some lower ter-
mites (Isoptera: Kalotermitidae): Lack of congru-
ence among different data sets.
Luykx, P. and R. M. Syren. 1979. The cytogenetics
of Incisitermes schwarzi and other Florida ter-
mites. Sociobiology 4: 191-209.
Miller, E. M. 1949. A Handbook on Florida Ter-
mites. Technical Series. University of Miami Press.
Miami. 30 pp.
Ortiz, E. 1958. El valor taxonomico de las llamadas
razas chromosomicas de Gryllotalpa gryllotalpa
(L.). Publ. Inst. Biol. Apl. Barcelona 27: 181-194.
Snyder, T. E. 1933. Calcaritermes in the United States.
Proc. Entomol. Soc. Wash. 35: 67-69.
Walker, F. 1853. List of the specimens of Neurop-
terous insects in the collections of the British Mu-
seum. Part 3, Termitides: 501-529. Trustees for
British Museum. London.
Weesner, F. M. 1965. The Termites of the United
States, A Handbook. The National Pest Control
Association. Elizabeth, NJ. 70 pp.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 286-287
NOTE
Rapid and Non-Destructive Gender Determination of
Nymphal and Adult Cryptocercus punctulatus Scudder
(Dictyoptera: Cryptocercidae)
The primitive xylophagous cockroach,
Cryptocercus punctulatus Scudder, is dis-
tributed throughout the eastern and north-
western United States. Closely allied to the
termites, family units of C. punctulatus live
within decaying logs (Nelepa. 1984. Behav.
Ecol. Sociobiol. 14: 273-279). Because of
its unique biology, C. punctulatus has be-
come a popular laboratory and field re-
search insect.
The gender of most cockroaches is easily
determined by the presence (males) or ab-
sence (females) of styles on the sub-genital
plate. In addition, the females of many blat-
tid species have divided sub-genital plates.
The Cryptocercidae, however, possess ex-
panded seventh tergal and sternal sclerites
that completely conceal abdominal seg-
ments 8-10, the ceici, and the male’s styles
(McKittrick. 1964. Cornell Univ. Agric.
Exp. Sta. Memoir 389. 197 pp.). Living C.
punctulatus tightly close their supra- and
sub-genital plates concealing the cerci and
styles. In addition, living males reportedly
have a distinctive odor and females have a
dorsal abdominal gland between tergites VI
and VII (Seelinger and Seelinger. 1983. Z.
Tierpsychol. 61: 315-333). Pinned speci-
mens have either closed genital plates or
slightly opened plates with shrunken or hid-
den styles. In such instances, gender deter-
minations have been based upon dissection.
Thus, confirmation of gender by the pres-
ence of styles is difficult and time consum-
ing. Herein, I describe a morphological
character of the subgenital plate that will
allow the rapid and non-destructive deter-
mination of gender of C. punctulatus.
Examination of living and preserved (n
= 43) female C. punctulatus revealed api-
colateral emarginations of the subgenital
plate and a subtruncate apical median
prominence (Fig. 1A). These characters were
readily visible on nymphs (second through
last instar) and adults. There were no emar-
ginations, but a narrowly rounded apical
median prominence on this area was pres-
ent in nymphal and adult male C. punctu-
latus (n = 54) (Fig. 1B). It 1s possible that
subgenital plate emarginations of Crypto-
cercidae were the precursor to the complete-
ly divided plates of the Blattidae. The gen-
der of a few (n = 4; 3.96%) preserved second
and third instar C. punctulatus nymphs
could not be identified by this character.
However, these nymphs were light brown
and not completely sclerotized. The entire
Fig. 1.
Subgenital plates of adult female (A) and
male (B) Cryptocercus punctulatus. Arrow indicates the
apicolateral emargination.
VOLUME 91, NUMBER 2
subgenital plate had shriveled, making the
apical area character ambiguous. With this
exception, the presence (female) or absence
(male) of an apicolateral emargination of
the subgenital plate is a diagnostic character
for gender determination in C. punctulatus.
I thank W. E. Clark for help with mor-
phological terminology and G. L. Miller for
287
the illustration. This is Alabama Agricul-
tural Experiment Station Journal Series No.
17-881833P.
Arthur G. Appel, Department of Ento-
mology and Alabama Agricultural Experi-
ment Station, Auburn University, Alabama
36849-5413.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 287-288
Sex and Deposition of the Holotype of
Bareogonalos canadensis (Harrington)
(Hymenoptera:
Prior to 1900 the yellowjacket parasitoid
Bareogonalos canadensis (Harrington) was
known only from the holotype collected in
1893 at Victoria, British Columbia (Har-
rington. 1896. Canad. Entomol. 28: 108),
and 27 specimens collected on Gabriola Is-
land in 1897 by Taylor (1898. Canad. Ento-
mol. 30: 14-15). The present deposition of
19 of these 28 specimens is known (Table
1). The holotype was reported lost by
Townes (1956. Proc. U.S. Nat. Mus. 106:
295-304) but Sarazin (1986. Canad. Ento-
mol. 118: 957-989) claimed that the holo-
type is in the Canadian National Collection
(CNC). From a study of the pre-1900 ma-
terial, I conclude that the true holotype is
in the Zoologisches Museum der Hum-
boldt-Universitaet (Berlin).
Harrington (1896) described the holotype
as a male. Taylor (1898) reversed the sexes
of his series and Harrington accepted this
mistake. Harrington erroneously stated that
the holotype was a female, and described
the “male” sex using three females (auto-
types) (Harrington. 1898. Canad. Entomol.
30: 15-16). Harrington’s error, though not-
ed by Schulz (1907a. Jn Wytsman, Genera
Insectorum 61: 24 pp.), has been perpetu-
ated by others, including Carlson (1979.
Trigonalyidae)
1197-1198. 7m Krombein et al. Catalog of
Hymenoptera in America north of Mexico.
Vol. I. Smithsonian: Wash., D.C. 1198 pp.)
and Sarazin (1986).
The deposition of all four males from
Taylor’s series is known (Table 1) but at the
Zoologisches Museum der Humboldt-
Universitaet (Berlin) a fifth male has labels
from Taylor’s series with a date (24 Oct
1897) on which only females were collected.
Schulz [1907b, (1906). Berl. Entomol. Zeit.
51: 303-333] said he received the “‘typisch-
en Parchen” (typical pair). This fifth male
fits the holotype description; apparently its
original labels have been switched with the
labels on one of the three autotypes to cor-
respond with Harrington’s 1898 statement
that the holotype is a female. The holotype
currently has labels that read: ““Gabriola Isd.
B.C., Taylor, 24-10-97 [Taylor’s handwrit-
ing]/Trigonalys canadensis Harrington,
Type ¢ [Harrington’s handwriting]/ Zool.
Mus., Berlin. [printed label].”’ The holotype
labels, now on the autotype, read: *‘ Victoria
V.I. [no date, Harrington’s handwriting]/
Trigonalys canadensis Harrington, Type 2
[Harrington’s handwriting]/Zool. Mus.,
Berlin. [printed label].
Two autotypes at the Canadian National
Collection (CNC, Ottawa), with locality and
288 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Specimens of Bareogonalos canadensis
collected from Gabriola Island, B.C., Canada, in Oc-
tober, 1897.
Date! Number and Sex! Deposition
21 Oct 3 at Canadian National Col-
lection (CNC) (1 with yel-
low paratype label’)
1 H. Townes, Gainesville,
Florida
9 females
22 Oct 3 females
3 males 2 at CNC
1 at Humboldt
1 at CNC
2 at CNC
1 at Humboldt
female at Humboldt with
Victoria label (no date)
actually is autotype from
this date*
(male at Humboldt labeled
with this date and locality
is holotype collected in
1893')
2 at CNC
H. Townes, Gainesville,
Florida
1 at CNC
1 at Washington State Uni-
versity
CNC (With red holotype
label’)
23 Oct
24 Oct
2 females
4 females
25 Oct 2 females
1 male
26 Oct 2 females
27 Oct 1 female
' This information from Taylor (1898) but corrected
for gender.
’ Type labels at CNC added to autotypes.
‘ Labels of these two specimens switched.
date labels from Taylor’s series, have ho-
lotype and paratype labels. The ‘holotype,’
which has a type label in Harrington’s hand-
writing, is dated 27-10-97 but this was ap-
parently interpreted as 27-IX-93 by the per-
son that added the holotype label and again
by Sarazin (1986). Harrington (1896) stated
the holotype was received by a collector in
September 1893, but did not say when it
was collected.
Harrington’s original “Victoria, V.I.” la-
bel has been placed on the holotype and the
24-10-97 label of Taylor has been placed on
the autotype at the Humboldt museum. The
two type specimens in the CNC should be
relabeled as autotypes.
I thank J. C. Miller (Oregon State Uni-
versity) and R. D. Akre (Washington State
University) for their support; L. Masner
(Canadian National Collection) and F. Koch
(Humboldt Universitaet, Berlin) for loans
of Bareogonalos canadensis; and H. Townes
(American Entomological Institute) for the
information in Schulz (1907b). P. Rossig-
nol, N. Anderson and others at Oregon State.
University helped clarify this paper and
make it more concise.
David Carmean, Department of Ento-
mology, Oregon State University, Corvallis,
Oregon 97331.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 289-290
NOTE
Pedipalpal Anomalies in Neobisium simoni
(L. Koch) and N. bernardi Vachon
(Neobisiidae: Pseudoscorpiones: Arachnida)
Pseudoscorpions have been found with
segmental anomalies involving the sclerites
(tergites and sternites) (Curci¢ & Dimitri-
jevic. 1982. Revue Arachnologique. 4, 143-
150. 1984. Arch. Sci. Belgrade, 36, 9P-10P.
1985. Revue Arachnologique. 6, 91-98.
1986. Actas X Congr. Int. Aracnol. Jaca/
Espana, 1, 17-23, and references cited
therein). A curious aberration other than
anomaly in sclerite structure has been re-
ported by Vachon (1947. Bull. biol. Fr. Belg.
81, 177-194), who recorded a protonymph
of Chelifer cancroides (Linnaeus) with the
right foreleg fused basally with the pedipalp.
In addition, Chamberlin (1949. Amer. Mus.
Novit. 1430, 1-57) reported one of the most
unusual anomalies yet observed in pseudo-
scorpions: in the holotype of Xenochelifer
davidi Chamberlin, the movable finger of
the left chela is greatly reduced, being only
half the normal length. The fixed finger is
apparently normal.
As far as the family Neobisiidae is con-
cerned, pedipalpal anomaly has been re-
corded in a single female of N. carpaticum
Beier (Curci¢é. 1980. Bull. Brit. Arachnol.
Soc. 5, 9-15). In this pseudoscorpion, the
fixed finger of the left pedipalpal chela is
greatly reduced, being two-thirds the nor-
mal length. Consequently, four distal
trichobothria (ist, est, it and ef) are missing
and there are also fewer teeth on this finger
in comparison to that of the right chela which
is normal. The movable finger is apparently
normal. In other Neobisiid species, malfor-
mations of other appendages (chelicerae,
walking legs) occur rarely, as was already
observed by Curéi¢ (1980). The aim of this
note 1s to express quantitatively and qual-
itatively the phenomena of pedipalpal ter-
atology in the species studied in order to
assess the pathomorphological traits of such
aberrations.
In a collection of pseudoscorpions made
by one of us (RND) at Passarole, near Mou-
lis (Ariége), France, during July 1987, one
female of Neobisium simoni (L. Koch) and
one tritonymph of N. bernardi Vachon with
abnormal pedipalpal chelae were found.
These were obtained by sifting leaf-litter and
humus in a mixed oak forest. In the speci-
mens studied, only the pedipalpal chelae
were anomalous, the other appendages and
abdominal sclerites were normal.
NEOBISIUM SIMONI: Female (Figs. 1
& 2). The fixed finger of the right chela of
the pedipalp is reduced, being only half the
normal length. Consequently, instead of four
distal trichobothria, a single tactile seta is
present (Fig. 1). Its relative position 1s more
basal than in any of the distal trichobothria:
Figs. 1-2. Neobisium simoni (L. Koch). Scale line
= 0.5 mm. (1) Right chela of the pedipalps, aberrant
female, (2) Right chela of the pedipalps, normal female.
290
N
“ne fie _ ; ¢
1/\ y\ 4
yi t 3 4 Yi 4\ \
* ¢ ¥ /
bid, & Lily i. §
Vy Hy A i ,
Now | \ AG we oH |
1 ep 4 nee ye!
t i + ° \ \ \ 3 4 . \
i} 1 { 3 { \
. ee : } \ e }
> = e |
fe | is } . \
iy a / \ of
=y ‘- es /
=p tf Hie x
\ i [ea / t
7 — — L
ee. ——_ Cun
] > A ve
\ ~ 4
/ pS /
Figs. 3-4. Neobisium bernardi Vachon. Scale line
= 0.5 mm. (3) Right chela of the pedipalps, aberrant
tritonymph, (4) Right chela of the pedipalps, normal
tritonymph.
therefore, its proper identification is not
possible. In addition, the deficient finger
carries 37 teeth only, whereas the normal
complement is 58-64 teeth (Fig. 2). The
movable finger is apparently normal and
carries 60 teeth, which falls within the nor-
mal range for the movable finger (54-60
teeth) in females of this species.
NEOBISIUM BERNARDI: Tritonymph
(Figs. 3, 4). The movable finger of the right
pedipalpal chela is reduced and attains two-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
thirds the normal length (Fig. 3). In addi-
tion, this finger carries two instead of three
trichobothria (seta ¢ is missing). The fixed
finger is normal. The anomalous finger has
25 teeth and the normal complement is 32-
38 teeth. They appear much smaller and
more close-set than in normal specimens.
The fixed finger carries 38 teeth (normal
range is 35-40 teeth). In both N. simoni and
N. bernardi studied left pedipalpal chelae
are normal.
The pedipalpal anomalies in different
species of Neobisium Chamberlin from the
family Neobisiidae have been found to date
in the adult (female) and tritonymph stages
(N. carpaticum, N. simoni, N. bernardi). No
deficiencies have been noted in the deuto-
nymph or protonymph stages.
It appears probable that the origin of the
pedipalpal anomalies analyzed should be
sought in some irregularity of the ontogenic
(? postembryonic) process.
We are grateful to Dr. Christian Juber-
thie, Director of the Laboratoire souterrain
in Moulis, for his collaboration and per-
mission to collect pseudoscorpions in the
vicinity of Moulis.
B. P. M. Curéi¢ and R. N. Dimitrijevié,
Institute of Zoology, Faculty of Science, 16,
Studentski Trg, YU-11000 Beograd, Yu-
goslavia.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 291-292
NOTE
Ticks of the Subgenus /xodiopsis: First Report of
Ixodes woodi from Man and Remarks on
Ixodes holdenriedi, a New Junior Synonym of
Ixodes ochotonae (Acari: Ixodidae)
Recently, this laboratory received a par-
tially engorged nymphal /xodes tick that had
been removed from the neck of a child (age
and sex not stated) on 5 June 1967 at Fran-
cis E. Warren Air Force Base, Laramie Co.,
Wyoming. No previous effort had been made
to determine the specific identity of this
specimen, probably because the only avail-
able key to the Jxodes nymphs of North
America, that of Cooley and Kohls (1945,
Natl. Inst. Health Bull. 184: 1-246), is out
of date (nymphs were known for only 26 of
the 41 species treated by Cooley and Kohls;
since then, several taxa have been synony-
mized and 6 new species have been de-
scribed). Careful comparison with material
in the U.S. National (formerly Rocky
Mountain Laboratories) Tick Collection has
shown that the Wyoming specimen is /xodes
woodi Bishopp, a member of the morpho-
logically close-knit subgenus [xodiopsis Fi-
lippova, which comprises 7 species, all chief-
ly parasites of small rodents and insectivores
(Robbins and Keirans 1987, J. Med. Ento-
mol. 24: 310-314). This is the first known
instance of human parasitization by J. wood.
In North America, the subgenus /xodiop-
sis 1s often referred to as the “Jxodes an-
gustus group,” after Ixodes angustus Neu-
mann, a widespread parasite of cricetid
rodents. Other Nearctic members of the /.
angustus group include (besides J. woodi)
Ixodes eastoni Keirans and Clifford, to date
known only from southwestern South Da-
kota and the border area of northeastern
Wyoming, where it parasitizes cricetine and
microtine rodents; /xodes ochotonae Greg-
son, a parasite of ochotonid lagomorphs and,
to a lesser extent, cricetines in the montane
West; and Ixodes soricis Gregson, a char-
acteristic tick of western soricid insecti-
vores. Another taxon that would appear to
fit the definition of subgenus /xodiopsis is
Ixodes holdenriedi Cooley, which is known
only from the holotype and paratype, both
females taken on the pocket gopher Tho-
momys bottae (Eydoux and Gervais), So-
noma Co., California, 25 March 1945. It
seems remarkable that during more than 40
years no further specimens have come to
hand in an area that has been particularly
well worked by acarologists and public
health personnel. Examination of the orig-
inal description and of the type material
(now in poor condition) indicates that /.
holdenriedi is conspecific with, and there-
fore a junior synonym of, J. ochotonae.
Though Spencer (1963, Proc. Entomol.
Soc. Brit. Columbia 60: 40) published a rec-
ord of J. soricis from a girl, all other reports
of Ixodiopsis ticks from man pertain to
Ixodes angustus. As early as 1937, Cham-
berlin (Stn. Bull. Oregon Agric. Exp. Stn.
349: 1-34) noted that J. angustus will def-
initely feed on man, but he did not cite spe-
cific examples. Later, Cooley (1946, J. Par-
asitol. 32: 210) described 3 cases of human
parasitization by J. angustus in the Pacific
Northwest, Gregson (1956, Publ. Dep.
Agric. Canada 930: 1-92) noted 2 cases from
southwestern British Columbia, and Spen-
cer (op. cit.) added 3 more records from that
Province. In addition to these published
records, there are 4 specimens in the Fred
C. Bishopp Collection (now merged with the
National Tick Collection) that were re-
moved from humans: | partially engorged
2 from the scalp of a boy, Sandy River,
Multnomah Co., Oregon, 7 August 1934
(RML 118942); 2 partially engorged °° from
292
the arm of a 14-year-old girl, Forest Grove,
Washington Co., Oregon, 8 August 1932
(RML 118959); and | partially engorged 2
“taken from the shoulder of a man who
complained of rheumatic pains .... Tick
was firmly attached. Rheumatic symptoms
soon subsided after removal of tick,” Van-
couver, Clark Co., Washington, 11 July 1938
(RML 118964). There are also 4 unpub-
lished records in the National Tick Collec-
tion itself: | engorged 2 from a child, Kirk-
land, King Co., Washington, August 1947
(RML 24023); 1 engorged 2 from a young
boy, Washington [State], 20 July 1956 (RML
33925); 1 partially engorged 2? from the head
ofa young girl, Seattle, King Co., Washing-
ton, ca. 2 August 1977 (attachment site be-
came inflamed on 12 August) (RML
105346); and 1 engorged 2 from an other-
wise undefined human host, Juneau, Great-
er Juneau Borough, Alaska, | August 1953
(RML 118623).
The nymph of /. woodi reported herein
(RML 118594) may be described as follows
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(measurements in millimeters): Length of
body from scapular apices to posterior mar-
gin 2.157, greatest width 1.395; length of
capitulum from palpal apices to cornua api-
ces 0.297, width at level of cornua 0.261;
palpi 0.245 long, 0.091 wide, segment I ven-
trally without an anterior spur but with a
prominent posterior spur that projects lat-
erally; hypostome broken; scutum 0.598
long, 0.581 wide; left tarsus I missing, right
0.287 long, 0.121 wide.
Thanks to Jerome Goddard, U.S. Air
Force School of Aerospace Medicine, Brooks
Air Force Base, San Antonio, Texas, for
sending this most significant specimen.
Richard G. Robbins, Department of
Health and Human Services, Public Health
Service, National Institutes of Health, Na-
tional Institute of Allergy and Infectious Dis-
eases, Y Department of Entomology, Mu-
seum Support Center, Smithsonian
Institution, Washington, D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 293-294
Book REVIEW
Lepidoptera Anatomy. By John L. Eaton.
Wiley-Interscience (John Wiley & Sons),
New York, 1988, 257 p. $49.95.
The need for a comprehensive, revised
text on comparative anatomy of Lepidop-
tera has long been realized. Several impor-
tant morphological studies, particularly by
German and Russian authors, appeared in
the early decades of this century. Most of
these classics are no longer readily available,
or are unavailable in English, and recent
studies are so specific in the anatomical areas
discussed as to be of limited use as general
references. Unfortunately, the recent vol-
ume entitled Lepidoptera Anatomy by John
Eaton fails this need in several respects. Most
importantly, prospective purchasers of this
rather small but expensive volume ($49.95)
should be informed that Eaton’s book is not
a review of general lepidopteran anatomy
as its title implies. Instead it is primarily a
description of all stages of the sphingid,
Manduca sexta (L.), which has been the pri-
mary and almost exclusive focus of Eaton’s
research over the years. As a result, much
of the text reads like a laboratory guide for
the study of the tobacco hornworm moth,
with little reference to other species. Only
in few chapters and largely because of ne-
cessity (e.g. in the chapter on sound pro-
duction), are other Lepidoptera discussed.
In his preface, Eaton states that he has
chosen Manduca sexta as a model for the
Lepidoptera. The “archetype” or ““model”
approach for studies involving such vast
subjects certainly represents a logical and
often necessary method of treatment. Crit-
icism arises in the choice of the models and,
especially in this case, how many models
should be included to provide an adequate
representation for the Lepidoptera. Selec-
tion of one of the most specialized members
(a sphingid) of the order is not likely to result
in arepresentative example. Ata minimum,
it probably would have been necessary to
discuss examples from the Micropterigidae,
Incurvarioidea, Tineidae, Pyralidae, Noc-
tuidae, and probably Papilionidae in order
to present a meaningful review of the ana-
tomical diversity within the order.
The text is divided into | 1 chapters, which
vary considerably in scope and quality. Not
surprisingly, those chapters which had large-
ly been treated previously by Eaton in the
Annals of the Entomological Society of
America (e.g. the larval and adult muscu-
lature, nervous and endocrine systems) are
the best researched and most informative.
The remaining chapters, dealing mostly with
external morphology, are clearly written but
brief and are generally inferior to references
currently available. Literature references are
included at the end of each chapter. These
also vary in coverage according to chapter
and frequently do not provide an adequate
representation of the best references. The
most conspicuous omissions in the text are
the lack of references to larval chaetotaxy
and scale morphology. Considering the im-
portance of these structures for the order,
their absence constitutes a major oversight.
The subject matter is amply supported by
numerous, well labelled, line drawings. In
addition to a complete listing of all label
anagrams in a terminal appendix, some of
the more complex illustrations (e.g. fig. 4—
25) also are provided with anagram legends
which reduce page flipping appreciably. The
quality of illustrations, like the text, varies
among chapters, with those previously pub-
lished elsewhere generally being superior to
those prepared for the book. Several draw-
ings appear oversized and crudely executed
with a conspicuous lack of certain details or
accuracy (e.g. larval chaetotaxy). One rather
surprising Omission noted for a modern an-
atomical text is the absence of electron mi-
294
crographs, particularly SEM photos. Details
shown by such photographs are sorely need-
ed in support of the text dealing with sen-
sory structures and trophic organs. AI-
though no SEM photographs of eggs are
shown, the drawings for this brief chapter
reportedly were based on SEM studies.
In summary, the main criticism of this
book is its misleading title and all that it
implies. I would recommend the text as a
reference primarily for beginning students
Asa Fitch and the Emergence of American
Entomology. By Jeffrey K. Barnes. Bull.
No. 461, New York State Museum, Al-
bany, New York, 1988, viii and 120 pp.
$12.50 + $1.00 postage and handling,
from New York State Museum Publica-
tions, 3140 Cultural Education Center,
Albany, NY 12230.
With a marvelous grasp of the history of
the mid-nineteenth century in New York,
Barnes has written a fine appreciation of Asa
Fitch and the beginnings of the science of
entomology in North America. Fitch’s life
is sketched in a series of nine chapters 1n-
terwoven with three more that set his life
within the larger context of the times.
The early part of the nineteenth century
in New York when Fitch came of age was
optimistic and progressive. Barnes places
Fitch in a period of expanding economy,
growth of educational and scientific estab-
lishments, great migration, cheaper travel,
religious ferment, and new agricultural de-
velopments. Because of more extensive
farming and the need to grow crops more
efficiently, insect damage became more no-
ticeable. Albany, the capital of the richest
state, was a center of scientific activity (for-
eign scientists arriving in Boston asked di-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of Lepidoptera anatomy and especially for
those interested in internal anatomy. For
those primarily concerned with external
anatomy, they would be best advised to
search elsewhere for one of several superior
references currently available.
Donald R. Davis, Department of Ento-
mology, National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 294-295
rections to Albany), due largely to its po-
sition at the confluence of two rivers and at
the beginning of the new Erie canal.
Little was known about American insects
before 1845. Although Thomas Say had al-
ready described about 1500 species, those
insects, with a few exceptions, were not crop-
related. The study of insects lagged behind
other natural history studies and people had
strange notions concerning insect control.
But it was apparent to many that insects
were becoming more important to the de-
velopment of agriculture and that little was
known about how to fight them effectively.
Fitch, a product of his times, a man of social
purpose deeply interested in science, was
well-placed by his education, interests, and
integrity to be influential in the growth of
the new field of entomology.
Fitch was born in 1809, the same year as
Abraham Lincoln. We learn of his early years
on a farm, his education in the natural sci-
ences, travels, medical education. He en-
tered the practice of medicine, but lacked
respect for the profession and eventually
quit. While still young, Fitch returned to life
as a farmer. Because he was well-to-do he
was able to devote much of his time and
energy to insect study. His first papers, writ-
ten as early as 1845 while still an amateur,
VOLUME 91, NUMBER 2
were excellent treatments of insects inju-
rious to plants. During 1846-8 he was paid
to make a collection of insects for the State
Cabinet of Natural History, a collection that
included plant damage and which was even-
tually placed in the State Museum. Largely
due to his demonstrated knowledge and with
alittle help from well-placed friends, he won
in 1854 the post of State Agricultural So-
ciety Entomologist. Fitch continued in that
capacity for 19 years, during which he wrote
14 annual reports and many smaller papers.
Fitch was also greatly committed to ed-
ucation. He always had in mind that science
should “‘relate to the common purposes of
life.” Previous scientific reports in the state
had been criticized for being unintelligible
to ordinary citizens and of little practical
value. But Fitch’s reports can still stand as
models because they were useful to both
scientists and farmers and simply written.
Fitch carefully observed the life cycles of
many insects for the first time. As he told
the young Henry Comstock, then looking
for guidance, the best way to begin the study
of insects is to observe their behavior. Fitch’s
recommendations for the control of insects,
including critical growing times, selection
for early or late plant varieties, and biolog-
ical control (he was the first to suggest get-
ting parasites from abroad to fight immi-
grant pests), were highly influential. His
reports continued to be in demand decades
295
after they were written. Even in Europe they
were popular because of their practical ap-
plication. C. V. Riley rightly called Fitch
the father of economic entomology in North
America.
The book has a large page (10%4” by 812”),
two column format, is well illustrated with
copies of photographs and prints and with
some of Barnes’ own photos of historic
places, and is printed on excellent stock.
Each of the 12 regular chapters has its own
reference notes immediately following. Two
appendices follow. The first is a list of Fitch’s
entomological publications. The second ap-
pendix is an appreciation of Fitch’s taxo-
nomic work, with a discussion of his col-
lections and a catalog of his taxonomic
names and type specimens. The list of his
extant types in 3 classes, 15 orders, and 107
families is hierarchically arranged, com-
plete, most valuable, and reflects a great
amount of careful museum work by Barnes.
In this section I noticed a typographical
error, possibly done purposely, in the same
way that an otherwise perfect, handmade
oriental rug has a tiny flaw woven into it to
ward off the evil eye.
Raymond J. Gagné, Systematic Ento-
mology Laboratory, Agricultural Research
Service, % USNM NHB 168, Washington,
D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 296-297
Book REVIEW
The Mayflies of Florida, Revised Edition.
By Lewis Berner and Manuel L. Pescador.
University Presses of Florida. 1988, xvi
+ 415 pp. ISBN 0-8130-0845-X. $35.00/
hard, from 15 NW St., Gainesville, FL
32603.
The original edition of The Mayjlies of
Florida by Lewis Berner was published 1n
1950, and the numerous detailed observa-
tions it contained were an invaluable source
of data on the natural history of Ephemer-
optera. It has been out-of-print for a num-
ber of years, but fortunately Berner, with
the aid of his new coauthor Pescador, has
provided an updated edition. The format
remains basically the same, but new data
and recent references have been added and
discussions have been more compartment-
alized. A total of 71 species are treated in
this edition, a significant increase of 23 since
1950. Much added data were undoubtedly
contributed by the aquatic entomologists at
Florida A&M University, particularly with
regard to studies in the Florida panhandle
region. There are now 28 plates of very good
habitus drawings (not numbered as figures),
174 numbered figures, and 27 distribution
maps, an impressive increase over the 24,
88, and 19, respectively, of the earlier edi-
tion.
Chapters include an introduction that
contains a generalized account of the biol-
ogy of the various life stages, morphological
adaptations of larvae, higher classification,
and taxonomic characters. This is a clear
and concise discussion that non-specialists
will appreciate; however, it would have been
more useful if figure citations had been lib-
erally cited to accompany the adaptation
and character discussions. A chapter com-
paring the Florida fauna with more north-
ern ones emphasizes the relative non-sea-
sonality of many Florida populations. An
excellent chapter entitled Zoogeography
deals with distributions in the context of the
geological history of the peninsula along with
descriptions of the many available aquatic
habitats. This is valuable information for
those interested in any of the freshwater bio-
ta of Florida. A species key to the adults
(mainly males) and larvae makes up the next
chapter. It reflects the present state of
knowledge about generic and specific dif-
ferences and is well illustrated. The key is
followed by a synoptic list of species and
finally the individual species treatments.
The taxonomy is essentially up-to-date,
the most notable exception being in the
family Baetidae, whose genera are undergo-
ing considerable revision. The validity of a
few of the species that the authors recognize
is in some doubt. Of the 71 species treated,
6 are apparently endemic to Florida. The
doubtful species are among these endemics,
some of which may arguably be ecopheno-
typic variants. The great strength of the book
remains the wealth of original observational
data that can be gleaned from the topics
treated under each species. These include
taxonomy (diagnostic but not descriptive);
distribution; ecology (mainly a discussion
of specific habitats); seasonality and life his-
tory (often including emergence and repro-
ductive behavior); and behavior (mainly
larval orientation and feeding). Searching
for information about certain phenomena
without knowing to which species it may
pertain can be a real problem with this type
of format. In this book, however, there is,
happily, a very complete index to subject
matter. Citations available since 1950 are
quite complete, with very few pertinent be-
havioral and ecological publications missed.
For those workers who found the original
edition of this book useful, this revision will
better serve them. For the newer generation
of researchers and students who have not
VOLUME 91, NUMBER 2
been able to obtain the book previously, the
new edition is most worthy of their atten-
tion. And for entomologists, naturalists, and
aquatic biologists in the Southeast, the book
should definitely be considered for their ref-
erence libraries.
297
W. P. McCafferty, Department of Ento-
mology, Purdue University, West Lafayette,
IN 47907.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 298-303
OBITUARY
Frederick William Poos, Jr.
1891-1987
Frederick William Poos, Jr., a member of
the Entomological Society of Washington
since 1923, President of the Society in 1945
and Honorary Member since 1966, died
from surgical complications on June 28,
1987. Dr. Poos lived in northern Virginia.
His wife died in 1974. He is survived by
two sons: Frederick William Poos III, Long
Island, New York and George Ireland Poos,
Fort Washington, Pennsylvania.
Dr. Poos was born in Potter, Kansas on
November 12, 1891, the son of Frederick
W. and Dena (Steinhage) Poos. He grew up
in Kansas and attended the University of
Kansas where he received his A.B. in 1915
and A.M. in 1916. He worked for the Bu-
reau of Plant Industry in Florida from 1916
to 1917 and from 1917 to 1922 for the Bu-
reau of Entomology, USDA, in Charlottes-
ville, Virginia. During this time he co-au-
thored a number of papers on stem-boring
insects of wheat and corn. He married Edna
M. Ireland in 1919. In 1922 he moved to
Sandusky, Ohio as assistant entomologist in
charge of the European Corn Borer Station.
At this time he worked on his Ph.D., which
VOLUME 91, NUMBER 2
he received from Ohio State University in
1926. His Ph.D. thesis treated the biology
of the European corn borer and two closely
related species in Ohio. From 1926 to 1928
he worked at the Virginia Truck Experiment
Station in Norfolk, Virginia on insects at-
tacking spinach, potatoes, narcissus and
roses. In 1928 he returned to work for the
USDA, Bureau of Entomology and Plant
Quarantine at the experimental farm in Ar-
lington, Virginia.
Prior to 1928 Dr. Poos had made im-
portant contributions to the field of eco-
nomic entomology. His early work was on
the biology and control of the European corn
borer, larger corn stalk borer, wheat
strawworm and wheat jointworm. His rec-
ommendations for cultural and biological
control still have merit. He also did note-
worthy work on the life history and control
measures for the Hawaiian beet webworm
on spinach. He developed an effective spray
calendar for use in a five state area. Upon
his return to the USDA experimental farm
in 1928 he began an outstanding career in-
volving various insect pests, especially those
of potatoes and peanuts. He discovered the
insect vector, the corn flea beetle, of Stew-
art’s disease of corn. This important break-
through led to important control measures
for the vector and the disease. Dr. Poos, in
collaboration with Hurd-Karren was the first
in the United States to discover the systemic
action of insecticides in plants. Dr. Poos
also was well known for his work on leaf-
hoppers and in particular the potato leaf-
hopper. He was the first to discover that this
insect migrates from southern to northern
states each year. He discovered a number
of practical control measures for pests of
peanuts. He identified the cause of a disease
of peanuts, that was uncertain for years, was
actually the tobacco thrips. Another of his
fields of research was insects attacking soy-
beans.
When, in 1941, the property on which the
experimental farm was located was needed
299
for construction of a large military head-
quarters, later known as the “Pentagon,”
Dr. Poos moved to Beltsville, Maryland to
continue his research. He worked at Belts-
ville until his retirement in 1957. His re-
search at Beltsville included studies on in-
jurious insects attacking legume and grass
crops. In addition to his work on the use of
new insecticides to control pests, he also was
responsible for studies on the effect of in-
secticides on dairy cows and their milk.
In 1954 he received the Superior Service
Award from the USDA and in 1963 he re-
ceived a citation of merit from the Ento-
mological Society of America. In 1982 Dr.
Poos received international acclaim in a very
unusual manner. The well-known piece of
entomological equipment called the “‘aspi-
rator’’ is known in England as a “pooter.”
The origin of the term “pooter” came to
light in the October 1982 issue of Antenna,
a British Entomology Journal, where it was
disclosed that the term is derived from the
name “‘Poos.”’ Dr. Poos, the first to use the
aspirator in collecting leafhoppers, de-
scribed it in a 1929 article. British ento-
mologists coined the term “‘pooter” to in-
dicate the device used by Dr. Poos.
Dr. Poos had been a member of the Pres-
byterian Church of the Pilgrims since 1929,
where he had served as an elder. He was a
member of numerous societies including the
American Association for the Advance-
ment of Science, Entomological Society of
America (honorary member), American
Association of Economic Entomologists,
Biological Society of Washington, Kansas
Entomological Society, Kansas, Ohio, and
Washington (DC) academies of science, Sig-
ma Xi, and Cosmos Club in Washington,
DC. He also served as editor for the Journal
of Economic Entomology. He was always
an active person both professionally and in
civic activities. He was active and alert even
as he went into surgery on June 25, 1987.
Many entomologists in the Washington area
knew him well and often visited with him.
300
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
His death marks the loss of one of the great
contributors to entomology in the service
of American farmers and consumers.
Thomas E. Wallenmaier, Room 608,
Federal Building, Hyattsville, Maryland
20782.
PUBLICATIONS OF F. W. Poos
. Phillips, W. J.,G. W. Underhill, and F. W. Poos.
The larger corn stalk-borer in Virginia. Va. Agr.
Expt. Sta. Tech. Bull 22: 3-30, June 1921.
and F. W. Poos. Life-history studies of
three jointworm parasites. Jour. Econ. Ent. 21:
405-426, Dec. 1921.
and . A lamp for taxonomic work
in entomology. Jour. Econ. Ent. 14: 504-506, 1
fig., Dec. 1921.
and . Five new species belonging
to genus Harmolita Motschulsky (Isosoma Walk-
eret Auct.). Ka. Univ. Science Bull. 14: 349-359,
pls. 36-37, Oct. 1922.
and . The wheat strawworm and
its control. U.S. Dept. Agr. Farmers’ Bull. 1323:
1-10, May 1923.
. Poos, F. W. A new pest of spinach in Virginia.
Va. Truck Expt. Sta. Bull. 56: 491-497, July 1,
1926.
. Phillips, W. J. and F. W. Poos. Two hymenop-
terous parasites of American jointworms. Jour.
Agr. Res. 34: 473-488, March 1, 1927.
. Poos, F. W. Biology of the European corn borer
(Pyrausta nubilalis Hubn.) and two closely relat-
ed species in northern Ohio. Ohio Jour. Science
27: 47-94, March 1927.
. An insect borer damaging roses. South-
ern Planter 88(11): 15, June 1927.
. Virginia-grown narcissus bulbs. Florists’
Review 60: 31-33, June 1927.
and C. A. Weigel. The bulb flies of nar-
cissus with special reference to the bulb industry
in Virginia. Va. Truck Exp. Sta. Bull. 60: 571-
594, July 1, 1927.
. Poos, F. W. and H. S. Peters. The potato tuber
worm. Va. Truck Expt. Sta. Bull. 61: 597-630,
Octil; 19272
. McWhorter, F. P. and F. W. Poos. Spray calendar
for Virginia, West Virginia, Maryland, North
Carolina and South Carolina. American Produce
Grower 2(4): 6-7, April 1927.
and . Spray calendar for Virginia,
West Virginia, Maryland, North Carolina and
South Carolina. (Revised). American Produce
Grower 3(2): 6, 8, Feb. 1928.
15%
and . Recognizing diseases and in-
sects mentioned in the spray calendars. American
Produce Grower 3(3): 5, 12, March 1928.
. Poos, F. W. Bulb sterilizers used in Virginia. Flo-
rists’ Review 62: 35-36, May 24, 1928.
. Hot water treatment in bulbs. Florists’
Review 62: 35-36, Oct. 11, 1928.
. Experiments with narcissi. Florists’ Re-
view 62: 33-34, Oct. 18, 1928.
. An annotated list of some parasitic in-
sects. Proc. Ent. Soc. Wash. 30(8): 145-150, Nov.
1928.
. Leafhopper injury to legumes. Jour. Econ.
Ent. 22: 146-153, Feb. 1929.
. Leafhopper injury to legumes. (Author’s
Abstract). Jour. Wash. Acad. of Sciences 20(6):
116-117. March 19, 1930.
. Anew method of distributing Emmpoasca
fabae (Harris). Jour. Econ. Ent. 23(4): 770, Aug.
1930.
and F. F. Smith. A comparison of ovi-
position and nymphal development of Empoasca
fabae (Harris) on different host plants. Jour. Econ.
Ent. 24: 361-371, 11-12, April 1931.
24. Smith, F. F. and F. W. Poos. The feeding habits
30.
of some leafhoppers of the genus Empoasca. Jour.
Agr. Res. 43: 267-285, Aug. 1, 1931.
25. Poos, F. W. and C. M. Haenseler. Injury to va-
rieties of eggplant by the potato leafhopper, Em-
poasca fabae (Harris). Jour. Econ. Ent. 24: 890-
892, Aug. 1931.
. Poos, F. W. Biology of the potato leafhopper,
Empoasca fabae (Harris) and some closely re-
lated species of Empoasca. Jour. Econ. Ent. 25(3):
639-646, June 1932.
and R. B. Deemer. Is the absorption of
copper by certain crop plants influenced by cli-
matic, soil, or other factors? Jour. Econ. Ent. 26:
648, June 1933.
. Poos, F. W. Leafhoppers of the genus Empoasca
(Homoptera, Cicadellidae) in Virginia. Proc. Va.
Acad. Sci. 1932-1933: 36-37.
. Four new species of Empoasca (Homop-
tera, Cicadellidae). Proc. Ent. Soc. Wash. 35(8):
174-179, 1933.
and N. H. Wheeler. On the hereditary
ability of certain insects to transmit diseases and
to cause diseaselike injuries to plants. Jour. Econ.
Ent. 27(1): 58-69, 1934.
and H. L. Westover. ‘Alfalfa Yellows.”
Science 79(2049): 319, April 6, 1934.
. Poos, F. W. A stand used in photographing ob-
jects from above. U.S.D.A., Bur. Ent. ET-16, May
1934.
. A leafhopper injurious to peanuts. Pea-
nut Journal and Nut World 13(8): 9, 13, June 7,
1934.
VOLUME 91, NUMBER 2
34.
35.
36.
37.
38.
39.
40.
41.
43.
44.
45.
46.
47.
48.
49.
50.
Elliott, Charlotte and F. W. Poos. Overwintering
of Aplanobacter stewarti. Phytopathology 25(1):
32, January 1935.
Poos, F. W. and Charlotte Elliott. Bacterial wilt
of corn and its insect vectors. (Abs.). Phytopa-
thology 25(1): 32, January 1935.
Poos, F. W. Under heading of notes and exhi-
bition of specimens: Plants of Empoasca fabae
(Harris) and their probable significance. (Au-
thor’s abstract). Proc. Ent. Soc. Wash. 37(8): 170,
1935.
. New host plants of the potato leafhopper,
Empoasca fabae (Harris) and their probable sig-
nificance. Jour. Econ. Ent. 28(6): 1072-1073,
1935.
, N. H. Wheeler, and J. W. Scrivener.
Methods and apparatus used in identifying large
numbers of leafhoppers of the genus Empoasca.
U.S.D.A. Bur. Ent. & Plt. Quar. ET-72, 3 pp.,
Feb. 1936.
and H. W. Johnson. Injury to alfalfa and
red clover by the potato leafhopper. Jour. Econ.
Ent. 29(2): 325-331, illus., 1936.
and Charlotte Elliott. Certain insect vec-
tors of Aplanobactor stewart. Jour. Econ. Ent.
29(2): 325-331, 1936.
Hurd-Karrer, A. M. and F. W. Poos. Toxicity of
selenium-containing plants to aphids. Science
84(2176): 252, 1936.
. Poos, F. W. and E. T. Batten. Greatly increased
yields of peanuts obtained in attempts to control
potato leafhopper. Jour. Econ. Ent. 30(3): 561,
1937,
Phillips, W. J. and F. W. Poos. The wheat
strawworm and its control. U.S.D.A. Farmers’
Bull. 1323, May, 1923. Revised (by junior au-
thor) Oct. 1937.
Poos, F. W. Insects in relation to diseases of ce-
real and forage crops. Jour. Econ. Ent. 31(1): 24-
38, 1938.
Batten, E. T. and F. W. Poos. Spraying and dust-
ing to control the potato leafhopper on peanuts
in Virginia. Va. Agr. Exp. Sta. Bull. 316: 1-26,
April 1938.
Poos, F. W. and J. W. Scrivener. A convenient
cage for determining field populations of the po-
tato leafhopper. U.S.D.A. Bur. Ent. & Pl. Quar-
antine ET-121, June 1938.
Poos, F. W. Control potato leafhoppers on pea-
nuts. Southern Planter 100(6): 15, June 1939.
. Control potato leafhoppers on peanuts.
Extension Entomologists 3(2): 4-5, June 1939.
. Host plants harboring Ap/anobactor
stewarti without showing external symptoms af-
ter inoculation by Chaetocnema pulicaria. Jour.
Econ. Ent. 32(6): 881-882, 1939.
Elliott, Charlotte and F. W. Poos. Seasonal de-
56.
Ds
58.
59.
60.
61.
63.
64.
65.
66.
67.
301
velopment, insect vectors, and host range of bac-
terial wilt of corn. Jour. Agr. Res. 60(10): 645-
686, 1940.
. Poos, F. W. The locust leaf miner, Chalepus dor-
salis Thumb. as a pest of soybean. Jour. Econ.
Ent. 33(5): 727-728, 1941.
. Phillips, W. J. and F. W. Poos. The wheat joint-
worm and its control. Farmers’ Bulletin 1006: 1-
12, illus., revised by jr. author, Sept. 1940.
. Poos, F. W. On the Causes of Peanut ‘‘Pouts.”
Jour. Econ. Ent. 34(5): 727-728, 1941.
. Peanut “pouts” caused by thrips. The
Southern Planter 102(12): 12, illus., 1941.
. The potato leafhopper. A pest of alfalfa
in the eastern states. U.S.D.A. Leaflet No. 229:
1-8, 1942, revised 1952.
. Control of hay insects in the eastern
United States. Mimeographed by Bureau of
Entomology & Plant Quarantine, U.S.D.A.,
March 1943.
Poos, F. W. and Nancy H. Wheeler. Studies on
host plants of the leafhoppers of the genus Em-
poasca. U.S.D.A. Tech. Bull. 850, May 1943.
Poos, F. W. and L. A. Hetrick. Tetralopha scor-
tealis (Led. ) A new insect pest of lespedeza. Jour.
Econ. Ent. 38(3): 312-315, 1945.
Poos, F. W. DDT to control corn flea beetle on
sweet corn and potato leafhopper on alfalfa and
peanuts. Jour. Econ. Ent. 38(2): 197-199, April
1945.
. The control of tobacco thrips on seedling
peanuts. Jour. Econ. Ent. 38(4): 446-448, Aug.
1945,
Potts, S. F., T. E. Bronson, R. Latta, and F. W.
Poos. Atomized concentrated sprays containing
DDT to control pea aphid and clover leaf weevil.
Jour. Econ. Ent. 38(4): 497, Aug. 1945, scientific
note.
. Grayson, J. W. and F. W. Poos. Southern corn
rootworm as a pest of peanuts. Jour. Econ. Ent.
40(2): 251-256, illus., 1947.
Poos, F. W., J. M. Grayson, and E. T. Batten.
Insecticides to control tobacco thrips and potato
leafhopper on peanuts. Jour. Econ. Ent. 40(6):
900-905, 1947.
Poos, F. W. and E. T. Batten. Use of DDT dust
mixture is now recommended for peanut pests
control. Peanut Jour. and Nut World 27(6): 32,
1948.
Poos, F. W. and E. T. Batten. Control of peanut
pests. Southern Planter 109(6): 16, 1948.
Poos, F. W., S. F. Potts, L. D. Anderson, and J.
W. Brooks. Tests with mist blowers to control
various insects. U.S.D.A., Bur. Ent. & Pl. Quar.
E-751m, 8 pp., July 1948.
Poos, F. W. and Nancy H. Wheeler. Some ad-
ditional host plants of three species of leafhoppers
302
68.
69.
70.
71.
13:
74.
TD:
76.
TT:
78.
19):
80.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of the genus Empoasca. Proc. Ent. Soc. Wash.
51(1): 35-38, 1949.
Shepherd, J. B., L. A. Moore, R. H. Carter, and
F. W. Poos. The effect of feeding alfalfa hay con-
taining DDT residue on the DDT content of cow’s
milk. Jour. Dairy Science 32(6): 549-555.
Poos, F. W. Insects injurious to alfalfa. Report
of the 11th Alfalfa Improvement Conference: 75,
1948.
Poos, F. W., T. N. Dobbins, and R. H. Carter.
Sampling forage crops treated with organic in-
secticides for determination of residues. U.S.D.A.
Bur. Ent. & Pl. Quar. E-793, 12 pp., January
1950.
Ely, R. E., L. A. Moore, R. H. Carter, and F. W.
Poos. The DDT, toxaphene, and chlordane con-
tent of milk as affected by feeding alfalfa sprayed
with these insecticides. BDIM-Inf. 85, December
1949.
; Ely; Ro Ey Ae Moore, R. Hi: Carter, H: D:
Mann, and F. W. Poos. The effect of dosage level
and method of administration of DDT on the
concentration of DDT in milk. BDIM-Inf. 104,
June 1950.
Moore, L. A., R. H. Carter, and F. W. Poos.
Insecticide studies with dairy cattle. Milk and
Food Technol. Jour. 1 2(2): 103-104, March-April
1949.
Poos, F. W., T. N. Dobbins, E. T. Batten, and
G. M. Boush. Tests with benzene hexachloride
for the control of insects attacking peanuts. 1946-
1949. U.S.D.A. Bur. Ent. & Pl. Quar. E-820, 16
pp., May 1951.
Poos, F. W. Control of the garden webworms in
alfalfa. U.S.D.A. Leaflet 304, July 1951.
Kirkpatrick, M. E., B. M. Mountjoy, L. C. Al-
bright, Fred W. Poos, and C. E. Weigel. Flavor
and odor of cooked potatoes as affected by use
of lindane and benzene hexachloride as insecti-
cides. The American Potato Jour. 28: 792-798,
Dec. 1951.
Poos, F. W. and W. L. Howe. Control of the
southern corn rootworm on peanuts. U.S.D.A.
Bur. Ent. & Pl. Quar. EC-23, 7 pp., June 1952.
Ely, R. E., L. A. Moore, R. H. Carter, H. D.
Mann, and F. W. Poos. The effect of dosage level
and various methods of administration on the
concentration of DDT in milk. Jour. Dairy Sci-
ence 35(3): 266-271, March 1952.
Poos, F. W., T. N. Dobbins, R. H. Carter, R. E.
Hubanks, R. E. Ely, and L. A. Moore. Effects of
drying procedures, plant growth, and weathering
on insecticide residues on sprayed alfalfa.
Poos, F. W. Relative importance of various in-
sects harmful to alfalfa in Eastern Canada and
the United States. Dept. of the 13th Alfalfa Im-
provement Conference, 1952: 34-36.
81.
83.
84.
85.
86.
87.
89.
90.
Ol.
93.
94.
95:
. Get that grasshopper! The Windrow 2(4):
4-5, September 1952.
. Poos, F. W. and T. L. Bissell. The alfalfa weevil
in Maryland. Jour. Econ. Ent. 46(1): 178, May
1953.
Ely, R. E., L. A. Moore, P. E. Hubanks, R. H.
Carter, and F. W. Poos. Results of feeding me-
thoxychlor sprayed forage and crystalline me-
thoxychlor to dairy cows. Jour. Dairy Science
36(3): 309-314, March 1953.
Poos, F. W. and W. L. Howe. Control of the
southern corn rootworm on peanuts. USDA Bur.
Ent. and Pl. Quar. EC-23, 7 pp., April 1953.
Ely, R. E., L. A. Moore, P. E. Hubanks, R. H.
Carter, and F. W. Poos. Studies of feeding aldrin
to dairy cows. U.S.D.A. BDI-Inf-156, June 1953.
Poos, F. W. The meadow spittlebug—How to
control it. U.S.D.A. Leaflet 341, July 1953.
Carter, R. H., P. E. Hubanks, F. W. Poos, L. A.
Moore, and R. E. Ely. The toxaphene and chlor-
dane content of milk from cows receiving these
materials in their feed. Jour. Dairy Science 36(1 1):
1172-1177, November 1953.
. Poos, F. W. Insects as related to seed production.
South. Pasture and Forage Crop Impr. Conf. Rpt.
10: 36-38, 1953.
Gilpin, G. L., R. A. Redstrom, H. Reynolds, and
F. W. Poos. Flavor of peanut butter as affected
by aldrin, chlordan, dieldrin, heptachlor, and
toxaphene used as insecticides in growing pea-
nuts. Jour. Agr. and Food Chem. 2: 778-780,
July 21, 1954.
Poos, F. W. Soil insecticides dig out pests; Ap-
plication beneath the surface halts grubs hitting
forage crops below the belt. South. Seedsman
17(2): 20, 81, Feb. 1954.
Ely, R. E., L. A. Moore, R. H. Carter, P. E. Hu-
banks, and F. W. Poos. Excretion of dieldrin in
the milk of cows fed dieldrin-sprayed forage and
technical dieldrin. Jour. Dairy Science 37: 1461-
1465, Dec. 1954.
. Beattie, J. H., F. W. Poos, and B. B. Higgins.
Growing peanuts. USDA Farmers’ Bull. 2063,
54 p., May 1954.
Ely, R. E., L. A. Moore, P. E. Hubanks, R. H.
Carter, and F. W. Poos. Studies of feeding aldrin
to dairy cows. Jour. Dairy Science 37: 294-298,
March 1954.
Ely, R. E., L. A. Moore, P. E. Hubanks, R. H.
Carter, and F. W. Poos. Excretion of heptachlor
epoxide in the milk of dairy cows fed heptachlor-
sprayed forage and technical heptachlor. Jour.
Dairy Science 38: 669-672, June 1955.
Poos, F. W., J. L. Allsion, and K. W. Kreitlow.
The clover root borer (Hylastinus obscurus) as a
vector of southern and northern anthracnoses of
VOLUME 91, NUMBER 2
96.
Oe
98.
red clover. Plant Dis. Rptr. 39: 183, Feb. 15,
1955.
Yeomans, A. H. and F. W. Poos. Spray distri-
bution with boom sprayers. U.S.D.A., ARS-33-
8, 7 p., June 1955.
Poos, F. W. Studies of certain species of Chae-
tocnema. J. Econ. Ent. 48: 555-563, Oct. 1955.
Poos, F. W. Some forage insect problems in the
western United States. Joint Comt. Grassland
Farming Proc. 1955: 41-43.
99.
100.
101.
102.
303
Poos, F. W. Citation of merit. Bull. Ent. Soc.
Amer. 9(4): 272, Dec. 1963.
App, B. A., J. McGuire, and F. W. Poos. Francis
Marion Wadley 1892-1969. Proc. Ent. Soc. Wash.
72(2): 270-271, June 1970.
Poos, F. W. Walter Harrison Larrimer, 1889-
1970. J. Econ. Ent. 64(1): 345, Feb. 15, 1971.
Vance, A. M., A. S. Hoyt, and F. W. Poos. Clyde
Monroe Packard, 1889-1971. J. Econ. Ent. 65(5):
1531, Oct: 16, 1972.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 304-306
OBITUARY
Donald Joyce Borror
1907-1988
With extreme sadness I learned of the
death of Dr. Donald J. Borror, Professor
Emeritus of Entomology at The Ohio State
University, at age 80, on April 28, 1988.
He was born on August 24, 1907 in Shep-
ard, Franklin Co., Ohio, as the second son
of the Reverend Charles H. Borror. He at-
tended Otterbein College and received his
B.S. degree in 1928, the M.S. degree in
Entomology in 1930 at The Ohio State Uni-
versity, and his Ph.D. degree in Entomology
at the Ohio State University in 1935; for
the latter two degrees he worked on the sys-
tematics of dragonflies. He was a member
of the faculty of the Department of Zoology
and Entomology at The Ohio State Uni-
versity until his retirement in 1978, except
for duty with the Navy during World War
II. At The Ohio State University he taught
general and systematic entomology, insect
morphology, ornithology, and bioacoustics.
He was senior author of a textbook “An
Introduction to the study of Insects” and of
‘A Field Guide to the Insects.”’ He has pro-
duced several records and cassettes of bird
songs, and founded the Ohio State Borror
Laboratory of Bioacoustics.
Dr. Borror was a Fellow of the Entomo-
logical Society of America, Chairman of the
teaching section (1955), Assistant Managing
Editor of the Entomological Society of
America Annals (1943-44), and received the
North Central Branch Award of Merit in
1974. He was a Fellow of the American As-
sociation for the Advancement of Science
and of the Ohio Academy of Science, a
member of the Society of Systematic Zo-
ology, Sigma Xi, the American, Wilson and
Cooper Ornithological Societies, and the
National Audubon Society
My association with Dr. Borror began
when I entered graduate school at The Ohio
VOLUME 91, NUMBER 2
State University in 1957. New students in
Entomology were told by older students
what to expect of various instructors and
their classes; these sessions invariably con-
centrated on Dr. Borror and his classes and
were a great aid in preparation for what lay
ahead. The demands placed on the student
by Dr. Borror were greater than those 1m-
posed by any other college instructor that I
have ever had. He expected his students to
work as hard and efficiently as he did. It is
true that not all of Dr. Borror’s students
relished the experience, but for me it was
exactly what I was there for, and I reveled
in it.
In my first meeting with Dr. Borror, I
claimed a good knowledge of beetles. A look
of scepticism followed, whereupon he im-
mediately arose, walked over to a shelf,
pulled out a box of assorted beetles, pointed
to one, and said, ‘““What family is this one?”
One by one, I correctly named the family
to which each belonged. Dr. Borror was one
to base a conclusion on sound evidence only,
not on hearsay.
Those who assisted in preparation of
specimens for his morphology classes were
always amazed at his ability to recall details
of the morphology collection. Typical pre-
class instructions to the assistant by Dr.
Borror would go as follows: ““Now we have
just four specimens of this family, and I
collected two of them in Maine, I want you
to put out two for the class to examine.”
The assistant would invariably find that Dr.
Borror’s memory was accurate. This may
not seem impressive to a non-entomologist,
but there are nearly 580 families of insects
in North America, and over 90% of them
were represented in the morphology collec-
tion.
One graduate student enjoyed putting the
teaching style of the instructors in graph
form. A dot at each end of a piece of paper
represented the subject matter to be taught.
The line drawn from one side of the paper
to the other represented the various paths
taken by the instructors during the lecture.
305
The line of an instructor who enjoyed an
initial chat with students started to one side
of the dot, then approached and joined the
mid region to end up on the final dot. It was
amusing to see this student plot various
manners of lines (meandering, jagged, etc.),
then say, who is this? We would often cor-
rectly respond, ““Oh, that’s Dr. - - - -.”” The
line for Dr. Borror’s style was immediately
guessed by one and all: a solid, perfectly
straight line from one dot to the other.
A graduate department is only as good as
its instructors, and to me Dr. Borror made
the systematic and morphology section of
Entomology at The Ohio State University
truly outstanding. I do not see how better
training could have been given than that
which he provided. He was by far the great-
est academic influence in my life.
I was very fortunate to have been a grad-
uate student at The Ohio State University
when Dr. Borror was approached by a rep-
resentative of Houghton Mifflin and asked
to do an Insect Guide for the Peterson Field
Guide Series. He was not one to turn down
a challenge, but needed an artist to do the
drawings that he could not handle. It is dif-
ficult to express the extreme feeling of sat-
isfaction and joy experienced when Donald
Borror asked me to work with him on the
insect guide. Work on the insect guide got
started in 1964; the work was completed in
1967. Field guide authors must invariably
do the work in the midst of other activities,
and it typically takes many years to produce
a guide. It is rare that one is produced as
rapidly as the 3'2 years that 1t took Donald
Borror and I to do this one.
A reviewer of the insect guide predicted
that the book would sell more copies than
any other book on insects ever written; that
reviewer's prediction has doubtless been
borne out. For the period of 1970 to June
15, 1988, the insect guide has sold 412,790
copies, a level of sales never expected by
anyone connected with the project.
The association with Donald Borror gives
me, I feel, a full understanding of the intel-
306
lectual capabilities of true genius. I was not
alone in being much impressed with him.
In a letter of October 12, 1964, Roger Pe-
terson remarked on his construction plans
for the insect guide as follows: ““What a first-
rate worker you are! A real pro! I have looked
over your construction plans for the insect
field guide with great interest, and they are
going to work out just fine.” Dr. Borror’s
handling of the work on the insect guide was
truly masterful. Helen Phillips of Houghton
Mifflin referred to him as the most profi-
cient field guide author with whom she had
ever worked, and noted that she had found
only three typographical errors in more than
1500 pages of typing that he did for the
insect guide.
I enjoy reflecting on encounters with
Donald Borror in the halls of the Botany
and Zoology building at The Ohio State
University. He is striding along at near top
speed, gives you a warm grin, says ““How-
dy,” then continues unslowed to the task at
hand. The image is of a strong, determined,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and quiet (even shy) individual, and a true
leader who set a high standard of excellence
for colleagues and students alike in all work
he did.
To have been a student of, and worked
with Donald Borror was a grand intellectual
experience. It was, for me, the best of all
possible times. My association with him was
one that I will cherish as much as any that
I have ever had. I say with great pride that
I had the privilege and honor of having
known and worked with a truly great man.
He will be missed by all who knew him.
Richard E. White, Systematic Entomol-
ogy Laboratory, Plant Sciences, Agr. Res.
Sery., USDA, % U.S. National Museum of
Nat. Hist., Washington, D.C. 20560.
Photo courtesy of Frank W. Mead, Fla. Dept. Agr.
and Consumer Serv., Div. Plant Ind., Gainesville.
ENTOMOLOGICAL SOCIETY OF WASHINGTON
New MEMBERS FOR 1988
Masaki Abe
Alfred P. Arthur
Art Borkent
David Carmean
Christopher H. Dietrich
Robert M. Eisenberg
John D. Glaser
Harold J. Harlan
Lawrence E. Hurd
Edward A. Lisowski
Hans K. Loechelt
Robin Trevor Lubbert
Terry D. Miller
Stephen R. Moulton II
Mark A. Muegge
Allen L. Norrbom
Roy A. Norton
Robin J. Rathman
Eric Wellington Riddick
Thomas E. Rogers
Sara S. Rosenthal
John D. Sedlacek
Gary J. Steck
Brian M. Wiegmann
Richard C. Wilkerson
Total new members for 1988: 25
Total membership as of 1 December 1988:
547
Submitted by Geoffrey B. White, Member-
ship Chairman, 944th Regular Meeting, |
December 1988.
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 307-310
SociETY MEETINGS
941st Regular Meeting— May 5, 1988
The 94Ist Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Gene Wood in the
Naturalist Center, National Museum of
Natural History, at 8 p.m. on 5 May 1988.
Eighteen members and three guests were
present. Minutes of the previous meeting
were read and approved with one alteration.
No old business was transacted, nor were
there any applicants for membership.
F. C. Thompson distributed handouts de-
scribing the location and program of this
year’s annual banquet, which will be jointly
sponsored by our own Society, the Pest Sci-
ence Society of Washington, and the Mary-
land Entomological Society. This year the
Master of Ceremonies will be Ronald J.
McGinley, Chairman, Department of Ento-
mology, Smithsonian Institution.
R. G. Robbins distributed lists of Chinese
papers on ticks and mosquitoes that have
been translated by his wife Fu-meei. Among
these translations are several major biogeo-
graphic surveys, such as Teng Kuo-fan’s
“Ticks of Tibet” and Ma Su-fang’s “Studies
on the Anopheles (A.) sinensis group of mos-
quitoes in China.” Persons interested in ob-
taining either lists or translations should
contact the Recording Secretary at the ad-
dress on the inside front cover of this issue
or call (301) 423-5693 during regular busi-
ness hours, E.S.T.
The speaker for the evening was Lance
A. Durden, Research Collaborator, De-
partment of Entomology, Smithsonian In-
stitution, whose talk was entitled ‘Project
Wallace: Bioendemism, Mammal Ectopar-
asites and Some Other Insects in Sulawesi.”
Dr. Durden illustrated his presentation with
many beautiful slides as well as an exhibit
of insects that he had collected in northern
Sulawesi. Don R. Davis of the Smithsoni-
an’s Department of Entomology was de-
lighted to discover in this collection a single
specimen of the rare and unusual tineid moth
Ischnuridea virginella Sauber, on which he
has recently published. Until Davis collect-
ed four specimens on Mindanao in 1965,
this species had only been known from the
damaged female type, also collected on
Mindanao in 1882. Like the Durden ex-
ample, all subsequent specimens have been
taken on Sulawesi by Project Wallace staff.
Though a member of the clothes moth fam-
ily Tineidae, [schnuridea is atypical in pos-
sessing an extremely long, piercing ovipos-
itor that is thought to be an adaptation for
depositing eggs into the pithy stems of living
Musa (i.e. bananas, etc.). Dr. Durden has
graciously donated his specimen to the
Smithsonian Institution.
Dr. Davis also announced that he is or-
ganizing an entomological delegation to the
People’s Republic of China, which will de-
part on or about 9 September of this year.
Participants will tour most major entomo-
logical collections in mainland China as well
as the panda research station. About half of
each day will be spent visiting points of nat-
ural or historical interest, but there may also
be opportunities for field work. All ento-
mologists, regardless of afhliation, will be
warmly welcomed and each is invited, at
his option, to deliver a short talk on his area
of expertise. Though systematic entomol-
ogy will be emphasized, Don 1s also inter-
ested in hearing from anyone with experi-
ence in biocontrol. The cost per person is
$3550 round trip from San Francisco, but
up to 80% of this amount may be tax-de-
ductible.
Mignon Davis complimented Jil Swear-
ingen, Warren Steiner, and Doug Suther-
land for bringing such a tempting array of
refreshments to our final meeting before the
308
summer recess. In happy anticipation of this
repast, the meeting was adjourned at 9:10
p.m.
Richard G. Robbins, Recording Secretary
942nd Regular Meeting— October 6, 1988
The 942nd Regular Meeting of the En-
tomological Society of Washington was
called to order by President Gene Wood in
the Naturalist Center, National Museum of
Natural History, at 8 p.m. on 6 October
1988. Twenty members and 14 guests were
present. Minutes of the May meeting were
read and approved.
D. H. Messersmith announced that he has
been appointed to chair a committee to study
the feasibility of affording affiliate status to
other entomological organizations for the
purpose of publishing summaries of their
minutes or brief notices of their activities
in our Proceedings. M. S. Collins strongly
endorsed this idea. However, T. J. Spilman
reminded the membership that he had
voiced a similar suggestion in the early
1960’s but that there had been no follow-
through. Dr. Messersmith would welcome
additional comments on such intersocietal
cooperation.
Membership Chairman G. B. White read
the names of the following applicants for
membership: Masaki Abe, Systematic
Entomology Laboratory, USDA, Smithson-
ian Institution; David Carmean, Depart-
ment of Entomology, Oregon State Univer-
sity, Corvallis; Harold J. Harlan,
Crownsville, Maryland; Robin Trevor Lub-
bert, Beltsville, Maryland; Mark A. Muegge,
Department of Entomology and Plant Pa-
thology, University of Tennessee, Knox-
ville; Eric Wellington Riddick, Washington,
D.C.; Thomas E. Rogers, Kenner, Louisi-
ana; Sara S. Rosenthal, Bozeman, Montana;
Brian M. Wiegmann, Department of Ento-
mology, University of Maryland, College
Park; and Richard C. Wilkerson, Walter
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Reed Biosystematics Unit, Smithsonian In-
stitution.
W.E. Bickley asked whether nominations
had been received for a third Honorary
Member. President Wood replied that nom-
inations would be considered at a forthcom-
ing meeting of the Executive Committee.
The President then asked to borrow any en-
tomological cartoons sensu “The Far Side”
that members or their friends would like to
see exhibited at this year’s National Con-
ference of the Entomological Society of
America, to be held in Louisville, Ken-
tucky. Original drawings should be sent to
Dr. Wood no later than mid-October.
J. H. Fales displayed and discussed the
lepidopterological fruits of his summers
afield: striking aberrations of several mid-
Atlantic butterfly species; and a series of the
late season dark form of Junonia coenia
Hiibner, which has reddish hind wings ven-
trally. Fales also exhibited a specimen of
Euphyes ruricola metacomet (Harris), taken
for the first time in Rock Creek Park, Wash-
ington, D.C., on 10 August of this year. On
19 September, Fales was at Plum Point, Cal-
vert County, Maryland, where he collected
a single Danaus plexippus (Linnaeus)
marked with white paint along the ventral
external margins of the left fore- and hind
wings. He has added this specimen to his
personal collection of marked butterflies and
other insects, which was also shown to the
membership.
The speaker for the evening was Candy
Feller, Research Collaborator, Department
of Entomology, Smithsonian Institution,
who is well known as a former free-lance
artist at the National Museum of Natural
History and is now a graduate student at
Georgetown University. Her talk, entitled
“Insects on Mangrove Cays: Preliminary
Studies in Belize,” revealed the remarkable
diversity of insects and other creatures as-
sociated with these polyphyletic trees and
shrubs that fringe tropical tidal shores.
T. J. Spilman announced the recent death
of Jack E. Lipes (18 February 1924-17 Sep-
VOLUME 91, NUMBER 2
tember 1988), USDA-APHIS, a much ad-
mired member of this Society and our Pres-
ident in 1981.
Visitors were introduced and the meeting
was adjourned at 9:25 p.m. Refreshments
followed.
Richard G. Robbins, Recording Secre-
tary, Geoffrey B. White, Membership Chair-
man
943rd Regular Meeting— November 3, 1988
The 943rd Regular Meeting of the En-
tomological Society of Washington was
called to order by President Gene Wood in
the Naturalist Center, National Museum of
Natural History, at 8:10 p.m. on 3 Novem-
ber 1988. Eighteen members and 6 guests
were present. Minutes of the October meet-
ing were read and approved.
Mignon Davis circulated a sign-up sheet
for members who would like to volunteer
to bring refreshments to our meetings.
President Wood announced that the So-
ciety had twice received the name of Alan
Stone for Honorary Membership and that
the Executive Committee had unanimously
agreed to nominate Dr. Stone in recognition
of his distinguished career in mosquito sys-
tematics. Dr. Wood next placed this nom-
ination before the ESW audience, where it
was approved by acclamation.
Membership Chairman G. B. White read
the name of the following applicant for
membership: Allen L. Norrbom, System-
atic Entomology Laboratory, USDA,
Smithsonian Institution.
President Wood, assisted by D. M. An-
derson, unveiled a provisional slate of of-
ficers for 1989: President-Elect, vacant:
Treasurer, Norman E. Woodley; Editor,
Hiram G. Larew; Associate Editor, vacant;
Recording Secretary, Richard G. Robbins;
Corresponding Secretary, John M. King-
solver; Program Chairman, Warren E. Stei-
309
ner, Jr.; Membership Chairman, Geoffrey
B. White; Custodian, Anne M. Wieber. Dr.
Wood then announced the immediate for-
mation of a Nominating Committee to fill
the vacancies in this list.
At the request of President Wood, R. G.
Robbins noted this Society’s heartfelt con-
cern for the health of Robert Traub, the
world’s foremost authority on Siphonaptera
(fleas) and an ESW member since 1947. Of
Dr. Traub’s 156 peerless publications on
ectoparasites, 9 have graced the pages of our
Proceedings. The officers and members of
the Entomological Society of Washington
take this opportunity to wish Dr. Traub a
full and speedy recovery from his current
illness.
Gaye Williams exhibited elegant copper
wire sculptures of a mosquito and a scor-
pion, both crafted by the Kentucky artist
Ken Carns.
R. G. Robbins projected a series of color
slides showing males and females of 10 Af-
rican tick species in the genus 4mb/yomma
that are known to be capable of transmitting
heartwater (Cowdria ruminantium), a dead-
ly rickettsial disease of livestock that occurs
throughout sub-Saharan Africa and has be-
come established in the eastern Caribbean.
The slides were made directly from a set of
beautiful drawings by A. Olwage, who il-
lustrated the vectors of heartwater for a 1986
symposium published by the Onderstepoort
Journal of Veterinary Research (54: 161-
546, September 1987). A copy of this work
was also exhibited.
The speaker for the evening was Marc
Epstein, Postdoctoral Fellow, Department
of Entomology, Smithsonian Institution,
whose talk was entitled ““Lepidopteran Slugs:
A Natural History of the Limacodidae.”
Drawing on his extensive field experience,
Dr. Epstein reviewed the morphological and
developmental diversity 1n this pantropical
family of some 1200 species. Limacodid lar-
vae are ontogenetically remarkable: emerg-
ing from scale-like eggs, they are polypha-
gous but prefer late season, tannin-enriched
310 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
foliage and undergo 7 strikingly dissimilar
instars before pupating in solitary or com-
munal pyriform cocoons.
Ben Pagac appealed for new subscribers
to Cicindela, a quarterly journal devoted to
tiger beetles that is now in its 20th year but
whose future may be in jeopardy. Several
back issues were displayed to the member-
ship.
Visitors were introduced and the meeting
was adjourned at 9:20 p.m. Refreshments
followed.
Richard G. Robbins, Recording Secretary
REPORTS OF OFFICERS
Treasurer’s Report
SUMMARY FINANCIAL STATEMENT FOR 1988
Special
General Publications Total
Fund Fund Assets
Assets: November 1, 1987 $13,839.27 $66,051.13 $79,890.40
Total Receipts for 1988 60,899.98 7,105.05 68,005.03
Total Disbursements for 1988 53,152.66 0.00 53,152.66
Assets: October 31, 1988 21,586.59 73,156.18 94,742.77
Net Changes in Funds $ 7,747.32 7,105.05 14,852.37
Norman E. Woodley, 7reasurer
CORRESPONDING SECRETARY'S
REPORT FOR 1988
Eight letters were written thanking guest
speakers, 20 were sent to contributors to our
Special Publication Fund, eight were sent in
reply to requests for information, and 25
letters welcomed new members. The post-
age costs were $16.00.
John M. Kingsolver, Corresponding Sec-
retary
Epitor’s REPORT
Fifty-eight articles, nine notes and eleven
book reviews were published in The Pro-
ceedings in 1988 for a total of 532 pages.
The Society paid page charges for four of
the articles, and covered the cost of pub-
lishing all book reviews.
The variety of topics covered remained
great. Many articles dealt with systematics
of a group, while others discussed mor-
phology, host-relations, or behavior. Al-
though most authors were from the U.S.,
three first authors were from foreign coun-
tries.
The Editor wishes to thank the nearly sev-
enty reviewers who assisted in improving
all manuscripts. The Publications Commit-
tee and Book Review Editor were of special
assistance during the year, as was the pre-
vious Editor.
Hiram G. Larew, Editor
ee
MEMBERSHIP LIST OF THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
The Bylaws of the Entomological Society
of Washington require the Corresponding
Secretary to publish a membership list every
three years. The previous list was published
February 1, 1986 with 632 members; the
present list contains 547 members from
every state in the union except Alaska, Ar-
kansas, and Maine. The largest represen-
tation is in Maryland (91), followed by The
District of Columbia (51), California (40),
Florida (25), Virginia (22), and Pennsyl-
vania (14). The figures for jurisdictions in
the Washington, D.C. area are slightly
skewed since several members receive their
Proceedings at an office address. Twenty-
two other countries in every continent ex-
cept Antarctica are represented.
The format used in this list follows that
of the 1986 list. Names of Honorary Mem-
bers are capitalized, those of Emeritus
Members are italicized, and Life Members
are distinguished by an Asterisk (8) follow-
ing the date they joined the Society. Dates
of election to Honorary or Emeritus status
are entered in parentheses. In 1988, Dr.
Curtis W. Sabrosky was elected to Honorary
President, and Dr. Louise Russell and Dr.
Alan Stone were elected to Honorary mem-
bership.
I thank Dr. and Mrs. Wayne Mathis for
their assistance in the preparation of this
list. Any corrections to the list can be sent
to the Corresponding Secretary at the ad-
dress on the inside front cover of this issue.
Corrections will be read at the next meeting
of the Society, and will be published in the
Proceedings by the Recording Secretary.
Abe, M. 1988 District of Columbia
Abercrombie, J. 1975 New York
Adams, J. R. 1963 Maryland
Adams, M. S. 1983 New York
Adamski, D. 1984 Mississippi
Adler, P. H. 1986 South Carolina
Adler, V. E. 1961 (1987) Maryland
PROC. ENTOMOL. SOC. WASH.
91(2), 1989, pp. 311-317
Aitken, T. H. G. 1957 (1984) Connecticut
Aldrich, J. R. 1983 Maryland
Anderson, D. M. 1954 Maryland
Anderson, L. D. 1944 (1989) California
Anderson, W. H. 1937 Maryland
App, B. A. 1952 Florida
Appel, A. G. 1983 Alabama
Arduser, M. S. 1985 Missouri
Armitage, B. J. 1983 Alabama
Arnaud, P.H. 1955 California
Arnett, R. H., Jr. 1980 Florida
Arthur, A. P. 1988 CANADA
Ashlock, P. D. 1958 Kansas
Austin, D. F. 1977 Florida
Bacon, N. 1982 Pennsylvania
Baker, E. W. 1944 Maryland
Baker, G. T. 1987 Mississippi
Ball, G. E. 1948 CANADA
Barber, K. N. 1985 CANADA
Barnes, J. K. 1979 New York
Barnum, A. 1956 Utah
Barr, A. R. 1951 California
Barr, W. F. 1948 Idaho
Barrows, E. M. 1976 District of Columbia
Batra, S. W. T. 1985 Maryland
Baumann, R. W. 1973 Utah
Beal, R. S., Jr. 1958 Colorado
Bechtel, R. C. 1960 Nevada
Beck, T. R. 1979 Maryland
Becker, E. C. 1951 CANADA
Becker, V. O. 1987 BRAZIL
Bell, R. T. 1955 Vermont
Bellinger, R. G. 1972 South Carolina
Bergman, P. W. 1966 Virginia
Berry, R. L. 1972 Ohio
Bezark, L. G. 1974 California
Bicha, W. 1981 Ohio
Bickley, W. E. 1949* Maryland
BISSELL, T. L. 1941 (1983) Maryland
Blackburn, V. L. 1984 Maryland
Blom, P. E. 1986 Idaho
Boese, J. L. 1977 District of Columbia
Boettcher, R. A. 1955 District of Columbia
Bohart, R. M. 1944 California
Bohnsack, K. K. 1958 California
Boldt, P. E. 1987 Texas
Borkent, A. 1988 CANADA
Bouseman, J. K. 1988 Illinois
312 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Bowen, T. W. 1980 North Carolina
Brigham, W. U. 1986 Illinois
Brou, V. A. 1985 Louisiana
Brown B. V. 1982 CANADA
Brown, F. M. 1975 Colorado
Brown, H. P. 1977 Oklahoma
Brown, R. L. 1979 Mississippi
Brushwein, J. R. 1987 Florida
Bueno-Soria, J. 1977 MEXICO
Burger, J. F. 1975 New Hampshire
Burke, H. R. 1981 Texas
Burks, B. D. 1938 Arizona
Burns, J. M. 1975* District of Columbia
Burrows, W. L. 1983 West Virginia
Butler, L. 1966 West Virginia
Byers, G. W. 1984 Kansas
Calabrese, D. M. 1981 New Jersey
Carlson, R. W. 1970* Maryland
Carmean, D. 1988 Oregon
Carpenter, J. M. 1984 Massachusetts
Carroll, J. F. 1977 Maryland
Casdorph, D. G. 1977 California
Cave, R. 1977 HONDURAS
Chapin, J. B. 1973 Louisiana
Charpentier, P. O. 1962 Connecticut
Chemsak, J. A. 1987 California
Chilson, L. M. 1977 Maryland
Chittick, H. A. 1946 New York
Christiansen, T. A. 1986 Wyoming
Clarke, W. E. 1975 Alabama
Clarke, J. F. G. 1936 District of Columbia
Clement, S. L. 1987 Washington
Cochran, D. G. 1981 Virginia
Coffman, C. C. 1980 West Virginia
Cohen, N. Y. 1983 Maryland
Cokendolpher, J. C. 1987 Texas
Connell, J. G. 1988 Indiana
Contreras-Ramos, A. 1986 Alabama
Cook, D. R. 1952 Michigan
Cooper, K. W. 1955 California
Corpus, L. D. 1985 Mississippi
Coulson, J.R. 1961 Maryland
Couri, M. S. 1987 BRAZIL
Courneya, P. 1986 Texas
Courtney, G. W. 1985 CANADA
Covell, C. V., Jr. 1986 Kentucky
Craig, G. B. 1954 Indiana
Cross, H. F. 1954 Georgia
Cur¢ic, B. P. M. 1982 YUGOSLAVIA
Currado, I. 1978 ITALY
Darling, D. C. 1981 CANADA
Darsie, R. F., Jr. 1949 South Carolina
Davidson, J. A. 1957 Maryland
Davis, D. R. 1961 District of Columbia
Davis, J. R. 1985 Texas
Davis, L. G. 1952 Virginia
Davis, M. M. 1979 District of Columbia
Deeming, J. C. 1974* UNITED KING-
DOM
Dietz, L. L. 1982 North Carolina
Dennis, S. 1979 Colorado
Deyrup, M. A. 1979 Florida
Dicke, F. F. 1933 Iowa
Dietrich, C. H. 1988 North Carolina
Dodson, G. 1987 AUSTRALIA
Donnelly, T. W. 1962 New York
Downes, W. G. 1981 Connecticut
Downes, W. L. 1985 Michigan
Doyen, J. T. 1983 California
Dozier, H. L. 1952* South Carolina
Drea, J. J. 1984 Maryland
Drummond, R. O. 1954 (1987) Texas
Durden, L. A. 1987 Maryland
Easton, E. R. 1985 South Dakota
Edmunds, G. F., Jr. 1951 Utah
Ehler, L. E. 1987 California
Eikenbary, R. D. 1979 Oklahoma
Eisenberg, R. M. 1988 Delaware
Elias, M. K. 1972 Maryland
Emerson, K. C. 1952 Florida
Enns, W. R. 1960 Missouri
Erwin, N. 1986 Maryland
Erwin, T. L. 1972 District of Columbia
Evans, E. W. 1984 Utah
Evans, H. E. 1948 Colorado
Evans, W. G. 1957 CANADA
Evenhuis, N. L. 1980 Hawaii
Fairchild, G. B. 1934 Florida
Fales, J. H. 1944 Maryland
Fee, F. D. 1983 Pennsylvania
Ferguson, D. C. 1969 District of Columbia
Fisher, E. M. 1977 California
‘isk, F. W. 1968 (1988) Florida
Flint, O. S., Jr. 1961 District of Columbia
Fluno, J. A. 1957 Florida
Foote, B. A. 1958 Ohio
VOLUME 91, NUMBER 2
Foote, R. H. 1950 Virginia
Forattini, O. P. 1956 BRAZIL
Foster, J. R. 1853 Maryland
Fox, I. 1936 Puerto Rico
Franclemont, J. G. 1947 New York
Freidberg, A. 1979 ISRAEL
Freytag, P. H. 1979 Kentucky
Gagne, R. J. 1966* District of Columbia
Gentry, J. W. 1958 Florida
Gerberg, E. J. 1953 Maryland
Ghorpade, K. 1986 INDIA
Gibson, L. P. 1981 Ohio
Giles, F. E. 1981 Maryland
Gill, G. D. 1958 Michigan
Gimpel, W. F., Jr. 1970 Maryland
Glaser, J. D. 1988 Maryland
Godfrey, G. L. 1971 Illinois
Goeden, R. D. 1982 California
Gordh, G. 1975 California
Gordon, R. D. 1968 District of Columbia
Gorham, J. R. 1974 District of Columbia
Grabowski, W. B. 1970 New Mexico
Grace, J. K. 1987 CANADA
Gregg, R. E. 1945 Colorado
Grissell, E. E. 1979 District of Columbia
Grogan, W. L. 1974 Maryland
Gunther, R. G. 1981 Illinois
Guzman, L. E. P. 1988 CHILE
Habeck, D. H. 1957 Florida
Hacker, J. D. 1971 West Virginia
Hagen, K. S. 1949 California
Haile, D. L. 1986 Pennsylvania
Haines, K. A. 1952 Virginia
Halstead, J. A. 1983 California
Hamilton, S. W. 1982 Tennessee
Hannemann, H.-J. 1979 EAST GER-
MANY
Hanson, P. 1985 COSTA RICA
Hansson, C. 1985 SWEDEN
Harbach, R. E. 1972 District of Columbia
Hardy, A. R. 1974 California
Harlan, H. J. 1988 Maryland
Harman, D. M. 1966 Maryland
Harris, S. C. 1979 Alabama
Harrison, B. A. 1976 District of Columbia
Harrison, S. J. 1985 Maryland
Haskins, C. P. 1945 District of Columbia
Hawkins, B. A. 1983 UNITED KINGDOM
313
Hayes, D. K. 1970 Maryland
Hendricks, H. J. 1987 Virginia
Hendrickson, R. M., Jr. 1978 Delaware
Henry, C. S. 1975 Connecticut
Henry, T. J. 1975 District of Columbia
Heppner, J. B. 1974 Florida
Heraty, J. M. 1986 Texas
Hespenheide, H. A., III 1981 California
Hevel, G. F. 1970 District of Columbia
Heydon, S. L. 1986 Maryland
Hodges, R. W. 1960* District of Columbia
Hoebeke, E. R. 1980 New York
Hoffmann, C. H. 1945 Maryland
Hoffman, K. M. 1986 South Carolina
Holzbach, J. E. 1983 Ohio
Holzenthal, R. W. 1985 Minnesota
Hoover, G. A. 1985 Pennsylvania
Hopla, C. E. 1961 Oklahoma
Hopper, H. P. 1978 District of Columbia
Howden, H. F. 1948 CANADA
Huang, Y.-M. 1968 District of Columbia
Hung, A. C. F. 1981 Maryland
Hurd, L. E. 1988 Delaware
Huryn, A. D. 1984 Georgia
Husband, R. W. 1973 Michigan
Imai, E. M. 1983 Maryland
Irwin, M. E. 1976 Illinois
Ivie, M. A. 1984 Montana
Jackson, D. L. 1966 Ohio
Jacobi, H. 1987 Maryland
Jakob, W. L. 1977 Colorado
Jenkins, J. 1987 Michigan
Johnson, J. B. 1987 Idaho
Johnson, M. D. 1973 Indiana
Johnson, N. F. 1980 New York
Johnson, P. J. 1984 Wisconsin
Jones, R. H. 1955 Colorado
Jones, S. R. 1985 Pennsylvania
Joseph, S. R. 1957 Maryland
Kaster, C. H. 1979 Michigan
Keirans, J. E. 1984 District of Columbia
Kelley, R. W. 1982 South Carolina
Kennedy, J. H. 1977 Texas
Kethley, J. B. 1874 Illinois
Kim, K. C. 1983 Pennsylvania
Kingsolver, J. M. 1963 Maryland
Kirchner, R. F. 1981 West Virginia
Kitayama, C. 1974 California
314 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Kittle, P. 1975 Alabama
Kliewer, J. W. 1983 Virginia
Knipling, E. F. 1946 (1985) Maryland
Knisley, C. B. 1981 Virginia
Knudson, E. C. 1982 Texas
Knutson, L. V. 1963* District of Columbia
Korytkowski, C. A. 1982 PANAMA
Kostarab, M. 1978 Virginia
Krombein, K. V. 1941* District of Colum-
bia
Kurczewski, F. E. 1970 New York
Lago, P. K. 1984 Mississipp1
Lamb, N. J. 1978 Florida
Lambdin, P. L. 1974 Tennessee
Larew, H. G. 1982 Maryland
LaSalle, J. 1982 California
Lassman, G. W. 1946 MEXICO
LaTorre, L. T. 1984 West Virginia
Lavigne, R. 1972 Wyoming
Levesque, C. 1985 CANADA
Lewis, P. A. 1974 Ohio
Lewis, R. E. 1958 Iowa
Lien, J. C. 1967 REPUBLIC OF CHINA
Linam, J. 1962 Colorado
Linkfield, R. L. 1959 New Jersey
Lippert, E. A. 1986 CANADA
Lisowski, E. A. 1988 Illinois
Loeschelt, H. K. 1988 Washington
Longair, R. W. 1985 CANADA
Lubbert, R. T. 1988 Maryland
Lyon, R. J. 1961 California
Mabry, J. E. 1954 Florida
MacDonald, J. F. 1984 Indiana
MacKay, W. P. 1982 Texas
MacRae, T. C. 1987 Missouri
Magner, J. M. 1953 Missouri
Maier, C. T. 1976 Connecticut
Main, A. J., Jr. 1965 Connecticut
Maldonado-Capriles, J. 1947 Puerto Rico
Mangan, R. L. 1977 Texas
Manglitz, G. R. 1956 (1989) Nebraska
Manley, D. G. 1984 South Carolina
Manuel, K. L. 1983 North Carolina
Marsh, P.M. 1960 District of Columbia
Marshall, S. 1982 CANADA
Masner, L. 1967 CANADA
Mason, H. C. 1949 (1973) Maryland
Mason, W. R. M. 1970 CANADA
Mathieu-Veillard, J. M. 1982 MEXICO
Mathis, W. N. 1976 District of Columbia
Mawdsley, J. 1986 Virginia
McCabe, T. L. 1977 New York
McCafferty, W. P. 1968 Indiana
McComb, C. V. 1956 Virginia
McDaniel, B. 1964 South Dakota
McDonald, F. J. D. 1983 AUSTRALIA
McGovran, E. R. 1937 (1973) Maryland
McGuire, J. U., Jr. 1954 (1980) PUERTO
RICO
McMurtry, J. A. 1986 California
McPherson, J. E. 1985 Illinois
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Menke, A. S. 1969 District of Columbia
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Miller, D. D. 1987 Maryland
Miller, G. L. 1981 Alabama
Miller, J. 1987 Oregon
Miller, R. M. 1974 SOUTH AFRICA
Miller, R. S. 1981 Ohio
Miller, S. E. 1980 Hawaii
Miller, T. D. 1988 Idaho
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Mitter, C. 1987 Maryland
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Molineaux, M. J. 1986 Maryland
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Moron, M. A. 1985 MEXICO
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Moser, J. C. 1973 Louisiana
Moulton, S. R., I] 1988 Maryland
Muegge, M. A. 1988 Tennessee
Munroe, E. G. 1986 CANADA
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Murdoch, W. P. 1966 Pennsylvania
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Ng, Y.-S. 1986 Virginia
VOLUME 91, NUMBER 2
Nichols, S. W. 1985 Iowa
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Paggi, A. C. 1987 ARGENTINA
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315
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316
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
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VOLUME 91, NUMBER 2
Wilson, N. 1957 Iowa
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317
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VOLUME 91, NUMBER 2 319
Return Proof to:
John R. Doe
315 State St.
Meriden, CT 06420
(203) 555-1212
A new species of Xus (Order: Family)
injurious to hollies, Ilex spp. (Aquifoliaceae)
oe and John Smith
Connecticut 06420;
(IRD) Resident Biologist, 315 State St., Meriden,
(JS) Entomologist, Hartford, Connecticut 06540.
City Parks,
is described,
illustrated,
Abstract.- Xus albus, a new species of
compared with ...
Key Words. Distribution, ornamental shrub, damage, leaf roller
Figure Legends
1-4
Figs. 1, Habitus. 2, Male genitalia (lateral view).
Xus albus.
» Larva. 4, Pupa.
ey
Fig. Damage to holly leaves.
Literature Cited
» J. and J. Smith. 1970 Holly Insects, Jones and Case. New York,
NY. 38pp-
Smith, J. and J. R. Doe 1967 A list of insects injurious to hollies
Soc. Va 38: 54-68
(lex spp.). Proc. Entomol.
(The above citations are fictitious.)
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CONTENTS
(Continued from front cover)
ROTH, L. M.—The cockroach genus Margattea Shelford, with a new species from the Krakatau
Islands, and redescriptions of several species from the Indo-Pacific region (Dictyoptera:
Blattaria: Blattellidae)
SHAFFER, J. C.—Two new phycitine moths of the genus Coenochroa (Lepidoptera: Pyralidae)
from Brazil
SHAFFER, J. C. and E. MUNROE—Type material of four African species of Notarcha Meyrick,
with designations of lectotypes and changes in synonymy (Lepidoptera: Crambidae: Pyr-
austinae)
YOUNG, D. C., B. C. KONDRATIEFF, and R. F. KIRCHNER—Description of male Ostro-
cerca Ricker (Plecoptera: Nemouridae) using the scanning electron microscope
NOTES
APPEL, A. G.—Rapid and non-destructive gender determination of nymphal and adult Cryp-
tocercus punctulatus Scudder (Dictyoptera: Cryptocercidae) 286
CARMEAN, D.—Sex and deposition of the holotype of Bareogonalos canadensis (Harrington)
(Hymenoptera: Trigonalyidae) 287
CURCIC, B. P. M. and R. N. DIMITRIJEVIC—Pedipalpal anomalies in Neobisium simoni (L.
Koch) and N. bernardi Vachon (Neobisiidae: Pseudoscorpiones: Arachnida) 289
ROBBINS, R. G.—Ticks of the subgenus /xodiopsis: First report of Ixodes woodi from man and
remarks on /xodes holdenriedi, a new junior synonym of Ixodes ochotonae (Acari: Ixodidae) 291
BOOK REVIEWS
DAVIS, D. R.—Lepidoptera Anatomy
GAGNE, R. J.—Asa Fitch and the Emergence of American Entomology
McCAFFERTY, W. P.—The Mayflies of Florida, Revised Edition
OBITUARIES
WALLENMAIER, T. E.—Frederick William Poos, Jr. 1891-1987
WHITE, R. E.—Donald Joyce Borror 1907-1988
NEW MEMBERS FOR 1988
SOCIETY MEETINGS AND REPORTS OF OFFICERS FOR 1988
MEMBERSHIP LIST
INSTRUCTION TO AUTHORS
VOL: 91 JULY 1989 NO. 3
(ISSN 0013-8797)
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
of WASHINGTON
PUBLISHED
QUARTERLY
CONTENTS
FORATTINI, O. P. and M. A. M. SALLUM—Redescription of Culex (Melanoconion) delpontei
Duret, 1968 and Cx. (Mel.) pereyrai Duret, 1967, from southern Brazil ................ 473
HUSBAND, R. W.—Two new species of Coccipolipus (Acari: Podapolipidae) parasites of Chilo-
corus spp. (Coccinellidae) from Vera Cruz and Morelos, Mexico and Florida and Wisconsin,
LUIS WANS hae 3 eB es Te aie ail aie ee enn Re aR i gn a el
LAGOY, P. K. and E. M. BARROWS—Larval-sex and host-species effects on location of at-
tachment sites of last-instar bagworms, Thyridopteryx ephemeraeformis (Lepidoptera: Psy-
CHIDZG) PEER eer OMe eRe cr eee Pete are SRN eke win. Bile by eee he oy a. 468
LEE, K. M., W. W. WIRTH, and K. L. CHAN—A new species of Dasyhelea midge reared from
drainsinesinpapore (Diptera? Ceratopoponidde),.¢ 2322 cc ck ce ete etic cence eee snes 452
LEIDY, N. A. and H. H. NEUNZIG—Taxonomic study of the larvae of six eastern North
American Dioryctria (Lepidoptera: Pyralidae: Phycitinae) ...........................-. 325
NAKAHARA, S.—A new species of A/eurotulus (Homoptera: Aleyrodidae) ................ 436
NEUNZIG, H. H.—New records of Phycitinae from Bolivia including a new species of Peadus
URE DICODIelA ve VT allGac) Eee Re Te ele Siar oe ces tes Meche oho Blac ans Bases 355
NEUNZIG, H. H. and N. A. LEIDY—A new species of Dioryctria (Lepidoptera: Pyralidae:
BUY CIsINAe) MLOM tHe SONtMeASLCENI WO NILCC StALES Was cite lass: sh avectte le ev 'aydiese «nie waists comin eles 321
POLHEMUS, J. T. and P. J. SPANGLER—A new species of Rheumatobates Bergroth from
Ecuador and distribution of the genus (Heteroptera: Gerridae) ........................ 421
ROTH, L. M.—Sliferia, a new ovoviviparous cockroach genus (Blattellidae) and the evolution
okovoviviparity 1 Blattaria\(Dictyoptera)) <. .....6 sos... od os ocd Acic,c ewe Baud betas. 441
SCHWARTZ, M. D.—Polymerus castilleja, a new mirine plant bug from California and Oregon,
with remarks on generic characters of the genus Po/ymerus Hahn (Heteroptera: Miridae)... 461
SHAFFER, J. C.—Review of Goya Ragonot and description of a new species, G. simulata from
Parana, Brazil (Lepidoptera: Pyralidae: Peoriinae) ...........000-....0-cece cece eceees 398
SHAFFER, J. C. and E. MUNROE—Type material of two African species of Herpetogramma
and one of Pleuroptya (Lepidoptera: Crambidae: Pyraustinae) ......................... 414
SPANGLER, P. J.—A new species of Neotropical water bug, Paravelia biae, from Brazil (Het-
PEGE LAsay CLUGAO) Ea eae eee OT Lr hs ck ak et AAMC Rc ale ota hv deslelee va Hedle cs 360
(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1989
F. CHRISTIAN THOMPSON President NORMAN E. WooDLey, 7reasurer
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Title of Publication: Proceedings of the Entomological Society of Washington.
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PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 321-324
A NEW SPECIES OF DIORYCTRIA
(LEPIDOPTERA: PYRALIDAE: PHYCITINAE)
FROM THE SOUTHEASTERN UNITED STATES
H. H. NEUNzIG AND NANcy ANTOINE LEIDY
Department of Entomology, North Carolina State University, Raleigh, North Carolina
27695-7613.
Abstract. — Dioryctria taedivorella, a new species of coneworm, is described, illustrated
and compared with Dioryctria merkeli Mutuura and Munroe.
Key Words:
Hedlin et al. (1981), in their investigation
of cone and seed insects of North America,
included nine species of Dioryctria Zeller in
the southeastern United States. Recently, as
a result of collecting and rearing efforts in
North Carolina, and with the assistance of
USDA Forest Service entomologists who
supplied material from other southeastern
states, we have been able to study larger
series of Dioryctria and conclude that ad-
ditional undescribed species occur in the re-
gion. In 1964, Neunzig et al., in publishing
on Dioryctria in North Carolina, used the
name zimmermani (Grote) for a species
feeding in loblolly pine (Pinus taeda L.).
They indicated, however, that the identifi-
cation of this species was tentative and that
additional taxonomic study was needed.
Mutuura and Munroe (1979) were of the
opinion that this North Carolina material,
as well as specimens from elsewhere in much
of the Southeast, belonged to their Dioryc-
tria merkeli. Having compared the type of
D. merkeli, on loan from the Canadian Na-
tional Collection, with North Carolina spec-
imens and other Dioryctria that are sup-
posedly merkeli, we concluded: (1) the name
merkeli should be restricted to certain pop-
ulations of Dioryctria mainly feeding as lar-
vae in slash pine (Pinus e/liotii Engelmann)
Dioryctria, coneworm, distribution, loblolly pine
in northern Florida, southern Georgia and
southern Mississippi, and (2) the similar,
more northern and more widespread species,
associated with loblolly pine, 1s new.
Dioryctria taedivorella, Neunzig and
Leidy, New SPECIES
Figs. 1-6
Diagnosis. —Dioryctria taedivorella is
most similar to Dioryctria merkeli. D. tae-
divorella, however, is a darker species with
the antemedial and postmedial lines less
distinct. These lines are chiefly white in D.
merkeli and mostly gray in D. taedivorella.
Also, the dorsum of the thorax of D. tae-
divorella 1s mostly brown to reddish brown,
whereas the dorsum of the thorax of D. mer-
keli is mostly pale gray with few or no brown
or reddish brown scales.
Description. — Head: frons mostly brown
or fuscous, some scales white or tipped with
white or gray, vertex reddish brown or
brownish red. Labial palpus reaching above
vertex in both sexes, mostly brown, fuscous
or black with varying amounts of white, gray,
reddish brown or brownish red. Maxillary
palpus squamous, fuscous, white and gray.
Antenna of male very weakly serrate with
abundant sensilla trichodea. Collar: brown
and reddish brown. Thorax: dorsum brown
322
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. Dioryctria taedivorella. 3.5 x 1. Habitus, holotype. 2. Male genitalia, most of left valva and aedeagus
omitted. 3, 4. Right valva, paratypes. 5. Aedeagus. 6. Female genitalia.
to reddish brown, sometimes with a few gray
scales and usually with patches of fuscous
or black scales; tufts of reddish brown scales
posteriorly. Forewing: above with distinct,
strongly raised scales forming subbasal, an-
temedial and postmedial patches; addition-
al smaller patches of raised scales at base of
wing and on discal spot: ground color brown;
antemedial line obscure, formed of white-
upped gray scales (some scales suffused with
red); postmedial line similar in color to an-
temedial line (medial part of line sometimes
includes patch of fuscous or black-tipped
scales); basal, subbasal, medial and terminal
area with numerous reddish brown (usually
rust-colored) scales; a few black scales in
basal area (particularly part of small patch
of raised scales), additional black scales ba-
sally and distally bordering antemedial line
and expanding into broad dark patch along
costa, basally and distally bordering post-
medial line and forming large dark costal
patch but weakly developed near inner mar-
gin, and forming terminal line. In addition
to white-tipped gray scales of transverse
lines, small patches of similarly colored
scales near costa in basal half, in medial area
and basad of terminal line; undersurface of
male with no contrastingly-colored scales,
or with a few brownish red scales near costa.
Length of wing 12.0-15.0 mm. Hindwing:
above, smoky gray, conspicuously darker
along margins. Male and female genitalia:
(Figs. 2-6) essentially like those of D. mer-
keli.
Type material.—Holotype, 4, USA, N.
CAROLINA, Lenoir Co., Kinston, US 70
Byp & Neuse R., 1-IX-1983, N. A. Leidy,
Pinus taeda cone, emerg. 13-IX-1983, gen-
italia slide 895 HHN, in USNM. Paratypes
25 6, 51 9. USA, N. CAROLINA, Lenoir
Co., Kinston, US 70 Byp & Neuse R., 28-
VIT-1983, 14-VHI-1983, N. A. Leidy, Pinus
taeda cone, emerg. 8-IX-1983, 17-[X-1983,
6-X-1983, 12-X-1983, genitalia slides 872,
883, 893, 903 HHN (1 4, 3 2); USA, N.
CAROLINA, Robeson Co., Lumberton, NC
2114 mi W of I-95, J. B. Lattay Forest Tree
Nur., 27-VII-1983, N. A. Leidy, host: Pinus
taeda cone, emerg. 5-[X-1983, 10-IX-1983,
VOLUME 91, NUMBER 3
X-1983, genitalia slides 885, 887, 891 HHN
(3 8); USA, N. CAROLINA, Robeson Co.,
nr Lumberton, J. B. Lattay For. Nur., NC
211 4 mi W of I-95, 22-VII-1985, N. A.
Leidy & M. Maynor, in Pinus taeda cone,
emerg. 2-IX-1985 to 1-X-1985 (7 4, 8 Q);
323
-_———— 1mm ———1
USA, N. CAROLINA, Granville Co., Lew-
is, US 15 0.15 mi N of SR 1424, 18-VII-
1985, N. A. Leidy, in Pinus taeda cone,
emerg. 19-VIII-1985, 22-VIII-1985 (3 2);
USA, N. CAROLINA, Wayne Co., Golds-
boro, Claridge State For., SR 1326 1.5 mi
324
N of US 70, 15-VII-1985, N. A. Leidy, in
Pinus taeda cone, emerg. 13-IX-1985 (1 8);
USA, N. CAROLINA, Onslow Co., nr
Richlands, NC 24 at SR 1230, 16-VII-1985,
N. A. Leidy, in Pinus taeda cone, emerg.
21-IX-1985 (1 6); USA, N. CAROLINA,
Onslow Co., nr Maysville, Hofmann For-
est, 25-VI-1985, NA Leidy & DJ Lodge, in
Pinus taeda cone, emerg. | 1-IX-1985 (1 3);
USA, S. CAROLINA, Berkeley Co., Fran-
cis Marion Seed Orchard, col. 23-27-VI-
1983, 2nd year cones Pinus taeda (1 4, 2 2);
USA, GEORGIA, Putnam Co., col. 9-VII-
1985, emerg. 16-IX-1985, Pinus taeda (1
2); USA, ALABAMA, Greene Co., Wey-
erhaeuser Co., col. 1985, Pinus taeda (2 4,
10 2); USA, ALABAMA, Greene Co., Wey-
erhaeuser Co., col. 15-IX-86, emerg. 26-IX-
86, 3-X-86, Pinus taeda (9 2); USA, ALA-
BAMA, Greene Co., Weyerhaeuser Co., col.
15-IX-86, Pinus taeda (8 4, 14 2); USA,
MISSISSIPPI, Perry Co., Erambert Seed
Orchard, col. 16-20-VI-1986, 2nd yr. cone
Pinus taeda (1 2). Paratypes deposited in
USNM, NCSU and BMNH.
Distribution and life history.—Known
from Virginia south to northern Florida and
west to eastern Texas. The principal host is
loblolly pine. Neunzig et al. (1964), under
the name D. zimmermani, gave a detailed
account of the biology of D. taedivorella.
Comments.—Although Mutuura and
Munroe (1979) stated that their D. merkeli
included populations associated with lob-
lolly pine throughout most of the south-
eastern United States, they restricted their
type series of D. merkeli to moths reared
from slash pine growing in northern Flori-
da, southern Mississippi, and southern
Georgia.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ACKNOWLEDGMENTS
We are especially grateful to H. O. Yates,
USDA, Forest Service, for his interest and
support. G. L. DeBarr and C. W. Fatzinger,
USDA, Forest Service, and E. P. Merkel,
USDA, Forest Service, retired, generously
made available Dioryctria from Alabama,
Florida, Georgia, Mississippi and South
Carolina. We thank the following for as-
sisting in the collection of Dioryctria in
North Carolina: T. Hardin, Federal Paper
Board Co., Lumberton; D. Lodge, formerly
N.C. State University, Raleigh; M. Maynor,
Federal Paper Board Co., Lumberton; D.
Sparkman, Federal Paper Board Co., Lum-
berton; G. Turner, Claridge State Forest,
Goldsboro; W. Wick, Champion Intl. Corp.,
Deppe. J. D. Lafontaine, Biosystematics
Research Centre, Ottawa, Canada, sent the
holotype of D. merkeli for study. Research
funds were provided in part by the USDA,
Forest Service. This is paper no. 11919 of
the Journal Series of The North Carolina
Agricultural Research Service, Raleigh,
North Carolina 27695-7643.
LITERATURE CITED
Hedlin, A. F., H. O. Yates, D. Cibrian-Tovar, B. H.
Ebel, T. W. Koerber, and E. P. Merkel. 1981.
Cone and Seed Insects of North American Coni-
fers. Joint publication: Environment Canada, Ca-
nadian Forest Service; U.S. Department of Agri-
culture, Forest Service; Secrateria de Agricultura
y Recursos Hiratilicos, México. 122 pp.
Mutuura, A. and E. Munroe. 1979. American species
of Dioryctria (Lepidoptera: Pyralidae) V. Three
new cone-feeding species from the southeastern
United States. J. Georgia Entomol. Soc. 14: 290-
304.
Neunzig, H. H., E. D. Cashatt, and G. A. Matuza.
1964. Observations on the biology of four species
of Dioryctria in North Carolina (Lepidoptera: Phy-
citidae). Ann. Entomol. Soc. Amer. 57: 317-321.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 325-341
TAXONOMIC STUDY OF THE LARVAE OF SIX EASTERN
NORTH AMERICAN DIOR YCTRIA
(LEPIDOPTERA: PYRALIDAE: PHYCITINAE)
NANcy ANTOINE LEIDY AND H. H. NEUNZIG
Department of Entomology, North Carolina State University, Raleigh, North Carolina
27695-7613.
Abstract.—Larvae of six eastern North American species of Dioryctria Zeller were
examined using both a stereo light microscope and a scanning electron microscope (SEM).
Detailed descriptions and a key to the species, supplemented with electron micrographs,
are presented for the last instar larvae of D. abietivorella (Grote), D. disclusa Heinrich,
D. pygmaeella Ragonot, D. clarioralis (Walker), D. amatella (Hulst), and D. taedivorella
Neunzig and Leidy.
Key Words:
The genus Dioryctria Zeller is mostly
Holarctic in distribution with a few species
also occurring in the northern tropics. Fif-
teen species are known from eastern North
America. Larvae feed exclusively on coni-
fers.
The adults of the North American species
of Diorycia have been the subject of exten-
sive taxonomic study beginning with Hein-
rich (1956) and Munroe (1959). In 1969,
Mutuura, working in conjunction with sev-
eral others, began a series of papers follow-
ing up on Munroe’s work (Mutuura, Mun-
roe and Ross 1969a, b, Mutuura and Munroe
1972, 1973, 1974, 1979, Mutuura 1982,
Mutuura and Neunzig 1986). Additional
work on the adults has been done by Scha-
ber and Wood (1971), Coulson et al. (1972),
Blanchard and Knudson (1983) and Neun-
zig and Leidy (1989).
Although Dioryctria larvae have been
briefly covered in various publications on
forest pests, detailed descriptions of the lar-
vae are few. MacKay (1943) described and
figured the larvae of D. reniculelloides Mu-
tuura and Munroe (as D. reniculella Grote),
Pyralidae, Phycitinae, Dioryctria, immatures, coneworm, taxonomy
and briefly differentiated it from D. abieti-
vorella (Grote) (as D. abietella Denis and
Schiffermuller). Farrier and Tauber (1953)
described and illustrated the larvae of D.
disclusa Heinrich, and Lyons (1957) also
described D. disclusa as well as D. cambi-
icola (Dyar) and D. abietivorella (as D. abie-
tella). Neunzig et al. (1964), gave detailed
descriptions of six species occurring in the
southeastern United States: D. amatella
(Hulst), D. ebe/i Mutuura and Munroe (as
D. abietella), D. clarioralis (Walker), D. dis-
clusa, D. taedivorella Neunzig and Leidy (as
D. zimmermani (Grote), and D. pygmaeella
Ragonot. Schaber (1981) published on the
larvae of D. taedae Schaber and Wood,
however, larvae of D. taedivorella and D.
yates Mutuura and Munroe were probably
included in the described material.
Only two studies of Dioryctria have made
use of scanning electron microscopy (SEM).
Bradley et al. (1982) used SEM to examine
the eggs of three species, D. disclusa, D.
amatella and D. ebeli. The early-instar lar-
vae of D. reniculelloides were examined by
Spies and Dimond using SEM (1985).
326
Fig: 1.
Part of mesothorax including D and SD
pinacula of D. disclusa, dorsolateral aspect. (150).
TP, tonofibrillary platelets; A, area shown enlarged in
Figs. 22-27.
In this paper we update the descriptions
of the last-instar larvae of six species by
utilizing the conventional light microscope
and the scanning electron microscope to
reexamine the known diagnostic characters
and to discover additional species differ-
ences.
Materials and methods. — Dioryctria lar-
vae were collected in North Carolina from
Pinus taeda L. (loblolly pine), P. palustris
Miller (long-leaf pine), P. echinata Miller
(short-leaf pine), Taxodium distichum (L.)
(bald cypress), and Abies fraseri Pursh (Poi-
ret) (Fraser’s fir). Representative specimens
of six species known to occur in eastern
North America were obtained. Portions of
each collection were preserved as larvae and
the remainder reared to obtain adults. De-
scriptions of the color of the larvae were
prepared immediately after fixing. The width
of the head was measured at the widest point,
and the length from the distal margin of the
clypeus to the most posterior point of the
epicranium. The median breadth of the
spinneret was measured across its lateral
surface.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Larvae to be examined using SEM were
dissected in 40% ethanol. The head was re-
moved, separated laterally at the point of
mandible attachment, the mandibles re-
moved, and the dorsal and ventral halves
retained intact for the next several steps.
Abdominal segments 9 and 10 and the left
half of the thorax were also removed. The
four pieces of tissue were brushed with a
soft brush to remove loose debris, placed in
a 50:50 solution of 10% Tween® 20 and
40% ethanol, and sonicated for five min-
utes. The tissue was then rinsed in 40% eth-
anol, brushed again, and transferred to flow-
through capsules of Porex® Porous Plastic
(Porex Technologies). It was then dehy-
drated in a graded series of ethanols (40%,
70%, 95%, 100%, 100%, 100%) and critical
point dried using CO, in a Tousimis® Sam-
dn®-PVT-3B. The dried tissue was then
placed on studs using SPI® Conductive
Carbon Paint (colloidal graphite in isopro-
panol 20% solids) and allowed to dry at least
two days. Excess structures and tissue were
removed using a hypodermic needle and all
surfaces carefully brushed with a soft brush.
The tissue was then coated with gold-pal-
adium in a Technics Hummer V® sputter
coater to a thickness of 25 nm, and observed
with a JEOL JSM-T200® Scanning Elec-
tron Microscope (15 kV, WD 8, spot size
9). Micrographs were made using Polaroid ®
Type 55 P/N film. Setal nomenclature as
given in Stehr (1987) is used for the body
setae (Fig. 1). The terminology used for the
mouthparts follows Crumb (1929) (Figs. 2,
3). The term tonofibrillary platelet is from
Neunzig (1979). Terminology for the integ-
ument texture follows Eady (1968), Harris
(1979), and Torre-Bueno (1978). A pair of
structures on the epipharynx located mesad
and slightly ventrad of epipharyngeal seta 1
were found to have not been named in the
literature. They are herein referred to as epi-
pharyngeal sclerites (Fig. 2). The margins of
these sclerites are not clearly defined in elec-
tron micrographs but are generally visible
under the light microscope and consistently
VOLUME 91, NUMBER 3
Fig. 2.
Epipharynx of D. amatella (180 x): ESc, epi-
pharyngeal sclerites; ESh, epipharyngeal shield.
bare of spines, except occasionally on the
mesal margin.
Common names of most species follow
the Entomological Society of America
(1982). The name fir coneworm for D. abie-
tivorella is from Hedlin et al. (1981). Hedlin
et al. (1981) also used the name loblolly pine
coneworm for D. merkeli Mutuura and
Munroe. In light of recent work (Neunzig
and Leidy 1989), we suggest that the name
loblolly pine coneworm would be better
suited to D. taedivorella, as D. merkeli ap-
pears to feed chiefly on slash pine, P. e/liotii.
Scientific and common names of host plants
follow Radford et al. (1968).
All intact immatures and adults have been
placed as voucher specimens in the North
Carolina State University Insect Collection
(NCSU).
Dioryctria abietivorella (Grote)
(fir coneworm)
Figs. 4, 10, 16, 22, 28
Pinipestis abietivorella Grote, 1878, Bulle-
tin of the U.S. Geological and Geograph-
ical Survey of the Territories, 4: 701.
General.—Length 19.4 mm to 21.4 mm.
Head reddish brown in epicranial area,
shading to somewhat darker reddish brown
near mandibles and hypostoma; tonofibril-
lary platelets indistinct; hypostoma reddish
brown; antennal segments brown; mandi-
327
Fig. 3.
Hypopharynx of D. disclusa, dorsal aspect
(300 x): PmA, premaxillulary area; L, lingua; MX, lobes
of the maxillulae; G, gorge of the maxillulae; B, blade
of the maxillulae.
bles reddish brown. Prothoracic shield
translucent yellowish white anteriolaterally
and mesally, remainder medium brown.
Prespiracular plate yellowish brown, dark
brown on posterior half of dorsal and ven-
tral margins; tonofibrillary platelets medi-
um to dark brown. Remainder of prothorax
with brown granules dorsally and laterally,
granules indistinct ventrally; hypodermal
pigmentation yellowish white; tonofibril-
lary platelets yellowish white, shiny; pinac-
ula light brown. Mesothorax, metathorax
and abdomen with brown granules dorsally
and laterally, granules indistinct ventrally;
pinacula brown dorsally, paler brown to in-
distinct laterally and ventrally; tonofibril-
lary platelets yellowish white, shiny. Fused
D2 pinacula of abdominal segment 9 yel-
low. Anal shield yellow mesally, brown lat-
erally. Mesothoracic SD1 pinaculum form-
ing ring, dark brown with large pale center.
328
Eighth abdominal SD1 pinaculum forming
ring, medium brown with small pale center.
Setae brown basally, pale distally. Thoracic
legs light brown laterally, yellowish white
mesally. Head: Range of length and width
respectively (in mm): 1.55-1.60, 1.88-1.95.
Labrum with distal margin strongly in-
dented. Epipharynx (Fig. 4) bare mesally
from center to epipharyngeal shield, re-
mainder with fine pale spines (lateral spines
very short, in rows; mesal and basal spines
longer, not in rows); epipharyngeal sclerites
reddish brown; epipharyngeal shield some-
what narrow, pale brown. Hypopharynx
(Fig. 10) with premaxillulary area bare; lobes
of maxillulae bearing moderately slender
spines; lingua bare; blades of maxillulae
simple; gorge of maxillulae bare; mentum
reddish brown, slightly darker anteriome-
sally; arms of mentum reddish brown. Spin-
neret slender, approximately 8.5 x as long
as median breadth. Thorax: SD1 on me-
sothorax 1.35 mm to 1.73 mm long. Integ-
ument of mesothorax areolate-rugose ven-
trade of D1 &2 (Fig. 16); aerolate anterior
to D1 &2 (Figs. 16, 22). Abdomen: Seta D2
of abdominal segment 2 approximately
to 2 dorsoventral diameter of segment. SD 1
of segment 8, 1.18 mm to 1.83 mm long.
Crochets of abdominal prolegs biordinal,
arranged in a circle. Number of crochets on
abdominal prolegs: seg.3: 42-62, seg.4: 52-
56; seg.5: 54-62; seg.6: 48-64; seg. 10: 43-
47. Perianal region (Fig. 28) with short,
slightly stout pale spines, spines dorsad of
anus slightly longer and more slender than
those laterad.
Materials examined.— Four larvae, USA
N. Carolina: Mitchell Co., Bakersville, 28-
X-1982, 21-VII-1983, 3-VIII-1983, W. Ay-
ers. All larvae collected from Abies fraseri.
Comments.—In N.C. the larvae feed in
the apical terminals and young branches of
Fraser’s fir, leaving a mass of frass, webbing,
and needle fragments at the feeding site
(Leidy and Neunzig 1986).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Dioryctria disclusa Heinrich
(webbing coneworm)
RIgSe oh ellie 23h 9,
Dioryctria disclusa Heinrich, 1953, In Far-
rier and Tauber, Iowa State College Jour-
nal of Science, 27: 495.
General.— Length 18.5 mm to 22.3 mm;
moderately slender, at times slightly fusi-
form. Head reddish brown, sometimes with
darker brown maculations; tonofibrillary
platelets indistinct to dark brown; hypo-
stoma dark reddish brown posteriorly, black
anteriorly; antennal segments brown; man-
dibles reddish brown. Prothoracic shield
pale yellowish brown anteriorly, pale me-
sally, remainder medium brown; tonofi-
brillary platelets dark brown. Prespiracular
plate pale yellowish brown, dark brown on
posterior half of dorsal and ventral margins;
tonofibrillary platelets dark brown. Re-
mainder of prothorax with gray granules
dorsally and laterally; granules present ven-
trally but usually somewhat less prominent;
hypodermal pigmentation buff, tonofibril-
lary platelets gray to buff, somewhat gran-
ular; pinacula light brown to buff. Meso-
thorax, metathorax and abdomen with gray
granules dorsally, granules less distinct lat-
erally and ventrally, or with gray granules
dorsolaterally, granules less distinct mesally
and laterally, appearing striped; hypoder-
mal pigmentation buff; pinacula of meso-
thorax and metathorax pale brown to buff
pinacula of abdomen buff, indistinct except
in contrast to granules; tonofibrillary plate-
lets gray to buff, granular. Fused D2 pi-
nacula of abdominal segment 9 yellowish
brown. Anal shield yellowish brown. Meso-
thoracic SD1 pinaculum forming ring, light
brown with large pale center. Eighth ab-
dominal SD1 pinaculum forming ring, light
brown with small pale center. Setae brown
basally, pale distally. Thoracic legs medium
to dark brown laterally, pale brown to yel-
lowish white mesally. Head: Range of length
VOLUME 91, NUMBER 3
Figs. 4-9. Dioryctria spp. Epipharynx: 4. D. abietivorella (150); 5. D. disclusa (150); 6. D. pygmaeella
(150); 7. D. clarioralis (100 x); 8. D. amatella (100 x); 9. D. taedivorella (100 x).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ci» |
un \.
=
Figs. 10-15. Dioryctria spp. Hypopharynx, dorsal aspect: 10. D. abietivorella (200 x); 11. D. disclusa (200 x );
12. D. pygmaeella (200 =); 13. D. clarioralis (150); 14. D. amatella (150); 15. D. taedivorella (150 x).
VOLUME 91, NUMBER 3 331
Figs. 16-21. Dioryctria spp. Part of mesothorax including D and SD pinacula, dorsolateral aspect: 16. D.
abietivorella; 17. D. disclusa; 18. D. pygmaeella, 19. D. clarioralis; 20. D. amatella., 21. D. taedivorella. (100 x)
332 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
b
Figs. 22-27. Dioryctria spp. Integument of mesothorax anterior to D1&2, dorsal aspect (see Fig.1 for exact
location): 22. D. abietivorella. 23. D. disclusa; 24. D. pygmaeella, 25, D. clarioralis; 26. D. amatella, 27. D.
taedivorella. (1500 x)
VOLUME 91, NUMBER 3
»
ase
i
Kf
Figs. 28-33. Dioryctria spp. Perianal region, caudal aspect. abietivorella; 29. D. disclusa:
pygmaeella; 31. D. clarioralis, 32. D. amatella: 33. D. taedivorella. <)
334
and width respectively (in mm): 1.43-1.60,
1.60-1.78. Labrum with distal margin mod-
erately indented. Epipharynx (Fig. 5) bare
mesally to base and anterior to epipharyn-
geal sclerites; bearing short fine pale spines
laterally and mesad of epipharyngeal scler-
ite; epipharyngeal sclerite indistinct: epi-
pharyngeal shield narrow, reddish brown.
Hypopharynx (Fig. 11) with premaxillulary
area and lobes of maxillulae bearing long,
moderately slender spines; lingua bare;
blades of maxillulae simple; gorge of max-
illulae bare; mentum reddish brown, black
anteriomesally; arms of mentum reddish
brown. Spinneret slender, approximately 7 x
as long as median breath. Thorax: SD1 on
mesothorax 1.13 mm to 1.55 mm long. In-
tegument of mesothorax areolate ventrad of
D1&2 (Fig. 17), squamiform anterior to
D1&2 (Figs. 17, 23). Abdomen: Seta D2 of
abdominal segment 2 approximately 4 dor-
soventral diameter of segment. SD1 of seg-
ment 8, 1.38 mm to 2.0 mm long. Crochets
of abdominal prolegs mostly biordinal, oc-
casionally partly triordinal, arranged in a
circle. Number of crochets on abdominal
prolegs: seg.3: 53-78; seg.4: 60-74; seg.5:
57-76; seg.6: 59-75; seg. 10: 46-65. Perianal
region (Fig. 29) with short, light brown
spines, spines dorsad of anus slightly longer
and more slender than those laterad.
Material examined. — Twelve larvae. USA
N. Carolina: Wayne Co., nr. Princeton, 30-
IV-1985, N. A. Leidy; Johnston Co., nr.
Princeton, 30-IV-1985, N. A. Leidy; Wayne
Co., Goldsboro, 30-VI-1985, N. A. Leidy;
Johnston Co., Clayton, 2-V-1985, N. A.
Leidy; Bladen Co., Elizabethtown, 13-V-
1985, N. A. Leidy; Wake Co., New Hill, 24-
V-1984, Leidy & Richmond; Wake Co.,
Apex, 21-V-1984, N. A. Leidy. All larvae
collected from Pinus taeda.
Comments.—The length of seta SD1 of
the mesothorax and abdomen is at consid-
erable variance from that reported by Neun-
zig et al. (1964) and figured by Farrier and
Tauber (1953). It is consistent, however, in
larvae from several dates and localities col-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lected in 1985 and 1986. Larvae feed in
ovulate cones leaving a mass of frass and
webbing over the entry hole.
Dioryctria pygmaeella Ragonot
(baldcypress coneworm)
Figs: (6, 12,118; 24; 30
Dioryctria pygmaeella Ragonot, 1887, Di-
agnoses of North American Phycitidae
and Galleriidae, 5.
General.—Length 11.1 mm to 15.1 (see
comments); moderately slender, at times
slightly fusiform. Head brown with darker
brown maculations or uniformly dark
brown; tonofibrillary platelets dark brown
to indistinct; hypostoma dark brown; an-
tennal segments dark brown; mandibles dark
reddish brown. Prothoracic shield pale an-
teriorly and mesally, remainder light to me-
dium brown with darker maculations or
uniformly dark brown; tonofibrillary plate-
lets dark brown to indistinct. Prespiracular
plate light to dark brown; tonofibrillary
platelets dark brown to indistinct. Remain-
der of prothorax with indistinct granules;
hypodermal pigmentation pale yellow over-
lain with longitudinal white and pinkish or-
ange stripes laterally, pale yellow ventrally;
tonofibrillary platelets indistinct; pinacula
dark brown at base of setae, remainder in-
distinct. Mesothorax, metathorax and ab-
domen with indistinct granules; hypoder-
mal pigmentation pale yellow overlain with
longitudinal white and pinkish orange stripes
dorsally and laterally, pale yellow ventrally;
pinacula dark brown at base of setae, re-
mainder indistinct; tonofibrillary platelets
indistinct. Fused D2 pinacula of abdominal
segment 9 dark brown at base of setae, re-
mainder pale yellowish brown to indistinct.
Anal shield dark brown at base of setae,
remainder pale yellowish brown. Mesotho-
racic SD1 pinaculum forming ring, dark
brown with large pale center. Eighth ab-
dominal SD1 pinaculum forming ring, dark
brown with large pale center. Setae brown
basally, pale distally. Thoracic legs dark
VOLUME 91, NUMBER 3
brown. Head: Range of length and width
respectively (in mm): 1.13—1.25, 1.28-1.55.
Labrum with distal margin moderately in-
dented. Epipharynx bare mesally to base,
bearing short fine pale spines laterally (Fig.
6), occasionally bare mesad of epipharyn-
geal sclerites, bearing short fine pale spines
mesally at base and laterally; epipharyngeal
sclerites indistinct; epipharyngeal shield
narrow, pale distally, brown proximally and
mesally. Hypopharynx with premaxillulary
area bare; lobes of maxillulae bearing mod-
erately slender spines distally, longer slen-
der spines proximally; lingua bare; blades
of maxillulae simple; gorge of maxillulae
bare (Fig. 12), occasionally bearing slender
spines distally; mentum brown, darker an-
teriomesally; arms of mentum brown. Spin-
neret slender, approximately 6x as long
as median breadth. Thorax: SD1 on me-
sothorax 1.25 mm to 1.60 mm long. Integ-
ument of mesothorax areolate-rugose ven-
trad of DI&2 (Fig. 18); tuberculate-
squamiform anterior to D1 &2 (Figs. 18, 24).
Abdomen: Seta D2 of abdominal segment 2
approximately '3 the dorsoventral diameter
of the segment. SD1 of segment 8, 1.30 mm
to 1.90 mm long. Crochets of abdominal
prolegs partly biordinal, partly triordinal,
arranged in a circle. Number of crochets of
abdominal prolegs: seg.3: 48-68; seg.4: 50-
69; seg.5: 50-70; seg.6: 50-72; seg.10: 33-
49. Perianal region with short, peg-like pale
to light brown spines (Fig. 30).
Material examined.— Fourteen larvae.
USA N. Carolina: Currituck Co., Coinjock,
14-VII-1986, N. A. Leidy; Currituck Co.,
Coinjock, 1 7-VIII-1978, L. Grimes; Bladen
Co., Singletary Lake State Park (collected
by permit) 23-VII-1985, N. A. Leidy. All
larvae collected from Taxodium distichum.
Comments.—The epipharynx and hypo-
pharynx show considerable variation in spi-
nulation; however, both variations occur in
larvae from the same site and collection date,
and other larval characters and associated
reared adults give no reason to suspect two
separate species. Three larvae had an over-
335
all length of 11.1 mm to 11.8 mm. These
larvae were borrowed material which had
been stored for several years in 80% ethanol
and may have dessicated and shrunk slight-
ly. Freshly preserved larvae ranged from
12.7 mm to 15.1 mm. All larval feeding
records are from bald cypress ovulate cones.
Dioryctria clarioralis (Walker)
(blister coneworm)
Figs. 7; 113, 195 25, 3i
Nephopteryx (sic) clarioralis Walker, 1863,
List of the Specimens of Lepidopterous
Insects in the Collection of the British
Museum, 27: 54.
General.— Length 20.1 mm to 20.2 mm,
moderately slender. Head reddish brown
with dark brown maculations (uniformly
dark brown); tonofibrillary platelets dark
brown (indistinct); hypostoma reddish
brown (dark brown); antennal segments
brown; mandibles reddish brown. Protho-
racic shield pale anteriorly and mesally, re-
mainder pale yellowish brown (dark brown);
tonofibrillary platelets dark brown. Prespi-
racular plate pale yellowish brown (dark
brown, pale on anterior and ventral mar-
gins); tonofibrillary platelets dark brown.
Remainder of prothorax with gray granules
dorsally and laterally, granules present ven-
trally but somewhat less prominent; hypo-
dermal pigmentation yellowish white, tono-
fibrillary platelets gray, somewhat granular;
pinacula yellowish white and indistinct to
pale brown. Mesothorax, metathorax and
abdomen with gray granules dorsally, gran-
ules less distinct laterally and ventrally:
hypodermal pigmentation yellow overlain
with pale rose giving an overall pale orange
appearance; pinacula pale yellow, indistinct
(pale brown); tonofibrillary platelets pale
gray, somewhat granular. Fused D2 pina-
cula of abdominal segment 9 pale yellow.
Anal shield pale yellow. Mesothoracic SD 1
pinaculum forming incomplete ring, pale
yellow (brown) with large pale center. Eighth
abdominal SD1 pinaculum forming ring,
336
pale yellow (brown) with large pale center.
Setae brown basally, pale distally. Thoracic
legs pale brown (dark brown) laterally, yel-
lowish white mesally. Head: Length and
width respectively (in mm): (specimen 1)
1.62, 1.75: (specimen 2) 1.50, 1.60. Labrum
with distal margin moderately indented.
Epipharynx (Fig. 7) bearing short slender
pale spines laterally, longer slender pale
spines mesally; epipharyngeal sclerites pale
reddish brown; epipharyngeal shield large,
pale brown. Hypopharynx (Fig. 13) with
premaxillulary area, lobes of maxillulae,
lingua, and gorge of maxillulae anteriorly
bearing pale spines, spines somewhat short
mesally, longer laterally and anteriorly; gorge
of maxillulae bare posteriorly; blades of
maxillulae simple; mentum brown, darker
anteriomesally; arms of mentum brown.
Spinneret slender, approximately 6 x as long
as median breadth. Thorax: SD1 on me-
sothorax 0.60 mm to 0.67 mm long. Integ-
ument of mesothorax areolate ventrad of
D1&2 (Fig. 19), areolate anterior to DI1&2
(Figs. 19, 25). Abdomen: Seta D2 of abdom-
inal segment 2 approximately , dorsoven-
tral diameter of segment. SD1 of segment
8, 0.60 mm to 0.73 mm long. Crochets of
abdominal prolegs mostly triordinal, ar-
ranged in a circle. Number of crochets on
abdominal prolegs: seg. 3: 76-80; seg. 4: 68—
70; seg. 5: 76-78; seg. 6: 80-82; seg. 10: 49-
51. Perianal region (Fig. 31) with coarse pale
(medium brown) spines.
Material examined.—Two larvae. USA
N. Carolina: Wake Co., 27-VI-1986, N. A.
Leidy; Wayne Co., Goldsboro, 14-I-1987,
J. A. Smith. Larvae collected from Pinus
taeda.
Comments. —Only two larvae of D. clar-
ioralis were obtained. One larva was reared
by placing a gravid light-trapped female in
a sleeve cage over a branch of P. taeda bear-
ing developing ovulate cones and allowing
her to lay eggs. This larva forms the basis
of the above description. A second larva
obtained in Jan. 1987, from damaged host
material has been included but varies con-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
siderably from the first larva in coloration
of the pigmented, sclerotized structures. It
is also more slender but head capsule and
setal measurements indicate a last-instar
larva. Variations are indicated parentheti-
cally above. The differences in size and pig-
mentation are probably due to the colder
average temperature at which this larva de-
veloped. Larvae feed in ovulate cones or
branch terminals leaving a blister of resin-
coated webbing and frass over the entry hole.
Dioryctria amatella (Hulst)
(southern pine coneworm)
Figs. 8, 14, 20, 26, 32
Nephopteryx (sic) amatella Hulst, 1878, En-
tomologica Americana, 3(7): 131.
General.—Length 19.5 mm to 25.4 mm;
moderately robust. Head light to medium
reddish brown; tonofibrillary platelets in-
distinct; hypostoma reddish brown; anten-
nal segments brown; mandibles dark red-
dish brown. Prothoracic shield pale brown
to yellow anteriorly, pale brown to brown
mesally with pale longitudinal stripe, re-
mainder dark brown; tonofibrillary platelets
dark brown. Prespiracular plate dark brown,
tonofibrillary platelets black, shiny. Re-
mainder of prothorax with brown granules
dorsally and laterally, granules present ven-
trally, but indistinct; hypodermal pigmen-
tation greenish white to pale green ventrad
of tonofibrillary platelets, remainder green-
ish white to pale green overlain with rose;
tonofibrillary platelets black, shiny; pina-
cula brown to indistinct. Mesothorax, meta-
thorax and abdomen with brown granules
dorsally, granules less distinct laterally, in-
distinct ventrally; hypodermal pigmenta-
tion greenish white to pale green ventrad of
tonofibrillary platelets, remainder greenish
white to pale green heavily overlain with
rose dorsally, rose less distinct laterally and
ventrally; pinacula dark brown dorsally to
pale brown ventrally; tonofibrillary platelets
black, shiny. Fused D2 pinacula of abdom-
inal segment 9 pale to medium yellowish
VOLUME 91, NUMBER 3
brown. Anal shield pale to medium yellow-
ish brown. Mesothoracic SD1 pinaculum
forming ring, dark brown with minute pale
center. Eighth abdominal SD1 pinaculum
not forming ring, unicolorous dark brown.
Setae brown basally, pale distally. Thoracic
legs pale mesally, dark laterally. Head: Range
of length and width respectively (in mm):
1.62-2.10, 2.05-2.35. Labrum with distal
margin slightly indented. Epipharynx (Fig.
8) at base with numerous pale, stout spines
mesally; epipharyngeal sclerite pale reddish
brown; epipharyngeal shield large, dark
brown. Hypopharynx (Fig. 14) with pre-
maxillulary area, lingua and lobes of max-
illulae bearing coarse spines; blades of max-
illulae simple; gorge of maxillulae bare;
mentum reddish brown, dark brown anteri-
omesally extending anteriorly to lingua;
arms of mentum brown. Spinneret relative-
ly stout, approximately 5x as long as me-
dian breadth. Thorax: SD1 on mesothorax
0.75 mm to 1.02 mm long. Integument of
mesothorax tuberculate ventrad of D1&2
(Fig. 20); spinose anterior to D1 &2 (Figs.
20, 26). Abdomen: Seta D2 of abdominal
segment 2 approximately 4 dorsoventral
diameter of segment. SD1 of segment 8, 0.77
mm to 0.97 mm long. Crochets of abdom-
inal prolegs mostly biordinal, arranged in a
circle. Number of crochets on abdominal
prolegs: seg. 3: 52-64; seg. 4: 54-62; seg. 5:
46-64; seg. 6: 48-60; seg. 10: 29-39. Per-
ianal region with coarse pale spines (Fig.
BD):
Material examined. —Sixteen larvae. USA
N. Carolina: Richmond Co., Rockingham,
4-VIII-1985, 6-VIHI-1986, N. A. Leidy;
Moore Co., Southern Pines, 4-VIII-1985,
14-VIII-1983, N. A. Leidy; Onslow Co.,
Richlands, 16-VII-1985, N. A. Leidy. Lar-
vae collected primarily from Pinus palus-
tris, a few collected from P. taeda.
Comments.—Although the perianal
spines of D. amatella (Fig. 32) are in size
actually very similar to those of D. taedi-
vorella (Fig. 33), they appear more slender
when viewed with the light microscope be-
337
cause of their pigmentation. Larvae of the
spring generation on P. palustris often feed
within the branch terminals, leaving masses
of pitch mixed with some frass over the
entry hole. Later generations on P. palustris
feed in the ovulate cones, leaving large
masses of pitch mixed with frass over the
entry hole. Larvae feeding in the ovulate
cones of P. taeda were not observed to leave
masses of pitch at the entry hole.
Dioryctria taedivorella Neunzig and Leidy
(loblolly pine coneworm)
Bigs ols 2 e233
Dioryctria taedivorella Neunzig and Leidy,
1989, Proc. Entomological Society of
Washington, 91: 321-324.
General.— Length 18.6 mm to 23.4 mm;
moderately robust. Head medium to dark
reddish brown occasionally with slightly
darker maculations; tonofibrillary platelets
indistinct; hypostoma dark reddish brown;
antennal segments brown; mandibles dark
reddish brown. Prothoracic shield pale yel-
low anteriorly and mesally, remainder dark
brown, occasionally paler brown posterior-
ly; tonofibrillary platelets dark brown. Pre-
spiracular plate brown to dark brown; ton-
ofibrillary platelets black. Remainder of
prothorax with dark brown granules dor-
sally and laterally; granules present ven-
trally but less prominent and pale; hypo-
dermal pigmentation pale green ventrad of
tonofibrillary platelets, remainder pale green
overlain with rose; tonofibrillary platelets
black, shiny; pinacula dark brown dorsally
to pale brown ventrally. Mesothorax, meta-
thorax and abdomen with dark brown gran-
ules dorsally and laterally, granules less dis-
tinct ventrally; hypodermal pigmentation
pale green ventrad of tonofibrillary plate-
lets, remainder rose; pinacula dark brown
dorsally to pale brown ventrally; tonofi-
brillary platelets black, shiny. Fused D2
pinacula of abdominal segment 9 yellowish
brown to reddish brown, occasionally dark-
er marginally. Anal shield yellowish brown
338
to reddish brown mesally, reddish brown to
dark brown laterally. Mesothoracic SD1
pinaculum forming ring dark brown with
minute, pale center. Eighth abdominal SD1
pinaculum not forming ring, unicolorous
dark brown. Setae brown basally, pale dis-
tally. Thoracic legs pale mesally, dark brown
laterally. Head: Range of length and width
respectively (in mm): 1.62-1.80, 1.95-2.18.
Labrum with distal margin slightly indent-
ed. Epipharynx (Fig. 9) at base with nu-
merous dark reddish brown tipped, stout
spines mesally; epipharyngeal sclerites red-
dish brown; epipharyngeal shield large, dark
reddish brown, nearly black slightly paler
mesally. Hypopharynx (Fig. 15) with pre-
maxillulary area and lobes of maxillulae
bearing coarse spines; lingua bare to a few
spines; blades of maxillulae simple; gorge
of maxillulae bare; mentum reddish brown,
black anteriomesally, extending anteriorly
to lingua; arms of mentum reddish brown.
Spinneret stout, approximately 4 as long
as median breadth. Thorax: SD1 on me-
sothorax 0.62 mm to 0.87 mm long. Integ-
ument of mesothorax tuberculate-rugose
ventrad of D1 &2 (Fig. 21); spinose anterior
to D1&2 (Figs. 21, 27). Abdomen: Seta D2
of abdominal segment 2 approximately %
dorsoventral diameter of segment. SD1 of
segment 8, 0.70 mm to 0.82 mm long. Cro-
chets of abdominal prolegs mostly biordi-
nal, arranged ina circle. Number of crochets
on abdominal prolegs: seg. 3: 42-60; seg. 4:
46-62; seg. 5: 39-60; seg. 6: 45-64; seg. 10:
35-49. Perianal region (Fig. 33) with coarse,
dark tipped spines.
Material examined.—Seventeen larvae.
USA N. Carolina: Onslow Co., Maysville,
25-VI-1985, Leidy & Lodge; Wayne Co.,
Goldsboro, 16-VII-1985, N. A. Leidy;
Granville Co., Lewis, 18-VII-1985, N. A.
Leidy; Onslow Co., Richlands, 16-VII-1985,
N. A. Leidy; Robeson Co., Lumberton, 27-
VII-1983, N. A. Leidy; Lenoir Co., Kinston,
14-VIT-1985, N. A. Leidy; Robeson Co.,
Lumberton, 22-VII-1985, Leidy & Maynor;
Robeson Co., Lumberton, 9-VII-1986, Lei-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
dy & Hardin. Larvae collected from Pinus
taeda.
Comments.—Larvae feed in ovulate
cones, occasionally leaving small amounts
of resin and frass at the entry hole. Damaged
cones turn brown before maturing and do
not open.
Discussion.—New information on the
larvae of Dioryctria, particularly with re-
spect to spinulation of the trophi and peri-
anal region, and the texture of the integu-
ment was revealed using SEM. The spines
of the epipharynx of D. pygmaeella (Fig. 6)
and D. disclusa (Fig. 5) were easily observed
using SEM, but difficult to see with the light
microscope, because they lack strong pig-
mentation. Neunzig et al. (1964), using con-
ventional methods, did not observe these
spines. Similarly, SEM revealed numerous
spines on the lobes of the maxillulae of D.
pygmaeella (Fig. 12) although Neunzig et
al. (1964) described the hypopharynx of D.
pygmaeella as “bare or with a few indistinct
spines.” The perianal region of D. pyg-
maeella was also described by Neunzig et
al. (1964) as “lacking distinct spines”; how-
ever, SEM showed considerable spinulation
(Fig. 30). Additionally, Neunzig et al. (1964)
separated D. amatella (Fig. 32) and D. tae-
divorella (Fig. 33) (as zimmermani) partly
on the basis of the appearance of the peri-
anal spines; however, electron micrographs
of this region in the two species showed little
difference. The variation observed with the
light microscope is apparently due to pig-
mentation, not relative size of the spines.
The appearance of other structures also
varied depending on the method of obser-
vation. The epipharyngeal shield was ap-
parent with either method, but the epi-
pharyngeal sclerites, quite prominent under
the light microscope on some species be-
cause of their pigmentation, were not dis-
tinct using SEM. Their location could be
established in species with the epipharynx
heavily spinose, such as D. clarioralis (Fig.
7), because of the absence of spines on the
sclerites. In species such as D. amatella (Fig.
VOLUME 91, NUMBER 3
8), however, with only a few spines in the
general area, there is no real indication of a
structure.
The tonofibrillary platelets of the meso-
thorax also appeared quite dissimilar de-
pending on the method of observation.
Those of D. taedivorella (Fig. 21) were ap-
parent using SEM because of their inden-
tation and difference in texture from the sur-
rounding integument, but they are even
more striking under the light microscope
because of their black pigmentation. In con-
trast, the tonofibrillary platelets of D. dis-
clusa (Fig. 17) were more distinct using SEM
than in light microscopy.
SEM also made more apparent major
variation in the texture of the integument.
The mesothorax anterior to setae D1 &2 of
D. amatella (Fig. 26) and D. taedivorella
(Fig. 27) is spinose, whereas it ranges from
areolate in D. abietivorella (Fig. 22) and D.
clarioralis (Fig. 25) to squamiform in D.
disclusa (Fig. 23) and tuberculate-squami-
form in D. pygmaeella (Fig. 24).
Mutuura and Munroe (1972, 1974) placed
the adults of Dioryctria in eight species
groups. Most of the species which we stud-
ied were assigned (Mutuura and Munroe
1972, 1974, 1979) to four of these as fol-
lows: abietella group: D. abietivorella; au-
ranticella group: D. disclusa; baumhoferi
group: D. pygmaeella; and zimmermani
group: D. amatella. D. clarioralis and D.
taedivorella were not included by Mutuura
and Munroe, but obviously the former be-
longs to the baumhoferi group and the latter
to the zimmermani group. Our study of
Dioryctria, although treating relatively few
species, gives evidence that most of the lar-
vae can also be placed in the proposed
groups. Both species of the zimmermani
group have very similar spinulation of the
epipharynx (Figs. 8, 9), hypopharynx (Figs.
14, 15), and perianal region (Figs. 32, 33),
as well as similar integumental texture (Figs.
26, 27). Additionally, each has black ton-
ofibrillary platelets and a unicolorous SD1
pinaculum on the eighth abdominal seg-
339
ment. The remaining species can be readily
separated from one another as larvae. There
are distinct differences in the texture of the
integument of D. abietivorella (Fig. 22) and
D. disclusa (Fig. 23). In addition, the pig-
mentation of the D1 and D2 pinacula of the
abdominal segments, and the length of seta
D2 of the abdominal segments further sep-
arate these two species. As mentioned, D.
pygmaeella has been placed in the baum-
hoferi group. Mutuura and Munroe (1972)
noted, however, that their placement of D.
pygmaeela was tentative because of differ-
ences in the male genitalia. Examination of
the larval characters also casts doubt on the
placement of D. pygmaeella in the baum-
hoferi group. Variation in the spinulation
of both the hypopharynx (Figs. 12, 13) and
perianal region (Figs. 30, 31) of D. pyg-
maeella and D. clarioralis is considerable.
Differences are also found in the texture of
the integument (Figs. 24, 25). Blanchard and
Knudson (1983) recently described a new
species from Texas, D. caesirufella, which
they feel is most closely allied with D. pyg-
maeella, Although larvae of D. caesirufella
were not available for this study, the dis-
similarity of the larvae of D. pygmaeella
and D. clarioralis suggest that D. pygmaeel-
la may be better placed with D. caesirufella
in a new species group, rather than in the
baumhoferi group.
Key To LAsT-INSTAR LARVAE OF EASTERN
NortTH AMERICAN SPECIES OF DIORYCTRIA
1. Tonofibrillary platelets of abdomen pale, dif-
fering in texture from, or only slightly darker
than, integument; pinaculum SD1 of 8th ab-
dominal segment forming ring with pale cen-
ter; integument not spinose anterior to D1 &2
of mesothorax (Figs. 16-19, 22-25) 2D
— Tonofibrillary platelets of abdomen black,
shiny; pinaculum SD1 of 8th abdominal seg-
ment not forming ring, unicolorous dark brown;
integument spinose anterior to D1&2 of me-
sothorax (Figs. 20, 21, 26, 27) 5
Pinacula D1 & D2 of abdomen brown, at least
at base of setae; seta D2 of abdominal seg-
ments long, 3 to 2 dorsoventral diameter of
segment 3
N
340
— Pinacula DI & D2 of abdomen usually pale,
appearing lighter than surrounding integu-
ment; seta D2 of abdominal segments short, 4
to 4 dorsoventral diameter of segment
3. Tonofibrillary platelets yellowish white, shiny,
distinct from integument; pinaculum SD1 of
8th abdominal segment with small pale center;
labrum with distal margin distinctly indented
(Fig. 4); overall length approximately 20 mm
SANS MACHA Since fore ae TEE epee abietivorella
- Tonofibrillary platelets indistinct; pinaculum
SD1 of 8th abdominal segment with large pale
center; labrum with distal margin weakly in-
dented (Fig. 6); overall length approximately
MS SMTA ss. ah crn ego esco Menta ce + pygmaeella
4. Pinaculum SD1 of 8th abdominal segment with
small pale center; pinaculum SD1 of meso-
thorax forming complete ring; seta D2 of ab-
dominal segments approximately 4 dorsoven-
tral diameter of segment; lingua and gorge of
mentum bare (Fig. 11) .... disclusa
— Pinaculum SD1 of 8th abdominal segment with
large pale center; pinaculum SD1 of meso-
thorax forming incomplete ring; seta D2 of ab-
dominal segments approximately ', dorsoven-
tral diameter of segment; lingua and gorge of
mentum spinose (Fig. 13) clarioralis
5. Perianal region with pale brown tipped spines,
spines appearing slender at low magnification;
epipharynx with pale, coarse spines at base;
hypopharynx with lingua bearing numerous
coarse spines (Fig. 14) ................ amatella
— Perianal region with dark reddish brown tipped
spines, spines appearing stout at low magnifi-
cation; epipharynx with dark reddish brown
tipped coarse spines at base; hypopharynx with
lingua bare or bearing at most a few coarse
spines (Fig. 15) taedivorella
ACKNOWLEDGMENTS
We thank the following people who helped
in obtaining study material either by pro-
viding specimens, assisting in the actual col-
lection, or allowing access to their land or
land under their supervision: W. Ayers,
Roan Valley Tree Farm, Mitchell Co., N.C.;
H. Barron, Edwards State Forest, Morgan-
ton, N.C.; L. Grimes, Meredith College, Ra-
leigh, N.C.; T. Hardin, Federal Paper Board
Co., Lumberton, N.C.; L. Jervis, N.C. State
Univ., Raleigh, N.C.; J. Killian, Mary
Washington College, Fredricksburg, Va.; D.
Lodge, formerly N.C. State Univ., Raleigh,
N.C.; M. Maynor, Federal Paper Board Co.,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Lumberton, N.C.; N. Potts, Highlands, N.C.,;
J. Richmond, N.C. State Univ., Raleigh,
N.C.; D. Rogers, Edwards State Forest,
Morganton, N.C.; D. Sparkman, Federal
Paper Board Co., Lumberton, N.C.; D. Ste-
phan, N.C. State Univ., Raleigh, N.C.; B.
Taylor, N.C. Div. of Parks and Recreation,
Raleigh, N.C.; G. Turner, Claridge State
Forest, Goldsboro, N.C.; and W. Wicks,
Champion Intl. Corp., Deppe, N.C.
The NCSU College of Agriculture and Life
Sciences, Center for Electron Microscopy
provided financial assistance for use of the
scanning electron microscope through
minigrants. We also thank C. Giles, for-
merly CALS Center for Electron Micros-
copy, NCSU, Raleigh, N.C., and L. Grimes,
Meredith College, Raleigh, N.C., for their
assistance in specimen preparation tech-
niques. This paper is part of a thesis sub-
mitted by N. A. Leidy in partial fulfillment
of the requirements of the M.S. degree in
entomology, North Carolina State Univer-
sity, Raleigh, N.C., and is paper No. 11471
of the Journal Series of the North Carolina
Agricultural Research Service, Raleigh, N.C.
27695-7643. The use of trade names in this
publication does not imply endorsement by
the North Carolina Agricultural Research
Service of the products named, nor criticism
of similar ones not mentioned.
LITERATURE CITED
Blanchard, A. and E.C. Knudson. 1983. A new species
of Dioryctria Zeller (Lepidoptera: Pyralidae) from
Texas. Proc. Entomol. Soc. Wash. 85: 116-120.
Bradley, E. L., M. W. MacGown, B. H. Ebel, and W.
W. Neel. 1982. Surface patterns of eggs of some
cone eating Lepidoptera of southeastern United
States. J. Ga. Entomol. Soc. 17: 255-259.
Coulson, R. N., A. Mutuura, and E. Munroe. 1972.
The Dioryctria species of loblolly pine in east Tex-
as, with comments on the occurrence of two new
species. J. Econ. Entomol. 65: 868-870.
Crumb, S.E. 1929. Tobacco Cutworms. USDA Tech.
Bull. 88. 180 pp.
Eady, R. D. 1968. Some illustrations of microsculp-
ture in the Hymenoptera. Proc. Roy. Entomol.
Soc. London (A) 43: 66-72.
Entomological Society of America.
1982. Common
VOLUME 91, NUMBER 3
Names of Insects and Related Organisms. F. G.
Werner, Chairman. College Park, Md. 132 pp.
Farrier, M. H. and O. E. Tauber. 1953. Dioryctria
disclusa Heinrich, n. sp. (Phycitidae) and its par-
asites in Iowa. Iowa State Coll. J. Sci. 27: 495-
507.
Harris, R. A. 1979. A Glossary of Surface Sculptur-
ing. Calif. Dept. Food Agric., Div. Plant Industry-
Lab. Serv. Occas. Papers Entomol. No. 28. 31 pp.
Hedlin, A. F., H. O. Yates, D. Cibrian-Tovar, B. H.
Ebel, T. W. Koerber, and E. P. Merkel. 1981.
Cone and Seed Insects of North American Coni-
fers. Joint publication: Environment Canada, Ca-
nadian Forest Service; USDA, Forest Service; Se-
crateria Agricultura y Recursos Hidraulicos,
México. 122 pp.
Heinrich, C. 1956. American Moths of the Subfamily
Phycitinae. U.S. Nat. Mus. Bull. 207. 581 pp.
Leidy, N. A. and H. H. Neunzig. 1986. New host
record and range extension for Dioryctria abieti-
vorella (Grote) (Lepidoptera: Pyralidae: Phyciti-
nae). Proc. Entomol. Soc. Wash. 88: 395.
Lyons, L. A. 1957. Insects affecting seed production
in red pine II. Dioryetria disclusa Heinrich, D.
abietella (D. and S.) and D. cambiicola (Dyar)
(Lepidoptera: Phycitidae). Can. Entomol. 89: 70-
79,
MacKay, M. R. 1943. The spruce foliage worm and
the spruce cone worm (Dioryctria spp., Lepidop-
tera, Pyralidae). Can. Entomol. 75: 91-98.
Munroe, E. 1959. Canadian species of Dioryctria Zeller
(Lepidoptera: Pyralidae). Can. Entomol. 91: 65-
12.
Mutuura, A. 1982 American species of Dioryctria
(Lepidoptera: Pyralidae) VI. A new species of
Dioryctria from eastern Canada and northeastern
United States. Can. Entomol. 114: 1069-1076.
Mutuura, A. and E. Munroe. 1972. American species
of Dioryctria (Lepidoptera: Pyralidae) III. Group-
ing of species: Species of the auranticella group,
including the Asian species, with the description
of a new species. Can. Entomol. 104: 609-625.
1973. American species of Dioryctria (Lep-
idoptera: Pyralidae) IV. The schuetzeella group
and the taxonomic status of the spruce cone moth.
Can. Entomol. 105: 653-668.
. 1974. Anew genus related to Dioryctria Zell-
er (Lepidoptera: Pyralidae: Phycitinae), with def-
inition of an additional species group in Dioryc-
tria. Can. Entomol. 106: 937-940.
1979. American species of Dioryctria (Lep-
idoptera: Pyralidae) V. Three new cone-feeding
species from the southeastern United States. J. Ga.
Entomol. Soc. 14: 290-304.
341
Mutuura, A., E. Munroe, and D. A. Ross. 1969a.
American species of Dioryctria (Lepidoptera: Py-
ralidae) I. Western Canadian species of the zim-
mermani group. Can. Entomol. 101: 1009-1023.
1969b. American species of Dioryctria (Lep-
idoptera: Pyralidae) I]. Western Canadian species
of the baumhoferi and ponderosae groups. Can.
Entomol. 101: 1042-1047.
Mutuura, A. and H. H. Neunzig. 1986. New Central
American Dioryctria and notes on previously de-
scribed species of the genus in the region (Lepi-
doptera: Pyralidae), pp. 8-18. Jn Cibrian-Tovar,
D., et al., eds., Cone and Seed Insects of the Mex-
ican Conifers. USDA, Forest Service, Southeast-
ern Forest Exper. Sta., Gen. Tech. Rep. SE-40.
110 pp.
Neunzig, H. H. 1979. Systematics of Immature Phy-
citines (Lepidoptera: Pyralidae) Associated with
Leguminous Plants in the Southern United States.
USDA Tech. Bull. 1589. 119 pp.
Neunzig, H. H., R. L. Rabb, B. H. Ebel, and E. P.
Merkel. 1964. Larvae of the genus Dioryctria
(Lepidoptera: Phycitidae) in the southeastern
United States. Ann. Entomol. Soc. Am. 57: 693-
700.
Neunzig, H. H. and N. A. Leidy. 1989. A new species
of Dioryctria (Lepidoptera: Pyralidae: Phycitinae)
from the southeastern United States. Proc. Ento-
mol. Soc. Wash. 91: 321-324.
Radford, A. E., H. E. Ahles, and C. R. Bell. 1968.
Manual of the Vascular Flora of the Carolinas.
UNC Press, Chapel Hill, N.C. 1183 pp.
Schaber, B. D. 1981. Description of the immature
stages of Dioryctria taedae Schaber and Wood,
with notes on its biology and that of D. disclusa
Heinrich (Lepidoptera: Pyralidae). Proc. Entomol.
Soc. Wash. 83: 680-689.
Schaber, B. D. and F. E. Wood. 1971. A new species
of Dioryctria infesting loblolly pine. Proc. Ento-
mol. Soc. Wash. 73: 215-223.
Spies, C. J. HI and J. B. Dimond. 1985. Separating
early larvae of spruce budworm, Choristoneura
fumiferana (Lepidoptera: Tortricidae) and spruce
coneworm, Dioryctria reniculleloides (sic) (Lepi-
doptera: Pyralidae) in surveys of overwintering
larvae. Can. Entomol. 117: 261-263.
Stehr, F. W. 1987. Immature Insects. Kendall/Hunt
Pub. Co., Dubuque, Iowa. 754 pp.
Torre-Bueno, J. R. 1978. A Glossary of Entomology.
Fifth printing. N.Y. Entomol. Soc. New York. 381
pp.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 342-345
A NEW GENUS AND SPECIES OF HISPINAE
(COLEOPTERA: CHRYSOMELIDAE) FROM CENTRAL AMERICA
C. L. STAINES, JR.
3302 Decker Place, Edgewater, Maryland 21037.
Abstract. —Fossispa, new genus, is described from Guatemala, Jamaica, and Mexico; F.
lutena n. sp. is described and designated the type species. A key to the genera of Uroplatini
with clavate antennae is presented.
Key Words:
The Neotropical Hispinae are poorly
known. The literature consists of scattered
species descriptions, faunal lists, and a few
biological notes. The last comprehensive
works were by Baly (1885) and Weise (1911).
Both of these are much out of date due to
the large number of species and genera de-
scribed since their publication.
Specimens of this species were first re-
ceived from M. A. Ivie as a genus near Bra-
chycoryna. A third specimen was later re-
ceived from S. M. Clark. From a search in
the collection at the U.S. National Museum,
eight additional specimens were found
amoung the unidentified Neotropical His-
pinae.
Measurements were taken with an ocular
micrometer. The total length is from the
anterior margin of the pronotum to the apex
of the elytra. Pronotal length is from the
base to the apex of the pronotum. Pronotal
width is along the midline. Elytral length is
from the elytral base to apex. Elytral width
was taken at the humeri. In the type des-
ignations, a slash (/) separates data on dif-
ferent labels.
Fossispa, NEw GENUS
Head: micropunctate; median sulcus
present; three short lateral sulci near each
eye; antennae inserted into quadrate pit; pit
Chrysomelidae, Hispinae, beetle, Uroplatini
divided by keel; carina around each eye:
antenna 8-segmented, clavate. Pronotum:
wider than long; completely margined at
sides; raised areas between punctures mi-
cropunctate. Scutellum: quadrate; micro-
punctate. E/ytron: with four discal costae,
3rd costa short and weak, costae 1, 2, and
4 unite apically; punctures in double rows,
with 8 rows of punctures basally, 10 rows
apically, basal rows 5 and 6 in longitudinal,
median depression, latter rows briefly ex-
®
WWW Tivae leo © ow
Fig. 1.
Elytra of Fossispa lutena.
VOLUME 91, NUMBER 3
pand to 4 rows behind middle of elytron; a
short weak costa between apical rows 6 and
7; costae and raised areas between punc-
tures micropunctate. Venter: rugose at sides.
Type species of the genus. — Fossispa lu-
tena, new species.
Etymology.—Fossispa, fossa = ditch or
trench plus ispa, Latin. The gender is neuter.
Comparative notes. — Fossispa belongs in
the tribe Uroplatini as shown by the 8-seg-
mented antenna. Fossispa is most similar
to Heptatomispa Uhmann. Fossispa differs
from Heptatomispa as follows:
Character Heptatomispa Fossispa
vertex of — not sulcate sulcate
head
antennae 7-segmented, 8-segmented;
segments segments |-
thick, | not 6 thin, 7-8
much expanded
narrower
than 7
pronotum highly arched; not arched; no
longitudinal —_ prebasal
prebasal impression
impression
present
scutellum longer than about as long
wide as wide
elytra three entire three entire
costae; plus one
costa 2 does incomplete
not unite costae;
with | & 3 costa 2
unites with
1&3
KEY TO THE GENERA OF UROPLATINI
WITH CLAVATE ANTENNAE
1. Antenna 7-segmented; last four segments
combined not longer than the three preceding
combined . ; : bcinee
Antenna 7- or 8- -segmented; last four seg-
ments combined longer than the three pre-
ceding combined
to
le
343
in
Elytral costae irregular, tuberculate
ee: Physocoryna Guérin
Elytral costae regular Corynispa Uhmann
First 5 antennal segments similar; segment 6
widened; club as long as 4 preceding segments
combined; antenna 7-segmented . Bruchia Weise
Loe ea
— Antenna not asabove .................. 4
4. Elytral costae irregular, fipercilate oa 5
— Elytral costae regular ....... aoe 6
5. Antennal segments 7 and 8 almost fused; seg-
ments | to 6 thick . Acritispa Uhmann
— Antennal segments 7 and 8 distinct
Octotoma Chevrolat (in part)
6. Third tarsal segment not obviously bilobed
Ee ea eae ee a Stenopodius Horn
— Third tarsal segment obviously bilobed 7
7. Each elytron on apical third with 8 rows
OMPUNGLUTES ae geet weyers yee ape ere 8
— Each elytron on apical third with 10 rows of
PUNGLUTES) sexs soe eicieeke cern wet ean eees 9
8. Antennal segment 7 as wide as 8
. Octotoma Chevrolat (fe part)
Antennal segment 7 narrower than 8
ae ee ee . Parvispa Uhmann
9. Base ofeach elytron with 10r rows of punctures
AES Brachycoryna Guerin
- Base of each elytron with 8 rows of punctures
Pe ee 10
10. Expanded neat rows on napical halfofely=
tra not divided by a costa; vertex of head not
SulCate =n tees ate Heptatomispa Uhmann
— Expanded puncture rows on apical half of ely-
tra divided by a costa; vertex of head sulcate
. Fossispa NEW GENUS
Fossispa lutena, New SPECIES
Head: black; micropunctate; median sul-
cus present, sometimes weak; occiput con-
cave, with 3 lateral sulci near each eye; an-
tennal pit quadrate, longer than wide; keel
sharper anteriorly; carina around each eye;
antenna 8-segmented; segments I-VI cylin-
drical, II widest and longest; VII expanded,
ring of setae on middle; VIII expanded, hir-
sute, rounded at apex; I-III glabrous; [V—
VI with some setae; mouthparts ventrally
directed. Pronotum: wider than long; com-
pletely margined at sides; lateral margins
parallel at apical *, then convergent; cov-
ered with coarse, deep punctures, weaker on
disc; raised areas between punctures micro-
punctate; yellow with variable brown mac-
ulae anteriorly and laterally; width 0.9 to
344
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
75
195 5 ; =e x “
| ;
ne
. i/ Ba | 20
= 5 F
Sice GS ae
SEN aly = ie
a / Sa
= (6
¢ \ 15
LEA ma
/ —a|'3
a i
de a 9s J a if \ 7
90 85 80 7s 70 <¥S]
Fig. 2.
Distribution of Fossispa lutena.
1.3 mm (avg. 1.1) (n = 10); length 0.6 to
0.8 (avg. 0.7). Scutellum: brown, quadrate;
longer than wide; micropunctate. E/ytron:
(Fig. 1). 8 rows of punctures at base, ex-
pands to 10 after middle; scutellar row of 3
punctures; punctures usually in double rows,
but may be confused; 3 complete discal cos-
tae plus one short, weak costa on apical third,
1, 2, and 4 united on apical fifth; suture
costate; elytral base explanate, expanded
over base of pronotum; humeral angles
strongly produced; raised areas on costae
and between punctures micropunctate;
margin serrate; apices cojointly rounded; at
middle, puncture rows 5 and 6 enter lon-
gitudinal depression, expand to two addi-
tional rows divided by a weak costa; small
depression between puncture rows 3 and 4
on basal third, 3rd interspace weakly cos-
tate; 4th costa highly carinate from humer-
us, the extent of projection variable, up to
the lateral or posterior margin, the area be-
tween costa and margin recurved; yellow
with scattered brown maculae; width 1.3 to
1.7 mm (avg. 1.5); length 2.0 to 2.7 (avg.
2.3). Legs: yellow; trochanter brown, an-
gular; femur widest in middle; tibia widest
at apex, row of setae on inner side and at
apex; each tarsus as follows—segments I and
II small, cylindrical, pad of setae beneath;
III bilobed, 4 length of IV, pad of setae
beneath; IV long, widening at apex; claws
divergent. Venter: black, prosternum punc-
tate in middle, rugose at sides, truncate at
apex; mesosternum smooth, shining in mid-
dle, rugose at sides; several punctures around
coxae; metasternum smooth, shining in
middle, rugose at sides; abdominal sterna
smooth in middle, rugose at sides, all with
scattered setae. Total length: 2.6 to 3.4mm
(avg. 3.0).
Type data.—Holotype: MEXICO: Vera
VOLUME 91, NUMBER 3
Cruz-Tampico, 11/25/72, F. Parker & D.
Miller; deposited in the U.S. National Mu-
seum. 10 Paratypes; GUATEMALA: Es-
quintla Prov., May 14, 1965, E. J. Ham-
bleton/ sweeping ex. Cymbopogon spp./ 64—
65. JAMAICA: Portland Par., Millbank, 10
August 1987, M. A. Ivie colr.; deposited in
the M. A. Ivie collection; St. Ann Parish,
Ocho Rios, 19 October 1985, J. A. Suey
colr.; deposited in the S. M. Clark collec-
tion; St. Catherine Parish—2 miles n. of
Guanabon Vale, 6-VI-1975, A. Harkins.
MEXICO: Chaiapas-Tempisque, 6/VII/63,
Whitehead & Woodruff: Guerrero-Barra
Vieja, DCD, 10/X/84, Seguva, Gillett, and
Miranda/ on leaf of Sida sp., coll. 78/ LPL
1012; Bara Nieyja, ACD, 23/1/85, Richardo
Sequta, ex. Sida acuta/ LP 1490; Barra Vie-
ja, 5/IX/86, Gillett & Miranda/ on Sida sp./
LP 889; Barra Vieja, ACA, 10/X/84, Se-
quta, Gillett, & Miranda/ on leaf of Sida
acuta/ LPL 1011. Unless otherwise noted
the paratypes are deposited in the U.S. Na-
tional Museum.
Distribution: Jamaica, Guatemala, and
Mexico (Fig. 2).
345
Etymology.—the specific ephithet, /utena
is from the Latin luteus meaning yellow,
and refers to the yellow dorsal color of the
pronotum and elytra.
Habitat.—adults have been collected from
Sida acuta Burm. and Sida sp. (Malvaceae)
leaves and by sweeping Cymbopogon spp.
(Poaceae).
Immature stages unknown.
ACKNOWLEDGMENTS
I thank S. M. Clark (West Virginia De-
partment of Agriculture), M. A. Ivie (Mon-
tana State University), and R. E. White (U.S.
National Museum) for the loan of the ma-
terial used in this paper. R. E. White and
M. A. Ivie commented on earlier drafts on
this manuscript. Illustration by Susan L.
Staines.
LITERATURE CITED
Baly, J. S.. 1885. Biologica Centrali-Americana. In-
secta. Coleoptera. Vol. VI. Part 2. Phytophaga
Hispidae. 1-124.
Weise, J. 1911. Coleoptera: Phytophaga, family
Chrysomelidae, subfamily Hispinae. In P. Wyts-
man Genera Insectorum fac. 125: 1-123. Brussels.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 346-349
BIOLOGY OF A PINE NEEDLE SHEATH MIDGE,
CONTARINIA ACUTA GAGNE (DIPTERA: CECIDOMYIIDAE),
ON LOBLOLLY PINE
JuLIE C. WEATHERBY, JOHN C. Moser, RAYMOND J. GAGNE, AND
Huey N. WALLACE
(JCW) and (HNW) USDA Forest Service, Forest Pest Management, 2500 Shreveport
Highway, Pineville, Louisiana 71360; Present Address (JCW) USDA Forest Service,
Forest Pest Management, 1750 Front Street, Boise, Idaho 83702; (JCM) USDA Forest
Service, Southern Forest Experiment Station, 2500 Shreveport Highway, Pineville, Lou-
isiana 71360; (RJG) Systematic Entomology Laboratory, PSI, Agricultural Research Ser-
vice, USDA, % National Museum of Natural History, Washington, D.C. 20560.
Abstract. —The biology of a pine needle sheath midge, Contarinia acuta Gagné is de-
scribed for a new host in Louisiana. This midge was found feeding within the needle
sheath on elongating needles of loblolly pine, P. taeda L. Needle droop and partial de-
foliation were evident on heavily infested trees. Overwintering C. acuta adults were first
detected emerging from the soil on April 30, 1984. The population progressed through
four generations between May 11 and September 17, 1984.
Key Words:
Two species of Contarinia are known to
cause needle droop on pines in North Amer-
ica. One, the introduced Contarinia baeri
(Prell), is found on Scots pine, Pinus syl-
vestris L., and on red pine, P. resinosa Ait.,
in northeastern North America. It was first
discovered in Europe in 1930 (Skuhravy
1973) and later reported on Scots pine in
Canada (DeBoo et al. 1973, Wilson et al.
1988). The second species, Contarinia acuta
Gagné, is found on slash pine, P. e/liottii
Engelmann, in southeastern United States.
Gagne and Beavers (1984) reported on C.
acuta and three other Contarinia species.
The last three were recovered from pitfall
traps only, so their role on slash pine 1s un-
known.
Contarinia acuta was found in 1971 to
cause needle droop and defoliation on lob-
lolly pine at the Erambert Seed Orchard in
Contarinia sp., pine needle sheath midge, pine seed orchards, loblolly pine
Brooklyn, Mississippi (Overgaard et al.
1976). Populations continued to build until
1975 when the population collapsed (Ov-
ergaard et al. 1976). An evaluation con-
ducted from March to September 1975 de-
tected three major larval population peaks:
the first peak in May, the second peak be-
ginning in late June, and the third peak be-
ginning in mid-to-late July (Overgaard et al.
1976). Similar damage was reported in 1975
from an orchard in McNair, Mississippi
(Overgaard et al. 1976). The next docu-
mented report of an outbreak of this species
occurred in 1983 at the Stuart Seed Orchard
in Pollock, Louisiana (Weatherby et al.
1983). During this outbreak, only loblolly
pines were infested, while slash, longleaf,
and shortleaf pines remained unaffected.
This paper reports on the biology of C. acuta
infesting loblolly pine in central Louisiana.
VOLUME 91, NUMBER 3
MATERIALS AND METHODS
Larval sampling procedures. Field studies
were conducted at the Stuart Seed Orchard.
In 1983, actively growing shoots from sus-
ceptible clones were sampled. Clones 5, 18,
20, 30, and 43 were selected from clones in
the Texas loblolly seed source. One ramet
of each clone was randomly selected on Au-
gust 23. Five shoots of new growth were
clipped from each ramet and 25 fascicles
were randomly selected from each shoot.
The fascicle sheath was removed and the
number of larvae per fascicle recorded. On
August 31, four different ramets of three of
the five original clones were randomly se-
lected and sampled. Five shoots of new
growth were selected and five fascicles per
shoot were removed for examination from
each of the sample ramets. The number of
larvae per fascicle was recorded. A final ex-
amination was conducted on October 19.
In 1984, sample trees were randomly se-
lected from a 52-acre block of mature lob-
lolly trees grafted from Louisiana seed
sources. Larval development was moni-
tored by sampling fascicles from sample
branches. After needle elongation began in
the spring, two sample branches were re-
moved from the upper portion of the can-
opy and one sample branch was removed
from the middle portion of each sample tree.
A total of 12 trees were sampled every 2
weeks from April 13 through August 28.
Sample branches from each tree were placed
in a plastic bag and transported back to the
laboratory. Five fascicles were removed
from the last growth flush on each branch,
the needle sheaths were removed, and the
needles were inspected under a dissecting
microscope. The numbers of first instar,
second instar, and third instar larvae per
fascicle were recorded. During periods when
multiple growth flushes and overlapping
generations coincided, five fascicles were re-
moved from each of the last two growth
flushes on each sample branch. Larval den-
sities were determined for each instar on the
last two growth flushes.
347
Prepupal sampling procedures. Prepupal
migration from trees to pupation sites in the
soil was monitored with sticky traps. Ply-
wood squares, 0.30 m by 0.30 m, were cov-
ered with white freezer paper and sprayed
with Tree Tanglefoot (Tanglefoot Compa-
ny, Grand Rapids, MI). These squares were
mounted horizontally on top of 0.91 m
stakes. Five sample trees were randomly se-
lected and four traps, one at each cardinal
point, were placed under the dripline of each
tree. Prepupal traps were installed on May
25 and monitored through September 17,
1984. Traps were inspected weekly and lar-
vae were counted and removed.
Adult sampling procedures. Adult flights
were monitored by using adult emergence
traps. Traps were constructed from 11.36 |
plastic wash tubs and 0.24 | glass jars. The
mouth of a jar was inserted into a hole on
the side of the tub and secured with a fitting
that was attached to the exterior side of each
tub approximately 2.54 cm above the bot-
tom. Each tub was inverted and one trap
was placed under the dripline of each sam-
ple tree. A total of five traps were placed in
the Louisiana loblolly seed source. The
emergence traps were installed on March
10, 1984, and the jars were inspected weekly
for emerging adults. Adult midges were re-
moved from each trap and placed in vials
containing 70 percent ethanol. These vials
were forwarded to RJG for identification.
The traps were relocated after the prepupal
migration of each generation in order to
capture adults. Adult trapping was termi-
nated on September 17.
RESULTS AND DISCUSSION
In 1983, the mean larval density on clones
5, 18, 20, 30, and 43 was 14.48 larvae per
fascicle on August 23. Populations de-
creased dramatically by August 31 to 2.51
larvae per fascicle and no larvae were de-
tected on October 19, indicating that the
late August generation was the overwinter-
ing generation (Table 1).
Population sampling conducted in 1984
348
Table 1. Comparison of mean larval densities re-
corded on the last three sampling dates at the U.S.
Forest Service Stuart Seed Orchard, Pollock, LA (1983).
Mean Number of Larvae per Fascicle
Clone Aug. 23 Aug. 31 Oct. 19
5 11.14 -_ =
18 17.82 2.48 0.00
20 17.64 - -
30 14.09 1.43 0.00
43 11.72 3.61 0.00
Mean 14.48 piss) 0.00
« — Indicates that samples were not taken.
showed that the population progressed
through four generations between April 30
and September 17. Adult emergence of the
1983 overwintering generation was detected
on April 30 and continued through May 18.
The second, third, and fourth adult flights
occurred between June 4 and June 18, June
25 and July 18, and August 8 and August
27, respectively.
Adult emergence from overwintering sites,
oviposition, and egg hatch of the first gen-
eration coincided with the beginning of
needle elongation on the first growth flush
of the trees in the Louisiana loblolly seed
source. During subsequent flight periods, fe-
males preferentially oviposited on the most
recent foliage. First and second generation
larvae primarily infested the first growth
flush. Third generation larvae infested both
second and third growth flushes. The last or
fourth generation larvae were found within
the fascicles of the fourth growth flush.
Mean larval densities per fascicle of first,
second, and third instars for each sample
date are listed in Table 2. These densities
were considerably less than those recorded
for the final generation in 1983. The pres-
ence of first instar larvae were first detected
on May 11. Three additional population
peaks occurred on June 11, July 13, and
August 10. Similar peaks in larval density
were detected for second instars. Consid-
erable reduction occurred in the population
densities between second and third instars
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 2. Means of larval population densities ob-
tained from branch samples taken from loblolly pines
in the Louisiana seed source at the U.S. Forest Service
Stuart Seed Orchard, Pollock, LA (1984).
Number of Larvae/Fascicle
Growth Ist Instar 2nd Instar 3rd Instar
x x x
Date Flush # Fasc x
April
13 Ist 225 0.00 0.00 0.00
30 Ist 180 0.00 0.00 0.00
May
11 Ist 180 ; 0.00 0.00
17 Ist 750 0.11 0.19 0.11
31 Ist 180 0.18 0.00 0.00
June
11 Ist 180 0.35 0.00 0.01
2nd 120 0.10 0.00 0.01
18 Ist 180 0.25 0.29 0.11
2nd 145 0.03 0.04 0.02
26 Ist 175 0.07 0.08 0.10
2nd 170 0.05 0.06 0.02
July
13 2nd 155 0.66 0.09 0.04
3rd 170 1.30 0.09 0.00
20 2nd 180 0.09 0.16 0.00
3rd 180 0.45 0.74 0.03
27 3rd 180 0.02 0.02 0.17
August
10 3rd 170 0.06 0.04 0.05
4th 70 0.99 Only 0.14
28 3rd 150 0.19 0.15 0.03
4th 15 0.93 1E27 0.07
(Table 2). This reduction could have been
a real decrease attributed to natural mor-
tality, or it could have been a result of poor
synchronization between sampling frequen-
cy and phenology of each generation.
Third instar larvae developed to the pre-
pupal stage within the fascicle sheath. Prior
to pupation, prepupae left the sheath and
fell to the ground. Pupation occurred in the
ground litter under the canopy. Peak*mi-
grations to the ground of the first, second,
and third generation prepupae were detect-
ed on May 29, June 27, and August 6 (1984),
respectively (Table 3). A fourth generation
prepupal peak was barely detectable due to
a rapid collapse in the population.
VOLUME 91, NUMBER 3
Table 3. Mean number of prepupae captured on |
sq. ft. sticky traps located under the dripline of loblolly
pines in the Louisiana seed source at the U.S. Forest
Service Stuart Seed Orchard, Pollock, LA (1984).
Mean
Mean Number Number of
of Prepupae Prepupae
Date per | Sq. Ft Date per | Sq. Ft.
May 29 4.10 July 16 0.20
31 0.20 18 0.00
June 4 0.50 20 0.00
6 0.05 25 0.10
8 0.05 27 0.20
11 0.25 August 3 0.15
13 0.00 6 2.70
15 0.15 8 0.10
18 0.10 10 1.40
20 1.00 13 0.00
22 0.40 17 0.00
25 1.65 22 0.00
27 11.25 24 0.00
30 6.55 28 0.00
July 2 0.40 September 4 0.20
4 0.20 6 0.05
6 0.30 10 0.00
i) 0.60 12 0.00
11 0.05 17 0.00
13 0.15
During the 1984 outbreak, larval mor-
tality between the second stadium and the
prepupal stage of the fourth generation was
high. Increasing populations of the natural
enemies, particularly Pyemotes emargina-
tus Cross, Moser, and Rack, were observed
during larval sampling. Cross, Moser, and
Rack (1981) discuss the biology of this mite
parasitoid that is known only from C. acuta.
In addition, several predaceous larvae of
Lestodiplosis (Cecidomyiidae) were found
within the fascicle sheaths with C. acuta
larvae.
ACKNOWLEDGMENTS
The authors thank personnel at the U.S.
Forest Service Stuart Seed Orchard for pro-
349
viding the field site for this research. Men-
tion of a proprietary or commercial product
within this publication does not constitute
recommendation or endorsement of the
product by the U.S. Department of Agri-
culture and does not imply its approval to
the exclusion of other products that also
may be suitable.
LITERATURE CITED
Cross, E. A., J. C. Moser, and G. Rack. 1981. Some
new forms of Pymotes (Acarina: Pyemotidae) from
forest insects, with remarks on polymorphism. Int.
J. Acarol. 7: 179-196.
DeBoo, R. F., L. M. Campbell, J. P. LaPlante, and L.
P. Daviault. 1973. Plantation research: VIII. The
pine needle midge, Contarinia baeri (Diptera: Ce-
cidomyiidae), a new insect pest of Scots pine. En-
viron. Can. For. Serv., Chem. Cont. Res. Inst.
Info. Rep. CC-X-41. 31 pp.
Gagné, R. J. and G. M. Beavers. 1984. Contarinia
spp. (Diptera: Cecidomyiidae) from shoots of slash
pine (Pinus elliott: Engelm.) with the description
ofa new species injurious to needles. Fla. Entomol.
67(2): 221-228.
Overgaard, N. A., H. N. Wallace, C. Stein, and G. D.
Hertel. 1976. Needle midge (Diptera: Cecido-
mylidae) damage to loblolly pines in the Erambert
federal seed orchard, Mississippi. USDA-FS,
Southeastern Area, S&PF, FIDM, Rep. No. 76-2-
13. 11 pp.
Skuhravy, V. 1973. “Needle blight” and “needle
droop” on Pinus silvestris L. in Europe and P.
resinosa Ait. in North America (Diptera: Ceci-
domyiidae). Z. Angew. Entomol. 72 (1972/73):
421-428.
Weatherby, J., J. Moser, D. Starkey, R. Gaar, and S.
Covington. 1983. Needle midge damage to lob-
lolly pines at the Stuart seed orchard, Pollock,
Louisiana, 1983. USDA-FS, Region 8, S&PF,
FPM, Rep. No. 84-2-1. 8 pp.
Wilson, L. F., F. J. Sapio, and G. A. Simmons. 1988.
Biology, injury, and control of the European needle-
bending midge (Diptera: Cecidomyiidae) on Scotch
pine in Michigan. Great Lakes Entomol. 21(3): 97-
104.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 350-354
GRYPOTES PUNCTICOLLIS (HOMOPTERA: CICADELLIDAE),
A PALEARCTIC PINE-FEEDING LEAFHOPPER
NEW TO NORTH AMERICA
A. G. WHEELER, JR.
Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn-
sylvania 17110.
Abstract.—Grypotes puncticollis (Herrich-Schaeffer), an Old World deltocephaline leaf-
hopper belonging to the small tribe Grypotini, was recently detected at Erie, Pennsylvania.
Nymphs and adults were abundant on Scotch pine, Pinus sylvestris L., and Swiss mountain
pine, P. mugo Turra. Populations also were found on Scotch pine in nearby Crawford
Co. and in three western New York counties: Allegany, Cattaraugus, and Chautauqua. It
is suggested that G. puncticollis is a relatively recently invader in North America and that
it was introduced with egg-infested nursery stock. Adult and nymphal characters facili-
tating recognition of this immigrant in the Nearctic fauna are provided. Two other leaf-
hopper species were collected on Scotch pine during the survey for G. puncticollis: Em-
poasca perlonga Davidson & De Long and Gyponana geminata (Osborn).
Key Words: Auchenorrhyncha, immigrant insect, Pinus sylvestris, Pinus mugo
Grypotes puncticollis (Herrich-Schaeffer)
is a Palearctic deltocephaline leafhopper be-
longing to the small Old World tribe Gry-
potini (4 spp.). This pine specialist, com-
mon throughout much of continental Europe
and England, ranges from Scandinavia south
to Spain and northern Africa (Algeria, Tu-
nisia) and east to the European USSR and
Turkey (Metcalf 1967). Scotch pine, Pinus
sylvestris L., Austrian pine, P. nigra Arnold,
and P. brutia have been reported as hosts
(Ossiannilsson 1983, Lodos and Kalkan-
delen 1985 and references therein). This ap-
parently univoltine species overwinters in
the egg stage; adults are present from July
to October or even early November. In Tur-
key, G. puncticollis has been listed as a pest
of P. brutia and P. nigra (Ossiannilsson
1983, Lodos and Kalkandelen 1985 and ref-
erences therein). Metcalfs (1967) world cat-
alogue may be consulted for references to
additional European literature containing
ecological notes.
Herein, G. puncticollis is reported as new
to the Western Hemisphere. North Amer-
ican locality records are listed and mapped,
and characters facilitating its recognition in
the Nearctic fauna are provided.
DISTRIBUTION AND Host PLANTS
The initial collection of G. puncticollis was
made on 21 July 1988 on the Mercyhurst
College campus, Erie, Pennsylvania, during
routine insect collecting on Pinus sylvestris.
Although I had collected previously on
Scotch pine in Pennsylvania (Wheeler and
Henry 1973, Wheeler 1987), I was unfa-
miliar with the brownish-yellow leafhopper
found on this conifer at Erie. Fourth and
fifth instars were present and adults were
abundant on a row of pines. I suspected that
this species was an immigrant, but Hamil-
VOLUME 91, NUMBER 3
351
Fig. 1.
Known North American distribution of the Palearctic leafhopper Grypotes puncticollis. Black dots
indicate established populations; counties in which limited surveys for G. puncticollis on Scotch pine were
negative are represented by open circles (negative sites in Ohio are not shown).
ton (1983) did not include any Scotch pine
feeders among Cicadellidae common to the
Old and New World and (1985) noted that
leafhoppers were not known from this plant
in Canada. K. Valley was able to eliminate
the unknown Scotch pine cicadellid as con-
specific with any listed in Oman (1949) or
Beirne’s (1956) leafhoppers of Canada and
Alaska. With access to European literature
and specimens, E. R. Hoebeke determined
the species as the Palearctic G. puncticollis.
After this immigrant was identified, sur-
veys were made in Pennsylvania, particu-
larly in the northwestern counties, and in
western New York and northeastern Ohio
to try to delimit its Nearctic range. The first
Erie collection was from Scotch pine, and
late instars and adults were found later at
the same site on dwarf and on upright,
shrubby cultivars of Swiss mountain pine,
P. mugo Turra. For all other collections list-
ed below, P. sy/vestris was the host; all col-
lections were made by the author in 1988
except for the 13 September collection from
Erie, which was made by H. G. Wolff. Spec-
imens have been deposited in the insect col-
lections of Cornell University (CUIC), Ith-
aca, NY; Pennsylvania Department of
Agriculture (PDA), Harrisburg; and U.S.
National Museum of Natural History
(USNM), Washington, DC.
NEW YORK: Allegany Co., SUNY—
Alfred State University, Alfred, 31 July;
Cattaraugus Co., Salamanca, 31 July;
Chautauqua Co., Fredonia and Jamestown,
31 July. PENNSYLVANIA: Crawford Co.,
Allegheny College, Meadville, 1 Aug.; Erie
Co., Edinboro, | Aug.; Mercyhurst College,
Erie, 21 July, 1 Aug., 13 Sept.; North East,
1 Aug.
352
Although G. puncticollis was common on
Scotch and Swiss mountain pines in Erie
Co., Pennsylvania, and present at one of
several localities sampled in Crawford Co.,
it was not collected on pines 1n several near-
by counties or in other areas of the state.
Abundant in western New York, it was not
taken in limited surveys in other counties
or in northeastern Ohio (Fig. 1).
The abundance of this leafhopper near
Lake Erie and failure to detect populations
elsewhere suggest a limited distribution in
North America and a relatively recent in-
troduction. If G. puncticollis had been in-
troduced early in the twentieth century be-
fore the United States implemented plant
quarantine legislation (Wheeler and Nixon
1979, Kim 1983), it probably would be more
widespread and have been detected much
earlier. If it were now widely distributed on
Scotch pine in Ontario and elsewhere in
eastern Canada, it probably would not have
been overlooked during recent studies of
Canadian leafhoppers and been included in
Hamilton’s (1983) review of holarctic Cica-
dellidae or in Hamilton and Langor’s (1987)
report on the fauna of Newfoundland and
Cape Breton Island. It seems reasonable to
assume that G. puncticollis was introduced
to the Lake Erie region with European ship-
ments of conifer nursery stock containing
its eggs. Opening of the St. Lawrence Sea-
way to ship traffic in the late 1950’s may
have been involved in this introduction. The
large volume of maritime shipping along
this waterway has been implicated in the
spread of an immigrant coccinellid, Cocci-
nella undecimpunctata L. (Watson 1979).
RECOGNITION FEATURES
A diverse leafhopper fauna is not char-
acteristic of pines in Pennsylvania. The coe-
lidine Neocoelidia tuberculata (Baker)
sometimes is common on native pitch pine,
Pinus rigida Mill., and Virginia pine, P. vir-
giniana Mill., and occasionally occurs on
cultivated red pine, P. resinosa Ait. (per-
sonal observation). Few leafhopper species
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
have adapted to introduced pines like P.
sylvestris; Hamilton (1985) did not list any
Canadian species from Scotch pine. A typh-
locybine, Empoasca perlonga Davidson &
De Long, was commonly encountered on
Scotch pine in the survey for G. puncticollis
in New York and Pennsylvania. Hamilton
(1985) reported jack pine, P. banksiana
Lamb., as its host plant. In Pennsylvania,
Gyponana geminata (Osborn) also was tak-
en on Scotch pine. This gyponine feeds on
jack pine in Canada (Hamilton 1985) and
is common on pitch pine in Pennsylvania
(personal observation).
From these pine-feeding species and oth-
er North American leafhoppers, G. puncti-
collis can be separated by the characters giv-
en by LeQuesne (1969) and Ossiannilsson
(1983): Head distinctly wider than prono-
tum, ocelli remote from eyes (about equi-
distant between eye and median line), and
anteclypeus narrow and apically bent cau-
dad. North American material generally
agrees with descriptions of European spec-
imens. The adult (Fig. 2) is 3.96-4.68 mm
long, brownish yellow, and shining with a
greyish-green tinge. Among salient features
that can be considered diagnostic are face
with 3 dark, often fuscous, transverse
streaks, the upper 2 arched and faint or in-
terrupted medially and extending laterally
to eyes; vertex with sinuate dark streaks be-
tween eyes and oblique dark spots or bars
from eye toward base; pronotum with dark
spots anteriorly; and scutellum with dark
transverse suture and usually 2 dark spots
anteriorly. Male and female genitalia are il-
lustrated by LeQuesne (1969) and Ossian-
nilsson (1983); Villiers (1977) provided a
color illustration of the adult.
Ossiannilsson (1983) briefly described the
last instar nymph as uniformly brownish,
with head much wider than the pronotum,
and with longer setae only on abdominal
tergites VII and VIII. Last instars from Erie,
Pennsylvania, have the head broader than
pronotum and lack dorsal setae except at
apex of abdomen; coloration, however, dif-
VOLUME 91, NUMBER 3
353
Fig. 2.
fers from Ossiannilsson’s characterization.
Nymphs from Pennsylvania range from pale
yellow without dorsal markings (perhaps re-
cently molted individuals) to brownish yel-
low with well-developed dark markings; the
midline is pale in all specimens. Some spec-
imens are suffused with pink, the roseate
tinge usually restricted to the wingpads and
apex of abdomen. Other prominent features
are antennae fuscous except basal segments
pale; face generally unmarked except for se-
ries of faint transverse lines; vertex usually
bearing dark markings, sometimes appear-
ing as nearly circular areas on either side of
midline; pronotum with dark spots similar
to markings of adult or nearly uniformly
brown; wingpads generally pale, area be-
tween bases of pads darkened; abdomen
usually infuscate, ranging from yellow with
dark markings to dark brown with scattered
pale spots; legs mostly pale, prominent dark
spots at base of hind tibial spines.
Specimens examined: 51 adults and 57
fifth instars from Mercyhurst College, Erie,
Grypotes puncticollis, adult habitus; scale bar = 1.0 mm.
Pennsylvania, 21 July and | Aug. 1988, on
Pinus mugo and P. sylvestris; deposited in
the collections of CUIC, PDA, and USNM.
ACKNOWLEDGMENTS
I am grateful to K. Valley (BPI, PDA) for
identifying Gyponana geminata, eliminat-
ing any native species as the suspected im-
migrant leafhopper on Pinus sylvestris, and
reading the manuscript; E. R. Hoebeke
(Dept. of Entomology, Cornell University,
Ithaca, NY) for identifying Grypotes punc-
ticollis and reading the manuscript; M. R.
Wilson [CAB International Institute of
Entomology, % British Museum (Natural
History), London] for confirming the deter-
mination of G. puncticollis, K. G. A. Ham-
ilton (Biosystematics Research Centre, Ot-
tawa, Ont.) for identifying Empoasca
perlonga, H. G. Wolff (BPI, PDA) for mak-
ing a follow-up collection at Ernie; J. F. Stim-
mel (BPI, PDA) for the photograph used in
Fig. 2; T. J. Henry (Systematic Entomology
Lab., USDA, % U.S. National Museum of
354
Natural History, Washington) for reading
the manuscript; and M. Lacey (Systematic
Entomology Lab., Taxonomic Services Unit,
USDA, Beltsville, MD) for sending speci-
mens to the British Museum for confirma-
tion.
LITERATURE CITED
Beirne, B. P. 1956. Leafhoppers (Homoptera: Cica-
dellidae) of Canada and Alaska. Can. Entomol. 88
(suppl. 2): 1-180.
Hamilton, K.G. A. 1983. Introduced and native leaf-
hoppers common to the Old and New worlds
(Rhynchota: Homoptera: Cicadellidae). Can.
Entomol. 115: 473-511.
1985. Leafhoppers of ornamentals and fruit
trees in Canada. Agric. Can. Publ. 1779/E. 71 pp.
Hamilton, K. G. A. and D. W. Langor. 1987. Leaf-
hopper fauna of Newfoundland and Cape Breton
Islands (Rhynchota: Homoptera: Cicadellidae).
Can. Entomol. 119: 663-695.
Kim, K. C. 1983. How to detect and combat exotic
pests, pp. 261-319. Jn C. L. Wilson and C. L.
Graham, eds., Exotic Plant Pests and North Amer-
ican Agriculture. Academic Press, New York.
LeQuesne, W. J. 1969. Handbooks for the identifi-
cation of British insects. Vol. II. Part 2 (b) He-
miptera, Cicadomorpha, Deltocephalinae. Royal
Entomological Society of London. Pp. 65-148.
Lodos, N. and A. Kalkandelen. 1985. Preliminary
list of Auchenorrhyncha with notes on distribution
and importance of species in Turkey. 17. Family—
Cicadellidae: Deltocephalinae: Grypotini, Goni-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
agnathini and Opsiini (Part 1). Turkiye Bitki Kor.
Derg. 9: 79-90.
Metcalf, Z. P. 1967. General Catalogue of the Ho-
moptera. Fascicle VI Cicadelloidea; Part 10 Eus-
celidae, Section II. Agric. Res. Serv., U.S. Dept.
Agric., Washington, D.C., pp. 1078-2074.
Oman, P. W. 1949. The Nearctic leafhoppers (Ho-
moptera: Cicadellidae): A generic classification and
check list. Mem. Entomol. Soc. Wash. 3: 1-253.
Ossiannilsson, F. 1983. The Auchenorrhyncha (Ho-
moptera) of Fennoscandia and Denmark. Part 3:
The family Cicadellidae: Deltocephalinae, cata-
logue, literature and index. Fauna Entomol. Scand.
Vol. 7, pt. 3. Scandinavian Science Press, Copen-
hagen. Pp. 594-979.
Villiers, A. 1977. Atlas des Hémiptéres. Généralités-
Heteéroptéres; Homoptéres-Thysanoptéres. So-
ciété Nouvelle des Editions Boubée & C*, Paris.
237 pp.
Watson, W. Y. 1979. North American distribution
of Coccinella u. undecimpunctata L. (Coleoptera:
Coccinellidae). Coleopt. Bull. 33: 85-86.
Wheeler, A. G., Jr. 1987. Scymnus (Pullus) suturalis
Thunberg: Southernmost records of an immigrant
coccinellid (Coleoptera: Coccinellidae) in the
United States. Coleopt. Bull. 41: 150.
Wheeler, A. G., Jr. and T. J. Henry. 1973. Camp-
tozygum aequale (Villers), a pine-feeding mirid
new to North America (Hemiptera: Mirdae). Proc.
Entomol. Soc. Wash. 75: 240-246.
Wheeler, A. G., Jr. and H. F. Nixon. 1979. Insect
Survey and Detection in State Departments of Ag-
riculture. Spec. Publ. Pa. Dep. Agric., Harrisburg.
28 pp.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 355-359
NEW RECORDS OF PHYCITINAE FROM BOLIVIA INCLUDING A
NEW SPECIES OF PEADUS (LEPIDOPTERA: PYRALIDAE)
H. H. NEUNZIG
Department of Entomology, North Carolina State University, Raleigh, North Carolina
27695-7613.
Abstract. —The following species are recorded from Bolivia for the first time: Hypargyria
definitella (Zeller); Hypsipyla grandella (Zeller); Hemiptilocera chinographella Ragonot;
Hyalospila clevélandella (Dyar),; Piesmopoda ragonoti (Dyar), Peadus bolivianus, new
species; Megarthria beta Heinrich; Stylopalpia fuscifrontella (Zeller), new combination;
Adelphia ochripunctella (Dyar), Caristanius pellucidellus (Ragonot); Eurythmasis ignifatua
Dyar; Unadilla erronella (Zeller), Baphala homoeosomella (Zeller), Edulica compedella
(Zeller).
Key Words:
Heinrich in 1956 recorded nine ‘species
of Phycitimae from Bolivia based on spec-
imens collected by T. Steinbach in the early
1900’s, on loan from the British Museum
of Natural History and the Janse Collection.
It appears that the phycitines Heinrich stud-
ied were only a part of those collected by
Steinbach. Recently, through the generosity
and assistance of John E. Rawlins, I have
had an opportunity to study additional
Steinbach phycitines from Bolivia in the
Collection of the Carnegie Museum of Nat-
ural History (CMNH). In addition to pro-
viding a more comprehensive list of the
Phycitinae occurring in Bolivia, the study
has established the correct generic place-
ment of Zeller’s Nephopteryx (sic) fuscifron-
tella, and brought to light a new species in
the genus Peadus Heinrich.
As best I can determine, Steinbach did
all of his collecting in east Bolivia. Except
for a very few, his collection labels read sim-
ply “P. del Sara” or “Prov. del Sara.” This
refers to a province in east Bolivia presently
known as Provincia del Gutiérrez, in the
Department of Santa Cruz. In the few in-
distribution, new combination, new species
stances where specimens were obtained at
localities other than Provincia del Gutier-
rez, or additional data were placed on the
label, this information has been included as
an annotation in the following list.
Hypargyria definitella (Zeller): Two males
and one female; collected in January and
December.
Hypsipyla grandella (Zeller): One female;
collected in July; 450 m.
Hemiptilocera chinographella Ragonot:
Two males and one female; collected in Jan-
uary, August and November: 450 m. One
female; collected in July; Sta. Cruz de la
Sierra [Dept. Santa Cruz], 450 m. One fe-
male; collected in December; Puerto Suarez
[Dept. Santa Cruz] 150 m.
Hyalospila clevelandella (Dyar): Two
males; collected in December.
Piesmopoda ragonoti (Dyar): One male;
[no collection date].
Megarthria beta Heinrich: One female;
collected in January. Heinrich (1956) was
not able to associate males and females
within the genus Megarthria. The few fe-
males that he studied and considered to be-
356 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Imm
+ Imm 4
Figs. 1-2. Stylopalpia fuscifrontella. 1, male genitalia, most of left valva and aedeagus omitted. 2, aedeagus.
Fig. 3. Peadus bolivianus, holotype.
VOLUME 91, NUMBER 3
>A Fe mm 4
357
Figs. 4-6. Peadus bolivianus. 4, Male genitalia, most of left valva and aedeagus omitted. 5, aedeagus. 6,
female genitalia.
long to two species within the genus were
simply named by him, alpha and beta. Ap-
parently based on similar distributions,
Heinrich suggested that a/pha was possibly
the female of Megarthria peterseni (Zeller).
It now appears equally possible that beta is
the female of peterseni because the present
study has demonstrated that beta also oc-
curs in South America in the same general
area where peterseni is found.
Stylopalpia fuscifrontella (Zeller), New
Combination: Two males and twelve fe-
males; collected in January, March, April,
November, December. Zeller in 1881 treat-
ed fuscifrontella as a Nephopterix. Heinrich
(1956) examined females of the species and
358
concluded that fuscifrontella did not belong
to Nephopterix, but he was unable to estab-
lish its correct generic placement because he
had no males available for study. My ex-
amination of the male genitalia of speci-
mens collected by Steinbach (Figs. 1, 2)
clearly shows the species to be a member of
Stylopalpia Hampson. The small valva, the
presence of a strongly sclerotized, large
clasper near the inner base of the valva, and
the lateral flanges of the aedeagus with their
clusters of large spines are particularly char-
acteristic for the genus.
Adelphia ochripunctella (Dyar): One fe-
male; collected in November. 4. ochri-
punctella has previously only been reported
from San Diego, California (Heinrich, 1956).
Nevertheless, based on the characteristic
wing markings and genitalia of the species,
there is no doubt that the species also occurs
in Bolivia.
Caristanius pellucidellus (Ragonot): One
female; collected in November.
Eurythmasis ignifatua Dyar: Two males
and one female; collected in January, No-
vember and December.
Unadilla erronella (Zeller): One male;
collected in December.
Baphala homoeosomella (Zeller): One
male; collected in December; 450 m.
Edulica compedella (Zeller): One female;
collected in August; Buena Vista [Dept.
Santa Cruz]; 400 m.
Peadus bolivianus Neunzig, NEW SPECIES
Figs. 3-6
Description.— Head: Pale-golden-brown
suffused with reddish brown or with reddish
brown and fuscous. Labial palpus reaching
slightly above vertex (longer in female than
male), pale golden-brown suffused with red-
dish brown and fuscous; 3rd segment darker
than other segments. Maxillary palpus
squamous, pale golden brown. Male anten-
na with elongate basal tuft of dark, ap-
pressed scales; sensilla trichodea (cilia) of
shaft near base about as long as width of
basal segments of shaft. Collar: Pale golden
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
brown dorsally, with small lateral patch of
reddish brown or reddish brown and fus-
cous. Forewing: Pale golden yellow above
dusted with white in costal half, particularly
near costa; costal half also with a few red-
dish brown scales interspersed with white
scales, and solid reddish brown patches,
sometimes with fuscous, at costa; anteme-
dial line absent; discal spots reddish brown;
postmedial line white or very pale golden
brown, moderately distinct; patch of red-
dish brown and fuscous scales preceding
postmedial line; terminal margin with row
of dark scales; undersurface without con-
trasting sex-scales. Length of wing 8.0-8.5
mm. Hindwing: Pale smoky brown basally,
dark brown along termen. Male genitalia
(Figs. 4, 5): Uncus and tegumen greatly re-
duced; gnathos not defined; valva appearing
partially divided longitudinally with inner
subbasal digitate element and costal half
strongly recurved distally; vinculum long,
almost 3 x as long as its greatest width; ae-
deagus spined at apex and with cornutus
consisting of a cluster of small spines. Fe-
male genitalia (Fig. 6): ductus bursae short,
posterior half with numerous microspines;
corpus bursae with numerous microspines
in posterior two-thirds; signum small, short,
thornlike.
P. bolivianus is easily separated from oth-
er species in Peadus on the basis of the very
elongate vinculum of the male genitalia of
bolivianus and the more abundant, and more
generally distributed, microspines in the
corpus bursae of the female genitalia of bo-
livianus.
Type material.—Holotype: 4; Bolivia:
Prov. del Sara, Nov. 1913, Steinbach: gen-
italia slide HHN 2373: in CMNH. Para-
types: 1 4, 1 2; same location and coliector,
Dec. 1911: genitalia slides HHN 2374,
2388a: in CMNH.
ACKNOWLEDGMENTS
I thank John E. Rawlins, Carnegie Mu-
seum of Natural History, Pittsburgh for pro-
VOLUME 91, NUMBER 3
viding the specimens. This is paper No.
11801 of the Journal Series of the North
Carolina Agricultural Research Service, Ra-
leigh, North Carolina 27695-7643.
LITERATURE CITED
Heinrich, C. 1956. American moths of the subfamily
Phycitinae. U.S. Natl. Mus. Bull. 207: 1-581.
359
Zeller, P. C. 1881. Columbische Chiloniden, Cram-
biden und Phycideen. Hor. Soc. Entomol. Rossi-
cae 16: 154-256.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 360-366
A NEW SPECIES OF NEOTROPICAL WATER BUG,
PARAVELIA BIAE, FROM BRAZIL
(HETEROPTERA: VELIIDAE)
PAuL J. SPANGLER
Department of Entomology, National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C. 20560.
Abstract. —A new species of veliid water bug, Paravelia biae, from Para, Brazil, is
described and compared with the species Paravelia platensis Berg and Paravelia williamsi
Hungerford, which it resembles; distinguishing characters are illustrated by line drawings
and scanning electron micrographs. The habitat is discussed and illustrated.
Key Words:
In 1986, several biologists from the
Smithsonian Institution participated in a
survey of the fauna of an area on the Xingu
River about 60 km south of Altamira, Para,
Brazil, that is scheduled for impoundment
following the construction of a large hydro-
electric dam on the river. The attractive
water-strider described below was collected
during the survey.
Parayelia biae Spangler, New SPECIES
Figs. 1-8, 15, 16
Macropterous holotype male (Fig. 1).—
Size: Length, 6.45 mm; greatest width, 2.50
mm.
Color: Black except coxae and base of ros-
trum slightly reddish brown and each hem-
elytron with a bright yellow macula near
base and another, oval in shape, on mem-
branous apex; therefore, with wings in nor-
mal position over abdomen, 3 yellow mac-
ulae readily visible. Cuticle covered with
dense yellowish-brown pubescence.
Head: Width between eyes, 0.50 mm. Cu-
ticle finely pubescent; with sparse, long,
slender, black setae among shorter, dense,
Heteroptera, Veliidae, Brazil, Paravelia biae n. sp.
Paravelia biae, male, habitus view, x 14.
Fig. 1.
VOLUME 91, NUMBER 3
400um
i
s
1
,
i we
7a
piri ieee
NE
—_<_
b
wn
o
haasl fo)
rg
reddish-brown setae. Rostrum extending to
anterior margin of mesocoxal cavities. An-
tennae with fine, dense, reddish-brown to
yellowish pubescence and a few longer,
darker setae interspersed; segment 1 ar-
-5. Paravelia biae, male: 2, Protibial grasping comb, x70. 3, Distal end of protibial grasping comb,
rotibial grasping comb and grooming comb, x 700. 5, Protarsal claws, x 170.
cuate, narrowest at base then distinctly
swollen and parallel sided on distal four-
fifths, distinctly thicker and a fifth longer
than segment 2; segment 2 more slender and
slightly longer than segment 3; segment 4
362
more slender and about a fifth longer than
segment 3.
Thorax: Pronotum narrowest apically,
sides diverging, moderately arcuate, strong-
ly gibbose laterally slightly before mid-
length; lateral margins behind gibbose area
broadly rimmed; with low median longi-
tudinal carina on meson; terminating in a
moderately long, robust, finger-like process
projecting posteriad. Cuticle with 2 sparse,
coarsely punctate, transverse rows on an-
terior seventh; disc with most coarse punc-
tures separated by 4 to 5 times their di-
ameter; also with an oblique row of coarse,
distinct punctures laterad of procoxae. Pro-
tibia with distal grasping comb extending
about three-fourths length of tibia (Figs. 2—
4) and grooming comb on apex (Fig. 4). Pro-
tarsal claws slender (Fig. 5).
Abdomen: Cuticle with fine, dense, short,
yellowish pubescence intermixed with
sparser, longer setae. Laterotergites mod-
erately reflexed above abdominal terga. Seg-
ment 7 (sixth visible) with a strong angular
gibbosity on each side of meson along pos-
terior margin of segment; ovate genital cap-
sule twice as long as segment 7 on midline.
Genitalia: Proctiger with posterior half
pubescent and heavily sclerotized at baso-
dorsal angle (Fig. 6). Clasper sinuate; with
cluster of setae basally and a row of evenly
spaced, erect stiffsetae on upper margin (Fig.
8).
Female.—Similar to male but lacks an-
gular gibbosities on posterior margin of ab-
dominal segment 7 and genital capsule is
replaced by an extensible ovipositor.
Comparative notes.—Paravelia bhiae is
most similar to Paravelia platensis Berg,
1883, and Paravelia williamsi Hungerford,
1930, but differs from both by being black
instead of chocolate brown, by the low pro-
notal carina extending from collar to base
where it ends in a finger-like lobe, and by
its more robust body that is thicker dorso-
ventrally. In addition, males of P. biae have
a gibbosity on each side of the meson of
sternum 7 that are separated by a very broad
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and shallow apicomedial emargination (Fig.
7) in contrast to males of P. p/atensis that
have a broad longitudinal keel on the meson
of sternum 7 (Fig. 14) and males of P. wil-
liamsi that have a gibbosity on each side of
the meson of sternum 7 that are separated
by a broad and deep apicomedial emargi-
nation (Fig. 13).
The male claspers of the known species
of Paravelia are distinctive and the obvious
differences between the right (Fig. 9) and
left clasper (Fig. 10) in Hungerford’s (1930)
illustrations of P. platensis seemed odd. Both
claspers of specimens of P. biae and all other
species of Paravelia that I have examined
are symmetrical. The illustration of asym-
metrical claspers included by Hungerford
suggested that one of the two claspers of his
specimen of P. platensis is aberrant. Al-
though Hungerford did not mention that,
he may have had the same suspicion and,
by luck or design, he included figures of both
claspers. By dissecting and clearing the gen-
ital capsule of a male P. platensis in the
Drake Collection in the National Museum
of Natural History, I found that the claspers
are symmetrical as illustrated in Figure 11
and closely approximate Hungerford’s fig-
ure of the right clasper (Fig. 9). Therefore,
I believe his figure of the left clasper (Fig.
10) illustrates an aberrant one.
Type data.— Macropterous holotype male
and allotype: BRAZIL: PARA: Altamira (60
km S), 52°22'W, 3°39’S, 10 Oct 1986, Ist
stream, trail 4, coll’n #19, P. Spangler & O.
Flint; deposited in the Museu de Zoologia,
Universidade de Sao Paulo, Sao Paulo, Bra-
zil.
Paratypes: Same data as holotype, 147 4,
267 2; same locality and data except: 13 Oct
1986, 5 4, 14 9; 19 Oct. 1986, in flight in-
tercept trap, 1 4, 2 2; 21 Oct. 1986, in flight
intercept trap, 1 6, 1 ¢. BRAZIL: PARA:
Itaituba: Sao Lonis, 15.3.41, leg. H. Sioli,
S16, 1 4, 1 2. Paratypes will be deposited in
the Museu de Zoologia, Universidade de
Sao Paulo, Sao Paulo; the American Muse-
um of Natural History, New York, New
®
Figs. 6-14. 6-8, Paravelia biae, male genitalia: 6, genital capsule, lateral view; 7, abdominal venter; 8, left
clasper. 9-11, Paravelia platensis, male genitalia: 9, right clasper; 10, left clasper; 11, left clasper (specimen in
Drake Collection). 12-13, Paravelia williamsi: 12, left clasper; 13, abdominal venter. 14, Paravelia platensis,
abdominal venter. Figures 9, 10, 12, 13, 14—after Hungerford, 1930.
364 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 15-16. Paravelia biae: 15, exposed after overturning leaves. 16, biotope; 60 km south of Altamira,
Para, Brazil.
VOLUME 91, NUMBER 3
York; the British Museum (Natural Histo-
ry), London; the California Academy of Sci-
ences, San Francisco, California; the Ca-
nadian National Collection, Ottawa; Institut
Royal de Histoire naturelles de Belgique,
Bruxelles; Instituto de Zoologia Agricola,
Facultad de Agronomia, Universidad Cen-
tral Venezuela, Maracay; Laboratorium
voor Zoologische, Oecologie en Taxono-
mie, Utrecht; the Muséum National de His-
toire Naturelle, Paris; Museo Argentino de
Ciencias Naturales, Buenos Aires; Univer-
sidad Nacional de La Plata y Museo, La
Plata; the National Museum of Natural His-
tory, Smithsonian Institution, Washington,
D.C.; the Snow Entomological Museum,
University of Kansas, Lawrence, Kansas;
Zoologische Staatssammlung, Miinchen
(Sioli’s specimens); and the collection of
John T. Polhemus, Englewood, Colorado.
Etymology.—This species is named biae
for Maria Beatriz (Bia) Riviero do Vallé,
conservationist and our kind and efficient
host.
Habitat.— Most of the specimens of this
new species were collected from a small
stream in the lowland tropical forest near
our base camp. The stream was about | .5—
2 m wide and about 15-45 cm deep for
much of its course as it meandered and
flowed slowly through the forest. The stream
was mostly shaded throughout the day but
the collection site of this veliid was exposed
to sunlight for about an hour at noontime.
The first few specimens were observed on
the water at the shoreline (Fig. 16). When
attempts were made to net them, they ran
onto the leaf-covered bank where the leaves
had to be brushed aside to find them (Fig.
15). In so doing, I was surprised to find them
so abundant under the leaves that many
crawled back under the leaves faster than
they could be picked up. I quickly obtained
220 specimens within an area of approxi-
mately 2 square meters. All specimens were
found no farther than 1.25 meters from the
water’s edge. Additional collecting in the 10
365
meters downstream, where the bank was
shaded, yielded some specimens but dis-
tinctly fewer than I found in the sunny area.
Following a heavy rain about a week after
the first collections were made, I found 5
specimens of this veliid alive on the rain-
water and greatly diluted ethylene glycol in
a yellow plastic trough under a flight inter-
cept trap operating about a half kilometer
from the stream; evidently the bugs flew
from the stream and were captured by the
trap. Presumably, the veliid’s behavior of
moving about on the surface film kept them
from drowning as other insects did when
they flew into the intercept trap panel and
fell into the trough.
Possibly, when the heavy rains begin after
the dry season, as happened during this
fieldwork, the veliids disperse from areas of
dense population to new habitats with less
competition for food.
It is also of interest to note that all spec-
imens collected were winged adults. Per-
haps breeding begins after the onset of the
rainy season when new habitats become
available and the presumed dispersal of the
adults occurs.
ACKNOWLEDGMENTS
For making the fieldwork possible, I thank
the following: the Consorcio Nacional de
Engenheiros Consultores S.A. (CENEC) for
financial support; Paulo Vanzolini, who
kindly arranged for our participation in the
project; and Maria Beatriz (Bia) Ribeiro do
Valle, whose efficient logistical arrange-
ments made the trip so enjoyable.
I also extend my thanks to the following
individuals for their contributions to this
article: John T. Polhemus for kindly and
constructively reviewing the manuscript and
providing the specimens collected by H.
Sioli of the Max Planck Institute (Pl6n);
Victor Krantz, Photographer, Smithsonian
Institution Photographic Laboratory for the
habitus photograph; Young T. Sohn, Sci-
entific Illustrator, Department of Entomol-
366 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ogy, for the art work; Robin Faitoute, Mu- Hungerford, H. B. 1930. Three new Velia from South
seum Specialist for the micrographs; Phyllis America. J. Kans. Entomol. Soc. 3: 23-27.
M. Spangler, for preparing the specimens
for study and typing the manuscript.
LITERATURE CITED
Berg, C. 1883. Addenda et emendana ad Hemiptera
Argentina. Anales Sociedad Ciencias Argentina 16:
119-120.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 367-388
NEEM (AZADIRACHTA INDICA A. JUSS): ORGANISMS
AFFECTED AND REFERENCE LIST UPDATE
J. DAviD WARTHEN, JR.
Research Chemist, United States Department of Agriculture, Agricultural Research
Service, Plant Sciences Institute, Insect Chemical Ecology Laboratory, Beltsville, Mary-
land 20705.
Abstract. —A compilation of neem literature for the last decade is presented that updates
a review by Warthen 1979. The compilation includes literature citations and tables that
list organisms affected by neem as well as activities and compounds.
Key Words:
gosa tree, Indian lilac
In 1979, Warthen published a review of
the neem literature that was entitled “4 zad-
irachta indica: A Source of Insect Feeding
Inhibitors and Growth Regulators.” The re-
view was very popular worldwide with sci-
entists inolved in neem research as well as
with others who were either interested in
neem or in the utilization of neem for insect
pest control.
During the last decade, the number of
neem publications has increased greatly.
Since 1979, three international neem con-
ferences have been held; proceedings
(Schmutterer et al. 1980, Schmutterer and
Ascher 1983, 1987) of these meetings and
attendance have increased with each sub-
sequent conference.
During this time period, Vikwood Ltd.
formulated a neem seed extract (Margosan-
O), for which EPA approval was granted for
use on ornamental plants. Presently, W. R.
Grace & Co. and Vikwood Ltd. are working
cooperatively to market this product and to
gain EPA approval for its utilization on edi-
ble plants as well.
Worldwide interest has been generated by
neem as attested to in the 194 new citations
in the last decade within this compilation.
Neem, Azadirachta indica, Melia azadirachta, M. indica, Meliaceae, Mar-
The National Research Council, Office of
International Affairs, recognizing this inter-
est and the utilization potential for third
world countries, called a meeting in April
1988 to consider the possibility of publish-
ing a book on neem for the National Acad-
emy of Sciences.
Organisms affected by neem are listed in
Table 1; the entries, addressed mainly to
entomologists, are alphabetized by Phylum,
Class, Order, Family, Genus, Species, and
then Literature Citation. Activities, Com-
pounds, and Literature Citations of neem,
without reference to organisms affected, are
listed in Table 2; entries are alphabetized
by first author of the Literature Citation.
This compilation is not an exhaustive
search of the literature for the last decade;
many pre-1979 literature citations appear,
and others (Warthen 1979, Schmutterer et
al. 1980, Schmutterer 1981, Schmutterer
and Ascher 1983, 1987, Olkowski 1987, and
Jacobson 1988) should be consulted for fur-
ther information and other references.
The compilation updates the review by
Warthen (1979) and lists organisms in three
Phyla, Molusca being new, that are affected
by neem along with 277 (194 new) literature
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
368
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VOLUME 91, NUMBER 3
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(WpR6l ‘Te 19 Waypay)
(Z861 448d)
(P861 ‘Te 12 ppeT)
(P861 ‘Te 12 ppeT)
(8L61 ‘Ie 19 ppeT)
(S61 ‘Te 19 ppeT)
(S61 ‘Te 19 ppeT)
(€L61 BIdny)
(pL6l [ed pue ueyoes)
(€L61 BIdny)
(€L61 BIdny)
(T861 NIYD)
(snowAuouy)
(ZR6I YINWAZ[YOM Pure eIIdI0g)
(Q€861 Oseliqay)
(snowAuouy)
(TR61 YINWAsTYOAA puke eIIIIIg)
(SL61 JaJannuyds pure s}a—a}s)
($L6] Jasza1nuyog pure sja—ajS)
(CL6I $19018)
(O861 Ploquiay pue sd919];NWIYIS)
(S861 JoInTYyoS)
(P861 JoIn[yos)
(T861 “Te 19 ploquiay)
(O861 ‘12 19 pjoquiay)
(ZS6] JosaNUYyos pue a8ue7)
(ZR6] JasauNWYS pue adueT)
(Z86] Jo121INWIYIS puke aduRT)
a _—_.\.Q°rtr88.
SUONEUD JIN e13IT
aFrv—XKro —————————————LKS
Q-ursosieyy
O-uRsosIe|y
unyoerpeze
unyoespeze
unyorsipeze
unyorsipeze
unyorirpeze
unyoriipeze
unyoepeze
unyorspeze
unyoespeze
unyoestpeze
jeudon
ouexosas
Ices
punodwo,)
STWI91SAS
e1oidiq 19piO
JOUQIYUT YIMOIs
qua]jadoi
quayjadai
luepognue
JoVe[NSII YIMOI3
1o.e[N3aI YIMOIS
ymois
JUdLIO19p
Jouqryur yMoss
1UdLIO}0p
quejoRe
ue OPINe
JUROR
juepeaynue
quej90101d
JUBSIXO}
luepogjnue
jue199101d
uononpoldol
AVLAQSUO]
juepsaynue
jueidnisip yIMoi3
Jouqryur you
JoVe[N3II YIMOIT
jueidnisip yMoi3
1SI319UAS
1s1319UAS
1SI319UAS
AWANDW
pieyourlg anaiys pzAulolT
pieyourlg apaios vzAuoiurT
(IsqiaH) WinauDIspI WinN1]OgGM
(ISqiaH) UinaUuDdjIsvd UiNI]OgGIU
(ISQI9H) WiNAUDISDI WNIOGIM
(sqioH) WinauDd]spd UNIO
“TY ADO O1gaua
uBWIMAN DIIUOCdD! DIjIdog
UBWIMN DIIUOdD! DILIdog
ueUIMIN DIIUOdD! I]]Idog
UBWIMIN DIIUOdD! vDIIdog
ueuIMan vaIUodnl vijIdog
ueuIManN voIUodDl pijjidog
“4 DIDAIIS D1YI1410]0
DYSUII, SMDINSUI DIYLIOJOH
OYSUII SLDINSUI DIYIIAIOJOH
(paeyourlg) vauinsunsuod v1yo1djojoyy
adoy Sadiidnd pjpuoup
SUdAq WNMOUDAS DULIAPOSOA |
(J) anzduo snjiydons
(-q) avzduo snpiydons
(J) apzduo snpiydong
(suaydajs) snaulsniiaf SajsajojdAuD
URSIN S1SaalDa DUYIDjIdT
ques[nyy Sisaaliva DUYID]IdT
lues[njy Sisaaiiva DUuYyoDIdy
quURS[N Sisaaiuva DUYID|Id|
URSIN SiSaaliva DUYODIIdT
uURS[N, SsSaaLiva DUYIDIIGT
lueS[NY S17saaliva DUYODIdT
luURS[NY SUsaaliva DUYIDIIdT
JURS[NJ] S1JSaAliDA DUYIDIIGT
ques[nj Sisaaiioa DUuYyoo|Id_
JuURS[NJY S17saalia DUYID/Id|
saioadg pur snuan
odepizAWolsy
oepizAWolsy
oepluoLIqoua
depluoLqoua]
depluoLiqauay
oepluoLiqoua]
oeprluUoliqoua |
OEPlORGRILIS
SEPIOLGRILIS
aeploeqgeieoss
aeploeqRIiedss
aeploeqeieos
oeploeqeieog
aeploeqeieog
oeploeqReiedss
aeploeqeieos
oeploeqeseos
oeploeqeieosg
oepnsouliog
depruolnoiny
oepluolnoin)
depruolnoiny
oepifnon,)
dePT[[aUINIOD
dePI[[OUINIOD
IEPI[[PUINIO)
dePI][PUIIN0
dEPI[[PUIII0)
depl]JoUlNI0ZD
dBPI][OUINIOD
dBPI][AUINIOZD
dEPI][EUINIOD
dept [eulo90
dEPI[[9UINI0)
Ajtwey
‘ponunuoy “| 31qeL
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
370
(ZTL61 900snNy) U9.LI91]9p “UBIg SNIDIISD{ snI4apSAG deplLOsOYyLAd
(6L61 BXIquy pue weyriqy) [epronsesut “4 snypjnsuis snosapskq depluosoyAd
(6L61 ByIquiy pue weyeiqy) JUdLII19p “4 Snyojnsuid snosapsaq dePLOIOYyLIAg
(6L6] Bylquiy pure weyeiqy) juepsajnue “4 snipjnsuis snosapshq depuosoyLig
(SL6I 819918) (4aQotq) UnIDApOND viUusaig depuIy
(WpQ6] ‘Te 19 Waypay) ul yoelIpeze yIMol3 (se[[eq) sniviasn{ snjjadoouc oeploesAT
(O86I ‘Te 19 Waypay) unyorlipeze Joidnistp yMoi3 (sey[eq) snipiosp{ snyjadoouQ oeproesA]
(O861 ‘Te 19 Waypay) unyoerlipeze juBpsojrjue (se[jeq) snipiosp{ snjjadooucy oepraesAy
(6L6I ‘Te 19 WaJpay) unyoelipeze Jouqiyur 3unjow (sel[eq) snipiosp{ snyjadooucQ oepraesA qT
(L86| JeAvuley) unyorlpeze uononpoldal (seed) snipiosp{ snyjadoouc aeprlaesAq]
(L86] JeARUIey) unyoriipeze uonuinu (sel[eq) smipiosn{ snyjadoouQ aepraesAq
b191d01919]H IapiO
(ZR6] JoJOINWIYIS pue suayars) (uURWIapaI AM) DIDIIdDI $1111D49D oepniuyda y
(Q8Z6I ‘Te 19 UoyUeM) uluuryes WUdLA ap Surpsay “J polsauiop DISH oeprosnyy
(LL6| O8Y pue uped) “YT volsauiop vosnpy aeplosnyy|
(pR6] SoAeH pure qnoqgery) unyorlpeze JoViyur Bunpour I9aQH) aq SI/PULUNIND DISNPY oeplosnyjy
(9861 Z1IQ9Z) ul YyoRiipeze Aes snypiospjanbuinb xan) depron)
(9861 Z1IQaZ) unyorsipeze uolsry] isuaydajs sajaydoupr depron)
(986I Z11Q97) unyoriipeze (pleqosy_L) 1080] sapay aeproyny
(9861 Z11Q9Z) unyoRriipeze (J) uddsan sapay oeproryny
(PR61 Z1IGQ9Z) unyoerlpeze aplonoasul (J) uddsan sapay oeproyny
(S86I ‘[e 19 NIYD) JUdLID19p (uOseJ-POOM) ANZA40 D1]0ISAC. depUAWOPIII)
(S861 ‘Te 19 NIYD) quajadal (UOSEI-POOM) ANZAU0 D1]OASIC) depuUAWOPIII
(S861 ‘Te 19 2dr) unyoriipeze jeuortsod1ao (UURWIOpdI AA) DUAN DIIINT oepuoydryye)
(S861 ‘Te 19 29ry) unyoelipeze juepsayriue (UURLaPaAI AA) VULNI DIIINT oepuoydiyye)
(PR6l ‘Te 19 GqaM) unyoeiipeze OTW91SAS (ssading) 11/0/14] DZAWOLMIT oepizAWwoisy
(ER6T ‘Te 19 GqaM) (ssading) 11/0/14] DZAWOLIT oepizAwoisy
(O¢R6T ‘Te 19 MaIeT) JUBOIXO} (ssoding) 1/Ofi4] DZAULOLIT aepizAWol3y
(QSR6I ‘8 19 More) aprlonoasul (ssading) 1/0/14] DZAWUOLIT depizAwolsy
(QS86] ‘[B 19 More) quaj[adai (ssading) 17/0/14] DZAWOIT aepizAwolsy
(@GR6] ‘[e 19 MoleT) (ssading) 1/0/14] DZAWOLIT aepizAWolsy
(p86 ‘Te 19 MaleT) DTUUdISAS (ssading) 12/0/14] DZAWOLIT oepizAwolsy
(S861 ‘Te 19 Z1UOP\-[apouy) Q-ursosieyy aplonsasul (ssading) 1/Ofi4) DZAUOLIT aepizAWwol3sy
(€86] AJOO]L pue s0U008e4) [O11U0901q (ssoding) 1/Ofi4)] DZAWOLINT aepizAWOl3sy
(p86l Te 19 GqqaM) unyoelipeze STWI31SAS pieyourlg apaips DZAWOLIT aepizAwolsy
ee
suonety siniesay7] punodwo> AMANDW satsadg pure snuay Aye
‘ponunuoy “| aqeL
371
VOLUME 91, NUMBER 3
(Z861 NYO)
(9261 ‘Ie 19 JUSTIA)
yaya [enpIsal
UOIUIRI /Jau119 SUSIUIO]]AYd
(AaT[NULIAYIYIS pue stuaqd) ViUoUa/as soIsp
aepIe][IOBIH,
aeplawuo0an
(ZR61 YINWZ[YOA Pur eIIOIIg) que1a101d (A9IAT[Q) ]/2/DA19I DSOAJONS aeprryoajayH
(ZR6] Sepueyoy pue 1Aaq) juepoagjynue (J91AI]Q) D]/A]/DA4dI DIOAIONS aepiryoajanH
(LL61 aq pue uesy) uejsoiq jURpsgyUe (491AT]O) D//2/94499 VEOAIONG sepiryss[eH
(snowAuouy) juepoojnue (191AT[Q) ]/A]DA4II DIOAIONS depiryoajayH
(7861 2YOPY pue oqny) unyoriipeze SISOYIUASOTG (s1opunes) vjJaiddssos puoydouljIad depliysajayH
(8961 ‘Te 19 Jaled) 1Ud.I919p Jong 1a/0ou DjovsuLpy oepmnoay
eiojdopide] 19plO
(L861 ‘Te 19 MoleT) aprlonsasul (sanajadaq) vjyjisnd vsnuay depluIpaiy Udy
(O861 ‘Te 19 BWueYs) “J vaafiyjau sidy aepidy
(Z86I1 ‘Te 19 pjoquiay) unyoriipeze Joye[n3a1 yMol3 "T vaafijjau sidy oepidy
(1861 NaddozD pue pjoquiay) unyoriipeze Joye[N3a1 YMoIs “7 vduaafijjau sidy oepidy
(1861 eddozD pue proquiay) unyoRiipeze juepaajiue “"T viafijau sidy oepidy
(O861 ‘Te 19 ploqway) "Ty viafiyjau sidy avpidy
elaidouswAp IapiO
(S861 UooyuTYS) unyoelipeze juepaayiue (11S) Suasn) vIvAIDdD]IN oeproeydjaq
(P861 ‘Te 19 Buaxes) (TRIS) Suasn] vIDAMDdY]IN aeproeydiaq
(T861 NIYD) ([g1S) Suasny DIpALDdD]IN aeploeydjaq
(SS6] UeYY pue eudxes) JBALAINS paonpal aeploeydjag
(T86] ‘Te 19 ueddeteyy) UOTSSTWISUBI] SNLITA (lueISIq) suarsaiia XIIJaIOYdaN deplJapeoly
(S861 ‘12 18 2pAaH) Joydnisip ymoi3 (ue\siq) Ssuarsadia X111910YdaN deplapeoty
(TL6I ‘Te 19 RAD) 1Ua11919p ("q) avavyduiaAu unydisojpdoyy oepriprydy
(S261 ‘Te 19 SyWUD) jua[edas (49z[NS) apaisiad snzApy oepiprydy
(S861 ‘Te 19 laUpNoD) JUBSIXO} (snipeuuay) lonqn) vISIMiag depIpOIATTY
(C861 ‘1B 19 1aLIpNoD) 1Ud1I91ap (snipeuuad) long) visiuag aepIposAa[Vy
eINdOWOH JIpPiO
(P86 ‘Te 19 BIdIeD) unyoriipeze 1uBpsgjnuR [BIS suxyoid sniupoyy oeplanpay
(PS6l Te 19 BloIeH) (RUT YORIIpeze-]A1I0e-/ Jouqryur sunjow [RIS SHxOAd snIUupOoYYy oeprlanpoy
(P861 Ploquoy pur eidIey) unyoersipeze 1uBpsajue [BIS smxiod sniupoyy oepllanpoy
(Qr861 Noy) unyorlpeze juepoaue (4) ms1ua0y snr1apsaq deplosoy.LAd
(@p861 [Noy) unyoespeze (4) usiuaoy snosapsdq aepluosoy.uhg
(ZL61 909sny) quo|jado1 “UBIg SNIDIOSD/ SnNILaPSAG aepluoooyg
(ZL61 aoosny) Jojdnisip yimoi3 “UBIS SNIDIISD{ SNI4APSAG aepLosoyAd
suonely oiniesaiy punodwo,) AWANDY saisadg pur snuay Aytwey
‘ponunuoy “| aqeL
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
372
(O861 ‘Te 19 Waypay) unyoesipeze Joidnisip yous (yg “gq “f) Dpsadisnaf viaidopody oepinioon)
(OR61 ‘TB 19 ULaJpay) unyoRiipeze juepsajiue (yw “gq ‘f) vpsadisnaf p1ajdopods depINIOON,
(L861 Bye) unyoeiipeze 1Ud19]ap BuIpaagy (yt “A “f) Dpsadisnaf p1ajdopody oepiniooN
(S861 ‘[B 19 997) aprlonoasul (yuws “gq f) ppsadisnal d1ajdopody aepinjoon
(7861 FXO pue oqny,) unyoriipeze SISOYIUASOIG (yt “gq “f) Dpsadisnsf p1ajdopody aepinjoon
(986[ ‘[e 19 JoyRyqeid) (douqnH) VNSIXA D4ajdopods depIniooN,
(L861 FIqQuiniy pue 10) sisualsUulany] Snij1Ivg JUBOIXO1 (49uqnH) ONsIXa D4ajdopods depInjoOON
(L861 Quins pur 1e0|\) UNDIULIOAR JUBOIXO} (douqnH) ONsIXA DAajdopods oepinjooN,
(S861 ‘Te 19 Laprq) unyoripeze juepsajrue (4gWeID) DIUDpIsa Diajdopody oepiniooN
(S861 ‘Te 19 Waprq) splououll| JUBpsajtUe (49WRID) DIUuDplia Dsajdopody depInIOON,
(€861 OH) juepoajnue uosduwiey SijsilunjD2 DIWDSAS depInjooN
(ERG [B19 JoONINWYIS) onouasoydiow (ZO¥[e AA) VIDANdAS Dijajppnas_ oepinjooN
(Q86] NzruYs) O-ursosieyy onauasoruisds (“T) ADIISSDAG DAISAUD PY oepinioon
(S861 NzruIYysS) unyoriipeze onauoso0ruiads (“T) ADIISSDAG DAJSOUID PY aepmniooN
(7861 NIYD) IIOOJ, NG]DUAA DIUDINIT oepinjooN
(ZR61 AYO pure oqny) unyoriipeze sIsayUASOIG (QIPpog) vaz siyjolapy] aepiniooN
(7861 AYO puke oqny) unyorlpeze IOIQIYU SISApdo (QIPpog) Yaz s1yjolay oepinjooN
($L6I ‘[e 19 OUURZ) 1Ud1I919p (4) Suadsadia S1yJONAH dPPINIOON
(ZL6] 909snNy) jua[jedal (4) Suarsaaia SIYJONAH oepinioon
(Z7L6| 209sny) Joidnisip yimoid ((4) Suarsadia SIYIONAH depmnjoon
(Z7L6] 200sny) 1ua1191]9p (4) SUaISasIA S1YIONA PL oepinjooN
(S861 ‘TB 19 997) Oprlonoasul (4) SuadsadiA SIYIONAH oepInioon
(7861 AYDOTY pure oqny) unyoriipeze SISOYyLUASOIG (4) Suarsaaia S1yIONaH depInjooNn
(T86I ‘[e 19 Joustay)) uluue[es 1UdLIOIOpP BuIpsay ([RANPSIOg) DUD/NSUI SOLID] oepinjooN
(1861 ‘[e 19 JoustayA)) unyoripeze 1UdLIa]ap BUIpsay ([RANPSIog) DUD/NSU! SDD] oepInjooN
(9g6| BUYSLTyY pue yeUIed) Aqyyiqeyoiey 339 (SNIDIGRS) Djjaijia SDD depinjooN
(986 BUuUSLTy pue yeuyIeg) uontsodiao (SNIOLIQRS) D/jaljiA SOLID T oepinjooN
(7861 NIYD) (QQUaND) DUISsdqn (vIajdopods) vSIyIDUp depIniooN
(CQ6] URIeYpl[einjY puke Ley) [O11U09 199sUI (‘q) vinuvl panyopy depiInjooN
($L6| JousIaJy pure e[[N1eyS) (J) 4pdsip DiujUuDUAT oepILyUeWAT
(QI L6| tuemiof pue ueypeig) JOy[eA VIDUN] S1jI0AdNZ OePILURWAT
(W[L6| tuemiof pue ueypeld) JOX[eA VIDUN] SIIOAdNA depiLUeWAT
(Q96] luemiof pue ueYypeld) qud.L919p JOxeA VIDUN] S1JI0AdN_ depILUeWIAT
(€96] ‘[e 19 uRYpeld) quajjada1 JOy[eA VIDUN] SIjI0AdN| depsUeWAT
(696 Log pue nqeg) 1Ud1I919p JayeA VIDUN] S1I0AdN depILNUPWAT
suOne UD ainjesaiiT punodwo> AWAIDY salsedg pure snuan Awe
‘ponunuoy “| 91qeL
373
(S861 ‘Te 19 UOseUTY) unyoriipeze aplonsasul (49UQNH) SHpIqnu vIULAISE oeprescg
(S861 ‘Te 19 UOseULY) uNYyoRIIpeze juepaaynue (s9uqNH) SYDjIgnu vIULISO aepyeidg
(S861 ‘Te 19 UOOsUTYS) unyoerspeze Jo\dnisip yMos aguany sijpoDUdnf DIULISE aepyeicg
(€861 OH) luepsajtue 1ouoseY Yjjaipindas Dydd jy oepryelAg
(O861 ‘Je 19 BPUeYS) JIJIZ Vjjaiuyany vusaydy oepreidg
(Z861 ‘Te 18 pjoquiay) unyoesipeze Joye[N3a1 YMoI3 JOJIZ Vjjatuyany visaydy aepresAg
(O86 ‘Te 18 pjoquiay) Jojdnisip yMos3s JO[[9Z Vjjaluyany vijsaydT oepresAdg
(ZT861 YINWIs|YOA pue PIIO1Oq) que199101d (19ye MA) DjJainv2 vINsaydy aepleicg
(1861 aduey] pue s9u0038e4) JONJIZ SIYJoUIg VIULOJOP1IOAD oepreidg
(E861 ‘Te 19 JoatNWY IS) oneuasoydiour (QoUeND) SIMUIpaU s1II0AI0]DYdDUD oepresidg
(LL61 eypeYyD) JUdLIO19p Jeyouodng Sijpuny]pIp) vAISDSI]UP oepryeidg
(T861 NIYD) ("q) andp. suaid depot
(CL6I ‘Te 19 OUURZ) Ua1I919p (J) avaissvag Sal aeplaig
(TL61 s0osny) qua|jadar ("T) avaissp.aq Sal dePLIig
(ZL6]1 VOOsnyY) Joidnisip yMols (“J) avaisspag Sidi dePLoid
(ZL61 90osny) JUdLIO19p ("]) ADIISSDAG S141 dePLdid
(6L6] deuypoy) unyorlpeze jua|[edor ("T) Snajowap oyidog sepruorided
(9861 ‘Te 19 JayeYyqrid) Jojensa1 yMoIs (gouqn}) 1 vIsn/doyot depinjooN
(QIZ6[ tuemiof pue ueYypesg) (‘4) vanqy viaidopods aepiniooN
(B[ 26] tuemiof pue ueypeid) (‘4) Dunit) vsaidopods depInlooN
(896] luemiof puke ueypeid) UdLII1Op ((4) panqy vaaidopods aePINIION
(C961 ‘Te 19 UeYypeig) jug|[odo1 (4) Dani vsaidopods depInjoOON
($L61 ‘Te 19 1ysof) 1Ud.LI9}9p (4) Dani vsaidopods oepiniooN
(SL61 Peseidewey pue rysof) juepoajnue (4) vanity viaidopods aepinjlooN
(€R61 BLOIYIP pure seiny) unyoespeze auoWIOY a[tusant (4) vanity viaidopods depINIoON
(Z861 MIYD) (4) pani) viajdopods aepinjoon
(C861 ‘1B 10 JousIay\) proe otund (JeANpsIog) s1/740}]1) viaidopods depInlooN
(E861 Joyosy pue JousIay)) Joje[nsa1 YIMoIs3 (jeaAnpstog) 81/0/0111) Diaidopods depInlooN
(T86I ‘Te 19 JaUsIay\) uruury[es luepoaynue ({BAnpsiog) 87/0.401]1] D41aj]dopody aepmnioon
(1861 ‘[B 19 JoUsIay\) unyoeiipeze 1UdLa1ap BUIpsay (JRANpsiog) s1/p40}11) Dsaidopods dePINIION
(S861 Pades-[q) quay[adar ({RAnpstog) s7/0./0)]1] p4aidopody oepINIOON
(S861 Pedes-1y) juBpaagyue (JRANpsIog) s1/p40}]1] Déaidopods oepInjooN
(B8/6] ‘Te 19 UsyUeM) unyoeiipeze luepoajnue (yg “yo “f) Dpsadisnuf v1ajdopody oepmniooN
(QE86T ‘Te 19 SaxoIS) Juepsgjnue (yilug “q [) vpiadisnuf viaidopods depInjooN
(BERS [eB 19 SaxOIS) Juepsogjnue (yitWg “q “f) Dpsadisnif D1ajdopody aepinjooN
(QPS6l ‘ek 19 Waypay) juepoagjnue (yg “qo “f) Dpsadisnuf viaidopody oepInjooN
SUONEID dN eIIIIT punodwo> AWAY saioadg pure snuay Ayre
‘ponunuo)y “T aqeL
VOLUME 91, NUMBER 3
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
374
(6L61 pewyy pure ped)
(€961 NeIND pue eyurs)
(L861 Y8ulg pue ysurs)
(Q1L61 tuemiof pue ueypelg)
(ET L6] tueMiof pue ueypelg)
(8961 tuemior pue UeYpesg)
(€961 ‘Te 19 URYpeIg)
(L961 ‘Ie 19 atAeq)
(TL61 ‘Te 10 [eAoy)
(1L61 SIMOT Pur [IID)
(IL61 Stay pur 11D)
(1L6] uesi0 pue yUoMaiINg)
(896 UesIOW Pur YUOMIIIINg)
(LL61 Uny)
(€961 NeIND pure eyurg)
(pR6] Joueg puke woyeys)
(p86] Joueg puke woyeYs)
(E861 ‘Te 19 pjoquiay)
(E861 ‘Te 19 pjoquiay)
(Q1L61 tuemiof pue ueypeig)
(81 6] tuemiof pue ueypesg)
(S96] luemiof pure ueYypelg)
(L86] Jeavuiey)
(L86] Jeavuiey)
(986| snowAuouy)
unyorspeze
OLUPTToU
unyoriipeze
unyorsiipeze
unyoesipeze
unyoepeze
unyoeslpeze
unyoesipeze
unyoespeze
unyoesipeze
unyoeipeze
unyoesipeze
sapronsad jueld
[O11U09 1sad
Aupiqeyoiey 382
1Ud11919p
quaj[adai
juepsgtue
JUd1I919p
OTWIS1SAS
1UdLI919p
1Ud11919p
1Ud11919p
quaL91ap
JOmuod 1sad
auowoy arusant
JOIARYyaq Jenxas
1o1e[n3a1 YIMoIs
Jouqryur you
quoLa1ap
uononpoldal
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376
Table 2.
Activity
pyrethrin stabilizer
miscella refining
deterrent
pharmaceutical
antifeedant
antifeedant
feeding inhibitor
repellent
biosynthesis
crop protection
feeding inhibitor
leaf development
antifeedant
natural resistance
Compound
neem
neem oil
azadirachtin
limonoids
azadirachtin
azadirachtin
kaempferol
nimbin
quercetin
B-sitoserol
azadirachtin
azadirachtin
azadirachtin
tetranortiterpenoid
azadirachtin
azadirachtin
meldenin
azadirachtin
nimbolide
gedunin
nimbolin b
nimbolin a
nimbolide
azadirachtin
meliacin
tetranortriterpenoid
salannolide
tetranortriterpenoid
nimbraflavone
nimbin
azadirachtin
nimbin
salannin
salannin
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Activities, compounds, and literature citations of neem without reference to insects.
Literature Citation
(Ahmed et al. 1976)
(Ahuja et al. 1976)
(Anonymous 1977)
(Anonymous 1979b)
(Anonymous 1980)
(Anonymous 1981)
(Anonymous 1982a)
(Anonymous 1982b)
(Anonymous 1982b)
(Anonymous 1985a)
(Anonymous 1985b)
(Arigrabu and Don-Pedro 1971)
(Aschenbach 1982)
(Basak and Chakraborty 1968)
(Basak and Chakraborty 1968)
(Basak and Chakraborty 1968)
(Basak and Chakraborty 1968)
(Bilton et al. 1985)
(Bilton et al. 1987)
(Broughton et al. 1985)
(Bruhn et al. 1984)
(Butterworth et al. 1972)
(Chadha 1986)
(Chiu 1983)
(Connolly et al. 1968)
(Doria 1981)
(Duke 1983)
(Ekong 1967)
(Ekong et al. 1969)
(Ekong et al. 1969)
(Ekong et al. 1969)
(Ekong et al. 1971)
(Fagoonee 1979)
(Forster 1983)
(Garg and Bhakuni 1984a)
(Garg and Bhakuni 1984a)
(Garg and Bhakuni 1984b)
(Garg and Bhakuni 1984b)
(Garg and Bhakuni 1984c)
(Gilbert 1982)
(Harris et al. 1968)
(Harzal 1977)
(Helson 1984)
(Henderson et al. 1963)
(Henderson et al. 1964)
(Henderson et al. 1968)
(Hoddy 1985)
(Jacobson 1980)
(Jacobson et al. 1983)
(Jacobson 1986)
(Jain 1983)
VOLUME 91, NUMBER 3 377
Table 2. Continued.
Activity
Compound
Literature Citation
general uses
antifeedant
antifeedant
antifeedant
antifeedant
control agent
antifeedant
insecticide
systemic
antifeedant
repellent
repellent
feeding inhibitor
antifeedant
general uses
general uses
general uses
17-8-hydroxyazadiradion
epi-azadiradion
1,3-diacetylvilasinin
3-deacetylsalannin
salannol
tetranortriterpenoids
meliacin
azadirachtin
azadirachtin
azadirachtin
azadirachtin
deacetylazadirachtinol
piperony! butoxide
neem toxin
tetranortripterpenoids
azadiradione
azadirone
epoxyazadiradione
gedunin
tetranortriterpenoids
azadirachtin
azadirachtin
azadirachtin
chemicals
nimbin
constituents
nimbidic acid
nimbidinin
nimbidic acid
nimbidinin
azadirachtin
azadirachtin
nimbin
nimbin
nimbin
nimbin
nimbin
nimbin
nimbinic acid
nimbinin
vepinin
polysaccharide
vilasinin
(Ketkar 1976)
(Ketkar 1979)
(Kraus and Cramer 1978)
(Kraus and Cramer 1978)
(Kraus and Cramer 1981)
(Kraus and Cramer 1981)
(Kraus and Cramer 1981)
(Kraus et al. 1981)
(Kraus 1983)
(Kraus et al. 1985)
(Kraus 1986)
(Kubo 1979)
(Kubo and Klocke 1981)
(Kubo et al. 1984)
(Lange and Feuerhake 1984)
(Larew 1985)
(Larew 1988)
(Lavie and Jain 1967)
(Lavie et al. 1971)
(Lavie et al. 1971)
(Lavie et al. 1971)
(Lavie et al. 1971)
(Madhusudanan et al. 1984)
(Malik and Naqvi 1984)
(Malik and Naqvi 1984)
(Mansour and Ascher 1983)
(Mitra et al. 1947)
(Mitra 1957)
(Mitra 1963)
(Mitra et al. 1970)
(Mitra et al. 1970)
(Mitra et al. 1971)
(Mitra et al. 1971)
(Morgan and Thornton 1973)
(Nakanishi 1975)
(Narasimhan 1959)
(Narayanan et al. 1962)
(Narayanan et al. 1964a)
(Narayanan et al. 1964b)
(Narayanan et al. 1964c)
(Narayanan and Pachapurkar 1965)
(Narayanan and Pachapurkar 1966)
(Narayanan et al. 1967)
(Narayanan et al. 1969)
(Nayak et al. 1978)
(Olkowski 1978)
(Pachapurkar and Kornule 1974)
(Parmar 1984a)
(Parmar 1984b)
(Parmar 1985a)
378
Table 2. Continued.
Activity
nitrogen fixing
antifeedant
antinuclear study
deterrent
general uses
general uses
growth regulator
antifeedant
repellent
repellent
spermicide
mode of action
general uses
contraceptive
poisoning
sunlight effect
cardiovascular
pests of neem
antifeedant
antifeedant
nitrogen fixing
phagorepellent
growth disruptor
systemic
Compound
chemical constituents
azadirachtin
nimbidin
tiglic acid
azadirachtin
vepaol
azadirachtin
azadirachtin
warburganal
sodium nimbidinate
bitter principle
nimbidin series
azadiradione
17-hydroxyazadiradion
nimocinol
tetranortriterpenoid
azadirachtin
fatty acids
azadirachtin
kaempterol-3-glucoside
myricetin-3-1-arabinoside
quercetin-3-galactoside
chemical constituents
azadirachtin
nimbidin
nimbin
azadirachtin
azadirachtin
azadirachtin
azadirachtin
azadirachtin
nimbin
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Literature Citation
(Parmar 1985b)
(Parmar 1985c)
(Parmar 1985d)
(Peterson 1983)
(Pickett 1985)
(Pillai and Santhakumari 1984)
(Pradhan et al. 1962)
(Quasim and Dutia 1970)
(Radwanski 1977a)
(Radwanski 1977b)
(Radwanski 1977c)
(Radwanski 1977d)
(Radwanski and Wickens 1981)
(Raman and Santhanagopalan 1978)
(Rembold 1984)
(Sankaran et al. 1984)
(Schroeder and Nakanishi 1987)
(Schwinger 1982)
(Schwinger 1982)
(Sharma and Saksena 1959)
(Siddiqui 1942)
(Siddiqui and Mitra 1945)
(Siddiqui et al. 1975a)
(Siddiqui et al. 1975b)
(Siddiqui et al. 1978)
(Siddiqui et al. 1984)
(Siddiqui et al. 1984)
(Siddiqui et al. 1986)
(Siddiqui et al. 1988)
(Sieber 1985)
(Sinha and Gulati 1969)
(Sinha et al. 1984)
(Sinniah and Baskaran 1981)
(Skellon et al. 1962)
(Stokes and Redfern 1982)
(Subramanian and Nair 1972)
(Subramanian and Nair 1972)
(Subramanian and Nair 1972)
(Thompson and Anderson 1978)
(Tirimanna 1983)
(Uebel et al. 1979)
(Uthamasamy et al. 1973)
(Verma 1974)
(Verma 1974)
(Warthen 1979)
(Warthen et al. 1984)
(Watanabe et al. 1981)
(Yamasaki et al. 1986)
(Zanno 1974)
(Zanno 1974)
(Zanno 1974)
(Ziffer et al. 1966)
VOLUME 91, NUMBER 3
citations; the review by Warthen (1979)
contained 105 citations. Within the Phylum
Arthropoda, 7 Orders are represented in
Class Insecta with Hymenoptera being a new
order affected by neem. Since the review by
Warthen (1979), additional species affected
by neem have been added to the literature:
11 in Coleoptera, 9 in Diptera, 4 in Het-
eroptera, 3 in Homoptera, 22 in Lepidop-
tera, and 2 in Orthoptera.
Although literature is extensive and pro-
gress in research has been accomplished, the
commercial development of a marketable
natural insect pest control agent has been
painstakingly slow. However, the novel
chemistry within these literature citations
pertaining to neem has been a stimulus to
chemists to synthesize active analogs that
might have the possibility for commercial
development as insect feeding deterrents or
growth regulators.
LITERATURE CITED
Abraham, C. C. and B. Ambika. 1979. Effect of leaf
and kernel extracts of neem on moulting and vi-
tellogenesis in Dysdercus cingulatus Fabr. (Het-
eroptera: Pyrrhocoridae). Current Science 48: 554—
556.
Adhikary, S. 1985. Results of field trials to control
the diamond-back moth, Plutella xylostella L., by
application of crude methanolic extracts and
aqueous suspensions of seed kernels and leaves of
neem, Azadirachta indica A. Juss, in Togo. Z. Ang.
Ent. 100: 27-33.
Ahmed, S. M., M. R. Gupta, and H. M. Bhavanagary.
1976. Stabilization of pyrethrins for prolonged
residual toxicity Part—II: Development of new
formulations. Pyrethrum Post 13: 119-23.
Ahuja, M. M., R. R. Gupta, K. N. Agrawal, and A. C.
Gupta. 1976. Miscella refining ofneem & mahua
oils. Indian J. of Technology 14: 257-259.
Ali, S. I., O. P. Singh, and U.S. Misra. 1984. Effec-
tiveness of plant oils against pulse beetle Ca/lo-
sobruchus chinensis Linn. Tropical Stored Prod-
ucts Information No. 47: 84/7.
Anonymous. Neem Cake Blended Urea for Nitrogen
Economy. Directorate of Non Edible Oils & Soap
Industry. Khadi & Village Industries Commission,
Bombay.
1977. Five years of NIAB. Nuclear Institute
for Agriculture and Biology 4/6/72-4/6/77.
1979a. Extract from Indian tree bugs crop
pests. Chemical Week 125: 50.
379
1979b. Neem tree seed extracts repel Jap-
anese beetles. Agricultural Research, USDA 27:
8-10.
1980. Firewood Crops. National Academy
of Sciences. Washington, D.C., p. 114-117.
. 1981. Natural products repel cucumber bee-
tle. Agricultural Research, USDA 30: 12.
1982a. Neem tree may be source of safe in-
secticides. The IRRI Reporter, June.
1982b. Science watch—lemon oil shrinks
bugs. The New York Times, May 4.
. 1985a. Pesticides, new weapon: It’s a natural.
Industrial Chemical News 6: 12-13.
1985b. Science watch—chemicals of neem
tree deter pests. New York Times, September 3.
1986. Natural neem kills cockroaches and
greenhouse pests. Agriculture Research, USDA,
ARS 34: 13-14.
Arigrabu, S. O. and S. G. Don-Pedro. 1971. Studies
on some pharmaceutical properties of Azadirachta
indica or Baba Yaro. African J. of Pharmacy &
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Arnason, J. T., B. J. R. Philogene, N. Donskov, M.
Hudon, C. McDougall, G. Fortier, P. Morand, D.
Gardner, J. Lambert, C. Morris, and C. Nozzolillo.
1985. Antifeedant and insecticidal properties of
azadirachtin to the European corn borer, Ostrinia
nubilalis. Entomol. Exp. Appl. 38: 29-34.
Asari, P. A. R. and D. Dale. 1977. Studies on the use
of antifeedants for protecting stored paddy from
Angoumois grain moth, Sitotroga cerealella. Bul-
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Aschenbach, J. 1982. Prefer toothbrush. Harrisville,
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Ascher, K. R. S. and R. Gsell. 1981. The effect of
neem seed kernel extract on Epilachna varivestis
Muls. larvae. J. Plant Diseases and Protection 88:
764-767.
Attri, B. S. 1977. Utility of neem oil extractive as
feeding deterrent to locust. Indian J. Entomol. 37:
417-418.
Babu, T. H. and Y. P. Beri. 1969. Efficacy of neem
(Azadirachta indica Juss) seed extracts in different
solvents as a deterrent to the larvae of Euproctis
lunata Wik. Andhra. Agr. J. 16: 107-111.
Basak, S. P. and D. P. Chakraborty. 1968. Chemical
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Batra, H. N. 1980. Preliminary field trials of seed
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gregarious locust hoppers in Rajasthan desert. Per-
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Bilton, J. N., H. B. Broughton, S. V. Ley, Z. Lidert, E.
D. Morgan, H. S. Rzepa, and R. N. Sheppard.
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Bilton, J. N., H. B. Broughton, P. S. Jones, S. V. Ley,
380
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Bruhn, A., M. Bokel, and W. Kraus. 1984. 4a,6a-
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tripterpenoid aus Azadirachta indica A. Juss (Me-
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Butterworth, J. H.and E. D. Morgan. 1968. Isolation
of a substance that suppresses feeding in locusts.
Chemical Communications, 23-24.
1971. Investigation of the locust feeding in-
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Chadha, M. S. 1986. Trends in the application of
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Chadha, S.S. 1977. Use of neem (Azadirachta indica
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gastra catalaunalis Dupon. (Lepidoptera: Pyrali-
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Chakravorty, D. P., G. C. Ghosh, and S. P. Dhua.
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Chari, M. S. and C. M. Muralidharan. 1985. Neem
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Chavan, S. R. 1983. Chemistry of alkanes separated
from leaves of Azadirachta indica and their larvi-
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
liaceous plants on rice gall midge (GM) oviposi-
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Connolly, J. D., K. L. Handa, and R. McCrindle.
1968. Further constituents of nim oil: The con-
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Coudriet, D. L., N. Prabhaker, and D. E. Meyerdirk.
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Devi, D. A. and N. Mohandas. 1982. Relative efh-
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Doria, J. 1981. Neem: The tree insects hate. Garden,
July/August, 8-11.
Duke, J. A. 1983. Azadirachta indica A. Juss, Me-
liaceae, neem tree. Personal communication, 1-5.
Egunjobi, O. A. and 8. O. Afolami. 1976. Effects of
neem (Azadirachta indica) leaf extracts on popu-
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growth and yield of maize. Nematologica 22: 125-
1132.
Ekong, D. E. U. 1967. Chemistry of the meliacins
(limonoids). The structure of nimbolide, a new
meliacin from Azadirachta indica. Chemical Com-
munications, 808.
Ekong, D. E. U., C. O. Fakunle, A. K. Fasina, and J.
I. Okogun. 1969. The meliacins (limonoids).
Nimbolin A and B, two new meliacin cinnamates
for Azadirachta indica L. and Melia azedarach L.
Chemical Communications, 1 166-1167.
Ekong, D. E. U., S. A. Ibiyemi, and E. O. Olagbemi.
1971. The meliacins (limonoids). Biosynthesis of
nimbolide in the leaves of Azadirachta indica.
Chemical Communications, 1117-1118.
El-Sayed, E. I. 1985. Neem (Azadirachta indica A.
Juss) seeds as antifeedant and ovipositional re-
pellent for the Egyptian cotton leafworm Spodop-
tera littoralis (Boisd.). Bull. Ent. Soc. Egypt, Econ.
Ser. No. 13: 49-58.
Fagoonee, I. 1979. The potential of natural products
in crop protection in Mauritius. Proc. of the Na-
tional Agricultural Production Conference, Mau-
ritius, Dec. 10-15, 201-217.
Fagoonee, I. and G. Lauge. 1981. Noxious effects of
neem extracts on Crocidolomia binotalis. Phyto-
parasitica 9: 111-118.
Fagoonee, I. and V. Toory. 1983. Contribution to the
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NOTES ON PLEUROLITHOBIUS OF TURKEY
(CHILOPODA: LITHOBIOMORPHA)
MARZIO ZAPPAROLI
Istituto di Difesa delle Piante dell’Universita della Tuscia, Via San Camillo de Lellis,
1-01100 Viterbo, Italy.
Abstract. —The Turkish centipedes belonging to Pleurolithobius Verhoeff, 1899 are listed
and discussed: P. jonicus Silvestri, 1896, previously known only through doubtful data,
and P. orientis (Chamberlin, 1952) (n. comb.), previously recorded only on the original
description, are fully redescribed and new faunistic data are given.
The following new synonymies are proposed: Turkobius Chamberlin, 1952 = Pleuroli-
thobius Verhoeff, 1899; Archilithobius integrior caducus Chamberlin, 1952 = Pleuroli-
thobius orientis (Chamberlin, 1952); Pleurolithobius atopior Chamberlin, 1952 = Pleu-
rolithobius orientis (Chamberlin, 1952).
Key Words: centipedes, Pleurolithobius
The object of this paper is to summarize
our knowledge of the species of the genus
Pleurolithobius Verhoeff, 1899 presently
known from Turkey (as politically consti-
tuted today). Two taxa are recognized for
this area: P. jonicus Silvestri, 1894 and P.
orientis (Chamberlin, 1957).
For each taxon, literature records for the
study area are reported, material examined
is listed geographically from N to S and from
W to E, geographical distribution is dis-
cussed, description of material examined
including taxonomical notes is provided.
The following abbreviations have been
used: MZ = coll. M. Zapparoli; W = coll.
R. V. Chamberlin, National Museum of
Natural History, Smithsonian Institution,
Washington; vil. = vilayet (= province).
The original labels accompanying the type
specimens are quoted integrally and indi-
cated in quotation marks (“ ”’).
Pleurolithobius jonicus Silvestri, 1896
?Pleurolithobius jonicus: Matic, 1980: 98.
Material examined.—1 @, vil. Canakkale,
Truva, m 70, 23.1V.1982, A. Vigna leg.
(MZ); 4 22, vil. Canakkale, dint. Ayvacik,
m 350, 23.1V.1982, M. Bologna leg. (MZ);
3 34, 4 22, ibidem, A. Vigna leg. (MZ): 4 44,
1 2, ibidem, M. Zapparoli leg. (MZ); 1 2, vil.
Izmir, Izmir, 15.1V.1973, V. Sbordoni leg.
(MZ): 1 4, 1 2, vil. Balikesir, dint. Havran,
50 km prima di Balikesir, m 450,
23.1V.1982, M. Zapparoli leg. (MZ).
Distribution.—Southern Italy (excl. Sic-
ily), southern Yugoslavia (Montenegro), Al-
bania, mainland Greece and Ionian Islands,
southern Bulgaria and western Anatolia.
Matic and Golemansky (1964) record P.
jonicus from Crete, but this record 1s doubt-
ful and needs confirmation.
Description.—Size 9.5-12.0 mm _ long,
1.25-1.27 mm broad at T. 10; color light
brown.
Head slightly wrinkled, broader than long
and broader than T. 1, posterior border
straight, posterior marginal ridge with me-
dian thickening, lateral marginal interrup-
390
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-4. Pleurolithobius jonicus Silvestri, 1896 (Ayvacik). 1, Right ocelli and organ of Témésvary (TS).
2, Dental margin of prosternum, ventral. 3, Right female gonopod, ventral. 4, Left female gonopod, dorsal.
Scales: |—0.2 mm; 2-4—0.1 mm.
tons present; antennae about a third of body
length with 32-41 articles, the terminal one
about 2-3 times as long as penultimate; ocelli
1+ 1,2 (Fig. 1), depigmented or slightly pig-
mented, postero-superior ocellus about the
same size of principal ocellus, organ of T6-
mdsvary of the same size or slightly larger
than secondary ocelli; prosternum (Fig. 2)
with 2+2 teeth, porodont setiform, lateral
to porodont the shoulders are absent or
barely distinct.
Tergites slightly wrinkled; T. 1 broader
than T. 3, almost rectangular with posterior
border straight or slightly sinuated; lateral
border parallel in TT. 3, 5, 7 and 8, slightly
convergent posteriorly in T. 10, posteriorly
convergent in TT. 12 and 14; posterior an-
gles rounded in TT. 3, 5, 7 and 8, angled in
TT. 10, 12 and 14; posterior border straight
or slightly sinuated in TT. 3, 5 and 7, sin-
uated in TT. 8 and 10, emarginate in T. 12,
straight or emarginate in T. 14; TT. 9 and
11 without triangular projections on the
posterior angles, T. 13 generally without tri-
angular projections, sometimes only slightly
produced; intermediate tergites with round-
ed posterior angles and straight posterior
border; T. 16 of the male with lobate pro-
jection at the posterior angles as figured in
Matic and Golemansky (1964: fig. 1).
Anterior legs with tarsal articulations dis-
tinct, see Table 1 (48) and Table 2 (22) for
spinulation; coxal pores 4,3,3,2 or 4,3,3,3
(68), 5,4,4,4 or 4,4,4,4 (22), circular, sepa-
rated from one another by a space equal,
greater or smaller than their own diameter,
females with proximal pore of coxae XII
geverally smaller; males with femur and tib-
ia XIII evidently thickened (see Matic and
Golemansky 1964: fig. 1), XIV legs not
thickened, male with distal end of tibia XV
with dorsolateral swelling bearing a tuft of
VOLUME 91, NUMBER 3
391
Table 1. Pleurolithobius jonicus Silvestri, 1896. Spinulation (*), 44; letters in parentheses indicate variable
spines.
Ventral Dorsal
G tr Pp F T C tr P F T
1 m _ - (p) a a
2 = — _ (m) m _ - (p) a(p) a
3 - — = (m) m = = (p) ap a
4 = — = (a)m m - _ (p) ap a(p)
5 = = = (a)m (a)m = = (p) ap ap
6 — — — am am — — (p) ap ap
78 = = (m) am am — — (p) ap ap
9 = = m(p) am(p) am = = (p) ap ap
10 = = m(p) am(p) am = = (m)p ap ap
11 = = m(p) am(p) am ~ - (m)p (a)p (a)p
12 - — mp am(p) am — _ mp p p
13 — (m) mp m m — _ (m)p p —_
14 — m mp m m - = mp p _
15 - m m(p) m — — mp p =
* C = coxa, tr = trochanter, P = prefemur, F = femur, T = tibia; a = anterior spur, m = medial, p = posterior.
some setae (see Matic and Golemansky
1964: fig. 1), male with DpP XV spine in-
serted on the latero-internal side of the ar-
ticle, females with DpP XV spine normally
positioned; females with XIII-XV legs
without special modifications; apical claw
of XV legs with accessory claw one-half or
two-thirds of the principal claw length; glan-
dular pores on XII-XV legs.
Male first genital sternite with 8-10 setae,
second genital sternite without setae, gon-
opods without apical setae.
Table 2.
See Table | for codes explanation.
Female gonopods (Figs. 3 and 4) with 3+3
long conical spurs, progressively longer and
larger from the internal one to the external,
internal spur generally lying behind the in-
termediate spur, apical claw narrow and
without lateral denticles; basal article with
a group of 6 relatively strong dorsolateral
setae and a group of 6-7 dorsomedial setae,
as strong as the dorsolateral setae, posi-
tioned near the insertion of the spurs, sec-
ond article with 7-9 dorsolateral setae ar-
ranged in two rows and two dorsomedial
Pleurolithobius jonicus Silvestri, 1896. Spinulation, 92; letters in parentheses indicate variable spines.
Ventral Dorsal
C 7 Pp F T ic tr P F T
l = = = (a) (a)(m) = = (p) a(p) a
2 = = ~ (a)(m) (a)m _ - (p) ap a
3 _ — _ am (a)m — _ (p) ap a
4-7 — - _ am am — = (p) ap ap
8 - - mp amp am — _ (p) ap ap
9 -_ _ mp amp am — - (m)p ap ap
10 _ - mp amp am - — mp (a)p ap
ll — -- mp amp am —- — mp p ap
12 = ~ mp amp am — = mp p (a)p
13 = (m) mp amp am = — mp p p
14 = m mp amp am - - mp p p
15 - m m(p) (a)m ~ - — mp p (p)
392
setae, apical claw with two dorsolateral se-
tae and one dorsomedial seta.
Remarks.—The only known record for
Turkey for this species is from Uskudar (vil.
Istanbul) (Matic 1980). However, the au-
thor doesn’t mention the sex or the number
of specimens recorded; this record is du-
bious since it is possible that it might be
referred to the next species, P. orientis, with
females apparently indistinguishable from
those of P. jonicus and type-locality (Polo-
nezkGy) very close to the locality of the Mat-
ic (1980) record.
The presence of P. jonicus in Turkey is
however confirmed by the material record-
ed here:
The ecology of P. jonicus is little known.
Minelli and Iovane (1987) stated for the
Italian populations the general preference of
this species for open habitats, from sea level
up to 250 m; Matic and Golemansky (1965)
define this species as a “eurybionte’’; more-
over Matic and Golemansky’s (1964, 1965)
Bulgarian records have been collected be-
tween 340-500 m. The Turkish specimens
here recorded were collected in calcareous
soils, between 50 and 450 m, in anthropized
habitat (Truva), arid open land with Quer-
cus gr. coccifera Linné and Juniperus sp.
(Ayvacik) and in pine-wood (Havran).
Pleurolithobius orientis (Chamberlin, 1952)
New ComBINATION
Turkobius orientis Chamberlin, 1952: 225.
Archilithobius integrior caducus Chamber-
lin, 1952: 236 syn. nov.
Pleurolithobius atopior Chamberlin, 1952:
254 syn. nov.
Material examined.—2 4¢, | 9, vil. Istan-
bul, Belgrat Ormani, m 100, Biiyiikdere,
17.V.1987, A. Vigna leg.; 2 46, 9 99, vil.
Istanbul, ibidem, M. Zapparoli leg. (MZ); 1
2 (Holotypus), “L. orientis, 2, type, Polo-
nezk6, 15.V.48,” “43-642” (W); 1 4 (here
formally designated as Lectotypus of Pleu-
rolithobius atopior Chamberlin, 1952), 1 4
(here formally designated Paralectotypus of
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Pleurolithobius atopior Chamberlin, 1952),
“Pleurolithobius atopior Ch., Types, Polo-
nezkéy, 15.V.48,” “51-794” (W).
Distribution.—Species formerly known
only for its type locality, Polonezkéy, and
Yalova, both on the Asiatic side of the vil.
Istanbul. P. orientis is here recorded for the
first time on the European side of Turkey.
This species is also known for the islands
of Kos and Leros (southern Sporades Ar-
chipelago) (M. Zapparoli, unpublished data).
Description of the holotypus.— Female.
Size 19 mm long, 2.7 mm broad at T. 10;
color chestnut.
Head smooth, 3.3 mm broad, 3.0 mm
long, posterior border almost straight, pos-
terior marginal ridge with median thick-
ening; antennae 5.5 mm long, with 39 ar-
ticles (right), left antenna mutilated, the first
3-4 proximal articles large, the next ones
are longer than broader, last article about
three times as long as penultimate; ocelli
1+1,2 depigmented, principal ocellus not
contiguous with the secondary ocelli, pos-
tero-superior ocellus larger than principal
ocellus; organ of Témésvary of the same
size as the postero-superior ocellus; the for-
cipules are absent probably because dis-
sected by Chamberlin, since they are drawn
in the original work (Chamberlin 1952: fig.
31).
Tergites wrinkled; T. | subrectangular,
narrower than T. 3, posterior border slightly
sinuate; lateral borders parallel in TT. 3, 5,
7 and 8, slightly posteriorly convergent in
T. 10, posteriorly convergent in TT. 12 and
14; posterior border slightly sinuate in TT.
3, 5 and 8, straight in T. 7, slightly emar-
ginated in TT. 10 and 12, emarginated in
T. 14; posterior angles rounded in TT. 3, 5
and 8, squared in TT. 7, 10, 12 and 14; TT.
9 and 11 without triangular projection on
the posterior angles, T. 13 with slight pro-
jection; intermediate tergite with posterior
borders almost straight.
Anterior legs with tarsal articulations dis-
tinct, see Table 3 for spinulation; coxal pores
5,4,4,4, circular and separated one from
VOLUME 91, NUMBER 3
393
Table 3. Pleurolithobius orientis (Chamberlin, 1952). Holotypus: spinulation. See Table | for codes expla-
nation.
Ventral Dorsal
c tr P F T C tr P F T
1 — = _ = a — - p a a
2 - - - - am — ~ p ap a
3-5 - - — am am - _ p ap a
6-7 - _ =— am am - — p ap ap
8-9 _ - m am am ~ = p ap ap
10 - — mp am am — _ mp p ap
1] - = mp amp am - _ mp p ap
12-13 - m mp amp am — — mp p p
14 — m amp amp am — — mp p p
15 = m amp am — = — mp p -
another by a space little larger than their
own diameter, the proximal porus relatively
smaller than the other; XV legs 4 mm long,
apical claw with accessory claw about one-
half as long as the principal claw; glandular
pores on XII-XV legs, those of femur and
tibia XIII sparse and evident.
Gonopods with 3+ 3 long spurs, the inner
one relatively shorter than the others, apical
claw narrow and without lateral denticles.
A redescription of the male, based on the
Lectotypus of P. atopior, follows.
Size 12.50 mm long, 1.75 mm broad at
T. 10; color chestnut.
Head as broad as long, posterior border
almost straight, posterior marginal ridge
with median thickening; antennae 5.0 mm
long, with 39 articles (left), right antenna
mutilated at the level of article 22, the prox-
imal 3-4 articles larger, the following ones
are as broad as long, last article about three
times as long as the penultimate; ocelli
1 +1,2, depigmented, the two posterior ocelli
about twice larger than the two anterior ones;
organ of T6mésvary smaller than one an-
terior ocellus; the forcipules are absent,
probably because they were dissected by
Chamberlin, since they are figured in the
original work (Chamberlin 1952: fig. 44).
Tergites wrinkled; T. | subrectangular,
smaller than T. 3, posterior border slightly
sinuate; lateral borders parallel in TT. 3, 5,
7 and 8, slightly posteriorly convergent in
T. 10, posteriorly convergent in TT. 12 and
14; posterior border slightly sinuate in T. 3,
sinuate in T. 5, straight in T. 5, slightly
emarginate in TT. 8, 10 and 12, emarginate
in T. 14; posterior angles rounded in TT. 3,
5, 7 and 8, squared in TT. 10, 12 and 14;
TT. 9 and 11 without triangular projections
on the posterior angles, T. 13 with feebly
projection; intermediate tergites with pos-
terior border almost straight; last tergite with
lateral border posteriorly convergent, pos-
terior border deeply concave, posterior an-
gles rounded, posteriorly projected and ris-
ing slightly.
Anterior legs with tarsal articulation: cox-
al pores 4,4,3,2, circular and separated one
from another by a space little larger than
their own diameter, proximal pore of XII
and XIII coxae smaller than the other and
near the next pore; femur and tibia of XIII
legs thickened (cf. Fig. 9), femur little thick-
er than tibia and with a dorsal median sul-
cus not reaching the ends of the article; XIV
legs little thickened but without special
structures; XV legs 3.5 mm long, tibia with
a dorsolateral lobe bearing some setae at the
distal end of the article, DpP spine inserted
on the internal lateral side of the article,
apical claw with accessory apical claw about
one-half as long as the principal claw; glan-
dular pores XII-XV legs, those of femur and
tibia XIII very evident and rather sparse.
First genital sternite with 17-18 setae rel-
394 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 5-12. Pleurolithobius orientis (Chamberlin, 1952) (Belgrat Ormani). 5, Dental margin of prosternum,
ventral. 6, Right ocelli and organ of Témésvary (Té). 7, Last male tergite, dorsal. 8, Last male tergite, lateral.
9, Male XIII femur and tibia, dorsal. 10, Male XV tibia, dorsal. 11, Left female gonopod, ventral. 12, Right
female gonopod, dorsal. Scales: 5—0.1 mm: 6—0.2 mm; 7-10—0.5 mm; 1 1-12—0.1 mm.
VOLUME 91, NUMBER 3
395
Table 4. Pleurolithobius orientis (Chamberlin, 1952). Spinulation, 64 (Belgrat Ormani); letters in parentheses
indicate variable spines. See Table | for codes explanation.
Ventral
Dorsal
1 _ = = =
2 = _ = =
3 — - — a
4-7 — = = am
8-9 _ — mp am
10 _ — mp am
11 = — mp amp
12 — m mp am(p)
13 = m mp am(p)
14 = m mp am(p)
15 — m m(p) (a)m
atively long; second genital sternite without
setae; gonopods short and with one apical
seta.
The specimens from Belgrat Ormani (Bii-
yukdere, vil. Istanbul) differ from the sam-
ples described above in the following char-
acters.
Size 14.0-18.5 mm long, color dark chest-
nut. Antennae with 36-42 articles; proster-
num (Fig. 5) with evident shoulders, rela-
tively smaller in males than in females; ocelli
as in Fig. 6.
Tergites, especially the posterior ones,
with numerous sparse, relatively long setae;
T. 9 without triangular projection to the
T C tr P F Th
m - ~ p a a
m - _ p a a
m _ = p ap a
am _ - p ap ap
am = = (m)p ap ap
am - _ mp ap ap
am = = mp ap ap
am - — mp ap ap
am _ _ mp p p
a - - mp p p
posterior angles, T. 11 generally without tri-
angular projections, very little projection
when present, T. 13 with very slightly tri-
angular projections. Male last tergite as in
Figs. 7 and 8.
See Tables 4 (43) and 5 (29) for spinula-
tion. Female with DpP normally posi-
tioned; legs with numerous dorsal sparse se-
tae; coxal pores 5,4,4,4 (22), 4,3,3,2 or 3,3,3,2
(64). XIV and XV legs of the only male with-
out multilation both 4.6 mm long, female
with XIV legs 3.9-4.5 mm long and XV legs
4.9-5.2 mm long; femur and tibia XIII and
tibia XV figured in Figs. 9 and 10.
Female gonopods (Figs. 11 and 12) with
Table 5. Pleurolithobius orientis (Chamberlin, 1952). Spinulation, 2° (Belgrat Ormani); letters in parentheses
indicate variable spines. See Table | for codes explanation.
Ventral Dorsal
( ir P F T C tr P F T
l — _ _ (a)(m) (a)m = = (p) a a(p)
2 = = — a(m) am — - (p) (a)p a(p)
3 -_ ~ — am am - _ (p) ap a(p)
4-5 = _ = am am — — (p) ap ap
6-7 _ = m(p) am(p) am _- — (p) ap ap
8 —_ mp am(p) am = — (m)p ap ap
9 _ — mp amp am — _ (m)p ap ap
10 _ = mp amp am _ - mp ap ap
11 _ (m) mp amp am = = mp ap ap
12 — m (a)mp amp am _ — mp p ap
13 _ m (a)mp amp am ~ _ mp p (a)p
14 - m (a)mp amp am =_ _ mp p p
15 — m amp am — _ — mp p -
396
3+3 long spur, exceptionally 4+ 3, progres-
sively longer from the internal one to the
external one, internal spur generally lying
behind the other two; apical claw narrow
and without lateral denticles; basal article
with a row of 6-7 dorsolateral setae and a
group of 10-11 setae positioned near the
insertion of the spurs, second article with
9-11 dorsolateral setae arranged in two rows,
and three dorsomedial setae, apical claw
with 2-3 dorsolateral setae and one dorso-
medial seta.
Remarks. — Chamberlin (1952) based his
description of P. orientis on two females,
one from Polonezkéy (Holotypus) and one
from Yalova; he designated this species as
the type species of the genus Turkobius
Chamberlin, 1952, which he described as
new and assigned to the family Gosibiidae.
In the genus 7urkobius, Chamberlin (1952)
includes eight species divided into two sub-
genera, Turkobius s. str. and Alibius Cham-
berlin, 1952.
Based on examination of the Holotypus,
T. orientis must be referred to the genus
Pleurolithobius as defined by Verhoeff
(1899), and the following new synonymy is
proposed: Turkobius s. str. Chamberlin,
1952 = Pleurolithobius Verhoeff, 1899 syn.
nov.
It is not the aim of this work to discuss
the identity of the other species referred by
Chamberlin (1952) to Turkobius; however
it is suitable to point out that these taxa
belong to Lithobiidae and are certainly un-
related to Pleurolithobius. In fact some taxa
belong to Lithobius s. str. (those described
under 7urkobius s. str., with the exception
of 7. orientis), and others are referable to a
taxon of subgeneric rank of the genus Lith-
obius Leach, 1814 (those described under
Alibius). The identity of Turkobius and the
species which Chamberlin (1952) included
in this taxon will be discussed in a paper
now in preparation.
Chamberlin (1952) has described, on the
basis of a female from Polonezkéy, Lithobi-
us integrior caducus. This taxon was origi-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
nally referred to the genus 4rchilithobius
Stuxberg, 1875 (now considered identical
with Lithobius) and was considered at spe-
cific rank in the key that the author (Cham-
berlin 1952) gave for the Turkish species.
The identity of L. integrior caducus can-
not be discussed on the basis of the type
specimens since the material has been lost
(J. Coddington in litt. 1987). The following
considerations are therefore based only on
the very incomplete original description.
The few characters given by Chamberlin
(1952) for L. integrior caducus (especially
the shape of the prosternum and the shape
of the apical claw of female gonopods) fall
well within the variability of P. orientis de-
scribed above, and the two forms are dif-
ficult to distinguish one from another.
Moreover, noting not only the precise co-
incidence of the type-localities of the two
taxa, but also the identity of the collecting
date of the samples, it is quite justifiable to
suppose that the female described as L. in-
tegrior caducus and the Holotypus of P. or-
ientis might both likely refer to the same
population.
Therefore, the following new synonymy
is proposed: Archilithobius integrior cadu-
cus Chamberlin, 1952 = Pleurolithobius or-
ientis (Chamberlin, 1952) syn. nov.
Another species described by Chamberlin
(1952) on material (two males) collected in
the same type-locality and on the same day
as P. orientis and L. integrior caducus is P.
atopior. Holotypus and Paratypus of this
species are not indicated in the original work;
however two specimens, here examined, la-
belled ‘Pleurolithobius atopior Ch., Types,
Polonezkéy, 15.V.48,” preserved in Cham-
berlin’s collection in the National Museum
of Natural History of Washington and
agreeing with Chamberlin’s description of
P. atopior, represent undoubtedly the syn-
typical series of this taxon. Therefore these
specimens are here formally designated re-
spectively as Lectotypus and Paralectotypus
of P. atopior.
Based on the study of the type speci-
VOLUME 91, NUMBER 3
mens of P. atopior it was possible to verify
the identity of the characters of this taxon
with those here described for the male of P.
orientis. Consequently, the following new
synonymy is proposed: Pleurolithobius ato-
pior Chamberlin, 1952 = Pleurolithobius or-
lentis (Chamberlin, 1952) syn. nov.
As Chamberlin (1952) already stated in
his brief discussion on the morphological
affinities of P. atopior, P. orientis is close to
P. jonicus and 1s distinguishable from this
species by the shape of the last tergite of the
male, without lobed projection on the pos-
terior angles. However, this character can-
not be used for identification of the females
of the two species that, as already stated,
are apparently indistinguishable one from
another.
On the basis of the material examined it
is, however, possible to show some char-
acters useful to distinguish easily, at least
for the Anatolian populations, the females
of P. jonicus from those of P. orientis. These
characters are: body generally smaller (length
9.5-12.0 mm), prosternal shoulder feeble or
absent (Fig. 2), T. 1 subrectangular and ter-
minal tergites without sparse setae in P. jon-
icus; body generally larger (length 14.0-18.5
mm), prosternal shoulder present (Fig. 5),
T. | trapezoidal and terminal tergites with
sparse setae in P. orientis.
The ecology of P. orientis is practically
unknown. This species has been collected
at very low altitudes (100-200 m); the sam-
ples from Belgrat Ormani were collected in
litter of mixed woodland composed essen-
tually of Acer spp., Quercus spp., Fagus ori-
entalis Lipsky and Castanea sativa Miller,
with an undergrowth characterized by Smi-
lax sp., Crataegus sp. and Erica sp.
Another species recorded for the Turkish
fauna and originally described under Pleu-
rolithobius 1s Lithobius argaeensis Attems,
1905, known for Ercjias Dagi (vil. Kayseri)
(type-locality) and in some localities of Iran
(Attems 1905, 1951, Brélemann 1921).
Therefore, the original generic classification
of this species 1s wrong and has not been
397
followed by the following authors. In fact,
Bro6lemann (1921) and Attems (1951) cor-
rectly considered this species under Mon-
otarsobius Verhoeff, 1905.
ACKNOWLEDGMENTS
I wish to thank Dr. J. A. Coddington, of
the National Museum of Natural History,
Smithsonian Institution, Washington, and
Prof. A. Vigna Taglianti, Director of the
Zoological Museum of the Dipartimento di
Biologia Animale e dell’Uomo of Rome
University “La Sapienza,” for enabling me
to examine specimens in their charge.
This article is no. 124 for research of the
Zoological Institute of Rome University in
the Near East. Research was carried out with
a grant from the Italian C.N.R. (Gruppo
Biologia Naturalistica).
LITERATURE CITED
Attems, C. G. 1905. Myriapoda. /n Penther, A. and
E. Zederbauer, eds. Ergebnisse einer naturwissen-
schaftlichen Reise zum Erdschias-Dagh (Kleina-
sien). Ann. K. K. Naturhist. Hofmus. 20: 163(1)-
167(5).
1951. Ergebnisse der Osterreichischen Iran-
Expedition 1949-50. Myriopoden von Iran. Sitz-
ungsb. Osterr. Akad. Wiss. Mathem.-Naturw. KI].
160: 387-426.
Brélemann, H. W. 1921. Myriapods from Mesopo-
tamia and Persia, collected by Mr. P. A. Buxton.
J. Bombay nat. Hist. 28: 157-161.
Chamberlin, R. V. 1952. On the Chilopoda of Tur-
key. Rev. Fac. Sci. Univ. Istambul ser. B 17: 183-
258.
Matic, Z. 1980. Chilopodes recueillis en Turquie, Li-
ban et Iran. Acta Zool. Bulgarica 15: 93-98.
Matic, Z. and V. Golemansky. 1964. Contribution a
la connaissance des Lithobiides (Chilopoda, Lith-
obiomorpha) en Bulgarie. Ann. Univ. Sofia 57:
99-106.
1965. Nouvelle contribution a la connais-
sance des Lithobiides (Chilopoda, Lithobiomor-
pha) en Bulgarie. Ann. Univ. Sofia 58: 13-28.
Minelli, A. and E. lovane. 1987. Habitat preferences
and taxocenoses of Italian centipedes (Chilopoda).
Boll. Mus. civ. St. nat. Venezia 37 (1986): 7-34.
Verhoeff, K. W. 1899. Beitraége zur Kenntnis palaark-
tischer Myriopoden. XI Aufsatz: Neue und wenig
bekannte Lithobiiden. Verh. Zool. Bot. Gesell. 49:
451-459.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 398-405
REVIEW OF GOYA RAGONOT AND DESCRIPTION OF A
NEW SPECIES, G. SIMULATA FROM PARANA, BRAZIL
(LEPIDOPTERA: PYRALIDAE: PEORIINAE)
Jay C. SHAFFER
Department of Biology, George Mason University, Fairfax, Virginia 22030.
Abstract. —Goya simulata is described from Curitiba, Parana, Brazil. Included are pho-
tographs of adult moths, head profiles, genitalia of both sexes, scanning electron micro-
graphs of male antennae, and a discussion of the genus with a key to the four known
species. G. simulata is compared with the male holotype of the presumably sympatric
sibling species, G. albivenella and with the female lectotype of its junior synonym G.
cancelliella.
Key Words:
tropical lepidoptera
While recently examining a series of Bra-
zilian pyralid moths from the collection of
Dr. Vitor Becker, I sorted out six specimens
from Parana which appeared to belong to
Goya albivenella Ragonot, 1888. This
species is known only from the male type
and, if the association is correct, 2 nontopo-
typic females. On examination of the gen-
italia it was clear that the 2 males in the
series were specifically different from alhbi-
venella and represented an undescribed
species. The 4 females are likewise distinctly
different in genital characteristics from
specimens presumed to be albivenella. The
new species is described below.
Goya Ragonot, 1888 was erected for two
species, including G. a/bivenella, described
from Goya, Corrientes, Argentina, and sub-
sequently designated as the type of the genus
(Hampson in Ragonot, 1901: 349). The oth-
er originally included species is African,
clearly peoriine, but of uncertain generic af-
finity as it is known only from the female
type, not yet associated with male speci-
mens. Shaffer (1968: 56) transferred Saluria
stictella Hampson, 1918 to Goya, and Ato-
Goya albivenella, G. cancelliella, G. stictella, G. ovaliger, taxonomy, neo-
pothoures ovaliger Blanchard, 1975 was
shifted to Goya (Blanchard, 1978: 55), with
Atopothoures Blanchard, 1975 becoming a
junior synonym of Goya.
The last redescription of the genus (Shaf-
fer 1968: 56) was based on the 2 species
known at the time. With the discovery of 2
additional species, the distinction between
specific and generic characters becomes
more apparent, and it seems worthwhile to
comment on these. In all 4 species the uncus
possesses a medial process of variable shape
and armament, but always with a pair of
posteriorly directed spines along the caudal
margin. These spines are large in the 2 North
American species, moderate in a/bivenella
(Fig. 18), and small in simud/ata (Fig. 13 s).
The paired lateral processes of the uncus are
rhomboidal, pad-like, sparsely setose, and
vary little from species to species. They are
unusual apomorphies unlike those of any
other known peoriine. The medial process
of the gnathos is unique in detail for each
species, but very close in the two North
American ones and subject to individual
variation (see below). Also, its appearance
VOLUME 91, NUMBER 3
399
Figs. 1-4. Goya simulata. 1, é holotype; 2, 6 holotype, head, lateral view; 3, 2 paratype coll. 25 XII 1974;
4, 2 paratype coll. 4 II 1975, head, lateral view. Scale bar = 2.0 mm (1, 3), 1.0 mm (2, 4).
on prepared slides varies much depending
on the extent to which it is compressed by
the coverglass. Compare, for example, Figs.
16 and 17. These appear quite different be-
cause the flaps seen on edge in Fig. 16 were
flattened in the other preparation and so
appear broadside. A stereoscopic view of
the actual preparations reveals them to be
essentially identical. Few peoriines have the
transtilla developed, but in Goya it is pres-
ent in all four species, incomplete and 1n the
form of simple narrow bars (Figs. 13, 18 tr).
The valve shows relatively minor interspe-
cific variation, always more or less parallel
sided, apically diagonally truncate, and un-
armed. The aedeagus is simple, variable in
shape, always short; it and the vesica are
always unarmed but for minute denticles.
The female genitalia of peoriines tend to
be simple and rather uniform with the ovi-
positor compressed, the ostial chamber
membranous and unmodified, and the sig-
num only rarely present. Most often it is not
possible to identify peoriine species on the
basis of the female genitalia. Those of Goya
generally fit the above pattern, but are dis-
tinct for each of the 4 species. Also, they
exhibit two apomorphies each diagnostic of
the genus. Most evident of the two 1s a large
dorsocaudal sac on the 8th segment. It ap-
pears somewhat different in each species,
but is in every case well developed. Similar
400
Figs. 5-12.
features appear in two other New World
peoriine genera. In Atascosa all 4 known
species have a well developed narrow dig-
itate invagination in this position (see Shaf-
fer 1976, Plate 14e, g, i), and females of
Coenotropa have paired short pouches (ibid.,
Plate 14b, c) in the same place. The mark-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Goya simulata, basal modification of male antennal shaft. 5-8, holotype, left antenna, showing
5, lateral; 6, anterior; 7, medial; and 8, posterior views. 9-12, paratype, left antenna partly denuded to show
surface enclosed by scale tufts, same perspective as in 8. Scale bar = 500 um (5-9), 50 um (10), 25 um (11), 5
um (12).
edly different development of these sacs
suggests an independent development for
each genus.
The other apomorphy is more easily
overlooked. The 7-8 intersegmental mem-
brane is dorsally thickened and folded so as
to form an anterodorsal pocket at the an-
VOLUME 91, NUMBER 3
Figs. 13-18.
terior of the 8th segment collar. This feature
is present in all four species and is illustrated
for simulata in Fig. 19 (ap).
Distributions of Goya spp.— Two species
are North American. G. stictella was de-
scribed from Andros Island, Bahamas and
is represented from Florida, Mississippi,
Arkansas, and Texas. Blanchard’s ovaliger
is so far known only from Texas. He and I
have collected both species by blacklight at
the Welder Wildlife Refuge near Sinton,
Texas. Either or both may range into Mex-
ico, but are yet unknown there. G. albivenel-
401
Male genitalia. 13-16, Goya simulata, holotype; 13, male genitalia (aedeagus omitted); 14,
aedeagus; 15, detail of vesica enlarged to show denticles; 16, medial process of gnathos; 17, paratype, medial
process of gnathos, compressed by coverglass; 18, G. albivenella, lectotype, s = uncus spine, sc = subscaphium,
tr = transtilla. Scale bar = 1 mm (13, 14), 0.05 mm (15), 0.2 mm (16, 17), 0.5 mm (18).
/a is known with certainty only from the
type locality, Goya, Corrientes, Argentina,
but the two female specimens from Espirito
Santo and Pernambuco, Brazil are likely
conspecific. The latter (Fig. 21) is the lec-
totype of cancelliella (Ragonot), a junior
synonym of albivenella, and is clearly dis-
tinct from females of simulata. G. simulata
is described from the state of Parana, Brazil.
The holotype and female paratypes are from
Curitiba and the male paratype is from Qua-
tro Barras, about 100 km to the east.
The 2 North American species, if in rea-
402 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 19-21.
dorsal view of eighth segment modification; 21, G. albivenella, lectotype of its junior synonym G. cancelliella
(Ragonot), J. Shaffer slide 1181, lateral view. ap = anterodorsal pouch. Scale bar = 0.5 mm (19), 0.2 mm (20,
21).
sonably good condition, are readily distin-
guished by wing pattern, whereas the genital
differences are minor and show overlapping
individual variation. The 2 South American
species are very close in wing pattern, and
larger series of specimens will have to be
examined to properly assess individual
variation. Fortunately, the two species have
distinctly different genitalia in both sexes.
Key TO SPECIES OF GOYA
1. Forewing with well developed spots or trans-
verse bands; ¢ genitalia with subscaphium
weakly sclerotized; aedeagus strongly flared ba-
sally in vertical plane; North American
— Forewing lacking well developed spots or
transverse bands, veins prominently traced with
white; 4 genitalia with subscaphium well scle-
rouized, prominent, shield shaped (Fig. 18sc);
aedeagus subcylindrical, not flared; South
American
Female genitalia. 19, Goya simulata, paratype, USNM slide 57903, lateral view; 20, same,
. Forewing with well developed continuous an-
temedial and postmedial lines, discocellular
spot a dark circle with pale center; medial pro-
cess of gnathos usually with >2 denticles per
SidG: axe cspiviethe aiuto ache a eee ovaliger
Forewing lacking developed antemedial line;
postmedial line discontinuous, formed of spots;
discal spot small, solid: large dark spot on 2nd
A posterior to cell; medial process of gnathos
usually with 0-2 denticles per side ...... stictella
3. dgenitalia with medial process of uncus bearing
numerous minute spines on ventral surface (Fig.
18); apical process of gnathos subquadrate;
valve lacking patch of setae near base of costa
(Fig. 18); 2 genitalia with apophyses of normal
length; 8th segment collar about as long as high:
forewing 3rd A white-scaled . albivenella
¥ genitalia with medial process of uncus smooth
but for pair of small spines on caudal margin
(Fig. 13); apical process of gnathos cordate and
strongly bilobed; valve with conspicuous patch
of setae near base of costa (Fig. 13); 2 genitalia
with apophyses long; 8th segment collar about
VOLUME 91, NUMBER 3
1.7 times as long as high; forewing 3rd A with
little or no white scaling ....... simulata
Goya simulata Shaffer, NEw SPECIES
Figs. 1-17, 19, 20
Diagnosis.—The single pair of very short
uncus spines (Fig. 13 s) and the well devel-
oped hair tuft at the base of the valve costa
(Fig. 13 diagonal arrow) are each unique to
simulata, as are the unusually elongate eighth
segment and apophyses of the female (Fig.
19).
Description.—Frons brown. Labial pal-
pus obliquely ascending in both sexes; in
male (Fig. 2) 2.6 times as long as eye di-
ameter, 2nd segment narrow, gradually ex-
panded distally (unrubbed condition), api-
cally truncate; 3rd segment short, narrow,
fusiform, set off from second; basal segment
white, 2nd and 3rd segments brown dor-
sally, white ventrally, demarcation irregu-
lar; female (Fig. 4) similar to male, but un-
usually long and slender, 3.3-3.7 times as
long as eye diameter. Maxillary palpus cy-
lindrical, approaching end of frons, usually
hidden by labials. Proboscis rudimentary.
Male antenna shaft sublaminate, basal
modification (Figs. 5-12) with segments
partly fused, posterior surface with tuft of
opposing bands of scales, unequal (cf. Figs.
5, 7), astride of and enclosing a surface that
appears smooth at low magnification (Fig.
9), but at higher resolution irregularly car-
inate (Fig. 10), very finely rugose and mam-
millate (Figs. 11, 12). Female antenna fili-
form, unmodified. Eye diameter 0.85 mm
(range: 0.75-0.95 mm). Ocellus well devel-
oped, elliptical, set obliquely to eye margin
and separated from it by its own length.
Vertex brown, bearing pad-like tuft of scales
shielding antenna scapes anteromedially.
Antenna sockets narrowly fringed with yel-
low scales laterally and posterolaterally. Oc-
ciput dark brown laterally; dorsally with
erect wall of moderately dark brown scales
behind antennae. Patagium and tegula with
mixture of brown, dark brown, and yellow-
ish brown scales.
403
Forewing radius 12-15 mm. Costa nar-
rowly pale yellow, this color broadening near
apex. Sc and radius traced with white;
ground grayish red between costa and Sc
and between Sc and radius. Ground of cell
moderate yellowish brown with varying
number of scattered black scales; this ground
continuing between R; and M, to apex, but
beyond cell with predominance of black so
as to form dark dash between upper outer
margin of cell and wing apex. A similar but
much smaller dash extends from base of Cu,
distally to near M,—M, divergence. Cubitus
traced with white, broadest near lower outer
angle of cell, continuing distally on M,, M,,
Cu,, and Cu,, broadest on M,. Ground be-
tween cell (Cu, distal to cell) and 2nd A
moderate yellowish brown. Ist A narrowly
traced with white distal to cell. 2nd A broad-
ly bordered on both sides with scattered
blackish scales, weakly traced with white in
some specimens (this feature might be more
prominent in fresh specimens). Ground
posterior to 2nd A varying from moderate
yellowish brown to grayish red. Termen with
conspicuous black spots between veins.
Basal ' of fringe yellow, distal 7s brown. R,
basally approximate to R;,;, contiguous in
some specimens. R,,, fused, stalked with
R, about half length of latter. M,,,; from
lower outer angle of cell, stalked nearly half
its length.
Hindwing with M,,, stalked with Cu,
nearly half length of latter.
Male genitalia (Figs. 13—17)—Uncus with
lateral lobes about twice as long as wide;
medial process rounded, smooth margined,
armed only with a single pair of divergent
short pointed cusps. Gnathos with medial
process apically emarginate to about 4 its
length and thus divided into a pair of lateral
lobes, these lying in the plane of the lateral
gnathos arms, each bearing ventral blade-
like process set at about 90 degrees to plane
of lateral gnathos arms (compare Figs. 16
and 17, the latter with these blades pressed
flat in the preparation). Valve with a poorly
defined cusp in midregion of cucculus, and
404
strong patch of setae near base of costa (Fig.
13 arrow).
Female genitalia (Figs. 19, 20)—Apophy-
ses long, straight, nearly equal in length, 2.0
times as long as 8th segment collar height
and 1.3 times as long as its length. Dorso-
caudal sac of 8th segment large, half as long
as total 8th segment length. Ostial chamber,
ductus bursae, and corpus bursae membra-
nous, unmodified, unarmed.
Types.— Holotype: ¢ (Figs. 1, 2, 5-8, 13-
16), labelled: “Curitiba, Parana BRASIL—
920m 20. XII. 1974 V. O. Becker Col.”’; “6
genitalia on slide 2223 J. C. Shaffer”; ‘““Ho-
lotype Goya simulata Shaffer, 1989”
[NMRJ]. Paratypes: 4, 4 2; 6 (Figs. 9-12,
17), labelled: ““Banhado, Quatro Barras, PR.
800 m. 7-2-70 V. O. Becker leg.”; ““Geni-
talia Slide By Jay Shaffer USNM 57902”
[USNM]. 4 2, same locality data as holo-
type; “16. XII. 1974” J. Shaffer genitalia
slide 2246 [NMRJ]; “14. I. 1975” undis-
sected [NMRJ]; “4. II. 1975” (Figs. 4, 19,
20) USNM slide 57903 [USNM}]; “25. XII.
1974” (Fig. 3) abdomen lost [USNM]. All
paratypes bear the label: “Paratype Goya
simulata Shaffer, 1989.”
Discussion.— This species 1s very similar
to albivenella in external characters. The
most distinct difference 1s the almost com-
plete lack of white marking of 3rd A of the
forewing in simu/ata contrasted with a con-
spicuous white tracing over this vein in al-
bivenella. Also, there is a conspicuous white
tracing over the forewing 2nd A in albive-
nella, a marking much less prominent in 2
of the 6 simulata types and absent in the
other 4. Lastly, the forewing ground color
posterior to the cell 1s yellowish brown in
albivenella, brown in simulata and in some
specimens of the latter grayish red posterior
to 2nd A.
The male genitalia of simulata and albi-
venella can be compared in Figs. 13 and 18
respectively. Three differences are note-
worthy. In al/bivenella the medial process of
the uncus is laterally and ventrally armed
with a multitude of minute stout spines, and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
bears posterolateral expansions each ter-
minating in a large posteriorly directed spine,
the two spines essentially parallel. By con-
trast, this process in simulata is lacking in
minute spines, curved along its posterior
margin, and armed only with a pair of short
divergent conical cusps. Secondly, the me-
dial process of the gnathos is markedly dif-
ferent in the two species, cordate and keeled
in simulata, quadrate in albivenella. Finally,
the well developed patch of setae at the base
of the valve costa in simulata (Fig. 13 di-
agonal arrow) is completely absent in a/bi-
venella.
The female genitalia of simulata differ
from those presumed to be albivenella in
the more elongate shape of the 8th segment
collar, 1.7 times as long as wide in simulata,
1.0 times in a/bivenella; in the much longer
anterior and posterior apophyses, and in the
longer posterodorsal sac of the 8th segment.
This comparison is based on an examina-
tion of the lectotype of cancelliella, a junior
synonym of albivenella. These features dis-
tinguish simu/ata from all other known
species of Goya.
In summary, si?mulata and albivenella can
readily be distinguished by clear differences
in the male genitalia of their holotypes and
it is on these differences that the distinction
between the two species presently rests. The
constancy of the differences in wing pattern
described above can only be evaluated when
a larger series of specimens of both species
becomes available for study. Likewise, a
good series of specimens of albivenella in-
cluding both sexes from one population
should resolve any questions as to the iden-
tity and characteristics of females of that
species.
Etymology.—The specific epithet is an
adjective derived from the Latin similis (like)
in reference to the external resemblance to
albivenella.
ACKNOWLEDGMENTS
I thank Dr. Vitor Becker for the privilege
of examining a portion of his collections and
VOLUME 91, NUMBER 3
Ms. Linda Trimmer for assistance with SEM
work. The Hitachi S-530 scanning electron
microscope used in this study was support-
ed in part by NSF Grant No. BSR-8511148.
The holotype and 2 female paratypes are
deposited in the National Museum, Rio de
Janeiro, Brazil [NMRJ]. The male paratype
and two female paratypes are in the Na-
tional Museum of Natural History, Smith-
sonian Institution, Washington, D.C. USA
[USNM].
LITERATURE CITED
Blanchard, A. 1975. Anew phycitine genus and species
(Pyraloidea). J. Lepid. Soc. 29(2): 95-97.
405
1978. Atopothoures A. Blanchard: A syn-
onym of Goya Ragonot (Pyralidae). J. Lepid. Soc.
32(1): 55-56.
Ragonot, E. L. 1888. Nouveaux genres et espéces de
Phycitidae & Galleriidae. Paris, 52 pp.
. 1901. InN. M. Romanoff, Mémoires sur les
Lépidopteéres, vol. 8, xli + 602 pp., color pls. 24—
57. [MS. completed by Sir G. F. Hampson.]
Shaffer, J.C. 1968. A revision of the Peoriinae and
Anerastiinae (Auctorum) of America north of
Mexico (Lepidoptera: Pyralidae). Bull. U.S. Nat.
Mus. 280, 124 pp.
1976. A revision of the neotropical Peoriinae
(Lepidoptera: Pyralidae). Syst. Ent. 1: 281-331.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 406-413
JAPANESE SAWEFLIES OF THE FAMILY BLASTICOTOMIDAE
(HYMENOPTERA: SYMPHYTA)
IcHiyt TOGASHI
Ishikawa Agricultural College, 1-308, Suematsu, Nonoichi-machi, Ishikawa Prefecture
921, Japan.
Abstract.
—The family Blasticotomidae of Japan is revised, and one new species, Blas-
ticotoma warabii sp. nov., is described and illustrated. A key to the Japanese genera and
species and illustrations of their taxonomically important characters are given. Food plants
of Runaria flavipes are newly described.
Key Words:
Blasticotomidae is a small family of Ten-
thredinoidea associated with ferns and oc-
curs only in the Palearctic Region. Smith
(1978) listed 8 world species and subspecies
in three genera and Shinohara (1983) added
two species from Taiwan. In Japan, the fam-
ily contains four species and one subspecies
of two genera, namely, Blasticotoma atra
Zhelochovtsev, 1934, B. nipponica Takeu-
chi, 1939, B. filiceti pacifica Malaise, 1931,
Runaria reducta Malaise, 1931, and R. fla-
vipes Takeuchi, 1939. Recently, I had an
opportunity to examine three specimens
which were captured on stems of bracken
fern, Pteridium aquilium (L.), in Hokkaido,
Japan. These specimens resemble B. nip-
ponica and B. filiceti pacifica, but they differ
from B. nipponica by the small tooth of the
claw and by the slender sawsheath, and from
B. filiceti pacifica by the maculation of the
forewing. I, therefore, believe that these
specimens are new to science. In this paper,
I describe and illustrate this species.
DIAGNOSIS OF BLASTICOTOMIDAE
First discoidal cell of forewing petiolate,
stigma of forewing large and nearly semi-
circular in shape, anal cell of forewing with
Blasticotomidae, Blasticotoma, Runaria
an oblique crossvein (Fig. 8). Antenna with
four (Fig. 6) or three segments.
Key TO THE JAPANESE GENERA
AND SPECIES
1. Antenna with three segments, tarsal claw with-
out inner tooth (Runaria) . Singigal ndrauberen ere 2
— Antenna with four segments, tarsal claw with
inner tooth (Figs. 20-23) (Blasticotoma)
2. Legs black with anterior four knees, all tibiae
and tarsi yellow .............R. reducta Malaise
— Legs yellow to reddish yellow, only coxae black
SAT er eae oer ce R. flavipes Takeuchi
3. Female
Riise Races arstss wee te 4
= Malet sa.dnrwes oat at ei tees ee 7
4. Forewing entirely infuscate (Fig. 2) .......... 5
— Forewing partially infuscate (Fig. 1) ......... 6
5. Legs black with all knees, tibiae, and tarsi red-
dish yellow (Figs. 13 and 14); inner tibial spur
of fore leg as in Figs. 18 and 19; sawsheath as
in Fig. 27 Atco end ee B. atra Zhelochovtsev
— Legs reddish yellow with all coxae and tro-
chanters black; inner tibial spur of fore leg as
in Fig. 17; sawsheath as in Fig. 26.
GE See cuatro B. filiceti pacifica Malaise
6. Claws with a large inner tooth (Fig. 21); hind
leg black with hind tibia yellow to reddish yel-
Yow except for apical portion; inner tibial spur
of fore leg as in Fig. 16; sawsheath as in Fig.
25; apical portion of forewing clear; lance as
in Fig. 32 . B. nipponica Takeuchi
— Claws witha small? inner tooth (Fig. 20); hind
leg yellow to reddish yellow; inner tibial spur
VOLUME 91,
NUMBER 3
I
Figs. 1-2.
of fore leg as in Fig. 15; sawsheath rather slen-
der (Fig. 24); apical portion of forewing infus-
cate (Fig. 8); lance as in Fig. 31
Sn Bane ere B. warabii sp. nov.
T Forewing uniformly infuscate; genitalia as in
Figs. 36 and 40; penis valve as in Fig. 44 ..
: . B. filiceti pacifica Malaise
- Forewitle rather dean Bo mieciities partially in-
US CALC BPN TS ene eto ny ce ceva ha hoy 8
8. Legs black with anterior four knees, all fhe:
and tarsi reddish yellow (Figs. 11 and 12); gen-
italia as in Figs. 37 and 41; penis valve as in
Fig. 45 : .. B. atra Zhelochovtsev
— Anterior four legs reddish yellow with all coxae
and trochanters black . 9
9. Hind femur and apical portion of hind tibia
black; genitalia as in Figs. 35 and 39; penis
valve as in Fig. 43 _ B. nipponica Takeuchi
— Hund femur and apical portion of hind tibia
reddish yellow; genitalia as in Figs. 34 and 38;
penis valve as in Fig. 42 B. warabii sp. nov.
Blasticotoma warabii Togashi,
NEw SPECIES
Figs. 1, 3-8, 15, 20, 24, 28, 31,
34, 38, 42
Female.—Length 9 mm. Body including
antenna black, but cenchri pale white, apical
half of mandible and apical portion of saw-
sheath reddish brown. Wings as in Fig. 8.
Legs reddish yellow, with following parts
black: all coxae and trochanters, and basal
portion of all femora.
Head transverse, postocellar area dis-
tinctly raised; lateral and interocellar fur-
rows nearly absent; postocellar furrow rath-
Dorsal view—1, Blasticotoma warabii sp.
407
nov. 2, Blasticotoma filiceti pacifica Malaise. (3 x )
er distinct (Fig. 3); OOL:POL:OCL =
1.0:0.8; frontal area nearly flattened; post-
orbital groove distinct and broad (Fig. 5);
postgenal carina distinct; median fovea as
in Fig. 5; lateral foveae comma-like; supra-
clypeal area slightly convex; clypeus nearly
flattened, frontal margin nearly truncate (Fig.
4); labrum small; malar space broad; man-
dibles tridentate (Fig. 4); antenna as in Fig. 6.
Thorax normal; mesoscutellum as in Fig.
7. Inner tibial spur of fore leg as in Fig. 15;
claw with a small inner tooth (Fig. 20). Ab-
domen normal; sawsheath as in Fig. 24; ra-
tio between length of sawsheath and basal
plate at about 1.0:2.1; lancet with 13 ser-
rulae (Fig. 28); lance with 8-9 serrulae (Fig.
31):
Sculpture.— Face rather shallowly, irreg-
ularly, and reticulately sculptured; occipital
margin coarsely and distinctly punctured;
clypeus rather evenly, shallowly, and dis-
tinctly punctured; labrum densely punc-
tured: pronotum shagreened, with distinct
and large punctures; posterior half of meso-
notal lateral lobes coarsely and distinctly
punctured (Fig. 7); mesoscutellum nearly
impunctate, but posterior margin with dis-
tinct punctures (Fig. 7); mesopleuron dis-
tinctly and evenly punctured; mesosternum
nearly impunctate, but with some distinct
punctures. Abdominal tergites shagreened,
but last tergite nearly impunctate, shining.
Male.—Length 7 mm. Coloration and
structures except for male genitalia as in
408 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Be
VOLUME 91, NUMBER 3
female. Male genitalia as in Figs. 34 and 38;
penis valve as in Fig. 42.
Habitat.—Japan (Hokkaido and Hon-
shu).
Holotype: Female, 13. VI. 1986, Hitsu-
jigaoka, Sapporo, Hokkaido (Type No.
2661, deposited in the Entomological Lab-
oratory of Kyushu University).
Paratypes: Two females, same date for
holotype; three males, 17. V. 1968, Sasari,
Kyoto Pref., T. Naito leg.
Remarks.—This new species closely re-
sembles B. nipponica but it differs from the
latter species by the small inner tooth of the
claw (in nipponica, the claw has a large inner
tooth, as in Fig. 32); by the shape of the
apex of the inner tibial spur of the fore leg
(in nipponica, the apex of the inner tibial
spur has both furcations subequal in size as
in Fig. 16); and by the characters of the
lancet, lance and penis valve.
Blasticotoma nipponica Takeuchi, 1939
Figs. 16, 21, 25, 29, 32, 35, 39, 43
Blasticotoma nipponica Takeuchi, 1939, p.
394.
Specimens examined.—Two males, 23.
V. 1954, Mt. Tomuro, Kanazawa, Ishikawa
Pref.; one female and one male, 10. V. 1969,
Yoshioka, Kawachi-mura, foot of Mt. Ha-
kusan, Ishikawa Pref.; one female, 12. V.
1978, Mt. Shiritaka, near Tsurugi-machi,
Ishikawa Pref.; one male, 24. V. 1984, Tsu-
rugi-machi, Ishikawa Pref.; one female, 31.
V. 1987, Kinome Path, Imajo-cho, Fukui
Pref.; one female, 5. VI. 1988, Mt. Roku-
manbe, near Mt. Hakusan, Ishikawa Pref.
(i
Figs. 3-27.
409
Supplementary notes.—Apex of inner
tibial spur of fore leg bifurcate (Fig. 16);
sawsheath rather short (Fig. 25); ratio be-
tween length of sawsheath and basal plate
at about 1.0:2.1; lancet with 13 serrulae (Fig.
29); lance with 11 serrulae (Fig. 32); male
genitalia as in Figs. 35 and 39; penis valve
as in Fig. 43.
Blasticotoma filiceti pacifica
Malaise, 1931
Figs. 2, 9-10, 17, 22, 26, 30,
33, 36, 40, 44
Blasticotoma filiceti pacifica Malaise, 1931,
p22:
Specimens examined.—(Hokkaido) One
male, 10. VII. 1967, Aizankei, foot of Mt.
Taisetsu, T. Naito leg.; two females, 23, VII.
1969, Mt. Rausu, Shiretoko Peninsula, T.
Naito leg.; one female, 19. VII. 1987, Ai-
zankei, foot of Mt. Taisetsu, T. Naito leg.
(Honshu) One female, 17. VI. 1968, Sasari,
Kyoto Pref., T. Naito leg.; two females, 29.
VI. 1972, Mt. Shiritaka, near Tsurugi-ma-
chi, Ishikawa Pref.; one female, 7. V. 1972,
Fukase, Okuchi-mura, foot of Mt. Hakusan,
Ishikawa Pref.; one female, 9. V. 1984, Tsu-
rugi-machi, Ishikawa Pref.
Supplementary notes.—Posttergite red-
dish brown to dark brown; apex of inner
tibial spur of fore leg bifurcate (Fig. 17);
sawsheath as in Fig. 26; ratio between length
of sawsheath and basal plate at about 1.0:
2.3; lancet with 12 serrulae (Fig. 30); lance
with 8-9 serrulae (Fig. 33); male genitalia
as in Figs. 36 and 40; penis valve as in Fig.
44.
Figs. 3-8. Blasticotoma warabii sp. nov.—3, head, dorsal view; 4, do, frontal view; 5, do, profile;
6, antenna; 7, mesoscutellum and cenchri; 8, forewing. (Scale: 1 mm.) Figs. 9-10. B. filiceti pacifica Malaise—
9, head, dorsal view; 10, mesoscutellum and cenchri. (Scale: | mm.) Figs. 11-14. Fore and hind legs, except for
tibiae and tarsi, of B. atra Zhelochovtsev—11, fore leg of male; 12, hind leg of male; 13, fore leg of female; 14,
hind leg of female. (Scale: 1 mm.) Figs. 15-19. Inner tibial spur of fore leg—15, B. warabii; 16, B. nipponica;
17, B. filiceti pacifica, 18 and 19, B. atra. (Scale: 0.1 mm.) Figs. 20-23. Hind tarsal claw—20, B. warabii; 21,
B. nipponica; 22, B. filiceti pacifica, 23, B. atra. (Scale: 0.1 mm.) Figs. 24-27. Sawsheath, lateral view—24, B.
warabil; 25, B. nipponica; 26, B. filiceti pacifica, 27, B. atra. (Scale: 1 mm.)
VOLUME 91, NUMBER 3
411
Fig. 46. Oviposition of Runaria flavipes, food plant: Polystichum tripteron. (1 x)
Blasticotoma atra Zhelochovtsev, 1934
Figs. 11-14, 18-19, 23, 27, 37, 41, 45
Blasticotoma atra Zhelochovtsev, 1934, p.
154.
Specimens examined.—(Hokkaido) One
female, 23. VII. 1969, Mt. Rausu, Shiretoko
Peninsula, T. Naito leg.; one female, 19.
VII. 1987, Aizankei, foot of Mt. Taisetsu,
T. Naito leg. (Honshu) Two males, 4. VII.
1964, Karasawa, foot of Mt. Hodaka, Na-
gano Pref., T. Naito leg.; one male, 11. VI.
1968, Uchinikaya, Ina, Nagano Pref., T.
Naito leg.
Supplementary notes.— Legs dark brown
to black, but in female, all knees, tibiae and
tarsi reddish yellow, and in male, anterior
four knees, all tibiae and tarsi reddish yel-
low (Figs. 11-14); inner tibial spur of fore
leg as in Figs. 18 and 19; male genitalia as
in Figs. 35 and 39; penis valve as in Fig.
45.
Remarks.—The structure of the lancet and
lance of this species are similar to those of
B. filiceti pacifica, but the shape of the inner
tibial spur of the fore leg and male genitalia
and penis valve show differences between
these two species.
Runaria reducta Malaise, 1931
Runaria reducta Malaise, 1931, p. 213.
Specimens examined.—One female and
one male, 10. V. 1969, Yoshioka, Kawachi-
mura, foot of Mt. Hakusan, Ishikawa Pref.;
one male, 25. V. 1972, Mt. Shibatake, Nu-
gata Pref., K. Baba leg.; three females, Mt.
Shiritaka, near Tsurugi-machi, Ishikawa
Pref.; two males, 13. VI. 1986, Hitsujigao-
ka, Sapporo, Hokkaido.
—
Figs. 28-45.
Figs. 28-30. Lancet—28, B. warabii; 29, B. nipponica; 30, B. filiceti pacifica. Figs. 31-33. Lance —
31, B. warabii; 32, B. nipponica; 33, B. filiceti pacifica. Figs. 34-37. Male genitalia, lateral view— 34, B. warabii;
35, B. nipponica; 36, B. filiceti pacifica, 37, B. atra. (Scale: | mm.) Figs. 38-41. Male genitalia, dorsal view—
38, B. warabit, 39, B. nipponica; 40, B. filiceti pacifica, 41, B. atra. (Scale: | mm.) Figs. 42-45. Penis valve—
42, B. warabii; 43, B. nipponica; 44, B. filiceti pacifica; 45, B. atra. (Scale: 0.5 mm.)
NIPPONICA
/
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
WARABI I
ANCESTRAL TYPE
412
ae
Ap
Manes
Fig. 47.
of the adult.
Runaria flavipes Takeuchi, 1939
Fig. 46
Runaria flavipes Takeuchi, 1939, p. 396.
Specimens examined.—Three females
and six males, 20-24. V. 1984, Mt. Shiri-
taka, near Tsurugi-machi, Ishikawa Pref.;
one female, 5. V. 1987, Mt. Shiritaka, near
Tsurugi-machi, Ishikawa Pref.; one female,
5. V. 1988, Tsurugi-machi, Ishikawa Pref.
Supplementary note.—In 1987, I ob-
served a female of this species ovipositing
on the stem of Polystichum tripteron (Fig.
46); thus this must be the host plant.
Phylogenetic relationship of Blasticotoma
occurring in Japan
The comparative morphology of the male
genitalia indicates that Blasticotoma can be
clearly divided into two groups: nipponica-
group and filiceti-group. Each group has a
definite character of the penis valve as shown
in Figs. 42-45. The penis valve of filiceti-
group has a short projection at the end of
the valviceps (Figs. 42, 44, and 45), but that
of nipponica-group has an elongate projec-
tion (Fig. 43). The number of the serrula of
Pictorial phylogeny of the genus B/asticotoma on the basis of the shapes of the external characters
the lance in the female also shows the cor-
responding distinction. Also, the shape of
the inner tooth of the tarsal claw is different
from species to species (Figs 20-23). Ac-
cording to the shape of the inner tooth of
the tarsal claw, fi/iceti-group is divided into
two subgroups, fi/iceti-subgroup and wara-
bii-subgroup (Fig. 47). I also observed the
differentiational tendency of wing macula-
tion (Fig. 47). The phylogenetic relationship
of Blasticotoma is illustrated in Fig. 47,
which is based on the above-mentioned ex-
ternal characters.
ACKNOWLEDGMENTS
I express my thanks to Dr. David R.
Smith, USDA, Washington, D.C., for his
reading through this manuscript and for kind
advice. I am indebted to Drs. K. Baba, Nii-
gata Pref., and T. Naito, Kobe University,
Kobe, for the loan of the valuable speci-
mens.
LITERATURE CITED
Malaise, R. 1931. Neue japanische Blattwespen. Zool.
Anz. 94: 201-213.
Shinohara, A. 1983. Discovery of the families Xyel-
idae, Pamphiliidae, Blasticotomidae, and Orus-
VOLUME 91, NUMBER 3
sidae from Taiwan, with descriptions of four new
species (Hymenoptera: Symphyta). Proc. Ento-
mol. Soc. Wash. 85: 309-320.
Smith, D. R. 1978. Hymenopterorum Catalogus, Pars
14, Symphyta. 193 pp. The Hague-Holland.
Takeuchi, K. 1939. A systematic study on the sub-
413
order Symphyta (Hymenoptera) of the Japanese
Empire (II). Tenthredo 2: 393-439.
Zhelochovtsev, A. 1934. Neue und bekannte Ten-
thredinoidea aus Ost-Sibirien. Arch. Mus. Univ.
Moscou 1: 150-154.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 414-420
TYPE MATERIAL OF TWO AFRICAN SPECIES OF
HERPETOGRAMMA AND ONE OF PLEUROPTYA
(LEPIDOPTERA: CRAMBIDAE: PYRAUSTINAE)
JAY C. SHAFFER AND EUGENE MUNROE
(JCS) Department of Biology, George Mason University, Fairfax, Virginia 22030; (EM)
Granite Hill Farm, R.R.#2 Dunrobin, Ontario KOA 1T0O, Canada.
Abstract. —Lectotypes for the African pyraustine species Botys mutualis Zeller and Botys
aegrotalis Zeller (Lepidoptera: Crambidae: Pyraustinae) are designated, illustrated, and
characterized, and the species transferred to Herpetogramma and Pleuroptya, respectively
(new combinations). The holotype of the related Botys verminalis Guenée is also illustrated,
and characterized, and the species transferred to Herpetogramma (new combination). The
3 species are considered distinct from one another and from the American Herpetogramma
bipunctalis (Fabricius) with which they were synonymized by Hampson, 1899.
Key Words:
African Pyraustinae, Pyralidae, lectotypes, Zeller types, Herpetogramma
bipunctalis, Herpetogramma mutualis, Herpetogramma verminalis, Pleu-
roptya aegrotalis
This paper grew out ofan attempt to 1den-
tify two sibling species of the pyraustine ge-
nus Herpetogramma for a faunal study on
the Crambidae of Aldabra Atoll (in prep.).
We had narrowed our search to three ex-
ternally similar African species: Botys ae-
grotalis Zeller, B. mutualis Zeller, and B.
verminalis Guenée. All three appeared to be
very close to the two Aldabra species, and
in fact were among 18 names synonymized
by Hampson (1899: 204) under Pachyzan-
cla bipunctalis (Fabricius), a pest species
commonly known as the southern beet web-
worm. While it is beyond the scope of this
paper to review that synonymy, most of
those 18 are deserving of separate specific
status, with bipunctalis itself being restrict-
ed to the Western Hemisphere. Our pur-
poses here are to illustrate types in order to
facilitate identification, to provide proper
generic assignment, and to designate lec-
totypes so as to promote stability for the
names.
We follow Minet’s (1981) separation of
the traditional family Pyralidae into Cram-
bidae and Pyralidae, this essentially along
the same lines as the division of the Pyr-
alidae into the Series Crambiformes and
Pyraliformes in the Moths of North Amer-
ica (MONA) volumes (Munroe in Domin-
ick 1972).
Through the courtesy of Mr. Bert Gus-
tafsson of the Swedish Natural History Mu-
seum [NHRM] we were able to borrow for
dissection two specimens placed under ae-
grotalis, there being no material found there
under mutualis. Each specimen bore a “‘Caf-
fraria” label (Fig. 4) typical of the material
Zeller (1852) examined in his study, and so
could be a type. There is also Zeller material
labeled as aegrotalis and mutualis in the
collection of the British Museum (Natural
History) [BMNH].
After a more detailed examination of the
two Stockholm specimens it became clear
that they were not conspecific and that nei-
Figs. 1-3. Adult moths. 1, Herpetogramma mutualis, lectotype; 2, H. verminalis, holotype; 3, Pleuroptya
aegrotalis, lectotype. Scale bar = 2 mm.
415
416
_ Paravicini Coll.
B.M. 1937-333.
| Holotype
Figs. 4-6.
6, Pleuroptya aegrotalis, \ectotype (1.6 x).
ther matched Zeller’s description of aegro-
talis. However, one of these (Figs. 1, 4) was
a better match for mutualis than the spec-
imen under that name in the British Mu-
seum, and so we designate this Stockholm
specimen as the lectotype. The other Stock-
holm specimen (J. C. Shaffer 4 genitalia slide
no. 1944) is conspecific with an Aldabran
species which we intend to describe as new.
Herpetogramma mutualis (Zeller),
New ComBInaTION, Revised Status
Figs. 1, 4, 7-10
Botys mutualis Zeller, 1852: 40-41.
Pachyzancla mutualis (Zeller). Hampson,
1899: 204. As synonym of P. bipunctalis
(Fabricius).
Lectotype, hereby designated, labeled (Fig.
4): “Caffraria”’; ““Riksmuseum Stockholm”:
“$ genitalia on slide 1943 J. C. Shaffer’;
“Lectotype Botys mutualis by J. Shaffer &
E. Munroe, 1989.” [NHRM]. Zeller indi-
cated that he had both sexes represented,
but did not specify the number of speci-
mens. As we have not discovered any other
specimens conspecific with the lectotype in
material known to have been studied by
Zeller, we conclude that either his type se-
ries was of mixed composition or some of
it has been lost.
Type locality: South Africa. Zeller (p. 41)
cites the type locality as: “Habitat in trac-
tibus fluviorum Limpoponis et Gariepis.”
This is roughly the region of the Transvaal
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Pyralidae
Pyralidae s Brit..Mus
Bri Mus. ae vie | Stide No. |
ide No. . eee 1?
14576 ae RELI
5
Specimen labels. 4, Herpetogramma mutualis, lectotype (1.0 x); 5, H. verminalis, holotype (1.6 x);
and Orange Free State. (See Shaffer & Mun-
roe, 1989: 272.)
The following points of agreement be-
tween Zeller’s description and the lectotype
(Fig. 1) are particularly noteworthy (italics
text adapted rather freely from Zeller’s de-
scription). Comparisons of the same char-
acters are made with the BMNH specimen
under mutualis, and with the BMNH lec-
totype (designated below) of aegrotalis (Fig.
3). The purpose of the comparisons with the
BMNH specimens is to demonstrate clearly
that neither of them matches Zeller’s de-
scription of mutualis.
a. Wing lines nodulose—This is clearly
seen in the postmedial line of the mutualis
lectotype. In the BMNH specimen under
mutualis the postmedial line is interrupted,
but not nodular, and in the aegrotalis lec-
totype it is continuous and clearly not no-
dular.
b. Posterior sinus (outward bulge) of post-
medial line less rectilinear than in aegro-
talis—The upper and lower outer angles of
this bulge form right angles in the aegrotalis
lectotype and approximate this in the
BMNH specimen under mutualis, but are
rounded in the lectotype of mutualis.
c. Small spot or thickened dot on trans-
verse vein—This is a reference to the discal
spot, elliptical in both the mutualis lecto-
type and the BMNH specimen under mu-
tualis, but conspicuously different in aegro-
talis with its lunate discal spot. This is one
of the most obvious features separating mu-
tualis and aegrotalis.
VOLUME 91, NUMBER 3
d. Orbicular spot halfway between ante-
medial line and discal spot —This feature fits
the lectotype, but in the BMNH specimen
under mutualis and in aegrotalis the orbi-
cular spot is at approximately * the distance
from the antemedial line to the discal spot.
e. Postmedial line thickened at costa—This
feature is evident in the mutualis lectotype,
but not in the BMNH specimen under mu-
tualis where the line stops short of the costa,
nor in the aegrotalis lectotype where the line
reaches the costa but is not thickened there.
f. Postmedial line nearly straight above
sinus —The mutualis lectotype and the
BMNH specimen under mutualis agree in
this feature, but in the aegrotalis lectotype
this portion of the line is concave.
g. Hindwing discal spot round—The mu-
tualis lectotype and the BMNH specimen
under mutualis agree in this, but in the ae-
grotalis lectotype the discal spot is in the
form of a short bar.
h. Hindwing with outward sinus of post-
medial line straight—The mutualis lecto-
type agrees with this and the BMNH spec-
imen under mutualis is very close, but in
aegrotalis the upper side is curved.
In the male genitalia of the lectotype (Figs.
7-10) the uncus is narrowly triangular with
the sides less divergent on the distal third
and the apex narrowly rounded; the distal
third is moderately setose with the setae
extending toward the base along the sides,
but not quite reaching the base. The inner
margin of the sacculus bears a small, low,
moundlike setose projection (Fig. 8) on its
inner margin, but no clasper is present. Two
kinds of armature occur on the vesica; just
posterior to the center of the aedeagus is
seen an irregular platelike sclerite presenting
a short digitate profile, ill defined anteriorly,
rounded posteriorly, tapering, acute at its
apex. This feature did not photograph well,
but is outlined in black ink in Fig. 9. On
the distal fourth of the aedeagus the unev-
erted vesica exhibits a ‘bottle-brush’ ar-
rangement (Fig. 10) of what appears to be
a very large number of slender apparently
417
spatulate setae densely arranged around a
central axis.
Herpetogramma yerminalis (Guenée),
NEw COMBINATION, Revised Status
Figs. 2, 5, 14-16
Botys verminalis Guenée, 1854: 348.
Pachyzancla verminalis (Guenée). Hamp-
son, 1899: 204. As synonym of P. bi-
punctalis (Fabricius).
Holotype, female, Sierra Leone, labelled:
““Holo-type”; ““Verminalis Gn. Sierra
Leone”; “‘Paravicini Coll. B.M. 1937-383.”:
“Holotype”; “@ Pyralidae Brit. Mus. Slide
No. 14576” [BMNH].
We illustrate the holotype (Fig. 2) to show
the wing pattern, which is different from
both those of mutualis and aegrotalis, and
we figure the genitalia, although Michael
Shaffer reports that the abdomen appeared
to have been glued on, so that there is some
doubt as to its correct association. None-
theless, the wing maculation is clearly that
of a Herpetogramma, and characters of the
genitalia agree well with those of known
species of Herpetogramma. Further study
of the genus may well show that the genitalia
figured are indeed those of verminalis.
Female genitalia (Figs. 14—16).—Poste-
rior apophysis straight, slender; anterior
apophysis about 2 as long as posterior,
broadened and distinctly downcurved near
base. Anterior half of eighth-segment collar
nude, posterior half bearing numerous fine
setae (on this specimen about 36 on right
side, about 28 on left side, about 16 ven-
trally). Ostial chamber funnel shaped; an-
terior half with numerous folds bearing mi-
nute subovate scales, each several
micrometers wide (these best seen on edge,
and then with difficulty). Ductus bursae
broadly joining corpus bursae; posterior end
forming smooth, unarmed, well sclerotized
collar (Fig. 14, c) nearly 2 x as wide as long;
anterior and adjacent to this collar the
membrane armed with numerous minute
sharp spines, these continued anteriorly,
418 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 7-13. Male genitalia. 7, Herpetogramma mutualis, lectotype; 8, enlarged to show basal detail of left
valve; 9, aedeagus; 10, distal end of aedeagus, enlarged. 11, Pleuroptya aegrotalis, \ectotype; 12, aedeagus; 13,
detail of vesica showing cornuti. Scale bar = 0.50 mm (Figs. 7 & 9, 11 & 12); 0.20 mm (Fig. 8); 0.05 mm (Figs.
10, 13).
VOLUME 91, NUMBER 3
419
Figs. 14-16. Herpetogramma verminalis, holotype. Fig. 14, female genitalia (c = sclerotized collar of ductus
bursae, ds = ductus seminalis, s = signum); Figs. 15-16, signum, enlarged, two focal levels, each rotated 90°
clockwise relative to Fig. 14. Scale bar = 0.50 mm (Fig. 14), 0.10 mm (Figs. 15-16).
gradually becoming shorter, broader, and
forming triangular denticles which extend
onto corpus bursae to a short distance be-
yond signum (many are seen in Figs 15, 16).
Corpus bursae with midregion bearing
weakly developed transverse signum about
half as long as corpus bursae width (Fig. 14,
s); at high magnification (Figs. 15, 16) sig-
num seen as a deep narrow groove armed
with minute triangular denticles on its mar-
gin and sides, and with longer spatulate
scales at its bottom and ends. Ductus sem-
inalis (Fig. 14, ds) from ductus bursae just
anterior to sclerotized collar; broadened and
funnel shaped at origin.
Pleuroptya aegrotalis (Zeller),
New ComBINaTION, Revised Status
Figs. 3, 6, 11-13
Botys aegrotalis Zeller, 1852: 38-39.
Pachyzancla aegrotalis (Zeller). Hampson,
1899: 204. As synonym of P. bipunctalis
(Fabricius).
Psara aegrotalis (Zeller). Klima, 1939: 384.
As distinct species.
Lectotype, hereby designated, labeled (Fig.
5): “Type”; “Botys aegrotalis Z. Caffr. 39.
Led. 85. Caffraria.” [handwritten]; “‘Zell.
Coll. 1884.”; “*¢ Pyralidae Brit. Mus. Slide
No. 17755”; “Lectotype by J. Shaffer & E.
Munroe, 1989.” The original description
was based on the male sex. The number of
specimens was unspecified and we have not
discovered additional Zeller material of this
species.
Type locality: South Africa. Zeller (p. 39)
cites the type locality as: ‘“‘Habitat in trac-
tibus fluviorum Limpoponis et Gariepis.”
This is roughly the region of the Transvaal
and Orange Free State. (See Shaffer & Mun-
roe, 1989: 272.)
As discussed above neither of the males
under aegrotalis in the Stockholm museum
matches the original description. We have
chosen the single male in the British Mu-
420
seum as the lectotype as it is a good match
for Zeller’s description, the following points
of agreement being particularly noteworthy
(Fig 3):
a. The forewing apex is acuminate.
b. The forewing termen (outer margin) is
oblique and slightly convex posteriorly.
c. The discocellular spot is a distinct lu-
nule. This is one of the most obvious fea-
tures of aegrotalis.
d. The outward sinus of the postmedial
line forms nearly a right angle at both top
and bottom ends.
e. The hindwing discocellular is marked
by a short bar rather than by a spot.
We also note that in contrast to mutualis
the postmedial line is not thickened at the
apex and is not nodular.
The male genitalia of the lectotype (Figs.
11-13) show that, despite its resemblance
in wing pattern to certain species of Her-
petogramma, aegrotalis does not belong to
that genus at all, but is a typical member of
the genus Pleuroptya Meyrick, 1890, where
we transfer it as a new combination (see
above). Among the distinctive features are
the short, distally truncate and medially
slightly excavated uncus, without special-
ized dorsal scaling or spines, the narrow,
bridgelike gnathos, the broad, medially
complete transtilla, the prominent saccus,
the large saclike coremata, and the ven-
trodistally oblique, clavate clasper, arising
subbasally from the subcostal ridge of the
valve. The vesica, withdrawn in the holo-
type within the clavate aedeagus, is densely
set with small but strong, triangular denti-
cles (Fig. 13).
Several probably related species occur in
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Africa, but we have not so far detected any
junior synonym that needs to be supplanted
by aegrotalis.
ACKNOWLEDGMENTS
We thank Mr. Bert Gustafsson of the
Naturhistoriska Riksmuseet, Stockholm for
the loan of Zeller material and Mr. Michael
Shaffer of the British Museum (Natural His-
tory) for the loan of genitalia slides and pho-
tographs of specimens and labels repro-
duced herein (Figs. 2, 3, 5, 6). We also
appreciate the helpful comments of two
anonymous reviewers.
LITERATURE CITED
Guenée, A. 1854. Deltoides et Pyralites. Jn J.B.A.D.
Boisduval and A. Guenée, Histoire naturelle des
insects: Species général des Lepidoptéres, 8: 448
pages, 10 plates. Paris: Librairie Encyclopedique
de Roret.
Hampson, G. F. 1899. A Revision of the Subfamily
Pyraustinae of the Family Pyralidae, Part II. Proc.
Zool. Soc. Lond. 1899: 172-291, figs. 86-161,
plates 16-19.
Klima, A. 1939. Pyralidae: Subfamily Pyraustinae II.
In F. Bryk, ed., Lepidopterorum Catalogus 89:
225-384. 's-Gravenhage: W. Junk.
Minet, Joel. 1981. Les Pyraloidea et leurs principales
divisions systematiques. Bull. Soc. Ent. Fr. 86:
262-280.
Munroe, E.G. 1972. In Dominick, R. B., et al., The
Moths of America North of Mexico, Fasc. 13.1A,
Pyraloidea (in part). The Curwen Press, London.
Shaffer, J. C. and E.G. Munroe. 1989. Type Material
of Four African Species of Notarcha Meyrick, with
Designations of Lectotypes and Changes in Syn-
onymy (Lepidoptera: Crambidae: Pyraustinae)
Proc. Entomol. Soc. Wash. 91(2): 265-273.
Zeller, P. C. 1852. Lepidoptera Microptera, quae J.
A. Wahlberg in Caffrorum Terra Collegit. Kongl.
Vetenskaps-Akademiens Handlingar for Ar 1852,
pages 1-120.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 421-428
A NEW SPECIES OF RHEUMATOBATES BERGROTH FROM
ECUADOR AND DISTRIBUTION OF THE GENUS
(HETEROPTERA: GERRIDAE)
JOHN T. POLHEMUS AND PAUL J. SPANGLER
(JTP) University of Colorado Museum, 3115 S. York, Englewood, Colorado 80110,
USA; (PJS) Department of Entomology, National Museum of Natural History, Smith-
sonian Institution, Washington, D.C. 20560.
Abstract. — Rheumatobates peculiaris, n. sp., is described from Ecuador and compared
to other species of the genus; distinctive characters are illustrated by line drawings. Ad-
ditional distributional data, mainly from Mexico and Mesoamerica, are given for other
species of the genus.
Key Words:
bution
Spangler, Froeschner, and Polhemus
(1985) provided a checklist of the species
and subspecies of Rheumatobates Bergroth,
a euryhaline genus restricted to the New
World. They recognized 30 species and 3
subspecies. Polhemus and Manzano (in
press) add a new species from marine hab-
itats in Colombia and Ecuador and report
distributions of the marine species of the
eastern tropical Pacific. In this paper we add
another new species, Rheumatobates pecu-
liaris, and report distributions of a number
of species, based mainly on material in the
Polhemus collection.
Rheumatobates peculiaris, NEw SPECIES
Figs. 1-5
Material examined.— Holotype, apterous
male; and apterous allotype: ECUADOR:
LOS RIOS: Babahoyo (20 Km N.), 22 June
1975, A. Langley, J. Cohen and P. Monnig,
Ecuador-Peace Corps-Smithsonian Insti-
tution Aquatic Insect Survey (USNM).
Paratypes: Same data as holotype, 4 apter-
ous males, 19 apterous females, 3 alate fe-
males, 35 nymphs (USNM, JTPC).
Heteroptera, Gerridae, New World, Rheumatobates peculiaris n. sp. distri-
Etymology. —The name peculiaris (Latin)
means strange, new and refers to the oddity
of the male.
Distribution. — Ecuador.
Diagnosis. — Rheumatobates peculiaris,
new species, vaguely resembles Rheuma-
tobates klagei Schroeder but the shape of
the male first antennal segment, shape and
armature of the fore and hind legs, and male
genitalia are unique and diagnostic. The very
long, highly modified coxae separate this
species from any other known species of the
genus.
Description. — Length, apterous male, 2.4
mm; apterous female, 2.9 mm. Ground col-
or of apterous male deep brown; antero-
medial part of pronotum, broad ovate patch
on mesonotum, entire sternum except basal
abdominal segments, pleura, basal *% of an-
tennal segment 1, basal *4 of fore femur,
coxae, most of abdominal segment VIII yel-
lowish to leucine. Color of apterous female
similar to male except with more extensive
light markings as follows: antennal segment
1 and basal * of segment 3, fore femur ex-
cept distal 45, fore tibia medially, mid and
422 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-5. Rheumatobates peculiaris Polhemus and Spangler, new species. 1, Antenna. 2, Posterior leg. 3,
Jenital segments. 4, Abdominal ventrite IX. 5, Fore leg.
hind femora basally, abdominal tergites me- I-IV: male, 0.79 mm; 0.14 mm; 0.47 mm;
dially yellowish. 0.40 mm; female, 0.40 mm; 0.11 mm; 0.40
Antennae, fore legs, hind legs of male mm; 0.36 mm. Male antennae with stiff se-
modified (Figs. 1, 2, 5). Antennal formula tae and bristles (Fig. 1); segment | slightly
VOLUME 91, NUMBER 3
bowed, widest medially, with stiff setae di-
rected mostly laterally; segments II and III
straight, with various bristles and setae di-
rected mainly ventrally but at various an-
gles; segment IV curved, with short ven-
trally directed setae.
Head of male short (0.47 mm), broad (0.54
mm across antennal tubercles); equally short
(0.47 mm) but narrower (0.47 mm) in fe-
male; with two long curved ocular setae.
Male genital segments strongly modified;
genital segment II strongly excavate be-
neath, with 2 (1 +1) anteromedially directed
structures apparently composed of stiff se-
tae (Figs. 3, 4). Connexiva of male narrow;
broader and flatter in female. Male fore fe-
mur modified, irregularly set with spines and
bristles along posterior margin; fore tibia set
with stout setae (Fig. 5). Female fore femur
set with slender setae beneath; fore tibia with
three long setae, two stout and one slender,
and many short appressed setae. Male mid-
dle femur set along entire length with se-
mirecumbent spines on inner side and me-
dium length decumbent setae along outer
side, with a dorsal row of about 17 long
decumbent setae on distal third. Male hind
coxae very long (0.68 mm), subequal to hind
femur (0.76 mm), carinate dorsally, exca-
vate laterally, strongly modified with a dis-
tal brush of posteriorly directed setae (Fig.
2); trochanter long (0.18 mm), with a me-
dially directed spine-like process composed
of stiff setae (Fig. 2). Female middle and
hind femora unarmed. Male and female fore
claws long, slender, similar.
Length (in mm) of legs of allotype:
Tarsal Tarsal
Femur Tibia ] 2
Anterior 0.79 0.47 0.07 0.22
Middle 223 2223 Al 0.43
Posterior 1.29 LOLS Os18i 0:32
Habitat.—The habitat of this species is
unknown.
Distribution.—The following distribu-
tional notes were derived mainly from data
423
with specimens in the Polhemus collection
(JTPC). The CL numbers refer to detailed
ecological notes made by JTP and cowork-
ers during 25 years of collecting in Mexico
and Mesoamerica. The bound journals are
in the Polhemus library. Abbreviations are
given only for collections other than JTPC.
The names of collectors D. A., J. T., and
M.S. Polhemus are abbreviated DAP, JTP,
and MSP respectively. Major additions to
the distributions given by Spangler,
Froeschner, and Polhemus (1985) are pre-
ceded by an asterisk. Two papers not cited
in the checklist by Spangler, Froeschner, and
Polhemus (1985) provide additional distri-
butional records; Drake and Hottes (1951)
record R. bonariensis (Berg) from *Bolivia,
*Paraguay, *Uruguay as well as those lo-
calities discussed below; Nieser (1970) re-
corded R. crassifemur esakii Schroeder
(1931) from *Suriname, R. k/agei Schroeder
(1931) from *Peru, and R. trinitatis (China
1943), from *Suriname.
Rheumatobates bergrothi Meinert, 1895
This species was originally described from
Grenada. Drake and Hottes (1951) list it
also from *Panama and the Virgin Islands.
Hungerford (1954) added Venezuela, and
Spangler, Froeschner, and Polhemus (1985)
listed El Salvador and Honduras without
definite locality data. We have the following
additional records from Mesoamerica.
EL SALVADOR: 13 mi. NW of Amatil-
lo, CL) 1259.22) Dec. 1969) JEP:
HONDURAS: 16 mi. S. San Lorenzo, CL
1310, 9 Jan. 1970, JTP:; 17 mi. E. of Jicaro
Galan, CL 1311, 9 Jan. 1970, JTP.
*NICARAGUA: near Lake Nicaragua,
CL 1308, 8 Jan. 1970, JTP.
PANAMA: San Carlos, CL 1290, | Jan.
1970, JTP; E. of Panama City, CL 1295, 2
Jan. 1970, JTP.
Rheumatobates citatus
Drake & Hottes, 1951
This species was originally described from
the state of Oaxaca, Mexico, and is also list-
424
ed from the adjacent states of Chiapas and
Guerrero (Hungerford 1954). All three of
those localities are close to the Pacific coast,
but we now have specimens from an inland
locality in Morelos, dispelling any notion
that this species is restricted to coastal hab-
itats. We have the following records from
Mexico and Mesoamerica.
*GUATEMALA: E. of Puerto San José,
CL 1251720 Deer 1969s aimP:
MEXICO: MORELOS: 20 mi. S. of Cuer-
navaca, CL 1047, 27 Apr. 1964, JTP & MSP;
OAXACA: Tequisistlan, CL 1066, 30 Apr.
1964, JTP & MSP; E. of Tehuantepec, CL
1067, 30 Apr. 1964, JTP & MSP.
Rheumatobates clanis
Drake & Harris, 1932
Hussey (1955) reported this marine
species as new for the United States, citing
Herring’s collections at Bayport, Florida, in
November 1947. Herring (1958) later hy-
pothesized that its presence there was the
result of hurricane transport, and ques-
tioned whether breeding populations were
established in the United States. In the Pol-
hemus collection we have specimens col-
lected at Pine Island (a few miles north of
Bayport) by H. C. Chapman in November
1952, indicating a long term presence on
the Florida west coast. In both March and
August, 1988, D. A. Polhemus (personal
communication) found a large and appar-
ently well-established breeding population
on brackish water near Everglades City,
Florida, establishing this species as a per-
manent part of the fauna of the United
States. We have the following records from
the Caribbean and Mesoamerica.
GUATEMALA: Santo Tomas de Costilla
(east coast), CL 1320, 11 Jan. 1970, JTP.
JAMAICA: St. James Greenwood, at
beach, 14 Mar. 1970, C. E. Aarons; Tre-
lawny, Martha Brae River, nr. Falmouth,
14 Mar. 1970, C. E. Aarons.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Rheumatobates crassifemur crassifemur
Esaki, 1926
The records from *Bolivia given by Hun-
gerford (1954) were overlooked by Span-
gler, Froeschner, and Polhemus (1985).
Rheumatobates creaseri Hungerford, 1936
This species was originally described from
a cenote in Yucatan, Mexico, and is addi-
tionally known from Guerrero (Hungerford
1954). We have records from both coasts
of Mexico.
MEXICO: *NAYARIT: San Blas, lower
reaches of a spring fed river, along man-
groves, CL 1208, 3 June 1966, JTP; *VE-
RACRUZ: Rio Blanco, La Tinaja, CL 504,
4 Jan. 1971, JTP & MSP.
Rheumatobates drakei Hungerford, 1954
There are specimens with the following
data from Suriname and Peru in the Snow
Entomological Museum that were appar-
ently overlooked by Professor Hungerford
when he described this species.
PERU: HUANUCO: Loc. Shapajilla
Jungle, 630 m, No. 3837, 26 July 1938, F.
Woytkowski (SEMC).
SURINAME: “Dutch Guiana,” upper
right Coppename River, Se kamp, Sept.
1943, D. C. Geijskes (SEMC).
Rheumatobates hungerfordi Wiley, 1923
In Hungerford’s (1954) monograph, he
gives a Utah locality for this species as
“Emery Co., Utah, Aug. 2, 1922, Mrs. Grace
Olive Wiley.”’ One of us (JTP) has collected
on the Colorado River in Grand County,
Utah, and the Green River in adjacent
Emery County, Utah, without finding this
species, and has long considered this record
suspicious. In her paper describing this
species Wiley (1923) gives the following
data: “Holotype: apterous male, collected
near Rock Island, Texas, Aug. 2, 1922
(Grace O. Wiley); in author’s collection.” In
1922, before air travel was common, it is
VOLUME 91, NUMBER 3
unlikely that she was in eastern Texas and
Utah on the same day. Mrs. Wiley undoubt-
edly collected these specimens in Texas,
perhaps at Cisco. There is a town of the
same name in Grand County, Utah, near
the Colorado River. The border between
Emery and Grand counties in Utah is the
Green River and it could be easy to misread
the map and place Cisco in Emery County.
The Colorado River (of Texas, not Utah)
flows a few miles east of Rock Island, another
confusing coincidence. Mrs. Wiley lists as
paratypes specimens taken in June near Cis-
co, Texas (in Eastland County, about 250
miles north of Rock Island), but does not
mention any from Utah collections. Utah
should be removed from the United States
distribution of this species. Although there
are published records of Rheumatobates
from Saskatchewan, Arizona, New Mexico,
and much farther west in Mexico, as far as
we can determine, the 100th meridian es-
sentially marks the western boundary of dis-
tribution of Rheumatobates in the middle
latitudes of the United States.
Rheumatobates hungerfordi is common
in central and eastern Texas and along the
east coast of Mexico, particularly in Vera-
cruz, and ranges southward to Belize. It is
less common in other southern parts of the
United States states but ranges as far north
as Missouri (Smith 1988). With his collec-
tions at Carlsbad, New Mexico (most likely
the Pecos River), and E] Paso, Texas, Drake
extended the known distribution westward
along the Rio Grande drainage (Drake and
Harris 1937). Drake and Hottes (1951) list
it from Arizona, but we have been unable
to confirm that record. They also give lo-
calities in the eastern Mexican states of
Puebla, San Luis Potosi, Tamaulipas, and
Veracruz. In addition to many collections
from Texas, we have the following records
from Mexico and Mesoamerica.
BELIZE: Mountain Pine Ridge, Rio On,
CL 644, 31 Dec. 1973, JTP.
MEXICO: *CHIAPAS: Montebello
425
Lakes, CL 1083, 3 May 1964, JTP & MSP.
*OAXACA: Valle Nacional, CL 506, 4 June
1971, JTP & MSP. VERACRUZ: Rio Blan-
co, La Tinajya, CL 504, 4 Jan. 1971, JTP &
MSP; Rio Paso de Ovejos, CL 513, 6 Jan.
1971, JTP & MSP; N. of Nuatla, CL 518,
7 Jan. 1971, JTP & MSP; 17 mi. S. of Tux-
pan, CL 521, 7 Jan. 1971, JTP & MSP; 5
mi. S. of V. Alatorre, CL 675, 8 Jan. 1974,
Jae:
Rheumatobates imitator (Uhler, 1894)
Spangler, Froeschner, and Polhemus
(1985) listed Martinique without specific lo-
cality data, which is: MARTINIQUE: Ri-
viere Salée S. ditch, 12 July 1967, P. W.
Hummelinck.
Rheumatobates mexicanus
Drake & Hottes, 1951
This species was originally described from
the state of Guerrero, Mexico, with addi-
tional localities recorded in the states of
Aguascalientes, Mexico D. F., Oaxaca,
Puebla, San Luis Potosi, Tamaulipas, and
Veracruz. This species is widespread and
common in Mexico, and also occurs farther
south. We have the following records.
*HONDURAS: 17 mi. E. of Jicaro Ga-
lan, CL 1311, 9 Jan. 1970, JTP.
MEXICO: *COLIMA: Melaque, CL
1226, 20 Nov. 1968, JTP; Santiago, CL
1227, 25 Nov. 1968, JTP; Cuyutlan, CL
1228, 26 Nov. 1968, JTP. GUERRERO:
Ixtapa, CL 1892, 27 Jan. 1985, JTP; large
lagoon N. of Pie de la Cuesta, CL 1046, 26
Apr. 1964, JTP & MSP. *JALISCO: Puerto
Vallarta, CL 731, 4 June 1975, JTP; Tena-
catita, CL 738, 10 June 1975, JTP. *NAYA-
RIT: Santa Cruz, CL 728, 8 June 1975, JTP;
San Blas, CL 1208 & CL 1209, 3 June 1966,
JEP; 7 mi NE of San Blas; Cl. 1025, 21
Apr. 1964, JTP & MSP. *SINALOA: Es-
cuinapa, CL 1023, 21 Apr. 1964, JTP &
MSP.
426 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Rheumatobates minimus Drake, 1958
In addition to the type locality in Peru,
we can add the following.
BRAZIL: Ponte Nova, 29 July 1967, R.
L. Usinger; Nova Teutonia, Santa Catarina,
May 1957, F. Plaumann.
Rheumatobates minutus minutus
Hungerford, 1936
Hussey (1955) reported this species as new
to the Untied States based on collections
from Lakeland and the Big Pine Key in Flor-
ida. Herring (1958) considered its presence
there to be the result of hurricane transport,
and questioned whether breeding popula-
tions were established in the United States.
In the Polhemus collection we have speci-
mens taken in the month of November by
H. C. Chapman at Christmas and Orlando,
Florida. D. A. Polhemus (personal com-
munication) has recently seen in March and
August an established breeding population
near Everglades City, so this species can be
considered a permanent resident of Florida.
It is common in Mexico and Mesoamerica
from which we have the following records.
*BELIZE: Beaver Dam Cr., CL 629, 27
Dec. 1973, JTP; 10 mi. W. of Stann Creek,
@Li639) 27, Dec. 1973. TDP:
COSTA RICA: 10 mi. S. of Palmar Sur,
CL 1281, 28 Dec. 1969, JTP; S. of San Isi-
dro del General, CL 1302A, 6 Jan. 1970,
JP:
*GUATEMALA: Peten, 3 mi. S. of Tikal,
CL 653), 2 Jans 1973, JP:
MEXICO: *CAMPECHE: Edzna, Pond,
CL 613, 15 Dec. 1973, JTP. *QUINTANA
ROO: Tulum, CL 621, 20 Dec. 1973, JTP.
*VERACRUZ: 16 mi. S. of La Tinaja, CL
505, 4 Jan. 1971, JTP & MSP; S. of Gu-
tierrez Zamora, CL 519, 7 Jan. 1971, JTP
& MSP; Papantla, CL 520, 7 Jan. 1971, JTP
& MSP: S. of Loma Bonita, CL 1333, 15
Jan. 1970, JTP; N. of Nuatla, CL 518, 7
Jan. 1971, JTP & MSP.
Rheumatobates minutus flavidus
Drake & Harris, 1942
The distribution of this subspecies abuts
that of the nominal subspecies in Costa Rica,
and we can find no isolating mechanisms.
In the series from Nuatla, Veracruz, Mex-
ico, and Tikal, Guatemala (see above), the
males and winged specimens key to minutus
minutus but the apterous females key to mi-
nutus flavidus, so we question the distinct-
ness of these subspecies. There appears to
be a gradation between the two forms in
Mesoamerica. We have the following rec-
ords.
COSTA RICA: San Vito de Java, CL
1284, 29 Dec. 1969, & CL 1286, 31 Dec.
1969; JEP:
*PANAMA: San Carlos, CL 1290, 1 Jan.
1970, JIP; Portobello, CL 1292, 1 Jan.
1970, JTP.
PERU: SAN MARTIN DEPT.: 8 km.
ENE of Tarapota, 550 m, 9 Jan. 1974, R.
T. Schuh (AMNH).
Rheumatobates petilus
Drake & Hottes, 1951
This species was originally described from
the state of Guerrero, Mexico, and is ad-
ditionally known from the state of Cam-
peche on the opposite coast (Hungerford
1954). We have specimens from a tidal es-
tuary emptying into the Gulf of Mexico.
MEXICO: *TABASCO: near Paraiso, Es-
tuary, CL 607, 12 Dec. 1973, JTP.
Rheumatobates spinosus
Hungerford, 1954
This species was reported from Bolivia
without specific locality data by Spangler,
Froeschner, and Polhemus (1985). We have
the following records.
BOLIVIA: Santa Cruz, 22 Oct. 1957, G.
Pinckert (also in USNM: see Drake, 1958);
Beni, Rio Itenez, opposite Costa Marques
(Brazil), 4-6 Sept. 1964, Bousemann & Lus-
senhop (AMNH).
VOLUME 91, NUMBER 3
Rheumatobates vegatus
Drake & Harris, 1942
Rheumatobates vegatus was described
from Cuba and has been reported addition-
ally from Florida and Puerto Rico. It is
common around mangroves in southern
Florida. Herring (1949) listed it from three
Florida localities and described it as a new
species, R. crinitus, which is now synony-
mized with R. vegatus. JTP collected it at
Coral Gables (Matheson Hammock Pk., CL
410, 4 Feb. 1968), where it was swarming
beneath the mangroves. In March and Au-
gust, 1988, D. A. Polhemus found it in large
numbers on salt water around mangroves
near Everglades City, Florida.
We found this species to be common in
mangrove habitats in Belize. JTP took a se-
ries from a limestone rimmed lagoon along
the Caribbean side of the Yucatan Peninsula
where freshwater from huge springs mingled
with the sea water.
*BELIZE: Cay Caulker, CL 624, 24 Dec.
97356 CL 628, 28 Dec 1973) JP:
STANN CREEK DISTR.: Twin Cays, West
Bay, red mangrove tidal flats, 11 Nov. 1987,
P. J. Spangler, R. A. Faitoute (USNM).
MEXICO: QUINTANA ROO: Xel-ho,
CL 619; 20 Dec: 1973; JTP.:
ACKNOWLEDGMENTS
We are indebted to the following: for fur-
nishing specimens and data: R. T. Shuh,
American Museum of Natural History, New
York (AMNH) and the late H. B. Hunger-
ford, Snow Entomological Museum, Uni-
versity of Kansas, Lawrence (SEMC); for
the gift of specimens: the late C. J. Drake,
the late R. L. Usinger, and N. Nieser. Spec-
imens are also deposited in the J. T. Pol-
hemus Collection, Englewood, Colorado
(JTPC) and the National Museum of Nat-
ural History, Smithsonian Institution
(USNM).
We thank Young T. Sohn, biological il-
427
lustrator, Smithsonian Institution, for the
illustrations.
LITERATURE CITED
China, W. E. 1943. A new genus and two new species
of Gerridae, subfamily Halobatinae (Hemiptera,
Heteroptera) from Trinidad. Proceedings Royal
Entomological Society London | 2(5-6, ser. B); 71—
80.
Drake, C. J. 1958. A new neotropical halobatid (He-
miptera: Gerridae). Bull. Brooklyn Entomol. Soc.
53: 100-102.
Drake, C. J. and H. M. Harris. 1932. An undescribed
water-strider from Honduras (Hemiptera, Gerri-
dae). Pan-Pacific Entomologist 8(4): 157-158.
. 1937. Notes on some American Halobatinae
(Gerridae, Hemiptera). Rev. Entomol., Rio de Ja-
neiro 7: 357-362.
1942. Notas sdbre ““Rheumatobates” com
descricao de uma nova espécie (Hemiptera, Ger-
ridae). Rev. Brasil. Biol. 2(4): 399-402.
Drake, C. J. and F. C. Hottes. 1951. Notes on the
genus Rheumatobates (Hemiptera: Heteroptera).
Proc. Biol. Soc. Washington 64: 147-155.
Esaki, T. 1926. The water-striders of the subfamily
Halobatinae in the Hungarian National Museum.
Annales Musei Nationalis Hungarici 23: 117-164.
Herring, J. L. 1949. A new species of Rheumatobates
from Florida (Hemiptera, Gerridae). Florida Ento-
mol. 4: 160-165.
1958. Evidence for hurricane transport and
dispersal of aquatic Hemiptera. Pan-Pacific Ento-
mol. 34: 174-175.
Hungerford, H. B. 1936. XIV Aquatic and semi-
aquatic Hemiptera collected in Yucatan and Cam-
peche, pp. 145-150. /n Pearse, A. The cenotes of
Yucatan, a zoological and hydrographic survey.
Carnegie Institution of Washington 457: 304 pages.
1954. The genus Rheumatobates Bergroth
(Hemiptera-Gerridae). Univ. Kansas Sci. Bull. 36:
529-588.
Hussey, R. F. 1955. Some records of Hemiptera new
to Florida. Quart. J. Florida Acad. Sci. 18: 120-
122:
Meinert, F. 1895. Rheumatobates bergrothi n. sp.
Saertryk af Entomologisk Medd. 5: 1-8, 2 pls.
Nieser, N. 1970. Gerridae of Suriname and the Am-
azon with additional records of other neotropical
species. Studies on the Fauna of Suriname and
other Guyanas 12: 94-138.
Polhemus, J. T. and M. del Rosario Manzano. (in
press). Marine Heteroptera of the eastern tropical
Pacific (Gelastocoridae, Gerridae, Mesoveliidae,
Saldidae, Veliidae). Jn D. Q. Arias, ed., Fairchild
428 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Commemorative Volume, The Insects of Panama
and Mesoamerica.
Schroeder, H. O. 1931. The genus Rheumatobates
and notes on the male genitalia of some Gerridae
(Hemiptera, Gerridae). Univ. Kansas Sci. Bull.
20(2): 63-98.
Smith, C. L. 1988. Family Gerridae Leach, 1815, the
water striders, pp. 140-151. Jn T. J. Henry and
R. C. Froeschner, eds., Synoptic Catalog of the
Heteroptera or True Bugs of Canada and Conti-
nental United States. E. J. Brill, Leiden; i-xviii +
1-958 pages.
Spangler, P. J., R. C. Froeschner, and J. T. Polhemus.
1985. Comments on a water strider, Rheumato-
bates meinerti from the Antilles and checklist of
the species of the genus (Hemiptera: Gerridae).
Entomol. News 96: 196-200.
Uhler, P. R. 1894. On the Hemiptera-Heteroptera of
the Island of Grenada, West Indies. Proceedings
Zoological Society of London 1894: 167-224.
Wiley, G.O. 1923. A new species of Rheumatobates
from Texas (Heteroptera, Gerridae). Canadian
Entomol. 55: 202-205.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 429-435
TWO NEW SPECIES OF COCCIPOLIPUS
(ACARI: PODAPOLIPIDAE) PARASITES OF CHILOCORUS SPP.
(COCCINELLIDAE) FROM VERA CRUZ AND MORELOS,
MEXICO AND FLORIDA AND WISCONSIN, U.S.A.
RosBert W. HUSBAND
Biology Dept., Adrian College, Adrian, Michigan U.S.A.
Abstract. — Coccipolipus oconnori n. sp. (Acari:Podapolipidae) is parasitic on Chilocorus
stigma (Coccinellidae) at several localities in Florida and near Sauk City, Wisconsin,
U.S.A. and Coccipolipus cactii n. sp. is parasitic on Chilocorus cacti from Vera Cruz and
Morelos, Mexico. The new species is closest to the African Coccipolipus chilocori Husband
1981. New distributional records of C. macfarlanei include Wenatchee, Washington and
Castelar, Argentina.
Key Words:
Mites of the genus Coccipolipus are sub-
elytral parasites of coccinellid beetles, with
the greatest diversity of known species oc-
curring in Africa (Husband 1972, 1981,
1984a, b). To date, only one species of Coc-
cipolipus has been collected from the beetle
genus Chilocorus, C. chilocori Husband,
1981. During studies of the phoretic asso-
ciations between the mite genus Hemisar-
coptes (Hemisarcoptidae) and Chilocorus,
B. M. OConnor and M. A. Houck of the
University of Michigan examined over 5000
museum specimens of Chilocorus. Collec-
tions representing two new species of Coc-
cipolipus from North American Chilocorus
species were recovered during the study.
These species, each restricted to a single
species of Chilocorus, are described below.
All beetles from which Coccipolipus speci-
mens were recovered are housed in the U.S.
National Museum of Natural History col-
lections unless otherwise noted. Insects are
vouchered with labels reading “Mites re-
moved, B. M. OConnor” and a voucher
number corresponding to the number on the
mite slides.
Acari, Podapolipidae, Coccipolipus, Chilororus, parasite, new species, mites
Measurements were made using a Wild
phase contrast microscope with a drawing
tube calibrated from a stage micrometer.
Terminology is based on Lindquist (1986).
All measurements are in micrometers (mm).
Coccipolipus oconnori, NEw SPECIES
Female (Figs. 1, 2).—Gnathosoma longer
than wide; length 57, width 41. Cheliceral
stylets slender, smooth, length 21. Pharynx
length 29, width 23. Gnathosoma usually
retracted into sclerotized chamber. Stig-
mata on slender stalks dorsolateral to
gnathosoma. Palps prominent. Tectum
covers gnathosoma, length 60, width 45.
Idiosoma—smooth, lightly sclerotized;
length 445-486, width 251-278. Fully de-
veloped females with anterolateral lobes
wider than posterior idiosoma. Lightly
sclerotized posteroventral internal triangu-
lar structure, length 105-148, width 85-125.
Legs—two pairs; anterior pair without suck-
er-like pad, with well developed hooked
spine and terminal spine, anterior femoral
setae, 5. Leg II with 2 terminal spines, an-
430
MN ing
0.1mm
Fig. 1. Coccipolipus oconnori n. sp., female, dorsal
aspect.
Fig. 2. Coccipolipus oconnori n. sp., female, partial
ventral aspect.
terior spine nearly as thick but about *4
length of posterior spine.
Male (Figs. 3, 4).—Gnathosoma length
22, width 25; dorsal setae microsetae, ven-
tral setae, length 2. Palps 2-segmented, dis-
tal segment truncate, basal segment with a
short seta. Cheliceral stylets length 8, about
Ys gnathosomal width. Idiosoma—length
120, width 100. Dorsum-prodorsal plate
narrows anteriorly, all setae microsetae.
Plates C and D fused, all setae microsetae;
Fig. 3. Coccipolipus oconnori n. sp., male, dorsal
aspect.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 4. Coccipolipus oconnori n. sp., male, ventral
aspect.
triangular aedeagus extends anteriorly be-
yond posterior margin of prodorsal plate.
Venter—slightly sclerotized apodemes | and
2 meet medially at sternal apodeme. Coxae
III fused, separate from coxae III. Legs—
leg setation as in Table |. Leg I with a ter-
minal stout claw and an anterolateral stout
spine with 2 adjacent thick setae as long as
the spine. Solenidion w reduced, scarcely
longer than width of setal socket. Solenidion
@ not apparent. Tibia I d seta length 19.
Femur II seta present. Tarsi III] with 2
spine-like setae, ventral spine with an ad-
jacent seta shorter than spine. Ambulacra
II, Hl 15, 12; with minute symmetrically
paired claws, length 1. One spine on tibia
I, 3 spines each on tibiae IT, III, spine lengths
4-5. Tarsus and tibia III longest setal length
6, genu III seta spine-like.
Larval female (Figs. 5, 6).—Gnathosoma
length 50, width 50; dorsal setal length 8,
less than % gnathosomal width, ventral se-
tae length 4. Palps 2-segmented, small seta
on each segment. Stylets smooth, slender,
length 30. Idiosoma—length 186, width 141.
Dorsum-prodorsal plate narrowed anterior-
ly; v, 4, sc, 99. Plates C and D fused. All
setae microsetae. Plate EF wider than long,
setae f length 10. Plate H somewhat trian-
VOLUME 91, NUMBER 3
Fig. 5. Coccipolipus oconnori n. sp., larval female,
dorsal aspect.
gular; setae h, length 11, h, length 168. Ven-
ter-apodemes poorly sclerotized; apodemes
1 and 2 meet medially at anterior sternal
apodeme. Coxae III separate from each oth-
er and from coxae I,II. Legs—leg setation
as in Table |. Leg I with 2 parallel, terminal
claws, solenidion w length 2. Thick tarsus I
pv’ - pv” setae at base of spine-like s seta.
Seta tc’ slightly less than twice length of seta
te”. Tarsi III] each with setae tc’ and u
subterminal, spine-like. Solenidion ¢@ no
longer than width of setal socket. Ambu-
lacra III] 28, 25, with minute symmetri-
cally paired claws.
Type data.—Holotype male: Enterprise,
Volusia County, Florida, U.S.A., from male
Chilocorus stigma (Say) (Coccinellidae) col-
lected May 25 (year unknown) by Hubbard
and Schwarz. Deposited in the United States
National Museum (U.S.N.M.) collection,
Washington, D.C. (BMOC 86-0703-18-1).
431
Fig. 6. Coccipolipus oconnori n. sp., larval female,
ventral aspect. Notation (after Lindquist 1986); pl,
primilateral; pv, primiventral; s, subunguinal; tc, tec-
tal; h, setae associated with plate H (6th opisthosomal
segment).
Paratypes (8 males, 20 larval females, 23
females)—same data as holotype. One fe-
male (BMOC 86-0703-31-43) and one lar-
val female (BMOC 86-0703-31-42) Hills-
boro Co., Florida, collected 7-12 April 1930
by J. C. Bowver on C. stigma. One larval
female (BMOC 8&6-0703-26-440) Brevard
Co., Florida on 19 April 1930 by R.A.
Schlernitsauer on C. stigma. Two females
(BMOC 86-0703-27-45, -46) Osceola Co.,
Florida between 7-12 April 1930 by M. M.
Smith on C. stigma. Two females (BMOC
86-0703-22-47, -48) Orange Co., Florida on
21 July 1929 by W. A. Miers on C. stigma.
One female (BMOC 86-0703-19-49) Sump-
Table 1. Total setae on legs of Coccipolipus spp. parasitic on Chilocorus spp. Solenidia are included.
Leg I Leg I Leg Il
iB G Ti Ta le G Ti Ta F G Ti Ta
C. oconnori n. sp. 3 5 6 8 1 ] 4 4 0 l 4 4
C. cacti n. sp. 3 3 6 8 1 1 4 4 0 1 4 4
C. chilocori H. 2 3 6 8 0) l 4 4 0 l 4 4
432
ter Co., Florida, 10-14 Feb. 1930 by B. L.
Smith on C. stigma. One female (BMOC
86-0703-16-50) Flagler Co., Florida on 21
Dec. 1929 by E. B. Webb on C. stigma. Two
males (BMOC 87-1023-3-1,-2), 3 larval fe-
males (BMOC 87-1023-3-3, -4, -5)) and 6
females (BMOC 87-1023-3-6, -7, -8, -9,
-10, -11) Sauk Co., Wisc., on 1 Sept. 1960
by G. Lockwood on C. stigma. Three males
(BMOC 86-0703-18-2, -3 and BMOC 87-
1023-3-1), 4 larval females (BMOC 86-
0703-18-9, -10, -11 and BMOC 87-1023-
3-3) and 4 females (BMOC 86-0703-18-25,
-26, -27 and BMOC 87-1023-3-6) are in the
collections of Adrian College, Adrian,
Michigan, U.S.A. Three males (BMOC 86-
0703-18-4, -5 and BMOC 87-1023-3-2), 3
larval females (BMOC 86-0703-18-12, -13
and BMOC 87-1023-3-4) and 3 females
(BMOC 86-0703-18-28, -29 and BMOC 87-
1023-3-7) are deposited in the Museum of
Zoology, the University of Michigan, Ann
Arbor, Michigan. The remaining paratypes
are deposited in the U.S.N.M., Washington,
DIC:
Etymology.—The species is named for Dr.
Barry M. OConnor of the University of
Michigan in tribute to his basic studies in
the field of acarology.
DIAGNOSIS
Of the 13 species of Coccipolipus, only 4
have femoral II setae and 3 of the 4, in-
cluding C. oconnori, have plates C and D
completely fused in the larval female stage.
C. oconnoriand the species described below
have h, setae (length 11) which are longer
than the distance between setae h,. The re-
maining species, C. coormani, has setae h,
(length 2) which are shorter than the dis-
tance between setae h,. C. oconnori has gna-
thosomal setae which are 20% shorter than
the species described below. Setae sc,, f,
femoral I |’ and tibial I 1’ are shorter in C.
oconnori than 1n the new species described
below.
Six of the 13 species of Coccipolipus have
males with three spine-like setae on tibia II
but only 3 species, including C. oconnori
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
have spine-like setae on genua III. C. ocon-
noriand C. cooremani share the 2 characters
above while C. bifasciatus has 2 spine-like
setae on tibia III in combination with spine-
like setae on genu III. The tibial III shortest
seta in C. oconnori is shorter (length 6) than
the basal width of tibia III (12) in contrast
to the same tibial seta (length 20) in C. coor-
emani in which basal width of tibia III is
14.
Adult females of C. oconnori and the
species described below have femoral I setae
(length 5) which do not extend beyond the
distal margin of genu I. Five Coccipolipus
spp. have femoral setae (lengths 13-18)
which extend beyond the distal margin of
tibiae I. The remaining 6 species lack fem-
oral I setae. Typically, anterior lobes of fe-
male C. oconnori do not extend beyond the
anterior margin of the gnathosoma and the
width of the width of the anterior idiosoma
is greater than the width of the posterior
idiosoma. The two characters contrast with
the condition in adult females of the species
described below.
Coccipolipus cacti, NEW SPECIES
Female (Figs. 7, 8).—Gnathosoma length
60, width 48: cheliceral stylets smooth, slen-
der, 23. Muscular pharynx length 35, width
26. Gnathosoma often retracted into a scler-
otized chamber. Stigmata on slender stalks
dorsolateral to gnathosoma. Palps promi-
nent. Tectum covers gnathosoma, length 60,
width 50. Idiosoma smooth, lightly sclero-
tuzed; length 260-560, width 200-375. Ful-
ly developed females with anterolateral lobes
less than or equal to widest part of posterior
idiosoma. Lightly sclerotized, triangular,
internal, posteroventral, structure, length
90-165, width 76-125. Legs—two pairs; an-
terior pair without sucker-like pad, with well
developed hook-like spine and terminal
spine, anterior femoral seta 5. Legs IT with
2 terminal spines, anterior spine '2 width
and length of posterior spine. Apodemes |
and 2 thin and weakly sclerotized but con-
spicuous.
Male (Figs. 9, 10).—Gnathosoma length
VOLUME 91, NUMBER 3
0.1mm
Fig. 7. Coccipolipus cacti n. sp., female dorsal as-
pect.
Fig. 8. Coccipolipus cactin. sp., female, partial ven-
tral aspect.
26, width 29; dorsal and ventral setae 2.
Conspicuous muscular pharynx. Palps
2-segmented, distal segment truncate, prox-
imal segment with a short lateral seta. Chel-
iceral stylets 8, about 4 width of gnatho-
soma. Idiosoma—length 140, width 124.
Dorsum-prodorsal plate narrow anteriorly,
all setae microsetae. Triangular aedeagus
extends beyond posterior margin of pro-
dorsal plate (not evident in Fig. 9 because
the specimen was flattened to observe other
structures). Venter-slightly sclerotized apo-
demes | and 2 meet medially at the sternal
apodeme. Coxae III fused, separate from
coxae III, coxal setal bases evident but
without setae. Legs—leg setation as 1n Table
1. Leg I with terminal stout claw and an-
teroventral subterminal spine. Solenidion @
not apparent. Tibial setae |’ are small thorns
no longer than diameter of setal sockets.
Tibia I d seta 22, femur II seta present. Tarsi
IL, III with 2 spine-like setae; ambulacra II, III
20,18, with minute, symmetrically paired
claws. One spine on tibia I, two spines on
tibiae II, III, spine lengths 5—6. Tibia III seta
14, tarsus III longest seta 10, about equal
to tarsus III basal width.
Larval female (Figs. 11, 12).—Gnatho-
soma length 44, width 48, dorsal setae 10,
ventral setae 8. Pharynx bulb-shaped. Palps
2-segmented, distal segment setal length 3,
proximal seta 1. Stylets smooth, slender,
length 40. Idiosoma—length 180, width 139.
A Ss . u
rm
433
) J
Fig.9. Coccipolipus cactin. sp., male, dorsal aspect.
Dorsum-prodorsal plate quadrate, nar-
rowed anteriorly; v, 5, sc, 127. Plates C and
D fused, all setae microsetae. Plate EF wider
than long, setae f 12. Plate H somewhat
Fig. 10.
pect.
Coccipolipus cacti n. sp., male, ventral as-
434
[ << Ss IX,
a Sh ah J
Oo \ /\ >
Fig. 11. Coccipolipus cactin. sp., larval female, dor-
sal aspect.
triangular; setae h, 11, setae h, 183. Venter-
apodemes weakly sclerotized, | and 2 meet
medially at anterior sternal apodeme. Cox-
ae III separated from each other and from
coxae I, II, all coxal setae microsetae. Legs—
leg setation as in Table 1. Leg I with 2 par-
allel, terminal claws, solenidion w no longer
than diameter of socket. Setae tc’ slightly
more than 2 times length of setae tc”. So-
lenidion # absent or vestigial. Ambulacra
of legs H,III] 22 and 30 respectively, with
minute paired claws.
Type data.—Holotype male: Cordoba,
Vera Cruz, Mexico, on Chilocorus cacti
(Linnaeus) (Coccinellidae) collected April
21, 1908 by F. K. Knab (BMOC 86-0707-
63-1). Deposited in the United States Na-
tional Museum, Washington, D.C. Para-
types (2 males, 19 larval females, 9 adult
females)—same data as holotype. One fe-
male (BMOC 86-0918-7-32), 3 miles S. Al-
puyeka, Morelos, Mexico, 3400 ft. elev.,
collected 8 April 1959 by H. E. Evans on
C. cacti; host beetle in the Cornell Univer-
sity Insect Collection. One male (BMOC 86-
0707-63-2), 3 larval females (BMOC 86-
0707-63-4, -5, -6) and 2 females (BMOC
86-0707-63-23, -24) are deposited in the
collection at Adrian College, Adrian, Mich-
igan. One male (BMOC 86-0707-63-3), 3
larval females (BMOC 86-0707-63-7, -8,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 12. Coccipolipus cacti n. sp., larval female,
ventral aspect.
-9) and 2 females (BMOC 86-0707-63-25,
-26) are deposited in the collections of the
Museum of Zoology, the University of
Michigan, Ann Arbor, Michigan. One fe-
male (BMOC 86-09 18-7-32) is deposited in
the Cornell University Insect Collection,
Ithaca, N.Y. The balance of paratypes are
deposited in the U. S. National Museum of
Natural History.
Etymology.—The species is named for the
host species, Chilocorus cacti.
DIAGNOSIS
Coccipolipus cacti is similar to C. ocon-
nori. Larval female C. cacti have longer
cheliceral stylets, gnathosomal setae, and sc;,
f, h,, femoral and tibial I I’ setae. Male C.
cacti have minute thorn-like tibial setae 1’
while male C. oconnori have typical slender
tibial I’ setae. Male C. cacti have typical
genu III setae while male C. oconnori have
spine-like genu III seta. Adult female C. cacti
have anterolateral lobes which extend be-
yond the anterior margin of the gnathosoma
and are less than or equal to the greatest
width of the posterior idiosoma. Leg II
spines in C. cacti, although not equal in
length are equal in width. In C. occonori,
the posterior tarsal spine is nearly twice the
thickness of the anterior tarsal spine.
VOLUME 91, NUMBER 3
DISCUSSION
Some species of Coccipolipus are wide
ranging in both host association within Coc-
cinellidae and 1n geographic distribution, e.g.
C. macfarlanei on 3 genera of coccinellid
beetles on 3 continents. In contrast, C. chil-
ocorl, C. oconnori and C. cacti are restricted
to the genus Chilocorus and each to a por-
tion of a continent. C. chilocori occurs on 5
species of Chilocorus in Central Africa, C.
cactl occurs on one species in Mexico and
C. oconnori occurs one one species in the
Eastern United States. This discussion is
limited to the 3 species of Coccipolipus as-
sociated with Chilocorus spp.
African C. chilocori are larger, have long-
er setae in general and longer leg setae spe-
cifically in females, males and larval females
than their counterparts C. cacti and C. ocon-
noriin North America. The idiosomal length
of larval female C. chilocori is about 1.2
times the length of the American species but
gnathosomal setae are 2 to 4 times longer
than the American species. Comparisons of
the longest tibial and tarsal III setae in males
and femoral setae in females give ratios of
2 to 6 times longer than in the American
species.
Although males ofall 3 species are similar
in that all idiosomal are microsetae, the ae-
deagus of C. chilocori extends farther for-
ward and 1s nearly equal to the width of the
gnathosoma. The aedeaguses of C. oconnori
and C. cacti are at most ¥4 the width of the
gnathosoma. Femoral I, II, III setal patterns
in male and larval female C. oconnori and
C. cacti are 3-1-0 whereas the pattern in the
African species is 2-0-0. Patterns on genua,
tibiae and tarsi are similar. Empodia are
conspicuously longer and more slender in
male and larval female C. chilocori than in
American species.
Since the genus Chilocorus is found in
both hemispheres and has a wide distribu-
tion pattern, it is likely that additional Coc-
cipolipus will be discovered. Ai this point,
it may be stated that there are differences
435
in numbers of setae, setal lengths and size
of structures between Coccipolipus from Af-
rican Chilocorus and Coccipolipus from
North American Chilocorus. The differ-
ences are consistant in the species studied.
Additional distribution records of
Coccipolipus spp.
In addition to records cited above and in
Husband(1984b), Coccipolipus macfarlanei
Husband 1972 is now known from Cocci-
nella transversoguttata (Fald.), 6 mi. N. of
Wenatchee, Washington, U.S.A. (Devin
Carroll, personal communication, 1983) and
from Eriopis connexa (Germ.) in Castelar,
Argentina (Alida Bolart, personal commu-
nication, 1983).
ACKNOWLEDGMENTS
Collections of mites from Chilocorus bee-
tles was funded by a grant from the National
Science Foundation to B. M. OConnor and
M. A. Houck (BSR 83-07711). I thank B.
M. OConnor for his review of the manu-
script.
LITERATURE CITED
Husband, R. W. 1972. A new genus and species of
mite (Acarina: Podapolipidae) associated with the
coccinellid Cycloneda sanguinea. Ann. Entomol.
Soc. Amer. 65: 1099-1104.
1981. The African species of Coccipolipus
with a description of all stages of Coccipolipus so-
lanophilae (Acarina: Podapolipidae). Rev. Zool.
Afr. 95(2): 283-300.
1984a. New Central African Coccipolipus
(Acarina: Podapolipidae). Rev. Zool. Afr. 98(2):
308-326.
1984b. The taxonomic position of Coccipo-
lipus (Acarina: Podapolipidae), a genus of mites
which are parasites of ladybird beetles (Coccinel-
lidae), pp. 328-336. /n Grifiths, D. A. and C. E.
Bowman, eds., Acarology, Volume 1. Ellis-Har-
wood Ltd., Chichester.
Lindquist, E. E. 1986. The world genera of Tarso-
nemidae (Acari: Heterostigmata): A morphologi-
cal, phylogenetic, and systematic revision, with a
reclassification of family-group taxa in the Het-
erostigmata. Mem. Entomol. Soc. Canada 136: l-
Dili
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 436-440
A NEW SPECIES OF ALEUROTULUS
(HOMOPTERA: ALEYRODIDAE)
SUEO NAKAHARA
Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, Belts-
ville, Maryland 2070S.
Abstract.—The pupal cases and third instar of Aleurotulus anthuricola, new species, are
described. This whitefly is apparently host-specific to Anthurium spp. and is endemic to
Colombia and several islands in the Lesser Antilles. It was detected in Hawaii in 1978.
Key Words:
Aleurotulus anthuricola, new species, 1s
apparently host-specific on Anthurium spp.
(Araceae) and endemic to Colombia and
several islands in the Lesser Antilles. In 1978
it was discovered in Hawaii on the inner
surface of leaf sheaths of anthurium plants.
Injury was not observed on infested plants
and therefore, anthuricola was of minor
concern to the anthurium flower industry in
Hawaii. In recent years however, infesta-
tions were found on the flower spathes and
although the damage is negligible, anthuri-
cola has become a major pest because of
problems with agricultural quarantine reg-
ulations. Infested flowers, unless treated,
cannot be exported from Hawaii to the con-
tinental United States and other uninfested
areas of the world. A description is pre-
sented here to provide a scientific name for
biological and control studies now in pro-
gress in Hawaii.
Measurements are based on 10 speci-
mens. The values are given first for the ho-
lotype followed by those for the paratypes
in parentheses. The terms pupae and pupal
cases are used interchangeably in this paper.
Aleurotulus, anthuricola, Aleyrodidae, whitefly, Anthurium spp., Hawaii
Aleurotulus anthuricola Nakahara,
New SPECIES
Figs. 1-7
Pupae and third instar larvae grayish
black; margins pale; with fringe of white,
filamentous wax. Often clustered in a mass
of white, filamentous wax on inner surface
of leaf sheaths and flower spathes and on
leaf petioles. Severe infestions also on outer
surface of sheaths and on flower spathes
(Figs. 6-7).
Pupa (Fig. 1): Pupal cases broadly oval.
Measurements of slide mounted specimens:
Length 880 (623-955) um, width 636 (438-
677) um.
Margin and submargin (Fig. 2): Margin
dentate, 13 (13-16) teeth per 100 um on
lateral margin. Teeth of two types: those
with elongate, triangular areas at bases;
smaller ones without triangular areas. Tra-
cheal pore area undifferentiated. Anterior
marginal setae 15 (12-20) um long, 14 (6-
18) teeth from midline; posterior marginal
setae 24 (17-30) um long, 34 (34-45) teeth
between setae. Submargin not differentiated
VOLUME 91, NUMBER 3 437
Figs. 1-5. Fig. 1. Pupa habitus (148 x). Fig. 2. Marginal and submarginal areas (390). Fig. 3. Vasiform
orifice and dorsal caudal area of pupal case (308 =). Fig. 4. Thoracic legs and antennae (230x). Fig. 5. Third
instar larva habitus (202 x).
438
Figs. 6-7.
of anthurium.
from dorsal disc by fold or furrow; with
short, transverse ridges; each ridge with
elongate triangular area extending into mar-
ginal tooth; proximal part of area with an
oval, pale spot. Micropore between ridges,
in irregular spaced row around case. Sub-
marginal setae 12-20 um long; 6 pairs on
cephalothorax, | pair on anterior abdomi-
nal segment. Small disc pores about | wm
in diameter mesad of ridges and submar-
ginal setae. Oval, transverse tubercle be-
tween caudal setae.
Dorsal disc: Derm smooth. Eyespots ab-
sent. Longitudinal molting suture extending
to anterior margin; transverse molting su-
ture extending caudolaterally and anteriorly
into subdorsum, lateral ends not attaining
level of its midpoint; sutures smooth; other
cephalothoracic segmental division ob-
scure. Submedial furrow on metathorax ex-
tending obliquely from suture. Cephalic se-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
«2
Fig. 6. Typical infestation in leaf sheath of anthurium. Fig. 7. Severe infestation on the petioles
tae 74 (49-84) um long; metathoracic setae
62-99 um long, in posterior '2 of segment;
eighth abdominal setae 64 (44-67) um long,
laterad of anterior margin of vasiform ori-
fice. Caudal setae on distal part of submar-
gin just mesad of submarginal ridges, 77
(49-77) um long, extending beyond poste-
rior margin, separated by 1.2-1.6 times
width of vasiform orifice; transverse tuber-
cle between setae. Shallow, oval, submedial
depressions on head, prothorax, mesotho-
rax and | pair each on abdominal segments
HI-VIII. Medial area of abdomen slightly
elevated: rachis slightly developed; medial
part of segment VII about '/2—¥4 times as long
as VI; segment VIII with medial tuberculate
area between small pockets. Vasiform ori-
fice (Fig. 3) subquadrate, 44 (35-42) um long,
47 (44-47) um wide, |.5—2.5 times its length
from posterior margin: rim thin, often cov-
ered by spinulose membranous inner wall
VOLUME 91, NUMBER 3
that may extrude over rim; bottom of orifice
open in anterior 73. Operculum irregularly
transverse, caudal part membranous and
spinulose, caudal margin rather straight or
irregular; 24 (20-24) um long, 37 (32-40)
um wide, occupying /2—% length of vasiform
orifice. Lingula spinulose; apical knob oval,
with 2 setae, exposed, included or protrud-
ing posteriorly beyond orifice. Disc pores
about 2 um in diameter and smaller asso-
ciated porettes distributed as follows: head
with | submedial, 3 subdorsal pairs; pro-
thorax with | submedial, 2 subdorsal pairs;
mesothorax with | submedial, 2 subdorsal
pairs; and metathorax with | submedial, 2
subdorsal pairs; abdominal segment I with
1 submedial pair, I] with 0-1 submedial
pair, III with | submedial and 1-2 subdorsal
pairs, segments IV—VIII each with | sub-
medial and | subdorsal pair.
Venter. Derm membranous, without dis-
tinct sculpture. Thoracic tracheal fold ob-
scure, abdominal tracheal fold with spi-
nules. Antenna (Fig. 4) not reaching anterior
thoracic spiracle, apex with a small, conical
point. Legs each with 2 distal setal bases
and 4 basal setae, mesothoracic and meta-
thoracic legs each with | distal seta. Eighth
abdominal setae 22 (22-35) um long, an-
terior of posterior abdominal spiracle.
Third instar larva (Fig. 5): Similar in shape
and morphological characters as pupal cases.
Length 537-562 um, width 394-418 um.
Margin and submargin: Margin broadly
crenulate, 12-16 crenulae per 100 um on
lateral margin. Anterior marginal setae 10-
12 um long, separated by 12-17 crenulae;
posterior marginal setae 12-24 long, sepa-
rated by 19-28 crenulae. Submargin similar
to pupal cases. Submarginal setae 7-10 um
long, 6 pairs on cephalothorax, | pair on
anterior abdominal segment. Row of small
disc pores mesad of setae.
Dorsal disc: Cephalic setae 24-53 um long,
metathoracic setae 49-67 um long, eighth
abdominal setae 37-42 um long; caudal se-
tae 54-67 um long. Vasiform orifice 27-32
um long, 35-37 um wide, anterior of pos-
439
terior margin by about its length. Opercu-
lum 17-20 um long, 27-35 um wide. Disc
pores and associated porettes distributed as
follows: cephalic segment with | submedial
pair; prothorax with 0-1 subdorsal pair;
mesothorax and metathorax each with 2
subdorsal pairs; abdominal segment I with
1 submedial pair; segments III-V with 0-1
submedial pairs; segments III and VI-VIII
each with | subdorsal pair; occasionally
present on segment IV.
Type material: Holotype pupal case, 11
pupal cases and 2 third instar larvae para-
types on slide labeled: Colombia, Anthur-
jum sp., 31-VII-78, F. D. Matthews at
Miami (78-7710). Other Paratypes: 6 pupal
cases with same data as holotype slide. CO-
LOMBIA: 7 pupal cases, Anthurium sp., 22-
II-72, J. C. Buff, at Miami (72-5009); 5 pu-
pal cases, Anthurium sp., 13-VII-73, E. B.
Lee, at Miami (73-13786): 9 pupal cases,
Anthurium sp., 20-III-79, G. Stone, at
Miami (79-2655): 5 pupal cases, 5 third in-
star larvae, Anthurium, | 3-XII-76, Froster,
at Miami (77-967); 10 pupal cases (2 slides),
Anthurium sp., 28-III-84, J. Russo, at Miami
(84-3835): Bitaco, 3 pupal cases, Anthurium
sp., 23-VII-58, F. T. Kenworthy (58-13126);
Sasaima, 6 pupal cases, 10-VIII-72, F. Mos-
quiera Paris (72-18229). BARBADOS: 12
pupal cases, Anthurium sp., 8-H-80, L.
Schroeder, at JFKIA (JFKIA 32840).
DOMINICA: 7 pupal cases, Anthurium sp.,
19-VI-81, E. B. Lee, at Miami (Miami
28783). GUADELOUPE: Duclos, 4 pupal
cases, Anthurium palmarum, 20-III-87, J.
Etienne (GR627). HAWAII: Aiea (Oahu),
23 third instar larvae (2 slides), Anthurium
sp., 29-VIII-78, M. Rabago (78-8071); 6 pu-
pal cases, 2 third instar larvae, 5-[X-78, L.
Nakahara. MARTINIQUE: 11 pupal cases
(2 slides), Anthurium, 8-III-71, J. C. Buff,
at Miami (71-15065); 8 pupal cases (2 slides),
Anthurium andreanum, \13-VIU-78, R. Sil-
vestre de Sacy; Morne-Rouge, 6 pupal cases,
Anthurium andreanum, 16-V-81, L. Paley;
St. Joseph, 26 pupal cases (5 slides), 4An-
thurium andreanum, 16-VI-81, L. Paley:
440
TRINIDAD: 9 pupal cases, Anthurium sp.,
10-IV-78, E. B. Lee, at Miami (Miami
17802). Holotype and paratypes in the al-
eyrodid collection of the U.S. National Mu-
seum of Natural History located at Belts-
ville, Maryland. Depositories of other
paratypes: British Museum (Natural His-
tory), London; California Dept. of Food and
Agriculture, Sacramento; Florida State Col-
lection of Arthropods, Gainesville; and Ha-
wail Dept. of Agriculture, Honolulu.
Etymology: The specific epithet is a com-
bination of the host plant, Anthurium, and
latin “cola” which means inhabitant. This
species is known only from Anthurium.
Comments: Of the four species currently
assigned to the genus A/eurotulus, only the
type-species, nephrolepidis (Quaintance
1900), which infests ferns in greenhouses 1n
several countries, is well documented. The
other three species (arundinacea Singh 1931,
maculata Singh 1931, and mundururu Bon-
dar 1923) are known only from their orig-
inal collections. Aleurotulus anthuricola has
well developed cephalic, metathoracic and
eighth abdominal setae, and grayish black
pupal cases; the corresponding setae of ne-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
phrolepidis are short and indistinct and the
pupal cases are pale.
ACKNOWLEDGMENTS
I thank A. H. Hara, University of Hawaii,
at Hilo, for the insitu photographs, and to
the following colleagues for their reviews of
the manuscript and constructive comments;
D. M. Anderson, Systematic Entomology
Laboratory, Washington, D.C., L. M. Rus-
sell (cooperative scientist) and M. B. Stoet-
zel, same laboratory, Beltsville, Maryland;
A. B. Hamon, Florida Dept. of Agriculture
and Consumer Services, Gainesville; and R.
J. Gill, California Dept. of Food and Ag-
riculture, Sacramento. Figures 1-5 were il-
lustrated by L. H. Lawrence, Systematic
Entomology Laboratory.
LITERATURE CITED
Bondar, G.
Bahia.
Quaintance, A. L. 1900. Contribution towards a
monograph of the American Aleurodidae.
U.S.D.A. Bur. Entomol. Tech. Ser. 8: 9-64.
Singh, K. 1931. A contribution towards our knowl-
edge of the Aleyrodidae (Whiteflies) of India. Mem.
Dept. Agric. India 12: 1-98.
1923. Aleyrodideos do Brazil. 183 pp.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 441-451
SLIFERIA, A NEW OVOVIVIPAROUS COCKROACH GENUS
(BLATTELLIDAE) AND THE EVOLUTION OF OVOVIVIPARITY
IN BLATTARIA (DICTYOPTERA)
Louis M. RotH
Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts
02138, U.S.A. (Correspondence: Box 540, Sherborn, Massachusetts 01770, U.S.A.)
Abstract. — Sliferia, a new ovoviviparous blattellid cockroach genus from the Seychelle
Islands is described. It includes S/iferia lineaticollis (Bolivar) n. comb., and Siliferia sp.
False ovoviviparity occurs in | genus of Pseudophyllodromiinae, | genus of Blattellinae
(Blattellidae), and practically all subfamilies and genera of Blaberidae. True ovoviviparity
is found in 2 blaberid genera of Panesthiinae (Geoscapheini). False viviparity is known
only in | species of Blaberidae (Diplopterinae). The evolution of ovoviviparity in the
Blattaria is discussed.
Key Words:
roaches
While studying some of Bolivar’s types
of cockroaches from the Seychelles, I found
that Blattella lineaticollis Bolivar is ovovi-
viparous and belongs in a new genus which
I describe here. The specimens were bor-
rowed from the British Museum (Natural
History) (BMNH).
Sliferia Roth, New GENus
Type species by selection: Blattella linea-
ticollis Bolivar.
Etymology.—The name honors the late
Dr. Eleanor H. Slifer (McIver 1987: 195),
insect morphologist and colleague.
Description (2 only; ¢ unknown).— Head
triangular (Fig. 1), strongly flattened (Fig.
4). Tegmina and wings fully developed.
Tegmina with discoidal vein unbranched,
cubitus and median veins and their branch-
es diagonal. Hind wing with discoidal and
median veins simple, cubitus vein straight
with complete branches, incomplete
Sliferia n. gen., Blattellidae : Pseudophyllodromiinae, ovoviviparity, cock-
branches absent, apical triangle small (Fig.
5). Anteroventral margin of front femur with
a row of piliform spinules terminating 1n 2
large spines (Type C;); pulvilli present on 4
proximal tarsomeres, tarsal claws strongly
asymmetrical, unspecialized, arolia present.
Ovoviviparous; ootheca, not rotated, is re-
tracted with eggs in original vertical posi-
tion.
Comments.— Unfortunately, the male is
unknown but needed to complete the ge-
neric description. The females are similar
to Balta in front femur armament, tarsal
claws, and wing venation. But, they can be
distinguished from that genus by the strong-
ly flattened triangular head, reminiscent of
the Australian genus Mediastinia Hebard
(1943: 9).
Only two species are known, the type
species and a taxon I did not name because
it differed principally only in color from /in-
eaticollis. Both are described below.
442
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1.0mm
Figs. 1-5.
1.0mm
SRA EAD BT Ee
2.0mm
Sliferia lineaticollis (Bolivar). 1-4. Female holotype. 1, Head. 2, Pronotum. 3, Subgenital plate
and protruding ootheca (ventral view). 4, Head and part of pronotum (lateral view). 5, Paratype, sex unknown,
hind wing.
Sliferia lineaticollis (Bolivar),
New ComMBINATION
Blattella lineaticollis Bolivar, 1924: 327;
Chopard, 1951: 143; Scott, 1933: 340;
Princis, 1969: 1007 (sp. incertae sedis);
Roth, 1985: 211 (sp. incertae sedis).
Material examined.— Holotype: Female
(carrying an ootheca internally), Mahé,
1908-9 Seychelles Exp., Percy Sladen Trust
Exped. Brit. Mus., 1926-203 (BMNH).
Paratype.—1 (terminal abdominal seg-
ments missing), same data as holotype
(BMNH).
Male. — Unknown.
Female.—Interocular space about same
as distance between antennal sockets (Fig.
1). Palpal segments 4 and 5 about equal,
each longer than segment 3. Tegmina and
wings extending beyond end of abdomen.
Hind wing with costal veins weakly thick-
ened distad; cubitus vein with 3 complete
VOLUME 91, NUMBER 3
and no incomplete branches, apical triangle
small (Fig. 5). Supraanal plate trigonal, apex
weakly indented.
Measurements (mm).—Length, 9.3;
pronotum length x width, 2.4 x 3.5; teg-
men length, 9.5.
Coloration.—Head with occiput pale, a
black transverse band on vertex then a whit-
ish hyaline area and a broad, uneven, brown
transverse band between antennal sockets,
remainder yellowish brown (Fig. 1). Pro-
notal disk yellowish brown with narrow dark
brown longitudinal lines demarcating 1t lat-
erally, broad lateral regions hyaline (Fig. 2).
Tegmina yellowish brown, hyaline, me-
diastine vein dark brown. Abdominal terga
light yellowish brown with dark infuscation
laterally on distal segments; terminal seg-
ments blackish brown; supraanal plate
blackish with a dirty white transverse stripe
basally, a similarly colored macula medially
near apex. Abdominal sterna with yellowish
brown lateral borders, remainder mottled
brownish. Cerci dark brown on proximal
segments becoming paler distad.
Ootheca.—The holotype 1s carrying a ful-
ly formed egg case in the upright position
(keel region dorsad) inside the abdomen,
with a small part of the ootheca protruding
beyond the end of the abdomen (Fig. 3). The
exposed keel region 1s flat and lacks raised
respiratory serrations. Those eggs seen
through the ventral hyaline regions of the
oothecal membrane are apparently unde-
veloped.
Comments.—Scott (1933: 340) and Cho-
pard (1951: 143) found this species in the
axils of leaves of screw pines (Pandanus sp.)
and the palm Verschaffeltia splendida.
The holotype was collected probably while
she was ovipositing and had not yet retract-
ed the ootheca completely into the brood
sac. Or, she may have completed oviposi-
tion but partly extruded the egg case when
she was killed. Several years ago (Roth 1985:
211) I noted that this female was carrying
an ootheca but I incorrectly stated that it
443
was rotated and I failed to point out that it
was carried internally.
Sliferia sp.
Figs. 6-8
Material examined.—Seychelles, La Di-
gue, | 2 (carrying ootheca internally),
711.1953, E. S. Brown, presented by
Comm. Inst. Ent. B.M. 1954-368, genitalia
slide 264 (BMNH).
Female.—Head exposed, strongly flat-
tened; palpal segments 3 and 4 equal, each
slightly larger than segment 5; interocular
space less than distance between antennal
sockets. Tegmina and wings fully developed
(glued together), extending beyond end of
abdomen. Front femur Type C,; pulvilli
present on 4 proximal tarsomeres, tarsal
claws strongly asymmetrical, unspecialized,
arolia present. Supraanal plate trigonal, apex
rounded (Fig. 6).
Measurements (mm).—Length, 9.9;
pronotum length x width, 2.3 x 3.3; teg-
men length, 9.2.
Coloration.—Head with occiput pale,
hyaline; edge angular where vertex leads into
flattened face, with a narrow, transverse
brown line, then a colorless hyaline area and
a small, poorly defined, light brown macula,
rest of face pale. Pronotal disk yellowish,
without markings, broad lateral regions
hyaline. Tegmina almost colorless hyaline.
Abdominal terga yellowish without dark
markings. Abdominal sterna pale, in part
hyaline. Cerci with ventral surface darker
than dorsal, inner lateral edges of segments
narrowly infuscated. Legs pale.
Ootheca.—The female was carrying an
ootheca that was partially collapsed and
protruding from the end of the abdomen.
Most of the egg case could be seen through
the hyaline parts of the abdominal sterna
and it occupied a large part of the abdominal
cavity (Figs. 6, 7). The ootheca was oriented
with its keel region dorsad and was not ro-
tated prior to being retracted into the brood
sac. I placed the abdomen in 10% KOH for
444
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
10mm
1.0mm
CF ELL LP?
ZH
~T.0mm _
Figs. 6-8. Sliferia sp., pregnant female from La Digue, Seychelles. 6, Abdomen (dorsal view). 7, Abdomen
(ventral view; most of the internal ootheca is visible through the abdominal sterna). 8. Ootheca removed from
abdomen and treated with KOH (dorsal view).
several minutes and the egg case floated out
of the abdomen and swelled (Fig. 8). It was
4.5 mm long, wider than high, and con-
tained 25 eggs, some of which showed early
embryonic development. The oothecal
membrane was thin, flexible, and complete-
ly covered the eggs. There were neither ser-
rations (respiratory tubules) in the keel re-
gion, nor calcium oxalate crystals.
Evolution of ovoviviparity and
viviparity in the Blattaria
In the two phyletic lines of Blattaria, ovo-
viviparity and viviparity arose in the su-
perfamily Blaberoidea but not in the Blat-
toidea. It was originally believed that
Ovoviviparity was found only in the Bla-
beridae (McKittrick 1964, Roth 1970). Re-
cently it was discovered in an African genus
Stayella Roth (Roth 1982a, 1984) (Blattel-
lidae: Blattellinae), and now in Sliferia
(Blattellidae: Pseudophyllodromiinae of
Vickery and Kevan 1983: 157 = Plectop-
terinae of McKittrick 1964). Internal in-
cubation of eggs by Blattaria evolved from
Oviparous ancestors (Fig. 9) and required
changes in physiology, morphology, and
oviposition behavior. The families and
subfamilies shown in Fig. 9 are principally
those of McKittrick (1964) except for the
Nocticolidae and Attaphilinae which she did
not study. I (Roth 1988: 300) discussed the
VOLUME 91, NUMBER 3
BLABEROIDEA
Panesthiinae TRUE OVOVIVIPARITY
BASE RIDAET
Many subfamilies
FALSE OVOVIVIPARITY
¥
OOTHECA RETRACT
74 CTED
Blattellinae
Ectoblinae
Nyctiborinae
FALSE OVOVIVIPARITY
7 4
BLATTELLIDAE
ae OOTHECA RETRACTED
Pseudophyllodromiinae
Anaplectinae
Attaphilinae
NOCTICOLIDAE
PRIMITIVE ROTATION
(NR)
POLYPHAGIDAE
OOTHECA
OOTHECA ABSENT
OOTHECA CARRIED
EXTERNALLY
OOTHECA CARRIED
EXTERNALLY
445
FALSE VIVIPARITY
Diplopterinae
~£OOTHECA RETRACTED
~ ADVANCED ROTATION
BLATTOIDEA
BLATTIDAE
CRYPTOCERCIDAE
DROPPED EARLY (DE)
NO ROTATION (NR)
OOTHECA FORMED
@®
OVIPARITY
EXTINCT ANCESTORS
Fig. 9.
taxonomic position of the former family and
McKittrick (77 Roth 1968c: 135) placed the
Attaphilinae in the Blattellidae.
Extinct ancestors.— Vishniakova (1968)
summarized the literature on fossil cock-
roaches. The oviparous females had exter-
nal ovipositors and have been found in the
Upper Carboniferous and Permian deposits
of Europe, Asia, and North America, and
Possible evolution of ovoviviparity and viviparity in the Blattaria.
in the Triassic and Jurassic deposits of the
USSR. Presumably the species laid eggs sin-
gly in plants or soil. With time, the ovipos-
itor was gradually reduced in length. Even-
tually a structure evolved hidden within the
vestibulum that was used to form an oothe-
ca containing a group of eggs. In some Ju-
rassic species, the ovipositor is very short
and the transition from long external ovi-
446
positors to short internal ones apparently
took place towards the end of the Mesozoic.
Oothecae have been described from the
Carboniferous, but the fossils only remotely
resemble an egg case (Roth 1967b: Fig. 16),
and most likely are imprints or fragments
of other organisms (Brown 1957, Vishni-
akova 1968). An ootheca from the Eocene
(Brown 1957, Roth 1967b: Fig. 15) appears
to be an authentic blattellid egg case.
McKittrick (1964) suggested that the prim-
itive cockroach ootheca was probably a
packet of eggs glued together like the oothe-
ca of the termite Mastotermes darwiniensis
Froggart (Roth 1967b: Fig. 17). Except for
two genera (possibly four) of ovoviviparous
Panesthiinae, cockroaches that have been
studied produce an ootheca.
Selection pressure.—According to Vish-
niakova (1968) the selection pressure for the
reduction of the external ovipositor and the
change from oviparity to internal incuba-
tion of the eggs was the appearance in the
Carboniferous to the Jurassic of predators
and parasites that destroyed eggs. A number
of Hymenoptera are known that destroy
cockroach eggs (Roth and Willis 1960: 234—
255). Some wasps can oviposit through a
deposited and hardened ootheca. For ex-
ample Tetrastichus hagenowii (Ratzeburg)
lays eggs in oothecae of Periplaneta amer-
icana (Roth and Willis 1954b: Plate I). Oth-
er wasps can Oviposit into the soft ootheca
before the wall hardens. For example, Evan-
ia appendigaster (Linn.) lays eggs in the
ootheca of P. americana (Kieffer 1912):
Zeuxevania splendidula Costa into the
oothecae of Loboptera decipiens (Germar)
(Genieys 1924). Anastatus floridanus Roth
and Willis oviposits through the soft walls
of the egg case of Eurycotis floridana (Walk-
er) while the ootheca is still carried by the
female and before it hardens and is depos-
ited (Roth and Willis 1954c: Fig. 9); the
female often buries the ootheca in sand and
once this is done the wasp usually can’t lo-
cate it for oviposition. Oothecae still at-
tached to females of Blattella dethieri Roth
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and Blattella armata (Princis) have con-
tained a single evaniid wasp (Roth 1985:
Figs. 92C, 102D, 102E). Schletterer (1889)
found the evaniid Brachygaster minutus
(Olivier) in the ootheca of Blattella ger-
manica (Linn.). Wasps that parasitize the
eggs of Blattella oviposit into the ootheca
while it is carried by the female. The cock-
roach continues to carry the egg case while
the eggs are being eaten by the parasitoid.
In classifying the placement of parasitic
wasp eggs in relation to host species, Clau-
sen (1972) erected the category: Egg placed
in the embryo while the latter is still within
the parent. Although this behavior was not
known to occur, Clausen believed that it
probably could occur. Evania appendigaster
was claimed to parasitize the eggs of the
blaberid Leucophaea maderae (Fab.)
(Schletterer 1889, Bordage 1896, Kieffer
1912). But after realizing that Leucophaea
is OVOviviparous, Bordage (1913) admitted
misidentifying the parasitized ootheca and
concluded that the developing eggs of Leu-
cophaea are protected against egg parasites
because they are carried within the female.
Sells (1842) reported that an ootheca of Leu-
cophaea maderae (= Blaberus maderae)
contained ninety six specimens of a small
chalcid wasp; some oothecae had a round
hole through the side of the capsule from
which the wasps had emerged. Sells state-
ment that the ootheca had a keel with 16
dentations indicates that the parasitized
oothecae probably were Periplaneta amer-
icana (Roth and Willis 1954b: Plate III).
It is true that eggs of ovoviviparous cock-
roaches have not been found to be parasit-
ized by wasps. The oviposition of some
wasps into an ootheca while it is still carried
by the female is evidence that wasps that
destroy the eggs of ovoviviparous species
probably will be found. I suspect that wasp
parasitoids will be found in species of Stay-
ella with oothecae similar to those of Blat-
tella but incubated internally. The Stayella
ootheca could be parasitized as it is being
formed and before it is retracted into the
VOLUME 91, NUMBER 3
brood sac. If the female cockroach extruded
the parasitized ootheca when the wasp ma-
tured, the adult parasite could easily escape
the host abdomen. Similarly, the eggs of
ovoviviparous Blaberidae could be parasit-
ized while the ootheca was being formed
before it was retracted into the uterus. Pro-
tection from wasp egg parasitoids could
probably be assured if the eggs were never
exposed to the outside, and this occurs only
in true Ovoviviparous genera Macropanes-
thia and Geoscapheus.
Laurentiaux (1951) suggested that the
ootheca appeared as an adaptive response
to climatic and hygrometric changes. The
oothecal membrane in the Blattoidea 1s ca-
pable of preventing desiccation of the eggs
even when exposed to very low humidities.
In the Blattellidae, the protective covering
varies in its ability to prevent water loss,
and in the Blaberidae, the greatly reduced
covering does not prevent the eggs from los-
ing water even in very high humidities.
These studies in water loss (Roth and Willis
1955b, Roth 1967b) indicate that an im-
portant selection pressure for the evolution
of internal incubation was the prevention
of egg death from water loss.
Ootheca formation.— Roth (1974) found
that there has been a divergence of the cen-
ters that control oviposition in the Blaber-
oidea and Blattoidea. The brain 1s not need-
ed for oviposition in the Blaberidae where
the control center for formation, 90° rota-
tion, and retraction of the ootheca into the
brood sac probably lies in the last abdom-
inal ganglion. In the Blattidae the brain is
needed to initiate egg case formation, but it
is unnecessary once the process has begun.
During oviposition, the female extrudes
some colleterial gland secretion as a soft
rubbery blob in a mold formed by the in-
tersternal membrane of the vestibulum. As
the eggs leave the gonopore they are guided
by the ovipositor valves into the blob of
secretion where, with micropylar ends dor-
sad, they line up, alternating sides, and pro-
trude from the end of the abdomen as a
447
double row (Roth and Willis 1954a: Pl. 5).
In oviparous species the newly formed
ootheca is pale and soft but hardens quickly
when exposed to air. The dorsal ovipositor
valves serve as a mold and impart charac-
teristic shapes to the keel. Each egg lies in
a cell surrounded by an air space connected
to a duct in the keel that leads to external
air. Oothecae of different species differ in
size, shape, surface texture, keel serration,
and the number of contained eggs (Roth,
1968b, 1971). Ovoviviparous species of
Blaberidae and species of Blattellidae that
carry their oothecae externally during em-
bryogenesis generally have a larger number
of eggs per ootheca but produce fewer oo-
thecae than oviparous species that drop their
egg cases shortly after forming them (Roth
1970: fig. 21).
The paired ovaries of cockroaches consist
of a variable number of ovarioles. Each
ovariole consists of a variable number of
oocytes and at oviposition supplies one yolk-
filled egg in the ootheca. Those oocytes con-
taining yolk at oviposition are found in a
region known as Zone V. Both the Blatto-
idea and Blaberoidea probably evolved from
taxa in which three or more oocytes con-
tained yolk in Zone V. In the Blaberoidea
there 1s an evolutionary trend towards a de-
crease in the number of oocytes in Zone V,
as well as the total number of oocytes per
ovariole. The Polyphagidae have two or
three oocytes in Zone V, and a large number
of oocytes per ovariole. Species of Blattel-
lidae have one or two oocytes in Zone V,
and show some reduction in total number
of oocytes. In ovoviviparous and vivipa-
rous Blaberidae there is a marked reduction
in the number of oocytes, and only one 0o-
cyte develops yolk at oviposition. Species
like Blattella that carry their oothecae ex-
ternally during embryogenesis have only one
oocyte in Zone V and few oocytes per ovar-
iole as in most Blaberidae (Roth 1968c).
During pregnancy in ovoviviparous and
viviparous taxa, the corpora allata are in-
hibited and yolk deposition does not occur
448
in the oocytes as long as there is an ootheca
in the uterus. Yolk inhibition also occurs in
pregnant Blattella. In ovoviviparous taxa
the colleterial glands show cyclical activity
corresponding to the ovarian cycle: active
during the preoviposition period, inactive
during gestation, and active again after par-
turition (Roth 1968b). In oviparous species
that drop their oothecae early, the colleterial
glands secrete protein continuously, there is
no distinct cycle related to oviposition, and
during the height of their reproductive pe-
riod females produce oothecae frequently.
In oviparous species, a hardened ootheca
is carried externally for various lengths of
time. It can be deposited shortly after its
formation or retained for the full gestation
period (Roth and Willis 1958: figs. 1-12,
30-37). When first deposited the eggs may
have sufficient water and yolk to complete
development (Blattidae), or they have suf-
ficient yolk but require water which is ob-
tained from the female (Blatte/la spp., Lo-
phoblatta spp.) or from the substrate (most
Blaberoidea, except Blaberidae) (Roth
1967b, 1968a, Roth and Willis 1955b).
Once the ootheca is formed, the female
may oviposit in one of the following ways:
No rotation, ootheca dropped early
(NR).—The original vertical position of the
ootheca is not changed and the egg case 1s
deposited shortly after its formation (Roth
and Willis 1954a: Pl. 5). The female may
try to hide the dropped egg case by burying
it or covering it with a mixture of saliva and
chewed substrate, and then she abdandons
it (McKittrick 1964). Examples are Nocti-
colidae (Chopard 1932: 489), some species
of Polyphagidae, Blattellidae (Attaphilinae,
Anaplectinae, many genera of Pseudophyl-
lodromiinae), all species of Blattidae and
Cryptocercidae (Roth 1968b).
Ootheca carried externally during gesta-
tion.—These oviparous species carry their
oothecae attached to the end of the abdo-
men until the eggs hatch. The ootheca can
be held in the original vertical position as
in Lophoblatta (Roth 1968a) or rotated 90°
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
as in Blattella (Roth and Willis 1958: PI.
25, figs. 7-12, Pl. 27, figs. 29-37).
Primitive rotation.—The sotheca is ro-
tated 90° and deposited shortly after its for-
mation. The egg case has a flange which is
held by the paraprocts and none of the an-
terior eggs are held within the vestibulum
in contact with the tissues of the female
(Roth 1970: fig. 5). Primitive rotation is
known only in some Polyphagidae, includ-
ing Arenivaga (Arenivaga) spp. (Roth 1967a)
and Latindia sp. (Roth 1971: 128).
Advanced rotation. — After the ootheca is
formed and still attached, it is rotated 90°
so the keel is lateral, to the left in Blaberidae,
and the right in most Blattellidae. Several
of the most anterior eggs, those leaving the
oviduct last, are in contact with the vestibu-
lar tissues of the female. This contact is nec-
essary for oviparous species like Blattella
spp. and Lophoblatta spp. whose eggs ob-
tain water from the mother during embryo-
genesis (Roth 1967a). Examples are Nycti-
borinae, Ectobiinae, most Blattellinae and
Pseuphyllodromuinae, and all Blaberidae
that form an ootheca.
Oothecal changes. — During the evolution
of ovoviviparity in the Blattellidae, changes
in the ootheca involved reduction in height
of the keel and finally loss of the keel, and
softening of the increasingly flexible protec-
tive membrane due to the gradual reduction
and finally loss of calcium oxalate crystals.
The oothecal changes in the Blaberidae were
greater than those that took place in the
Blattellidae (Blattella, Stayella, Sliferia). In
some blaberids the oothecal wall is rela-
tively thick and covers the eggs completely.
In others, the wall shows various degrees of
reduction and does not cover the micro-
pylar ends of the eggs. Although the keel is
absent in most blaberids, relicts of this
structure occur in Epilampra cribrosa Bur-
meister (Roth 1968b), for example.
Ootheca retracted.— After the ootheca is
formed, it is retracted into the brood sac
with or without being first rotated 90° (Roth
and Willis 1954a: figs. 74-79, 86-90).
VOLUME 91, NUMBER 3
McKittrick (1964: 116) believed that rota-
tion of the ootheca freed the keel from the
Ovipositor valve bases, which supposedly
block anterior movement of the vertical
ootheca. But, as noted above, in the Bla-
beridae the ootheca is reduced, keel serra-
tions are absent or rarely subobsolete so ro-
tation is not necessary to free its keel from
the valve bases (Roth 1967a). In the blat-
tellid S/iferia the ootheca lacks keel serra-
tions and is retracted in the vertical posi-
tion.
In the ovoviviparous Blaberidae, the eggs
enlarge in the brood sac during embryogen-
esis (Roth and Willis: 1955b, fig. 7A—D).
When stretched by a newly deposited oothe-
ca, the blaberid uterus presses against the
inner surfaces of the terga and sterna and
leaves room laterally for expansion (Roth
1967a: 85). Just before parturition the oo-
theca occupies practically the entire abdo-
men (Roth 1967a: 85). But even at this time
it is difficult to determine if the female is
pregnant because the abdomen hasn’t
stretched as much as it would have had the
ootheca been retracted into the uterus in the
vertical position. To determine pregnancy
one has to separate the supraanal and
subgenital plates and look for the terminal
end of the ootheca. Cockroaches generally
are relatively flat insects and tend to hide
in narrow crevices and spaces where they
can escape from predators. It 1s advanta-
geous for the eggs to be so oriented in the
uterus that their increase in size does not
markedly increase the height of the abdo-
men. In Blattella rotating the ootheca al-
lows the insect to hide in narrower crevices
than would be possible if the ootheca was
carried externally in the vertical position.
Wille (1920) measured the minimum gaps
within which various stages of B. germanica
could move. Gravid females | day before
oviposition needed a space of 4.5 mm; with
the ootheca in the vertical position, the width
decreased to 3.3 mm, and after the egg case
was rotated the female could move ina space
2.9 mm high. In S/iferia, the eggs probably
449
do not increase much in length during em-
bryogenesis and because its ootheca is wider
than high rather than higher than wide, ro-
tation is not necessarily advantageous and
the ootheca is retracted in the vertical po-
sition.
False ovoviviparity.—The ootheca 1s
formed and retracted into the brood sac.
Initially the eggs have enough yolk to com-
plete development but must obtain water
from the female during embryogenesis. The
marked similarity of the oothecae of Stay-
ella and Blattella (Blattellinae), and Sliferia
and Lophoblatta (Pseudophyllodromiinae),
strongly suggests that the ancestors of the
two ovoviviparous genera were Blattella-
like and Lophoblatta-like species in their
respective subfamilies. Cockroaches with
false ovoviviparity include two genera in
the Blattellidae, and all Blaberidae except
two truly ovoviviparous genera, and one vi-
viparous species.
True ovoviviparity.—No ootheca is
formed. During oviposition the eggs are
never exposed to the outside but pass di-
rectly into the vestibulum and brood sac
where they lie in a jumbled mass. At ovi-
position the eggs have enough yolk to com-
plete development, and take up only water
from the mother during embryogenesis
(Rugg and Rose 1984a). Examples include
Macropanesthia rhinoceros Saussure, Geo-
scapheus dilatatus (Saussure), and probably
species of Neogeoscapheus and Parapanes-
thia. All four of these genera were originally
placed in the Panesthiinae, tribe Geoscaph-
eini (Roth 1982b), but because of the ab-
sence of an ootheca Rugg and Rose (1984b)
placed them in the family Geoscapheinae.
Roth (1968b: 103) speculated that com-
plete loss of the oothecal membrane would
result in eggs being simply glued together in
a double row, presumably like the ootheca
of Mastotermes darwiniensis, and then could
be retracted into the brood sac. Rugg and
Rose’s discovery shows that some species
can deposit their eggs in the uterus without
first extruding them in a double row.
450
False viviparity.—The ootheca is very
small, containing about a dozen eggs, and
has an incomplete covering membrane
(Roth and Hahn 1964). Because of their
small size, hardly any of the eggs are seen
during oviposition, but the ootheca is ro-
tated and retracted into the brood sac. Ini-
tially the eggs lack sufficient yolk and water
to complete development (Roth and Willis
1955a). During embryogenesis the embryos
drink water and dissolved proteins and car-
bohydrates, synthesized and transported by
the brood sac (Stay and Coop 1973, 1974,
Ingram et al. 1977). Diploptera punctata
(Eschscholtz) 1s the only known example,
although other species in the genus probably
have the same type of reproduction.
True viviparity.—No example in cock-
roaches is known. In true viviparity, the
eggs would pass directly into the brood sac
without first being exposed externally, and
they would obtain solids and water from the
female during embryogenesis.
The discovery that ovoviviparity occurs
in two subfamilies of Blattellidae rather than
only in the Blaberidae as proposed by
McKittrick (1964) does not contradict her
basic thesis that ovoviviparity arose only in
the Blaberoidea and not in the Blattoidea.
In this regard there is no reason to revise
MckKittrick’s system.
ACKNOWLEDGMENTS
I thank Mrs. Judith Marshall for the loan
of the specimens from the British Museum
(Natural History), the Bureau of Flora and
Fauna, Australian Biological Resources
Study for partial support, and Dr. Frank
Carpenter, Harvard University, for a trans-
lation of Vishniakova’s Russian paper.
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PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 452-457
A NEW SPECIES OF DASYHELEA MIDGE REARED FROM
DRAINS IN SINGAPORE (DIPTERA: CERATOPOGONIDAE)
K. M. Lee, W. W. WIRTH, AND K. L. CHAN
(KML, KLC) Department of Zoology, National University of Singapore, Lower Kent
Ridge Road, Singapore 0511; (WWW) Systematic Entomology Laboratory, USDA, and
Florida State Collection of Arthorpods, 1304 NW 94th St., Gainesville, Florida 32606.
Abstract.—Male and female adults, pupae and larvae of Dasyhelea schizothrixi new
species are described from material collected from free-flowing open-channel concrete
drains in Singapore. Diagnostic characters are given to distinguish D. perfida, D. laeta,
and D. bullocki from the new species.
Key Words:
Biting midges of the genus Dasyhelea Kief-
fer are common and widespread in all re-
gions of the world. The aquatic or semi-
aquatic larvae are found in various habitats
that have at least a thin film of water in
which the larvae can survive. The larvae
are poor swimmers and slowly climb or push
through the substratum by using their
mouthparts and posterior anal hooks. Their
preferred habitats include wet moss or algae
along the shores of streams, lakes, ponds,
puddles and other water bodies, or wet rot-
ting plant materials such as sap from trees,
wet bark, and tree holes. Some species in-
habit rock pools, hot springs, or water with
high mineral content. Others have colo-
nized the intertidal zone along algae-cov-
ered rocks or algae growing on mud exposed
to tidal action in salt marshes.
Adult Dasyhelea midges are usually found
near breeding sites. Shrubs, plants near
water, and flowers are favorite resting places
of some species. Their feeding habits are
little known but some have been found feed-
ing on nectar from flowers and extrafloral
nectaries, honeydew, and sweet secretions
from plants (Downes and Wirth 1981). None
Dasyhelea schizothrixi, open concrete drains, Schizothrix algae
are known to be bloodsuckers. Some species
are important pollinators of Theobroma ca-
cao, Hevea brasiliensis, and other tropical
tree crops.
This is the first record of a Dasyhelea from
free-flowing open-channel concrete drains
in a city. The drainage system was imple-
mented for the control of Anopheline mos-
quitoes through the removal of all surface
water larval habitats. The midge larvae were
found living in Schizothrix algal beds grow-
ing on these drains.
Eggs, larvae, and pupae were collected
from the drains by removing a portion of
the Schizothrix algal bed with a blunt scal-
pel. Larvae and pupae were killed in hot
water and preserved in 70 percent ethanol.
Egg masses were allowed to hatch and the
larvae were individually reared through to
adulthood in the laboratory. Twenty-four
hours after emergence, the adults were killed
by immersion in 70 percent ethanol, and
transferred to glass microvials with the pu-
pal exuviae for storage in 70 percent etha-
nol. Adults were dissected and mounted on
slides according to the methods of Wirth
and Marston (1968) and Lee and Chan
VOLUME 91, NUMBER 3
(1985). Terminology follows that given by
Waugh and Wirth (1976) and Downes and
Wirth (1981).
Dasyhelea schizothrixi Lee and Wirth,
New SPECIES
Figs. 1-13
Male holotype.— Wing length 1.06 mm;
breadth 0.37 mm; costal ratio 0.50.
Head.—Brownish, proboscis and palpi
stramineous. Eyes with fine pubescence.
Antenna (Fig. 1) with lengths of flagellar
segments in proportion of 40-30-30-28-27-
25-25-25-25-52-55-50-82, antennal ratio
(segments 12-15/3-11) 0.98; proximal seg-
ments not fused; with dense long brown
plume setae; segment 12 binodose, 13
slightly so, 15 with long terminal papilla;
all segments sculptured. Palpus (Fig. 2)
moderately stout, with 14 club-shaped sen-
silla; lengths of segments in proportion of
28-38-23-25:; palpal ratio 1.5.
Thorax.—Brownish; two pale longitudi-
nal lines following the well-defined double
series of dorsocentral setae; double line of
acrostichal setae well defined, lateral setae
scattered. Scutellum (Fig. 3) with eight large
marginal setae and five smaller ones on disc.
Legs stramineous, knee spots blackish; fem-
ora with narrow, faintly brownish bands just
past midlength, fore tibia with a faint brown
band at midlength and narrow brown apex;
sparse vestiture of slender setae; tibiae with
bristly setae; fifth tarsomeres brownish.
Wing membrane whitish, radial veins
slightly infuscated; first radial cell not
formed, second radial cell faintly indicated
by obscure thickening of veins, apex of cell
truncated; macrotrichia long and slender,
decumbent, moderately sparse, forming
rows along veins and on membrane on
proximal 0.7 of wing; bare linees between
rows of macrotrichia. Halter deeply infus-
cated.
Abdomen.—Brown without segmental
bands; sparse vestiture of long slender brown
setae in transverse rows. Genitalia (Fig. 4):
Ninth sternum slightly broader and slightly
453
bilobate in midportion; ninth tergum slight-
ly broader than long, markedly tapering dis-
tally to a pair of well-developed apicolateral
processes. Gonocoxite slightly bulbous,
stout, about 1.5 times longer than broad,
without mesal hooklike process; gonostylus
slightly longer than gonocoxite, stout prox-
imad, tapering at midportion to slender,
slightly arcuate, blunt-tipped distal portion.
Aedeagus (Fig. 5) dark brown, slightly
broader at base than total length, bearing a
narrow bisinuate transverse sclerite proxi-
mally and a pair of elongate, pillar-like sub-
median processes caudally with bases nar-
rowly joined and with a transparent
membrane between except at extreme apex.
Parameres (Fig. 6) heavily sclerotized,
brownish, asymmetrical; basal apodemes
unusually slender, the posterior process also
slender in its entire length, slightly curved,
extending caudad nearly to tip of ninth ter-
gum.
Female allotype.— Wing length 1.00 mm;
breadth 0.44 mm: costal ratio 0.56.
Similar to male with usual sexual differ-
ences. Antenna (Fig. 7) with lengths of fla-
gellar segments in proportion of 34-30-30-
30-30-30-30-30-32-34-35-35-60, antennal
ratio (11-15/3-10) 0.80; segments ovoid,
slightly attenuated distally; last segment with
long terminal nipple as in male; all segments
with definite sculpturing. Palpus (Fig. 8)
stouter than in male, with 17 club-shaped
sensilla; lengths of segments in proportion
of 16-36-17-24, palpal ratio 1.4. Scutellum
(Fig. 9) with nine large marginal setae and
six smaller ones on disc. Wing with mem-
brane dusky and veins more brownish than
in male, second radial cell with oblique apex;
macrotrichia slightly more numerous than
in male, shorter and stiffer. Abdomen
brownish, especially on pleural membrane
which is dark brown and shagreened; setae
scattered and dense on pleura. Genital scler-
otization (Fig. 10) characteristic of the gri-
sea species group with sinuate caudolateral
arms on each side of gonopore and evenly
tapering anteromedian lobe. Spermatheca
454 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-6. Dasyhelea schizothrixi (male): 1, antenna; 2, palpus; 3, scutellum; 4, genitalia, ventral view; 5,
aedeagus; 6, parameres.
single, pyriform with neck tapering and cephalothorax dark brown; degree of pig-
slightly oblique; measuring 0.072 by 0.055 mentation on abdomen fainter posteriorly.
mm including neck portion. Integument shagreened with minute tuber-
Pupa.—Length 2.06 mm. Dorsum of cles that are coarse and thornlike on ceph-
VOLUME 91, NUMBER 3 455
0.100 mm
11
0.051 mm
Figs. 7-13. Dasyhelea schizothrixt, 7-10, female; 11-12, pupa; 13, larva: 7, antenna; 8, palpus; 9, scutellum;
10, genitalia, ventral view; 11, respiratory horn; 12, operculum; 13, last abdominal segment, ventro-posterior
view.
456
alothorax, particularly on operculum (Fig.
12); setae poorly developed. Respiratory
horn (Fig. 11) 0.30 mm long, transversely
ridged, marginally expanded and slightly
curved distally; outer edge of base with eight
large spiracles; 67 (60-72) small spiracles
form a dorsal loop near margin of respira-
tory horn: dorsal slit 0.16 mm long. Ab-
dominal segments each with a ring of low
scale-like tubercles near posterior margin,
most of these tubercles with a minute trans-
parent seta. Last abdominal segment with
two dorsal and two ventral lobes; dorsal lobe
with large dorsal spine and lateral spine;
ventral lobe stout and blunt in female but
tapering posteriorly in male pupa.
Larva.— Length 4.70 mm in fourth instar.
Head capsule dark brown with two con-
spicuous black pigment spots; body milky
except for slight brown pigmentation on
dorsum. Head and body setae inconspic-
uous, few. A pair of distinct, internal,
brownish pigment spots in each of the first
two thoracic segments. Terminal abdomi-
nal segment (Fig. 13) with two pairs of slen-
der hooks dorsally, four pairs of slender
hooks ventrally; each pair of hooks consists
of one small and one large hook; a band of
slender spicules immediately anterior to
hooks. Rectal gills grouped into two pairs
dorsally and two pairs ventrally.
Distribution. — Singapore.
Types.— Holotype male, allotype female;
Kent Ridge, Singapore, 2.x.1985, K. M. Lee,
reared from concrete drains with Schizo-
thrix algae. Paratypes: 9 males, 7 females,
4 larvae, 10 pupae, same data, but dates
16.v-8.1x.1984. Holotype and allotype
specimens are deposited in the National
Museum of Natural History, Smithsonian
Institution, Washington, D.C. Paratypes are
deposited in the Australian National Col-
lection, Canberra; B. P. Bishop Museum,
Honolulu, Hawau; British Museum (Nat-
ural History), London; Museum National
d’Histoire Naturelle, Paris; and Zoological
Reference Collection, National University
of Singapore, Singapore.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Discussion. —Dasyhelea schizothrixi be-
longs to a large group of closely related
species that characteristically breed in rock
holes or in drains, roof gutters, or similar
habitats. The group includes D. carolinensis
Tokunaga (1941) from the Caroline Islands,
D. hitchcocki Wirth (1976) from the Tonga
Islands and Samoa, D. /aeta (Johannsen)
(1931) from Sumatra, D. perfida (Johann-
sen) (1931) from East Java, D. pseudoinci-
surata Waugh and Wirth (1976) from the
southeastern United States, D. saxicola Ed-
wards (1929) [synonym D. /ithotelmatica
Strenzke (1956)] from Europe, D. sublettei
Wirth (1987) from the southwestern United
States, D. thompsoni de Meillon (1936) from
Transvaal, D. tugelae de Meillon (1936)
from Natal, and D. upsilon de Meillon and
Wirth (1987) from Transvaal.
According to its incomplete original de-
scription, Dasyhelea perfida, described from
water-sprayed leaves of Colocasia indica
near a waterfall, closely resembles D. schi-
zothrixi. Dasyhelea perfida has a yellowish
thorax with brownish vittae; yellow un-
banded legs; body length of 1.2 mm, and a
whitish halter with darker stem. The female
has large brownish spots on the interme-
diate pleura of the abdomen. The male gen-
italia are nearly identical with those of the
new species except that the ninth sternum
is not bilobed and the posterior process of
the parameres is distinctly bent backwards
at the tip. According to Mayer (1934) the
pupal respiratory horn of D. perfida differs
from that of D. schizothrixi in having a loop
of 49 spiracles around the margin of the
distal half (similar in arrangement to many
species of the genus) and eight spiracular
openings on the transversely ridged basal
portion.
Tokunaga and Murachi (1959) apparent-
ly incorrectly redescribed Dasyhelea perfida
from misidentified Micronesian material.
Their species differs from D. perfida in the
male genitalia: the apicolateral processes on
the ninth tergum are blunt; the aedeagus has
four distinct posterior processes, and the
VOLUME 91, NUMBER 3
posterior process of the parameres is broad-
ly expanded distally and has a blunt distal
point.
Dasyhelea laeta, reared from rock pools
in Sumatra, differs from S. schizothrixi in
its larger size (male body 2 mm, female 1.75
mm); in its shining dark brown mesonotum
with four darker brown vittae; in its pale
halteres, and in its brownish legs with under
sides of the femora yellowish and the tips
of the femora with preapical whitish rings.
In D. laeta, the pupal respiratory horn has
coarse scales on most of the midportion, 10
spiracles in a loop at the extreme tip, and
five widely spaced spiracles on the basal
portion.
Dasyhelea bullocki, from a rock hole near
a stream in Korea, belongs to the same group
as the new species, but differs in its more
distally attenuated antennal segments; in its
spermatheca with elongate oblique neck; in
its male genitalia with a stout, tapering, pos-
terior process on the parameres, and in its
single loop of spiracular openings on the
pupal respiratory horn.
Five closely related species that breed in
the pitchers of Nepenthes are more distantly
related to this group of species, differing in
details of the pupal respiratory horn, male
genitalia, and in the more elongate, bent
neck of the spermatheca. This group of
species includes D. ampullariae Macfie
(1934) from Malaysia, D. biseriata Wirth
and Beaver (1979) from Malaysia, D. con-
finis (Johannsen) (1931) from Sumatra, D.
nepenthicola Wirth and Beaver (1979) from
Malaysia, and D. subgrata Tokunaga (1961)
from West Irian.
LITERATURE CITED
Downes, J. A. and W. W. Wirth. 1981. Ceratopo-
gonidae, Pp. 393-421. Jn J. F. McAlpine et al.,
eds., Manual of Nearctic Diptera, Volume 1. Agric.
Canada Monogr. 27. 674 pp.
Edwards, F. W. 1929. British non-biting midges
(Diptera, Chironomidae). Trans. Entomol. Soc.
London 77: 279-430.
457
Johannsen, O. A. 1931. Ceratopogoninae from the
Malayan Subregion of the Dutch East Indies. Arch.
Hydrobiol. Suppl. Bd. 9: 403-448, 5 plates.
Lee, K. M. and K. L. Chan. 1985. The biology of
Dasyhelea ampullariae in monkey cups at Kent
Ridge (Diptera: Ceratopogonidae). J. Singapore
Nat. Acad. Sci. 14: 6-14.
Macfie, J. W.S. 1934. Report ona collection of Cer-
atopogonidae from Malaya. Ann. Trop. Med. Par-
asitol. 38: 177-194, 279-293.
Mayer, K. 1934. Ceratopogoniden-Metamorphosen
(C. Intermediae und C. Vermiformes) der
Deutschen Limnologischen Sunda-Expedition.
Arch. Hydrobiol. Suppl. Bd. 13: 166-202, 6 plates.
Meillon, B. de. 1936. Entomological studies. Studies
on insects of medical importance in South Africa.
Part HI. South African Ceratopogonidae. Part II.
Some new and unrecorded species. Publ. S. Afr.
Inst. Med. Res. 7: 141-207.
Meillon, B. de and W. W. Wirth. 1987. Subsaharan
Ceratopogonidae (Diptera). XII. New species and
records, mainly from South Africa. J. Entomol.
Soc. S. Afr. 50: 35-74.
Strenzke, K. 1950. XI. Anhang. Dasyhelea lithotel-
matica n. sp., Pp. 178-187. In A. Thienemann,
ed., Lunzer Chironomiden. Arch. Hydrobiol.
Suppl. Bd. 18: 1-202.
Tokunaga, M. 1941. Biting ceratopogonid midges
from the Caroline Islands. Annot. Zool. Jap. 20:
109-119, 1 plate.
1961. Notes on biting midges IT. Kontyu 29:
180-185.
Tokunaga, M. and E. K. Murachi. 1959. Diptera:
Ceratopogonidae. Insects of Micronesia 12: 103-
434.
Waugh, W. T. and W. W. Wirth. 1976. A revision
of the genus Dasyhelea of the eastern United States
north of Florida (Diptera: Ceratopogonidae). Ann.
Entomol. Soc. Am. 69: 219—247.
Wirth, W. W. 1976. A new species and new records
of Dasyhelea from the Tonga Islands and Samoa
(Diptera: Ceratopogonidae). Proc. Hawaiian Ento-
mol. Soc. 22: 381-384.
1987. A new species of Dasyhelea (Diptera:
Ceratopogonidae) from rock pools in the south-
western United States. J. N. Am. Benthol. Soc. 6:
72-76.
Wirth, W. W.and R.A. Beaver. 1979. The Dasyhelea
biting midges living in pitchers of Nepenthes in
Southeast Asia (Diptera: Ceratopogonidae). Annls.
Soc. Entomol. France (N.S.) 15: 41-52.
Wirth, W. W. and N. Marston. 1968. A method for
mounting small insects on microscopic slides in
Canada balsam. Ann. Entomol. Soc. Am. 61: 783-
784.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 458-460
A NEW SPECIES OF NEOBAPHION BLAISDELL, FROM IDAHO
(COLEOPTERA: TENEBRIONIDAE)
CHARLES A. TRIPLEHORN
Department of Entomology, The Ohio State University, Columbus, Ohio 43210.
Abstract.—Neobaphion alleni, a new species from Idaho, is described and illustrated.
Eleodes (Metablapylis) insolitus Doyen is made a synonym of Neobaphion papula Tri-
plehorn and Aalbu.
Key Words:
For almost 50 years the genus Neoba-
phion consisted of only two species, N. plan-
ipenne (LeConte 1866) and N. elongatum
Blaisdell (1933). In 1985, a third species, N.
papula Triplehorn and Aalbu was de-
scribed. The following new species was dis-
covered only a few months after the de-
scription of N. papula was published:
Neobaphion alleni, NEw SPECIES
Fig. 1
Holotype, female: Length: 15.2 mm;
width: 7.0 mm. Body stout, subopaque,
black.
Head subquadrate, °4 as long as broad,
flattened; clypeal suture entire but weakly
defined, epistomal margin truncate; surface
dull with numerous fine, rounded tubercles,
each bearing a short, dark colored seta api-
cally; eyes narrowly reniform with dorsal
lobe larger, more rounded than ventral lobe;
antennae short, stout, apical 6 segments
moniliform; relative lengths of antennal
segments (from base to apex): 10:4:18:10:9:
8:8:8:8:9:10.
Pronotum % as long as broad, broadest
in anterior half, with a faint transverse crease
in basal fifth; in dorsal view, lateral margin
strongly arcuate and distinctly explanate,
especially medially, marginal bead narrow
and finely denticulate from base to apex;
Tenebrionidae, Darkling Beetle, Neobaphion
anterior margin broadly and evenly concave
from side to side, angles acute, prominent;
base nearly straight, angles obtuse; disc fee-
bly convex, surface with tubercles similar
to those of head but larger, shiny, densely
and uniformly distributed, each with a short,
brownish seta originating on caudal side.
Scutellum triangular, acute caudally, fine-
ly sculptured.
Elytra moderately convex from side to
side, lateral margins subparallel, widest be-
hind middle, abruptly deflexed posteriorly
with pronounced caudal lobe; base slightly
concave with humeri obsolete; surface with
dense conspicuous shiny tubercles on disc
similar to those of pronotum and with larg-
er, denser and more spiculiferous tubercles
laterally, each with a short, pale seta di-
rected caudad.
Legs moderate in size, finely, densely,
muricately punctate; profemur slightly
emarginate subapically; protibial spurs
about equal in size; basal protarsomere with
dense tuft of golden setae interrupting plan-
tar grove.
Ventral surface alutaceous with coarse,
dense tubercles on hypomera and proster-
num and basal 4 abdominal sterna, less
coarse but equally dense on meso- and
metasterna, with smaller and scarcely evi-
dent tubercles on apical sternum; prosternal
VOLUME 91, NUMBER 3
459
Fig. 1. Adult Neobaphion alleni n.sp. 6.7 x.
process acute, horizontal. Genital segment
with coxites fused, coxites | and 2 triangular
in outline and concave ventrally, convex
dorsally, coxite acute apica!ly; gonostyle
minute with single long apical seta.
Allotype, male: Similar to female but
slightly smaller and more slender; Length:
13.0 mm; width: 5.7 mm.
Variation.—There is little variation
among the five specimens available. The al-
lotype male is the smallest and the paratype
from Oregon is the largest (L: 17.8 mm; W:
7.8 mm).
Diagnosis.—Neobaphion alleni closely
resembles N. papula and will run to that
species in our key (Triplehorn and Aalbu
1985, p. 591). It may be readily separated
from the latter by the explanate lateral mar-
gins of the pronotum and the lack of an
elytral sutural “keel.” In addition, the sub-
apical emargination of the profemur,
scarcely evident in N. alleni, is very pro-
nounced in both sexes of N. papula. The
genitalia of both sexes appear identical in
the two species.
Types.— Holotype, female: IDAHO:
Owyhee County, Brunneau Dunes, 14 April
1986, Albert Allen; a//otype male, same lo-
460
cality, 18 May 1986, Albert and Kip Allen;
paratypes: female, same locality, 13 May
1986, Albert Allen; (probably male), IDA-
HO, Ada County, 14 miles S. of Kuna, 15
April 1978, R.C. Biggam; female, ORE-
GON [Malheur County], 32 miles SW of
Vale, Twin Springs, 2 April 1969, Kenneth
Goeden. Holotype in California Academy
of Sciences, allotype in personal collection
of Albert Allen, paratypes in University of
Idaho, The Ohio State University and Or-
egon Department of Agriculture Collec-
tions.
Etymology.—I take pleasure in naming
this species for Albert Allen who sent me
the first specimen, made special efforts to
collect more, and provided valuable eco-
logical data.
Discussion.—Albert Allen has provided
interesting collection data on the three spec-
imens he captured. All were dug from ro-
dent burrows (probably kangaroo rat) at the
base of sagebrush plants on sand dunes by
Mr. Allen and his son Kip. One was found
along with two specimens of Eleodes ar-
matus LeConte and one in company with
E. longipilosus Horn, a common species on
Brunneau Dunes. Mr. Allen has collected
intensively in those dunes over 10 years but
had never encountered this Neobaphion un-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
til 1986. Despite diligent efforts by the Al-
lens, they obtained only two more speci-
mens. Obviously, it is a rare beetle or it has
been overlooked by individuals using con-
ventional collecting techniques.
It seems appropriate at this time to point
out an unusual circumstance involving syn-
onymy in Neobaphion. In 1985, Doyen (p.
230) described Eleodes (Metablapylis) in-
solitus from Esmeralda County, Nevada.
Also, in 1985, our paper (Triplehorn and
Aalbu), describing Neobaphion papula, from
Mineral County, Nevada, was published.
The journals in which the two descriptions
appeared were both mailed on 11 July 1985,
thus constituting simultaneous publication.
As the next reviser (this paper) I arbitrarily
select Neobaphion papula Triplehorn and
Aalbu as the valid name and am placing
Eleodes insolitus Doyen as a synonym of it.
Literature Cited
Doyen, J.T. 1985. New species of Eleodes from Cal-
ifornia and Nevada (Coleoptera: Tenebrionidae).
Pan-Pacific Entomologist 61(3): 230-235.
Triplehorn, C. A. and R. L. Aalbu. 1985. A review
of the genus Neobaphion Blaisdell, with descrip-
tion of a new species from Nevada (Coleoptera:
Tenebrionidae: Eleodini). Proc. Entomol. Soc.
Washington 87(3): 587-592.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 461-467
POLYMERUS CASTILLEJA, A NEW MIRINE PLANT BUG FROM
CALIFORNIA AND OREGON, WITH REMARKS ON GENERIC
CHARACTERS OF THE GENUS POL YMERUS HAHN
(HETEROPTERA: MIRIDAE)
MICHAEL D. SCHWARTZ
Scientific Assistant, Department of Entomology, American Museum of Natural History,
New York, New York 10024-5192.
Abstract.—The new species, Polymerus castilleja, is described and distinguished from
other Polymerus species. A male dorsal habitus and illustrations of the genitalia of both
sexes are presented. The small hyaline right paramere and the deeply cleft secondary
gonopore of the male genitalia, and the obsolete dorsal structure and median process of
the female genitalia are perceived to be synapomorphic for species of the genus Polymerus.
Key Words:
Curation of the Polymerus Hahn holdings
in the collections of the American Museum
of Natural History revealed 115 specimens
of a previously undescribed species. All the
specimens of this taxon were taken from
localities in California and Oregon on In-
dian paintbrush, and possessed marked sex-
ual dimorphism and enlarged claw bases.
All measurements are in millimeters. A
list of abbreviations of the specimen dep-
ositories are in the Acknowledgments.
Polymerus castilleja, NEW SPECIES
Figs. 1-11
Diagnosis. — Distinguished from all other
Polymerus species by the enlarged claw base
(Figs. 2, 3).
Description.— Male. Dorsal aspect. Fig-
ure |. Total length 4.40-5.40, length from
apex of tylus to cuneal fracture 3.10-3.90;
coloration variable, ranging from—dark
specimens with head, antenna, pronotum,
mesoscutellum, scutellum, clavus, anterior
half of embolium, distal two-thirds of cori-
um interiad of radius, and paracuneus black,
with frons mesially, vertex mesially and
Heteroptera, Miridae, Polymerus
contiguous with eye, carina entirely, scutel-
lum apically, embolium distally, distal one-
third of corium mesad of radius, corium
laterad of radius, cuneus, and veins of mem-
brane testaceous to light fuscous: to—light
specimens with base color orange or red tes-
taceous with dark color restricted to tylus,
lora, juga apically, frons with ‘U’ shaped
marking mesially, temporal area peripher-
ally, collum, first antennal segment apically
and basally, second segment apically, callus
and posterior lobe of pronotum diffusely,
mesoscutellum mesially and laterally, small
basomedial dash on scutellum, clavus in-
teriad of claval vein, corium diffusely dis-
tally between cubitus and radius and mem-
brane; vestiture with golden, recumbent,
sericeous setae, and black, suberect, simple
setae on corium and cuneus distolaterally:
surface structure smooth, except for weak
to moderately transversely rugulose prono-
tum and scutellum. Head. Triangular in
dorsal view; Width across eyes 0.95—1.00,
interocular width 0.41-0.43; tylus pro-
duced, anteocular length 0.30-0.33; anten-
nal fossa contiguous with eye; eye large,
Fig. 1. Polymerus castilleja, dorsal habitus of male (28 x).
462
VOLUME 91, NUMBER 3
Figs. 2, 3.
emarginate anteriorly, ventral margin ex-
tending ventrad of fossa in lateral view; an-
tenna beset with fine, fuscous to black, sub-
erect, simple setae, length of segment I 0.35-
0.41, II 1.70-2.05; length of labium 2.30-
2.63, reaching fourth to sixth abdominal
sternite. Pronotum. Trapeziform in dorsal
view; posterior width 1.40-1.64; posterior
margin broadly curved, lateral margin
rounded; callus smooth and weakly pro-
duced. Hemelytra. Macropterous, subpar-
allel sided, sometimes tapered posteriorly
in dorsal view. Legs. Tibiae longer than
femora; color variable—light specimens with
coxae, trochanters, and femora on basal half
and apically testaceous, femora with diffuse
stripes, tibiae, and first and second tarsal
segments orange red, third segment and
claws black:—dark specimens with coxae
and femora with diffuse spots or stripes and
apically, tibiae narrowly apically and ba-
sally, and tarsi dark fuscous to black; claws
broad, strongly produced basally (Figs. 2,
3). Ventral aspect. Color variable ranging
from black ground color with proepister-
num, propleura, and sternum entirely, epi-
meron bordering coxae, and posterior edge
of evaporative area of peritreme testaceous,
and with band ventrad of abdominal spi-
racles red fuscous—to testaceous ground
color with proepeisternum basally, propleu-
ra medially, sternum mesially, abdominal
Polymerus castilleja. Pretarsal structures (49 x). 2, End on view. 3, Lateral view.
sternites laterally and mesially dark fuscous
or black. Genitalia. Genital capsule: With
broad, truncate, and apically spinose tuber-
cle dorsolaterad of left paramere insertion
(Fig. 4). Left paramere: Broadly ‘U’ shaped,
surface without tubercles or spines; shaft ex-
tending beyond sensory lobe in lateral view;
sensory lobe small, flattened on lateral sur-
face; arm longer than shaft; shaft slightly
compressed, broader than cylindrical arm,
tapering to rounded apex (Fig. 6). Right par-
amere: Small, ovoid, with only apex and
base entirely sclerotized, median portion
hyaline; apex with small tubercle (Fig. 5).
Vesica: Ductus seminis: Cylindrical, grad-
ually expanded toward gonopore. Second-
ary gonopore: aperture with diffuse spinu-
lae, incomplete, left side with deep fissure
(Fig. 8). Membrane: Basal sclerite encircling
base of membrane, projecting posteriad of
ductus, and extending distally to form a
thinly sclerotized lobe of the membrane; two
lobal sclerites, one short and needlelike, one
much longer, broader and clawlike (Fig. 7);
portion of membrane adjacent to gonopore
with surficial spiculae which merge with ap-
erture of gonopore (Fig. 8).
Female.— Macropterous. Vestiture simi-
lar to, coloration usually lighter than (with
dark markings much less extensive), and
bodyform more robust than, male. Mea-
surements. Total length 4.70-5.80, length
464
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
C NG
g
57
AS
Figs. 4-10. Polymerus castilleja. Genitalic structures. 4-8, Male genitalia. 4, Genital capsule, dorsal view
(55x). 5, 6, Parameres, lateral view (110). 5, Right. 6, Left. 7, 8, Vesica. 7, Frontal view (110). 8, Detail of
secondary gonopore, frontal view (220). 9, 10, Female genitalia (110). 9, Right sclerotized ring, anterior
view. 10, Posterior wall, posterior view.
from apex of tylus to cuneal fracture 3.55-
4.40; width across eyes 0.98-1.15, interocu-
lar width 0.45-0.54; anteocular length 0.3 1-
0.43; length of antennal segment I 0.35-
0.40, IT 1.60-2.08; length of labrum 2.40-
2.88, reaching fourth to sixth sternite; pos-
terior width of pronotum 1.60-2.03. Gen-
italia. Sclerotized rings: Separate, subtrian-
gular, moderate size. Dorsal labiate plate:
Small, adhering and following curvature of
ramae anteriad, attaining lateral margins of
rings in dorsal view. Ventral labiate plate:
Not spanning rings, subquadrate, slightly
shorter than rings in dorsal view (Fig. 9).
Posterior wall: Inter-ramal sclerite: Narrow,
elongated laterally, sometimes disarticulat-
ed medially. Median process and dorsal
structure: Obsolete. Inter-ramal lobes: Large,
crescent shaped, extending ventrad to even
with ventralmost portion of dorsal labiate
plate in posterior view; surface structure with
needlelike spiculae (Fig. 10).
VOLUME 91, NUMBER 3
Etymology.—Named after the genus of
the host plant.
Holotype 6.—UNITED STATES. Ore-
gon. Linn Co., H. J. Andrews Experimental
Forest, | mi N of Frissel Pt., July 28, 1981,
coll. G. M. Stonedahl, ex Castilleja sp.; de-
posited at AMNH.
Paratypes.—UNITED STATES. Califor-
nia: A/pine Co., Ebbetts Pass, Aug. 6, 1974,
F. G. Andrews, K. S. Corwin 1? (CAFA).
Modoc Co.: 3 mi S of Bug Station, Hwy.
139, June 20, 1956, J. Schuh, sweeping Cas-
tilleja sp. 13 6, 9 2 (OSU); Fandago Pass
Summit, July 3, 1979, 1890 m, R. T. Schuh
& B. M. Massie, ex Castilleja sp. 9 6, 13 9
(AMNH). Shasta Co.: 1 mi S of Jct. of Rts.
89 & 299, 1219 m, July 9, 1980, R. T. Schuh
& G. M. Stonedahl, ex Castilleja applegatei
Fern. var. fragilis (Zeile) N. Holmgren 11
4, 32 2 (AMNH, CAS); “same locality and
host,” G. Stonedahl 11 6, 7 ?(AMNH). Sier-
ra Co., Sierraville, June 28, 1966, C. E.
Hawkins | 6, 1 2 (OSU). Siskiyou Co., 2.5
mi N of Medicine Lk. on Medicine Lake
Rd., July 18, 1985, G. M. Stonedahl & J.
D. Mclver, ex Castilleja sp. 5 6, 3? (AMNH).
Oregon: Jackson Co., Siskiyou Summit on
I-5, July 4, 1982, T. J. Henry & G. M. Stone-
dahl, ex Castilleja sp. 4 4, 2 2 (AMNH);
“same label data,” 4300 ft. 6 2 (USNM).
Klamath Co., Bly Mt., June 14, 1958, J.
Schuh | 4 (OSU). Lake Co.: Summer Lake:
June 16, 1938, Grey & Schuh 1 4, 1 2
(AMNH); Aug. 16, 1939 1 @ (AMNH);
““same date,” Schuh & Scott 1 2 (AMNH).
Warner Cyn. nr. Lakeview roadside, 2 mi
E of Hwy. 395, 5450 ft., July 19, 1971, Lat-
tin 1 @ (OSU). Linn Co.: ““with same label
data as holotype,” 9 4, 3 2 (AMNH, OSU);
Monument Peak, 8 mi ESE of Gates Sum-
mit, 4725 ft., June 16, 1960, J. D. Lattin 1
4 (OSU).
Additional specimens. —California: San
Luis Obispo Co., Arroyo Grande Creek, SW
Fig. 11. Distribution of Polymerus castilleja.
466
of San Luis Obispo, +160 m., May 18, 1985,
R. T. Schuh & B. M. Massie, ex grasses and
forbs 1 2(AMNH). Santa Barbara Co., Up-
per Oso Cmpgrd., off Rt. 154, 310 m., May
7, 1985, R. T. Schuh & B. M. Massie | 2
(AMNH). These two specimens were ex-
cluded from the paratype series because they
were not associated with male specimens,
and were collected at localities removed far
to the southwest of the paratypic localities.
Hosts. — All specimens where collected on
either Castilleja sp. or C. applegatei Fern.
var. fragilis (Zeile) N. Holmgren (Scroph-
ulariaceae). Hitchcock and Cronquist (1973)
summarized the distribution of applegatei
var. fragilis as occurring in sagebrush (Ar-
temisia spp.) and open coniferous wooded
slopes in central Idaho, across eastern Or-
egon to the southern Cascades and south to
northern Nevada and eastern California.
Distribution.—The Cascade region of
westcentral Oregon south to the Sierra re-
gion of northern California. The true range
of castilleja may be more extensive because,
as noted above, the range of at least one of
its host plants is more widespread than pres-
ently known for the plant bug (Fig. 11).
Discussion. — There are no keys to the Po-
/ymerus species which occur within the dis-
tributional range of castilleja. Polymerus
castilleja keys to hirtus Knight in Kelton
(1980), and basalis (Reuter) in Knight
(1941). However, castilleja is easily sepa-
rated from these and all other Po/ymerus
species by the enlarged clawbase. Addition-
ally, the labium of casti/leja is longer (2.30-
2.88) than either Airtus (1.75-1.89) or basa-
lis (less than 2.30), and it is questionable
whether basalis or hirtus actually occur
within the range of castilleja.
Polymerus basivittis (Reuter) was collect-
ed along with castilleja on Indian paint-
brush at the H. J. Andrews Experimental
Forest, Linn County, Oregon and Fandango
Pass, Modoc County, California. These two
species are distinguished by the length of
the labium and body conformation in ad-
dition to the different claw structure. The
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
labium of basivittis barely reaches the meta-
coxae, whereas the labium of castilleja
reaches from the fourth to the sixth abdom-
inal sternite. Both sexes of basivittis are
compact, ovoid bugs, while the sexes of cas-
tilleja are dimorphic with the males elon-
gate. Kelton (1980) reported that basivittis
has been collected on the widespread Ga-
lium boreale L. (Rubiaceae).
Superficially the female of castilleja has
the appearance of robustus Knight. How-
ever, the latter species is distributed in
southern California and possesses a labium
which barely reaches the metacoxae.
Schwartz (1987) included a preliminary
analysis of the genitalic attributes of many
representative mirine genera (234 species
encompassing 110 genera, including all the
North American genera). In light of that
study and from character information
gleaned from several species of Polymerus
dissected for the present paper, I will enu-
merate what I believe are autapomorphic
genitalic characters for the genus. I exam-
ined the genitalia of both sexes of the fol-
lowing species: basivittis, basalis (Slater
1950: pl. 3, fig. 5), diffusus (Uhler), nigritus
(Fallén), and testaceipes (Stal). The male
genitalia of three additional species were ex-
amined only through published illustra-
tions: atacamensis Carvalho and Carpin-
tero (Carvalho and Carpintero 1986, figs.
11, 13), chrysopsis Knight (Kelton 1959, fig.
32), and peruanus Carvalho and Meléndez
(Carvalho and Meléndez 1986, fig. 6).
Polymerus seems to be diagnosed by these
characters:
(1) The right paramere is relatively small
for a mirine, and only sclerotized at the apex
and base, with the remainder of the para-
mere hyaline (Fig. 5). This construction was
also noted by Kelton (1959) and illustrated
by Wagner (1973: fig. 324 b, e, h, 1, 0) for
asperulae (Fieber), brevicornis (Reuter), mi-
crophthalmus (Wagner), palustris (Reuter)
and wnifasciatus (Fabricius).
(2) The secondary gonopore is modified
from the simple coil-like, uninterrupted, ap-
VOLUME 91, NUMBER 3
erture of many mirine genera to a deeply
notched aperture with strong concentric
rings of minute spinulae (Fig. 8). The spi-
nulae of the gonopore are intermixed with
the spinulae of the adjacent portion of the
vesical membrane.
(3) The dorsal structure and median pro-
cess of the posterior wall are usually obso-
lete and flattened in the plane of the inter-
ramal sclerite, respectively (Fig. 10).
These genitalic features further define Po-
/ymerus, which is otherwise inadequately
diagnosed as medium sized, finely punctate
member of the Mirini, with densely distrib-
uted vestiture of shining light to golden se-
riceous simple setae. The hypothesis of the
genitalic autapomorphies of Polymerus is
tentative, as an understanding of the distri-
bution of the features listed above within
the tribe is limited.
ACKNOWLEDGMENTS
I thank the following individuals for pro-
viding material from collections in their care:
A. Asquith and J. D. Lattin, Systematic
Entomology Laboratory, Oregon State Uni-
versity, Corvallis (OSU); T. J. Henry, Sys-
tematic Entomology Laboratory, USDA, %
National Museum of Natural History,
Washington, D.C. (USNM); and R. T.
Schuh, American Museum of Natural His-
tory, New York (AMNH). I thank B. Hall,
Corvallis, Oregon, for the fine illustration
of the adult male of castilleja and J. D. Lat-
tin, through funds provided by NSF BSR-
8514325, for providing the financial sup-
467
port for it. I thank T. J. Henry, J. D. Lattin,
R. T. Schuh and an anonymous reviewer
for reading and commenting on the manu-
script. Financial support for field work and
technical assistance was received from NSF
Grants DEB 81-13401 and BSR 86-06621
to R. T. Schuh, AMNH.
LITERATURE CITED
Carvalho, J. C. M. and D. L. Carpintero. 1986. Mi-
rideos Neotropicais, CCLXXIV: descrigoes de
quatro espécies novas da América do Sul (He-
miptera). An. Acad. Brasil. Ciéne. 58: 291-296.
Carvalho, J. C. M. and E. Meléndez. 1986. Mirideos
Neotropicais, CCLXIX: descricoes de trés espé-
cies novas da Amazonia peruana (Hemiptera). Bol.
Museu Paraense Emilio Goeldi 2: 93-100.
Hitchcock, C. L. and A. Cronquist. 1973. Flora of
the Pacific Northwest, an illustrated manual. Univ.
of Washington Press, Seattle, 730 pp.
Kelton, L.A. 1959. Male genitalia as taxonomic char-
acters in the Miridae (Hemiptera). Can. Entomol.,
Suppl. 11, 91: 1-72.
1980. The Insects and Arachnids of Canada.
Part 8. The plant bugs of the prairie provinces of
Canada (Heteroptera: Miridae). Publication 1703.
Ottawa: Agric. Canada, 408 pp.
Knight, H. H. 1941. The plant bugs, or Miridae, of
Illinois. Bull. Illinois Nat. Hist. Surv. 22: 1-234.
Schwartz, M. D. 1987. Phylogenetic revision of the
Stenodemini with a review of the Mirinae (Het-
eroptera: Miridae). Ph.D. diss. New York: City
Univ., 383 pp.
Slater, J. A. 1950. An investigation of the female
genitalia as taxonomic characters in the Miridae
(Hemiptera). Iowa State College J. Sci., 25(1): 1-
81.
Wagner, E. 1973. Die Miridae Hahn, 1831, des Mi-
telmeerraumes und der Makaronesischen Inseln
(Hemiptera, Heteroptera). Teil 1. Entomol. Abh.
37 Suppl., 484 pp.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 468-472
LARVAL-SEX AND HOST-SPECIES EFFECTS ON LOCATION OF
ATTACHMENT SITES OF LAST-INSTAR BAGWORMS,
THYRIDOPTERYX EPHEMERAEFORMIS
(LEPIDOPTERA: PSYCHIDAE)
PETER K. LAGOY AND EDWARD M. BARROWS
(PKL) Environmental Strategies, 101 Metro Drive, San Jose, California 95110; (EMB)
Department of Biology, Georgetown University, Washington, D.C. 20057.
Abstract. —In last-instar bagworms, Thyridopteryx ephemeraeformis, on the deciduous
tree Robinia pseudoacacia, males more frequently attached their cases to leaves, and
females more frequently attached their cases to branches. Both males and females more
frequently attached their cases to branches, rather than leaves, of the evergreen Pinus
strobus. Diameters of branches that bagworms used as case-attachment substrates were
significantly related to bagworm sex and host species. Possible adaptive significances of
these phenomena are discussed.
Key Words:
Psychidae, Thyridopteryx ephemeraeformis, caterpillar behavior, Juniperus
virginiana, Pinus strobus, Robinia pseudoacacia
The bagworm, Thyridopteryx ephemer-
aeformis (Haworth), is a polyphagous her-
bivore which usually feeds on woody plants.
It occurs from the West Indies north to Ver-
mont, Michigan, and Minnesota and west
to Kansas and Texas, being most common
in southeastern United States according to
Davis (1964) and Longfellow (1980). Many
other workers including Riley (1869), Hase-
man (1912), Jones (1927), Jones and Parks
(1928), Barrows (1974), Barrows and Gordh
(1974), Kaufmann (1968), Kulman (1965),
Leonhardt et al. (1983), and Neal (1986)
have increased our knowledge of bagworm
biology. As background for our study, we
present a brief generalized summary of bag-
worm biology in the United States based on
these previous investigations.
In August to October, depending on lo-
cality, an inseminated female lays all of her
eggs in her pupal exuviae which remain in
the case that she made as a larva. Her egg-
laden case hangs on her host plant over the
winter, and her eggs hatch in late April to
early June. First-instar larvae emerge from
cases, construct conical cases of silk and
plant materials around themselves, and may
balloon, each by a silken thread, to a new
location. Many no doubt die due to pre-
dation, landing on unsuitable hosts, and
other factors. A larva that finds a suitable
host passes through five to eight instars
(Longfellow 1980), enlarging its case as it
grows, before it pupates in late summer to
early fall. Before pupation, a larva fastens
the anterior end of its case to a substrate
(usually a food plant), and turns 180 degrees
assuming a head-down position. Pupae
transform into adults in 2 to 3 weeks. Males
are typical of most kinds of adult Lepidop-
tera in having wings, legs, compound eyes,
antennae, and other adult characters. Fe-
males are essentially “egg bags” having re-
duced compound eyes, vestigial mouthparts
VOLUME 91, NUMBER 3
and legs, and no wings or antennae. After
insemination, a female lays her eggs in her
pupal exuviae, and then either dies in her
larval case or leaves it before dying. Most
bagworms, which are not killed by Homo
sapiens L., are killed by insect and other
parasites and invertebrate and vertebrate
predators before they reproduce.
Our study tests five hypotheses about case-
attachment behavior of bagworm last-instar
larvae on three hosts: the evergreen conifers
redcedar, Juniperus virginiana L., and white
pine, Pinus strobus L., and the deciduous
tree black locust, Robinia pseudoacacia
Ehrh. These hypotheses were prompted by
features of bagworm biology, its host-plant
architectures, and our preliminary field ob-
servations regarding case-attachment sites
(Barrows 1974 and later observations). Our
hypotheses are: (1) Female bagworms use
branches rather than petioles as case-at-
tachment sites on R. pseudoacacia (experi-
ment 1). (2) Males use branches rather than
petioles as case-attachment sites on R.
pseudoacacia (experiment 2). (3) Females
use branches rather than leaves as case-at-
tachment sites on P. strobus (experiment 3).
(4) Males use branches rather than leaves
as case-attachment sites on P. strobus (ex-
periment 4). (5) Bagworm case-attachment-
site diameter on R. pseudoacacia, P. strobus,
and J. virginiana is related to bagworm sex
and host species (experiment 5).
The bagworm’s range broadly overlaps
those of all three of its host plants that we
studied (Fernald 1950), and it can be locally
common on these species. Robinia pseudo-
acacia has alternate, petiolate compound
leaves from 20 to 36 cm long. Pinus strobus
usually has leaves in fascicles of five, each
7 to 12 cm long and persistent on a branch
for about 2 yr (Otis 1926, Ewers and Schmid
1981). Juniperus virginiana has small sessile
(petioleless), 1.5-12-mm long leaves which
persist on plants for 5 to 6 yr (Otis 1926).
Bagworms attach their last-instar cases to
branches, not individual leaves of J. virgin-
iana.
469
MATERIALS AND METHODS
Bagworms were sampled in Prince Wil-
liam, Fairfax, and Arlington Counties in
northern Virginia in September to October
1983 and 1984 before autumn leaf fall. We
measured attachment sites of bagworms,
that were up to 2.3 m above the ground and
on their hosts, from 25 trees of R. pseudo-
acacia, 10 trees of P. strobus, and 15 trees
J. virginiana. Bagworms occurred on other
parts of these trees which we did not sample.
The greatest diameters of pupal-case at-
tachment sites, which were nearly round to
elliptical in cross section, were measured
using Manostat® dial calipers accurate to
0.05 mm. We used data only from cases
whose former occupants could be sexed with
certainty. A male’s case had his pupal exu-
viae protruding from its distal opening or
was intact and empty with a distal tube with
a circular cross section through which he
obviously emerged. A female’s case con-
tained eggs in her pupal exuviae.
To test hypotheses 1-3, we used binomial
tests (BTs) to look for possible differences
between observed and expected attach-
ment-site frequencies. We designated ex-
pected frequencies as 50% on branches and
50% on leaves because each bagworm had
a hypothetical 50% chance of using one or
the other substrate by chance alone. We
tested hypothesis 5 by examining the 95%
confidence intervals of the differences be-
tween all possible pairs of means. This
method is more straightforward, gives more
information (the magnitudes of differences
between means and their 95% confidence
intervals), and makes each pairwise com-
parison at an alpha level of 0.05. Commonly
used simultaneous test procedures, e.g. the
Duncan’s multiple-range test, do not have
these advantages (Jones 1984). The SAS
computer package (SAS® Institute 1985)
was used to perform statistical analyses.
RESULTS AND DISCUSSION
Experiment 1.—In 1983, 88 of the 92 fe-
males collected from R. pseudoacacia were
470
Table 1.
confidence intervals, ranges, and sample sizes (N).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Experiment 5. Mean diameters of attachment sites of cases of six bagworm categories, their 95%
Mean (mm)
Males on:
Robinia pseudoacacia 1.46
Juniperus virginiana 1.70
Pinus strobus 2.09
Females on:
Juniperus virginiana 2.42
Pinus strobus 2.84
Robinia pseudoacacia 2.93
on branches; four were on petioles. In 1984,
297 of the 303 females collected from R.
pseudoacacia were on branches; 4 were on
petioles. Thus, female bagworms more fre-
quently attached their cases to branches
(both years, P < 0.0001, BT). Luther P.
Brown (personal communication), who
sampled bagworms in Maryland, did not
find any female cases on petioles of R.
pseudoacacia, but he sampled in November
after a substantial leaf drop could have oc-
curred. It may be reproductively advanta-
geous for a female last instar to attach her
case to a branch, rather than to a deciduous
leaf which would fall to the ground. First,
this site would keep her eggs above the
ground where they might have a lower prob-
ability of mammalian predation and fungus
infection (Barrows 1974, Berisford and Tsao
1975, Munte 1982). Second, her choosing
this site would put her first-instar offspring
in a place where they can readily find food
and from where they can balloon to other
host plants.
Experiment 2.—In 1983, of 36 males col-
lected from R. pseudoacacia, 33 were on
petioles and 3 were on branches. In 1984,
of 122 males, 89 were on petioles and 33
were on branches. Thus, males preferen-
tually attached to petioles (1983, P= 0.0003,
BT; 1984, P < 0.0001, BT). When we orig-
inally made our second hypothesis, we knew
of no reason why males should preferen-
ually attach to petioles rather than branches
95% CI (mm) Range (mm) N
0.08 0.75-4.50 122
0.10 0.95-3.60 6
0.14 0.90-4.00 69
0.06 1.20-4.90 328
0.10 1.70-5.50 171
0.10 1.65-9.05 303
of this plant. Because males leave no eggs
in their cases, it should be of no conse-
quence to their fitnesses (measured as num-
ber of offspring) if their empty pupal cases
fall to the ground with deciduous plant
leaves. However, possible advantages of pu-
pal-case attachment to petioles to male last-
instars and emerging males are worthy of
investigation.
Experiments 3 and 4.—In 1984, all 171
females and 86 of 89 males collected from
P. strobus were attached to branches rather
than leaves or leaf clusters. Thus, both sexes
preferentially attached to branches (both
sexes, P < 0.0001, BT). When we made our
third and fourth hypotheses, we assumed
that bagworms, attached to leaves or a group
of leaves, would usually overwinter on their
hosts because they were likely to attach to
at least some leaves that would not soon
dehisce. Under this hypothesis, case-attach-
ment location on P. strobus should have
little effect on bagworm fitnesses, but ac-
cording to our results, bagworms preferred
branches. Possible mechanistic explana-
tions for this behavior include bagworms
tend to attach their pupal cases to firm rod-
shaped substrates rather than to more flex-
ible single leaves or clusters of leaves, and
they are repelled by stimuli from leafy areas
just before attaching their pupal cases.
Experiment 5.— Data regarding bagworm
sex, host, attachment-site diameters, and
sample sizes are summarized in Table 1,
VOLUME 91, NUMBER 3
Table 2.
471
Experiment 5. All pairwise comparisons of six bagworm categories indicating the differences between
means and 95% confidence intervals of these differences for each pair.
MRS? MJV MPS FRS FJV
MJV 0.24 + 0.13
MPS 0.63 + 0.15 0:39) 0:16
FRS M47 = O112 1.23: = 0.33 0.84 + 0.16
FJV 0.96 + 0.10 0772 = OZ 0.33 + 0.14 O51 2 0.1
FPS 1.38 + 0.12 1.14 + 0.14 0.75 + 0.16 0.09 + 0.13 0.42 + 0.11
“MRS = males on Robinia pseudoacacia, MJV = males on Juniperus virginiana, MPS = males on Pinus
strobus,; FRS = females on R. pseudoacacia; FJV = females on J. virginiana; FPS = females on P. strobus.
Members of all pairs are different from one another at an alpha level of 0.05 except for FRS and FPS because
the 95% confidence interval of this pair’s difference between means contains 0).
and pairwise comparisons of differences be-
tween means are listed in Table 2. On all
three hosts, females attached their pupal
cases to significantly larger-diameter
branches than did males. Bagworms sam-
pled by L. P. Brown (personal communi-
cation) behaved similarly to the ones we
sampled on R. pseudoacacia and J. virgin-
jana. This may be due to the facts that fe-
males are larger than males, and larger bag-
worms use larger-diameter attachment sites
than do smaller ones (Brown, personal com-
munication); and on R. pseudoacacia, males
used petioles while females used branches.
Males used a significantly different mean
attachment-site diameter among all hosts,
and females used a significantly different
mean attachment-site diameter except be-
tween R. pseudoacacia and P. strobus (Table
2):
In conclusion, our study suggests some
other directions for future investigation in-
cluding: (1) behavior mechanisms that af-
fect bagworm choice of diameters and kinds
of case-attachment sites; (2) possible effects
of bagworm genetics, parasites, and site
availability, on this behavior; and (3) the
possible relationship between male attach-
ment-site location on R. pseudoacacia and
parasitism in subsequent bagworm gener-
ations. Parasitized cases on leaves fall to the
ground, and this possibly eliminates some
parasites from bagworm populations.
ACKNOWLEDGMENTS
We thank L. P. Brown (George Mason
University) for sharing his bagworm data
with us and R. S. Blanquet (Georgetown
University) and an anonymous reviewer for
insightful comments on preliminary manu-
scripts. This research is part of PKL’s MLS.
thesis.
LITERATURE CITED
Barrows, E.M. 1974. Some factors affecting the pop-
ulation size of the bagworm, Thyridopteryx
ephemeraeformis (Lepidoptera: Psychidae). En-
viron. Entomol. 3: 929-932.
Barrows, E.M.andG.Gordh. 1974. Insect associates
of the bagworm moth, Thyridopteryx ephemer-
aeformis (Lepidoptera: Psychidae). J. Kansas
Entomol. Soc. 47: 156-161.
Berisford, Y. C. and C. H. Tsao. 1975. Parasitism,
predation and disease in the bagworm, 7/yridop-
teryx ephemeraeformis (Haworth) (Lepidoptera:
Psychidae). Environ. Entomol. 4: 549-554.
Davis, D. R. 1964. Bagworm moths of the Western
Hemisphere. Bull. U.S. Natl. Mus. 244. 233 pp.
Ewers, F. W. and R. Schmid. 1981. Longevity of
needle fascicles of Pinus /ogaeva (bristlecone pine)
and other North American pines. Oecologia (Ber-
lin) 51: 107-115.
Fernald, M. L. 1950. Gray’s Manual of Botany. Eighth
Edition. American Book Company, New York.
1632 pp.
Haseman, L. 1912. The evergreen bagworm. Uni-
versity of Missouri. Agricultural Experiment Sta-
tion Bulletin. 104: 307-330.
Jones, D. 1984. Use, misuse, and role of multiple-
comparison procedures in ecological and agricul-
tural entomology. Environ. Entomol. 13: 635-649.
472
Jones, F.M. 1927. The mating of the Psychidae (Lep-
idoptera). Trans. Amer. Entomol. Soc. 53: 293-
312.
Jones, F. M. and H. B. Parks. 1928. The bagworms
of Texas. Texas Agric. Exp. Stn. Bull. 382. 36 pp.
Kaufmann, T. 1968. Observations on the biology and
behavior of the evergreen bagworm moth, 7hyr-
idopteryx ephemeraeformis (Lepidoptera: Psychi-
dae), in Kansas. J. Kansas Entomol. Soc. 47: 156-
161.
Kulman, H. M. 1965. Natural control of the bag-
worm and notes on its status as a forest pest. J.
Econ. Entomol. 58: 863-866.
Leonhardt, B. A., J. W. Neal, Jr., J. A. Klun, M. Schwarz,
and J. R. Plimmer. 1983. An unusual Lepidop-
tera sex pheromone system in the bagworm moth.
Science 219: 314-316.
Longfellow, S. N. 1980. Selected aspects of the bi-
ology and morphology of the bagworm, 7/hyridop-
teryx ephemeraeformis, [sic] (Haw.) in Kansas.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Ph.D. Dissertation. Kansas State University,
Manhattan, Kansas. 111 pp.
Munte, S-T. 1982. Larval behavior of the bagworm,
Thyridopteryx ephemeraeformis (Haworth) (Lep-
idoptera: Psychidae). M.S. Thesis. Georgetown
University, Washington, D.C. 50 pp.
Neal, J. W., Jr. 1986. Salient vestiture and morpho-
logical characters of the pharate female bagworm,
Thyridopteryx ephemeraeformis (Lepidoptera:
Psychidae). Ann. Entomol. Soc. Amer. 79: 814—
820.
Ous, C. H. 1926. Michigan Trees. The Regents of
the University of Michigan. 247 pp.
Riley, C. V. 1869. The bagworm, alias basket-worm,
alias dropworm, Thyridopteryx ephemeraeformis
Haworth. Amer. Entomol. 2: 35-38.
SAS® Institute, Inc. 1985. SAS® User’s Guide: Sta-
ustics, Version 5 Edition. SAS Institute, Inc., Cary,
North Carolina. 956 pp.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 473-485
REDESCRIPTION OF CULEX (MELANOCONION) DELPONTEI
DURET, 1968 AND CX. (MEL.) PEREYRAI DURET, 1967,
FROM SOUTHERN BRAZIL
OsWALDO PAULO FORATTINI AND MARIA ANICE MUREB SALLUM
School of Public Health, Department of Epidemiology, Av. Dr. Arnaldo, 715, CEP
01255, Sao Paulo, Brazil.
Abstract. — Adults of both sexes of Culex (Melanoconion) delpontei Duret and Cx. (Mel.)
pereyrai Duret, are redescribed and illustrated. Diagnostic characters for separating these
from similar species of the Spissipes Section of subgenus Melanoconion are given. Dis-
tribution and epidemiological aspects are considered.
Key Words:
Recent publication of arbovirus investi-
gation in Argentina (Mitchell et al. 1985,
1987a), report the isolation of several viral
strains from Culex (Melanoconion), many
obtained from Cx. de/pontei Duret. Various
strains were isolated including several be-
longing to the Venezuelan equine enceph-
alitis virus (VEE) complex. Culex delpontei
is now considered as important arbovirus
vector, but much about this species remains
unknown, including characters for identi-
fication.
During studies on the mosquitoes of the
Ribeira Valley and other regions of Sao Pau-
lo State, Brazil, several adults of Cx. del-
pontei and Cx. pereyrai Duret, were col-
lected. We take this opportunity to
redescribe them with the objective of
achieving a better characterization for pur-
poses of identification. Ideally types should
be examined, but, as far as is known, they
were retained in the collection of the author
of species and so of no easy access. By other
side, descriptions of these species are well
enough illustrated to leave no doubts about
their identities.
The terminology of Harbach and Knight
(1980) was utilized for the descriptions, ex-
Insecta, Diptera, Culicidae, Culex, Melanoconion, Brazil
cept for the wing veins which follow Belkin
(1962).
Culex (Melanoconion) delpontei Duret
Figs. 1-6, 10, 11 and 13
Culex (Melanoconion) paracrybda of Duret
1953: 119.
Culex (Melanoconion) delpontei Duret 1968:
8 (type not seen; type locality Argentina,
Chaco, Las Palmas; holotype deposited
in the collection of the author of specie
(Knight and Stone 1977)). Sirivanakarn
and Jakob 1981: 199.
Female.— Head: Antenna dark, length
about 2.00 mm; flagellum normal, whorls
normally with 6 setae. Proboscis entirely
dark-scaled; length 1.66-1.88 mm, mean
1.72 mm. Maxillary palpus entirely dark-
scaled; length 0.28-0.33 mm, mean 0.30
mm, about 0.17 of the proboscis length.
Vertex (Fig. 4) with narrow falcate scales in
a small median dark patch, between pale
whitish ones, broad appressed dingy white
in small lateral patch; forked scales dark.
Occipital region with some pale whitish fal-
cate scales. Cibarial armature (Figs. 1, 2, 3
474 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-9.
Adult structures. Culex delpontei
occipital region. 5, dorsal view of right wing. 6, ventral view of right wing. Culex pereyrai. 7, vertex and occipital
region. 8, cibarial teeth. 9, cibarial teeth showing thorn-shaped processes on the posterior border of the cibarial
bar.
and 10). Cibarial bar developed, moderately
chitinized, strongly concave, with about 22
small cibarial teeth arranged in single row,
with sizes gradually smaller laterally where
they may be visible in lateral profile; many
irregular chitinized folds on posterior bor-
der of cibarial bar where folds sometimes
1, female cibarial armature. 2,3, cibarial teeth. 4, vertex and
end as minute prickle; cibarial teeth laminar
shaped with two recognized parts, anterior
and posterior; the first one, including nearly
third or fourth total tooth dorsal length, as
a thin sagital plate; second one as transver-
sal plate, lozange or hexagonally roughly
outlined: minute irregularly disposed prick-
VOLUME 91, NUMBER 3
les may be seen at the margins of both parts.
Cibarial dome nearly circular, concave cap
covered by superficial denticles. Thorax:
Scutal integument brown or dark brown,
except yellow paratergite; scutum almost
entirely covered by uniform fine falcate dark
bronzy scales, with some colorless ones on
prescutellar area; scutal setae conspicuous
(acrostichal setae absent), brownish black
with reddish sheen; antealar setae, shiny
yellow to golden. Scutellar scales as scutal
scales; lateral lobe each with 3-5 large setae,
median lobe with 5,6 large setae. Ante-
pronotum dark, without scales, with scat-
tered dark and golden setae. Postpronotum
brown dorsally and yellow ventrally, with
narrow dark scales, like scutal scales, pos-
terodorsal margin with 3—5S upper dark and
lower golden setae. Pleural integument pale
yellow; with shiny golden yellow setae; pleu-
ral setae: about 9-11 upper proepisternal,
4-7 prealar, 5-8 upper mesokatepisternal,
8-10 lower mesokatepisternal, 5-7 upper
mesepimeral and 1 lower mesepimeral.
Pleura with scales on mesokatepisternum:
a small patch of pale spatulate scales on
lower posterior border and sometimes with
2,3 colorless spatulate scales on upper cor-
ner. Wing (Figs. 5, 6): Length 2.77-3.04
mm, mean 2.87 mm; mostly dark-scaled,
with a minute patch of clear scales on base
of costa; cell R, 3.48-5.17 of vein R3,;,
mean 4.37; cell M, 0.71-0.75 of cell R,;
subcosta intersects costa before furcation of
R,,;. Dorsal scaling: appressed spatulate
scales on costa, subcosta, R, R,, R,,;, distal
0.5 of M,,2, M34, Cu, Cu,, Cu, and basal
0.5 of 1A; linear plume scales on R,, R>,,
and M; inclined narrow spatulate scales on
R,, R;, proximal 0.5 of M,.., and distal 0.5
of 1A; remigium with appressed spatulate
scales and long golden setae distally. Ventral
scaling: appressed spatulate scales on costa,
subcosta, R,, R,,3, base of R,, base of R,
and on M; linear plume scales covering
nearly basal 0.5 of R,, on basal 0.3 of Cu,,
Cu, and middle of 1 A; inclined narrow spat-
ulate scales on distal 0.5 of R,, R5, Ry, Ry.s,
475
M, >, M;,,4, distal 0.7 of Cu, and distal 0.6
of 1A; Cu and basal 0.4 of 1A devoid of
scales. Halter: Yellow, capitellum slightly
darker at apex. Legs: Anterior surface of
forecoxa pale-scaled; anterior surface of
midcoxa with longitudinal patch of pale
scales; anterior surface of hindcoxa devoid
of scales. Antero- and posteroventral sur-
faces of foretrochanter pale-scaled; midtro-
chanter with antero- and posteroventral
surfaces pale-scaled; antero- and postero-
ventral surfaces of hindtrochanter pale-
scaled. Fore- and midfemora mainly pale-
scaled, posterior surface of forefemur with
indistinct longitudinal stripe of dingy pale
scales, posteroventral surface of midfemur
with dingy pale scales, hindfemur with com-
plete dorsal stripe of dark scales distally
widening and expanding onto anterior and
posterior surfaces at apex. Tibiae and tarsi
entirely dark-scaled. Abdomen: Tergum I
with median posterior patch of dark scales;
terga HI-VII dark-scaled with basolateral
patches of white scales; tergum VIII mainly
dark-scaled, with small basolateral patch of
white scales. Sterna II-IV_ white-scaled;
sterna V,VI mainly white-scaled, occasion-
ally with mixed dark scales distally; sternum
VII infrequently entirely white-scaled, gen-
erally with dark scales distally; sternum VIII
with lateral patches of white scales. Geni-
talia (Fig. 10): Tergum IX narrowed in mid-
dle, lobes each bearing 6-11 setae. Upper
vaginal lip narrow, distinct; lower vaginal
lip and insula indistinct; about 7,8 insular
setae in cluster. Upper vaginal sclerite dis-
tinct, inverted U-shaped, chitinized. Post-
genital lobe short, distally rounded, with 7—
13 setae on either side of midline, setae
mostly on ventral surface.
Male. — Like female except for sexual dif-
ferences as follow. Head: Antenna strongly
plumose, length about 1.78 mm. Proboscis
entirely dark. Maxillary palpus dark, length
about 2.52 mm, extending beyond tip of
proboscis by about apical 0.5 of palpomere
4 and all of palpomere 5; palpomeres 4 and
5 entirely covered by dark strong setae; pal-
476
pomere 3 with 8,9 setae at apex. Abdomen:
Tergum II entirely dark-scaled; tergum III
with small basolateral white patch; terga IV—
VII with basolateral white patches; tergum
VIII (ventral in position) mostly white-
scaled, with deep V-shaped median poste-
rior emargination and several long bristles
mixed with shorter setae (Fig. 11); sterna
predominantly white-scaled, with small
number of dark scales distally on sterna V—
VII; sternum VIII (dorsal in position) with
basolateral white patches. Genitalia (Fig. 11):
Ninth tergal lobe small, columnar shaped,
widely separate, bearing long and slender
setae on 0.5 of distal surface. Gonocoxite
stocky, outer margin convex, inner mod-
erately concave; ventrolateral surface with
strongly developed setae and small number
of scales, mesal surface with small setae in
indistinct rows extending from base to level
of subapical lobe, lateral surface with sparse
patch of slender setae (Isp) from proximal
area to level of subapical lobe, proximal part
of ventrolateral surface with scales; sub-
apical lobe distinctly divided, divisions
approximated; proximal division not divid-
ed, lengthened, with an apical infundibular
and hyaline expansion partially covering in-
sertions of setae a and b which are long,
enlarged and sinuous with another hyaline,
broad and hooked-falciform seta beyond
middle and 5-11 long, slender and curved
setae from base to level of insertion of
hooked-falciform seta; distal division with
2 approximated uneven arms, each bearing
an apical setae, proximal arm stronger, dis-
tally enlarged, bearing | long hooked seta h2
saberlike setae(s) (one larger than the oth-
er) inserted on prominent tubercle near
middle of arm, distal arm slender, cylindric,
bearing | stiff short nearly saberlike seta (s)
and 4 foliform setae (f) (2 similar and 2
slightly larger and dissimilar in size). Gono-
stylus slender, curved, moderately nar-
rowed distally, crest slightly wrinkled on
ventral surface before apical snout; gono-
stylar claw short, leaf-like broadest apically.
Phallosome with lateral plates and aedeagal
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
sclerites equivalent in length; aedeagal scler-
ite broad, curved and falciform in lateral
view, mostly hyaline, excluding mesal lon-
gitudinal sclerotinization and more heavily
sclerotinized area at angle of connection with
lateral plate, dorsal end narrowly fused to
base of lateral plate; lateral plate long, co-
lumnar shaped, apical process with blunt
and rounded apex, lateral and ventral pro-
cesses absent, dorsal process sclerotized and
separated from margin of aedeagal sclerite
by distinct angle; aedeagal sclerites con-
nected by dorsal aedeagal bridge. Proctiger
elongate; paraproct distally narrowed, ba-
sally expanded, base articulated with pos-
terolateral margin of tergum X, crown with
row of about 6 short, simple blades; cercal
sclerite long and narrow, sclerotized, broad-
est basally; 2,3 small cercal setae. Tergum
X large, rectangular, concavo-convex, dor-
sal surface concave.
Material examined. —Sixty specimens ex-
amined from several localities in Sao Paulo
State, Brazil, as follows: 41 92 (Experimental
Station, Pariquera-Acu, I.81, X1.81, XII.81,
1.82, 12; Bigua Road, Iguape, IX.82, X.82,
X1.82, 28; Santa Helena Farm, Sao Joao da
Boa Vista, X1.81, 1). 19 44 (Pariquera-Acu,
urban area, XI.79, 1; Pariquera-Mirim, Par-
iquera-Acu, V.85, VI.85, 2; Itapuan, Ita-
pitangui, Cananéia, V.80, XII.80, 1.81,
11.81, H1.81, 1V.81, X1.81, 11; Bigua Road,
Iguape, X.82, 2; Santa Helena Farm, Sao
Joao da Boa Vista, III.82. 3) Iguape, Can-
anéia and Pariquera-Acu, are located be-
tween 24.0°-25.0°S and 47.0°-48.0°W, Sao
Joao da Boa Vista is 22.0°S and nearly
47.0°W.
Distribution and bionomics. — Culex del-
ponte is reported here for the first time from
Southern Brazil. It has also been found in
Paraguay and Northern Argentina as next
related (Duret 1953, 1968; Sirivanakarn and
Jakob 1981; Mitchell et al. 1985). Its dis-
tribution is in the southern range of sub-
genus Melanoconion (Fig. 13): ARGEN-
TINA. — Chaco: Las Palmas (type locality);
Puerto Bermejo; Resistencia, San Fernando
VOLUME 91, NUMBER 3
Dept.; Antequera, | de Mayo Dept.; Cor-
rientes: Rincon de Vences, General Paz
Dept.; Santa Fe: Esperanza, Las Colonias
Dept.; San Justo, Sanjusto Dept. PARA-
GUAY.—Puerto-i, San Pedro Dept.; Tol-
do-cué, Concepcion Dept.; San Pedro, San
Pedro Dept.; Puerto Pinasco, Boqueron
Dept.
Little is known about the bionomics of
this mosquito. Swamps and riversides cov-
ered by aquatic plants, such as Pistia, may
shelter gravid females (Mitchell et al. 1987a).
It is assumed that these would also be suit-
able places for oviposition. In Argentina Cx.
delpontei was collected with chicken and
hamster baits, and one specimen from a
horse. Blood meal identification from en-
gorged females showed one fed on an am-
phibian, a few others on several mammal
species (mainly rodents), and two contained
a mixture of avian and mammal blood
(Mitchell et al. 1987a, b).
As mentioned above, several viral strains
were isolated from collected Cx. delpontei
in Northern Argentina. The Venezuelan
equine encephalitis virus subtype VI and
several Bunyaviridae were found in this
mosquito in the Chaco and Santa Fe Prov-
inces. These findings support it as a possible
vector in enzootic patterns of these viruses.
In addition, a disproportionate number of
virus isolates (18 of 40 in Chaco and 5 of
16 in Santa Fe), suggests the possibility of
transovarial transmission (Mitchell et al.,
1985, 1987a).
Taxonomic discussion. — Culex delpontei
was described by Duret (1968) who based
his description on adult males from Argen-
tina and Paraguay, including a specimen
which he later (1953) identified as Cx. par-
acrybda Komp. Later, Sirivanakarn and Ja-
kob (1981) partially described the female of
delpontei including a brief reference to the
cibarial armature. According to the classi-
fication proposed by Sirivanakarn (1982),
Cx. delpontei, Cx. paracrybda and Cx. pe-
reyrai, constitute the Paracrybda Group of
the Spissipes Section. Including the char-
477
acters recognized by Rozeboom and Komp
(1950) and Duret (1968) for Cx. paracryhbda,
the distinction among those three species
may be made as follows: Scutal and pleural
integuments and pleural setae are brown
with the scutal and pleural areas not sharply
contrasted in paracrybda, while there 1s a
well-marked contrast in delpontei and pe-
reyral where the pleural integument is yel-
low or yellowish. In pereyrai the pleural in-
tegument bears a pattern of dark spots on
the upper proepisternum, postspiracular
area, prealar knob, lower anterior surface of
the mesokatepisternum and an indistinct
spot on the upper corner of the mesokat-
episternum. In delpontei there 1s no pleural
pattern, and this species and pereyrai both
have shiny yellow to golden pleural setae.
The tarsi are entirely dark in de/pontei while
there are pale rings across joints of the tar-
someres and tarsomere 5 is entirely white
in paracrybda and pereyrai. Culex ocossa
Dyar and Knab also have pale pleural in-
tegument, but this species has a pattern of
darker areas on the prealar knob, postspi-
racular area and lower anterior surface of
the mesokatepisternum which separates it
from delpontel.
In the male genitalia the lateral plate (LP)
of both delpontei and paracrybda have only
the apical process which appears as a beak-
like hook at the apex in paracryhda and is
apically blunt and rounded in delpontei.
Apical, lateral and ventral processes of the
lateral plate are present in pereyrai. The dis-
tal arm of the distal division of the subapical
lobe (dSL) has four foliform setae clustered
at the apex in delpontei and paracrybda while
in pereyrai there is one apical foliform seta
and three subapical clustered foliform setae.
The proximal arm of the distal division of
the subapical lobe (dSL) is enlarged in de/-
pontei and narrowed in paracrybda. More-
over, the basal seta of the proximal arm is
foliform in pereyrai while it is saberlike in
delpontei and paracrybda. The slender setae
on the basolateral surface of the proximal
division of the subapical lobe (pSL) are
478
longer in de/pontei than in paracrybda and
pereyrai. The ninth tergal lobes are colum-
nar shaped in delpontei and paracrybda and
club shaped in pereyrai.
Culex (Melanoconion) pereyrai Duret
Figs. 7-9, 10, 12, 13
Culex (Melanoconion) pereyrai Duret
1967: 77 (type not seen; type locality Par-
aguay, Caaguazu, Cecilio Baéz; holotype de-
posited in the collection of the author specie
(Knight and Stone 1977)), Galindo 1969: 88
(tax.). Sirivanakarn 1982: 265 (tax.).
Female.—Head: Antenna dark, length
about 2.10 mm; flagellum normal, whorls
with 6 setae. Proboscis entirely dark-scaled,
length 1.63-1.81 mm, mean 1.73 mm.
Maxillary palpus entirely dark-scaled, length
0.27-0.37 mm, mean 0.31 mm, about 0.2
of proboscis length. Vertex (Fig. 7) with nar-
row falcate scales, predominantly pale whit-
ish laterally, dark in small median area, small
patch of broad appressed dingy white scales
along margin of eye; forked scales numer-
ous, dark; occipital region with some pale
whitish falcate scales. Cibarial armature
(Figs. 8, 9 and 10). Practically indistinguish-
able from that of Cx. delpontei. Surface of
cibarial bar not so noticeably folded and
sometimes with some thorn-shaped pro-
cesses on posterior border; cibarial teeth
double but posterior part frequently with
small thorn-shaped folds on ventral surface.
Thorax: Scutal integument brown, covered
with fine falcate shiny bronze-colored scales,
uniform in size, with some pale scales on
prescutellar area; scutal setae developed,
brownish black with reddish sheen; acros-
thical setae absent. Scutellar scales as scutal
scales, pale on lateral lobes and entirely dark
or mixed with some pale ones on median
lobe; lateral lobes each with 4 large setae,
median lobe with 6 long setae. Integument
of antepronotum similar to scutal integu-
ment, without scales and with some dark
setae. Postpronotal integument brown with
narrow dark scales similar to scutal ones;
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
posterolateral margin with 3—5 dark setae.
Pleural integument pale with distinct darker
spots on upper proepisternum, postspiracu-
lar area, prealar knob, lower anterior surface
of mesokatepisternum and an indistinct spot
on upper corner of mesokatepisternum.
Pleural setae yellowish with golden sheen,
brownish black on prealar knob: about 6—
11 upper proepisternal, 5,6 prealar, S—8 up-
per mesokatepisternal, 7-10 lower meso-
katepisternal, 4-8 upper mesepimeral and
1 lower mesepimeral. Pleuron with small
patch of pale spatulate scales on lower pos-
terior border of mesokatepisternum. Wing:
Length 2.80-3.13 mm, mean 2.96 mm;
scales dark; length of cell R, nearly 4.3 of
vein R,,;; cell M, nearly 0.8 length of cell
R,; subcosta intersects costa at level of fur-
cation of R,,;. Dorsal scaling: appressed
spatulate scales on costa, subcosta, R, R,,
R,.5, distal 0.8 of M,.5,M3.4, Cu, Cu,, Cu,
and proximal 0.5 of 1A; linear plume scales
on R,, R3,;, M and proximally on M,,,;
inclined narrow spatulate scales on R;, R,
and on distal 0.5 of 1A; remigium with ap-
pressed spatulate scales and 1,2 long distal
setae. Ventral scaling: appressed spatulate
scales on costa, subcosta, R,, R,,;, proximal
0.2 of R, and R;, M and proximal 0.2 of
M, 2; linear plume scales on proximal 0.5
of R,, proximally on R,,;, proximal 0.5 of
Cu,, Cu, and middle of 1A; inclined narrow
spatulate scales on distal 0.5 of R;, proximal
0.8 of R, and R,, R,,;, distal 0.8 of M,,>,
M,.4, distal 0.5 of Cu,, and distal part of
1A; Cu and proximal 0.5 of 1A without
scales. Halter: Entirely pale. Legs: Anterior
surface of forecoxa dark-scaled: anterior
surface of mid- and hindcoxae with longi-
tudinal patch of nearly colorless scales.
Antero- and posteroventral surfaces of
foretrochanter dark-scaled, mid- and hind-
trochanters with antero- and posteroventral
surfaces pale-scaled. Fore- and midfemora
mainly dark-scaled, posterior surface of fore-
femur with indistinct longitudinal stripe of
dingy pale scales, posteroventral surface of
midfemur with dingy pale scales, hindfemur
VOLUME 91, NUMBER 3 479
Cx.(Mel.) delpontei
ventral dorsal |
ventral dorsal Ct
| so Ss
rN
IX-Te
lateral
IsS
7
IX-Te
lateral
ete
Fig. 10. Female cibarial armature and genitalia of Culex delpontei and Cx. pereyrai. CA, cibarial armature.
Ce, cercus. CiB, cibarial bar. Ct, cibarial tooth. IsS, insular setae. PGL, postgenital lobe. UVL, upper vaginal
lip. UVS, upper vaginal sclerite. IX-Te, tergum IX. (Scale in mm.)
480 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Cx.(Mel.) delpontei
0.!
lateral
Vill-Te ——
BP
Figs. 11 and 12. Male genitalia of Culex delpontei and Cx. pereyrai. a, seta a of pSL. AeS, aedeagal sclerite,
b seta b of pSL. BP, basal piece. CSc, cercal sclerite. dSL, distal division of the subapical lobe. f, foliform seta.
Gc, gonocoxite. Gs, gonostylus. h, hooked seta of dSL. Isp, lateral setal patch. LP, lateral plate. Par, paramere.
VOLUME 91, NUMBER 3 481
Cx. (Mel.) pereyrai
lateral
HF wi | : | :
Fo
vill-Te
Ppr, paraproct. pSL, proximal division of the subapical lobe. s, saberlike seta. VIII-Te, tergum VIII. IX-te,
tergum IX. X-Te, tergum X. (Scale in mm.)
482
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
@ Culex (Mel) delpontei Durer
A Culex (Mel) pereyrai Duret
‘ San Pedro
\ Santa Fe } Corrientes
: }
| F
\ ae .
7 ii >
0 200 500 1000! ~ |
i 60° |
ae PAu @ I gquape [ |
Cananéia —} ; |
|
|
ATLANTIC | OCEAN |
J |
Fig. 13.
with complete dorsal stripe of dark scales
distally widening and expanding onto an-
terior and posterior surfaces at apex. All
tibiae with indefinite longitudinal dingy pale
stripe of scales on posterior surface. Ta-I,
and Ta-II, with indistinct longitudinal din-
gy pale stripe on posterior surface, more ev-
ident at apex, Ta-I,-Ta-I, and Ta-II,-Ta-II,
with indefinite pale bands on joints, Ta-I,
and Ta-II, paler, hindtarsus with distinct
narrow basal and apical white bands at joints
Distribution of Culex delpontei and Cx. pereyrai in South America.
of tarsomeres 1-4, Ta-III; entirely white.
Abdomen: Tergum I with median posterior
patch of dark scales, tergum II dark-scaled
with small basolateral patches of white
scales, terga HI-VIII dark-scaled with ba-
solateral patches of white scales, sometimes
appearing as narrow basal pale bands on
terga IV—VI, bands more evident on IV and
V. Sterna II-VII with broad basal white
bands, sternum II sometimes entirely white-
scaled; sternum VIII with lateral patches of
VOLUME 91, NUMBER 3
white scales, occasionally mixed with some
dark ones. Genitalia (Fig. 10): Tergum IX
narrow at middle, widened at posterolateral
margin to produce flat lobe bearing 7-9
slender setae. Upper vaginal lip distinct,
narrow; lower vaginal lip and insula indis-
tinct, insula with about 7-9 clustered setae.
Upper vaginal sclerite discrete, with in-
verted U-shape. Postgenital lobe short, dis-
tally with 10-17 setae on either side of mid-
line, mostly on ventral surface.
Male. —Like female except for sexual dif-
ferences as follow. Head: Antenna strongly
plumose; length about 1.77 mm. Proboscis
and maxillary palpus entirely dark, palpus
length about 2.62 mm, exceeding proboscis
tip by length of palpomere 5 and 0.9 of pal-
pomere 4; palpomeres 4 and 5 densely se-
tose; palpomere 3 with 6-8 apical setae. Ab-
domen: Tergum II entirely dark-scaled or
with few white scales on basolateral areas;
terga II-V with basal white bands; terga VI,
VII not examined; tergum VIII (ventral in
position) with basolateral white patches,
deep V-shaped median posterior emargi-
nation and several long bristles mixed with
shorter setae (Fig. 12); sternum II with white
scales mixed with some dark ones; sterna
IHI-V with basal white bands: sterna VI-
VIII not examined. Genitalia (Fig. 12):
Ninth tergal lobes small, slightly globose,
club-shaped, widely separated, bearing long
and slender setae. Gonocoxite stocky, outer
margin convex, inner moderately concave;
ventrolateral surface with strongly devel-
oped setae, mesal surface with small setae
in indistinct rows extending from base to
level of subapical lobe, lateral surface with
patch of sparse short and slender setae (Isp)
at proximal region up to level of subapical
lobe, proximal part of ventrolateral surface
with 1,2 scales; subapical lobe distinctly di-
vided, divisions approximated; proximal
division not divided, lengthened, with an
apical infundibular and hyaline expansion
partially covering insertions of setae a and
b (setae a and 4 long and sinuous) with
another hyaline, broad and hooked-falci-
483
form seta beyond middle, and 9-14 short,
slender and curved setae from base to level
of insertion of hooked-falciform seta; distal
division with 2 fairly well separated uneven
arms, proximal one stronger, bearing | long
hooked seta (/), | saberlike pointed seta (s)
and | shorter foliform seta (f) inserted on
prominent tubercle near middle of arm: dis-
tal arm slender, cylindric, bearing | stiff,
short, nearly saberlike seta (s) inserted near
middle of arm, 3 foliform setae (f) on distal
0.3 of arm, | longer nearly foliform seta (f)
inserted separately at tip ofarm. Gonostylus
slender, curved, moderately narrowed dis-
tally, crest slightly wrinkled on ventral sur-
face before apical snout; gonostylar claw
short, leaflike, apically broadest. Phallo-
some with lateral plates and aedeagal scler-
ites equivalent in length: aedeagal sclerite
broad, curved and falciform in lateral view,
more sclerotized area at angle of connection
with lateral plate and at mesal longitudinal
sclerotinization, dorsal end narrowly fused
to base of lateral plate; distal part of lateral
plate with apical, ventral and lateral pro-
cesses, apical process short, broad at base,
rounded at apex, ventral process curved lat-
erally, lateral process slender, nearly point-
ed and dorsolaterally directed, dorsal pro-
cess sclerotized and separated from aedeagal
sclerite margin by distinct angle; aedeagal
sclerites connected by dorsal aedeagal bridge.
Proctiger elongate; paraproct narrow dis-
tally, expanded basally, base articulated with
posterolateral margin of tergum X, crown
with row of 5—7 short simple blades. Cercal
sclerite long and narrow, sclerotized, broad-
est basally; 3,4 small cercal setae. Tergum
X large, rectangular, concavo-convex, dor-
sal surface concave.
Material examined.—Fifty-five speci-
mens examined from localities in Sao Paulo
State, Brazil, as follows: 25 22 (Experimental
Station, Pariquera-Acu, II.79, XII.79, [V.80,
VIII.80, XI.80, XII.80, 1.81, If.81, III.81,
IV.81, VI.81, 13; Bigua Road, Iguape, X.82,
X1.82, 12). 30 4¢ (Experimental Station,
Pariquera-Acu, 1.80, 11.80, 1V.80, XI.80,
484
11.81, I11.81, 1V.81, V.84, VII.84, V.85, 25;
Pariquera-Mirim, Pariquera-Acu, VII.84,
1.85, V.85, 3; Bigua Road, Iguape, X.76, 2).
Distribution and bionomics.— Until now,
Cx. pereyrai was known only from the male
holotype (type locality Paraguay, Caaguazu,
Cecilio Baéz). The collections reported here
extend its distribution to the localities in
southern Brazil (Fig. 13).
Nothing is known about the larval habi-
tats and little data are available about the
adult habits. The collections reported here
were made in patches of residual forests in
modified rural areas. A few specimens were
caught near houses.
Taxonomic discussion.— Culex (Mela-
noconion) pereyrai was described by Duret
(1967) based on one adult male from Par-
aguay. Additional specimens of this species
were unavailable until now. Sirivanakarn
(1982) placed this species in the Paracrybda
Group of the Spissipes Section where itself
constitutes the Pereyrai Subgroup. The dis-
tinction with Cx. delpontei Duret and Cx.
paracrybda Komp has already been dis-
cussed.
Since Cx. pereyrai has Ta-IIl; entirely
white, distinction must be made with other
species of the Spissipes Section that share
this character as Cx. epanastasis Dyar, Cx.
pedroi Sirivanakarn and Belkin, Cx. sac-
chettae Sirivanakarn and Jakob and CX. fae-
niopus Dyar and Knab. Including the char-
acters noted by Sirivanakarn (1978) and
Sirivanakarn and Belkin (1980), distinction
can be made as follows.
The pale pleural integument with a pat-
tern of darker spots, separates pereyrai from
epanastasis, pedroi and taeniopus which
have an entirely dark integument. From
sacchettae, which also has a pale pleural in-
tegument, distinction can be made by the
darker spots on the upper corner of the
mesokatepisternum and on the mesepime-
ron which pereyrai lacks. In addition, ‘ae-
niopus has a cluster of pale scales on the
upper corner of the mesokatepisternum,
while epanastasis and pedroi have clusters
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of white scales on the femoral apices, char-
acters absent in pereyrai.
The female cibarial armature of pereyrai
is quite similar to that of delpontei and Cx.
ocossa Dyar and Knab. The latter two
species are distinguished by the absence of
pale bands on the legs.
Finally, it is interesting to note that the
male genitalia of pereyrai is similar not only
to delpontei and paracrybda, but also to Cx.
adamesi Sirivanakarn and Galindo, Cx.
crybda Dyar, epanastasis, pedroi and Cx.
ribeirensis Forattini and Sallum. In all the
species the subapical lobe of the gonocoxite
has an infundibular process and one hook-
shaped seta on the proximal division and
the distal division is subdivided into two
arms, the proximal one with 3 setae and the
distal one with 4 setae.
ACKNOWLEDGMENT
This research was supported by Grant no.
86/0966-3, Fundacao de Amparo a Pes-
quisa do Estado de Sao Paulo-FAPESP and
Grant no. MVR-BR-2-84-6, National
Academy of Sciences, USA.
We express our sincere appreciation to
Daniel Marucci for preparing the scanning
electron micrographs.
LITERATURE CITED
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Duret, J. P. 1953. Las especies argentinas de Culex
(Melanoconion) (Diptera-Culicidae). Rev. Soc. Ent.
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. 1967. Dos especies nuevas de Culex (Mela-
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1968. Contribucion al conocimiento de los
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Galindo, P. 1969. Notes on the systematics of Culex
(Melanoconion) Taeniopus Dyar and Knab and
related species, gathered during arbovirus inves-
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89.
Harbach, R. E.and K. L. Knight. 1980 Taxonomists’
VOLUME 91, NUMBER 3
glossary of mosquito anatomy. Plexus Publishing,
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Daffner, C. B. Cropp, C. H. Calisher, R. F. Darsie,
Jr., and W. L. Jakob. 1987a. Arbovirus isola-
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the 1882-1983 epizootic of western equine en-
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36: 107-113.
Mitchell, C. J., T. P. Monath, M. S. Sabattini, H. A.
Christensen, R. F. Darsie, Jr., W. L. Jakob, and
J. F. Daffner. 1987b. Host-feeding patterns of
argentine mosquitoes (Diptera, Culicidae) collect-
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encephalitis. J. Med. Entomol. 24: 260-267.
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Rozeboom, L. E. and W. H. W. Komp 1950. A re-
view of the species of Culex of the subgenus Mela-
noconion (Diptera, Culicidae). Ann. Ent. Soc.
Amer. 43: 75-114.
Sirivanakarn, S. 1978. The female cibarial armature
of New World Culex, subgenus Melanoconion and
related subgenera with notes on this character in
subgenera Culex, Lutzia and Neoculex and genera
Galindomyia and Deinocerites (Diptera: Culici-
dae). Mosq. Syst. 10: 474-492.
Sirivanakarn, S. and J. N. Belkin 1980. The identity
of Culex (Melanoconion) taeniopus Dyar and Knab
and related species, with notes on the synonymy
and description of a new species (Diptera, Culic-
idae). Mosq. Syst. 12: 7-24.
Sirivanakarn, S. and W. L. Jakob 1981. Notes on the
distribution of Culex (Melanoconion) mosquitoes
in Northeastern Argentina (Diptera: Culicidae).
Mosq. Syst. 13: 195-199.
Sirivanakarn, S. 1982. A review of the systematics
and a proposed scheme of internal classification
of the New World subgenus Melanoconion of Cu-
lex (Diptera, Culicidae). Mosq. Syst. 14: 265-333.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 486-489
Book REVIEW
The Fleas of the Pacific Northwest, by
Robert E. Lewis, Joanne H. Lewis, and
Chris Maser. Oregon State University
Press, Corvallis, OR. [viii] + 296 pp.
1988. $49.95/cloth (alkaline paper). ISBN:
0-87071-355-8.
The fleas (order Siphonaptera) of the Pa-
cific Northwest are better known than those
of any other section of the continental
United States save California and Utah.
However, not since C. A. Hubbard’s Fleas
of Western North America (1947) has an
attempt been made to survey this region’s
rich fauna. Robert Lewis is a respected tax-
onomist who oversees a splendidly curated
flea collection at Iowa State University. To-
gether with his talented wife Joanne, he has
edited and published a helpful newsletter on
fleas since 1980. The present opus, initially
entitled The Fleas of Oregon and Their Im-
portance in Public Health (but expanded on
the eve of press time to include the sur-
rounding territory), owes its existence to the
Lewises’ long collaboration with Oregon
naturalist Chris Maser. It is at once a mon-
umental yet unaccountably incomplete
work.
Certainly the book invites perusal. Cloth-
bound in imitation leather, with lavish gold
lettering on the covers and spine, it opens
by paying due homage to Canada’s gentle-
man-scholar of Siphonaptera, the late
George Pearson Holland, whose beautiful
drawings of flea anatomy account for 198
of the 268 text figures in the book. There
follows an introduction to flea life history
and morphology that is concise but lucid
and includes a complete glossary of abbre-
viations used in the keys and illustrations.
There is also a brief but well-documented
discussion of flea-borne disease that covers
bacterial, rickettsial and viral agents, as well
as tungiasis and transmission of cestodes.
The body of this work (pp. 23-257) is
devoted to a systematic review of the 6 fam-
ilies, 54 genera, and 161 species and sub-
species of fleas that Lewis recognizes as
occurring 1n the Pacific Northwest. Surpris-
ingly, the boundaries of this region are never
defined—probably because the authors re-
alize that they are not dealing with a bio-
geographic unit—but the little dot distri-
bution maps that accompany each species
description imply that coverage embraces
not only all of Oregon and Washington but
also California north of San Francisco Bay,
the western strip of Idaho counties from that
state’s panhandle southward, and the whole
of Nevada except Clark County (i.e. every-
thing but Las Vegas). In no other work that
this writer has seen is “Pacific Northwest”
given such latitude. The maps themselves
are worthless, having been based on Maser’s
collecting itinerary and an arbitrary selec-
tion of literature records. The text’s best
features are its keys. I have tested the key
to families and find it easy to use, despite
the well-known absence of prominent fa-
milial characters in this order. Similarly,
keys to genera, species and subspecies em-
ploy structures that can be readily found by
a nonspecialist and that are defined and il-
lustrated in the introduction. The detailed
aedeagal terminology of Traub (1950, Field-
iana: Zool. Mem. |: 1-127) 1s referenced but
(fortunately for the layman) not discussed.
The book concludes with a depauperate bib-
liography of only 120 references (about half
the literature for this area), a woefully in-
complete host/flea index (Appendix I), a
lengthy discussion of how to mount fleas on
microscope slides coupled with an inade-
quate survey of collecting techniques (Ap-
pendix II), but a thoroughly orchestrated
index to all scientific names of flea taxa cited
in the text, including junior synonyms and
invalid names or combinations.
VOLUME 91, NUMBER 3
This work is fatally flawed on two counts,
of which the most damaging 1s its multi-
plicity of omissions. Contrary to Lewis’s as-
sertion (p. 18) that “there have been rela-
tively few publications dealing with the fleas
of the area,” the literature for the Northwest
is both extensive and diverse, ranging from
basic taxonomy to investigations of adult
and larval population dynamics, host spec-
ificity, and even population modeling.
Moreover, at least one graduate student at
Oregon State University successfully con-
ducted an intensive study of fleas associated
with small mammals throughout Oregon’s
vast Willamette Valley. When even the
theses on file at the school responsible for
publishing this book are not consulted, I can
only conclude that I am witnessing an ex-
ercise in arrogance rather than objective sci-
ence.
A review 1s not the forum for revising an
imperfect work, but some effort must be
made to plug a few of this book’s numerous
gaping lacunae. Thus, under Atyphloceras
multidentatus (C. Fox) (pp. 66-67), we find
no mention of the remarkable ecology of
this hystrichopsyllid in the northern Sierra
Nevada of Plumas County, California
(Jameson and Brennan, 1957, Ecol. Mono-
gr. 27: 45-54). There, A. multidentatus is
abundant on the deer mouse Peromyscus
boylii (Baird) and common on P. manicu-
latus (Wagner) in brushy field situations.
However, in coniferous forests (where P.
boylii rarely occurs), A. multidentatus is sel-
dom found on P. maniculatus but is fre-
quent on the microtine Clethrionomys cal-
ifornicus (Merriam). Therefore, while P.
maniculatus 1s a satisfactory host, the degree
to which it is parasitized appears to be large-
ly dependent on habitat. Similar observa-
tions have been made by Russian siphon-
apterists, but perhaps Lewis et al. do not
have access to translations. Further, 4. muil-
tidentatus is a capable vector of plague, hav-
ing been successfully infected experimen-
tally (Eskey and Haas, 1939, Publ. Hlth.
Rep. 54: 1467-1481) and found able to
487
transmit the disease (Prince, in Wayson,
1947, Publ. Hlth. Rep. 62: 780-791). This
flea should also be added to Lewis’s host/
flea index under the shrew Sorex vagrans
Baird and the vole Microtus canicaudus
Miller (Robbins, 1983, J. N.Y. Entomol.
Soc. 91: 348-354, and references cited
therein).
Lewis’s statement (p. 82) that Catallagia
charlottensis (Baker) is “not a conspicuous
element” in the flea fauna of the Northwest
is contradicted by Hubbard (op. cit.) and
other specialists familiar with this area
(Vernon J. Tipton, Brigham Young Uni-
versity, personal communication). In west-
ern Oregon, C. charlottensis can be collected
by the thousands from a broad range of small
mammals during all months of the year.
Once again, Jameson and Brennan’s obser-
vations on ecological segregation are ig-
nored, nor is there any indication of the role
of Catallagia species in plague transmission
(for which, see Anonymous, 1950, Publ.
Hlth. Rep. 65: 614; Allred, 1952, Gr. Basin
Nat. 12: 67-75; Beck, 1955, Brigham Young
Univ. Sci. Bull. 1: 1-37; and Kartman and
Prince, 1956, Amer. J. Trop. Med. 5: 1058—
1070). Add C. charlottensis to Lewis’s host/
flea index under Sorex vagrans, and add C.
sculleni Hubbard under Microtus canicau-
dus.
Another mistreated hystrichopsyllid is
Corrodopsylla curvata (Rothschild), which
is erroneously held to ‘show no particular
peak in seasonal abundance” (pp. 137-138)
when in fact samples from 3 different test
sites in western Oregon all showed spring
and early winter population maxima
(Faulkenberry and Robbins, 1980, Ento-
mol. News 91: 93-101). For some unknown
reason, Lewis cites Hansen’s (1964, Gr. Ba-
sin Nat. 24: 75-81) collection of C. curvata
from the Steens Mountain area of Harney
County, Oregon, without adding the host,
Sorex palustris Richardson, to his host/flea
index. Corrodopsylla curvata is also a reg-
ular accidental on Microtus canicaudus in
western Oregon.
488
Under Hystrichopsylla occidentalis Hol-
land (p. 64), we find the claim that “there
are no published records of this species being
associated with the transmission of dis-
ease.” However, as a rule, members of the
genus Hystrichopsylla are capable and effi-
cient plague vectors and frequently are in-
volved in maintaining epizootics (Eskey and
Haas, op. cit.; Prince, in Wayson, op. cit.;
Anonymous, 1950, Publ. Hlth. Rep. 65:
1174; and Kartman and Prince, op. cit.).
Add this species to the host/flea index under
Microtus canicaudus.
The ubiquitous ceratophyllid Aetheca
wagneri (Baker) is mistakenly missing from
Lewis’s host/flea index under Sorex va-
grans, Microtus canicaudus, and M. town-
sendii (Bachman), despite the existence of
scores of collections from these animals.
Worse is the fate of another ceratophyllid,
the northern (or European) rat flea, Nosop-
syllus fasciatus (Bosc), which was dismissed
from Maser’s surveys because “we were
concerned with the parasite fauna of en-
demic birds and mammals and no effort was
made to collect domestic rats and mice” (p.
200). In disowning this species, which has
long been recognized as a capable and mod-
erately efficient plague vector (McCoy, 1911,
Publ. Hlth. Bull. 43; Bacot and Martin, 1914,
J. Hyg. 13: 423-439; Bacot, 1915, J. Hyg.
14: 774-776; Eskey and Haas, 1940, Publ.
Hlth. Bull. Wash. (254): 1-83: and Bur-
roughs, 1947, J. Hyg. 45: 371-396) and
which may have played a secondary role in
maintaining the great 14th century Euro-
pean plague pandemic (Pollitzer, 1954,
Plague), the authors nullify their principal
justification for writing, namely that “‘pub-
lic health workers are exposed to fleas and
other disease vectors during the course of
their work” (Introduction, p. 1). Needless
to say, N. fasciatus should appear in the
host/flea index under several native mam-
mals, among them Peromyscus manicu-
latus, Microtus canicaudus, and M. town-
sendii.
Hubbard’s (op. cit.) claim that the lep-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
topsyllid Peromyscopsylla selenis (Roths-
child) is most numerous in the fall, winter
and spring has been repeatedly corrobo-
rated (Robbins, op. cit.). Lewis’s single late
summer peak (p. 143) is probably the result
of sampling error. Add Microtus canicaudus
to the host list on p. 142.
Readers seeking information on flea ecol-
ogy will have to look to other texts, for Lew-
is et al. make no effort to summarize the
wealth of published data on flea populations
in the Pacific Northwest. Only a smattering
of lost opportunities can be listed here,
among them: correlation of flea population
fluctuations with meteorological variables,
host phenology, or host attributes (e.g. sex,
size, age); application of uniform indices of
extensity and intensity, such as those intro-
duced by Janion (1968, Ekol. Polska (A) 16:
561-606) and refined by Lundqvist (1974,
Entomol. Scandinavica 5: 39-48); analysis
of flea sex ratios (an entire subset of ecto-
parasitological literature has developed
around this one subject!); and computerized
population modeling (Robbins and Faulk-
enberry, 1982, Entomol. News 93: 70-74).
Beyond these multifarious errors of omis-
sion lies a second, subtler failing: the ab-
sence of what in another context has been
called the “life spark.”” Those of us who have
had the unspeakably good fortune to sit
mesmerized at the feet of Robert Traub, the
world’s foremost authority on Siphonap-
tera, will know what I mean. Traub’s 85
peerless publications on fleas, which his
spiritual brother the late Karl Jordan called
“these jolly insects,” are alive with anec-
dotes drawn from all corners of the earth.
Who can tell how many graduate students
have been inspired by Traub’s work? His
writing is musical; there 1s Ketélbey in the
fauna of Egypt, Grofé in America. Of all the
omissions in Lewis’s book, it is this music —
this romance—that I miss most. Had Maser
taken the time to collect mammal nests and
extract their fauna using Berlese-Tullgren
funnels (a technique that was completely
overlooked), he would have found, among
VOLUME 91, NUMBER 3
other wonders, ticks in all life history stages,
phoretic deutonymphs of uropodid mites
attached to adult fleas via anal pedicels, in-
numerable flea larvae awaiting correlation
with adults of their species, nidicolous cara-
bid and staphylinid beetles that prey on flea
larvae and thereby regulate larval popula-
tions, and exquisite aphodiine scarabs (e.g.
Aphodius cribratulus A. Schmidt, A. hae-
morrhoidalis Linnaeus, and A. pardalis
LeConte) that complete their metamor-
phoses only in the nest chambers of bur-
rowing mammals. Music.
I said at the outset that this is a monu-
mental book: it is large, eye-catching, and
will probably achieve some recognition on
the entomological horizon. But if flea texts
could aspire to the status of skyscrapers, this
one would remind me of nothing more than
489
the World Trade Center in Lower Manhat-
tan, into whose brazen towers the demoli-
tion plans have already been built. And if I
were to summarize my feelings about this
book, I could do no better than defer to the
brilliant architecture critic Paul Goldberger
of The New York Times who, in addressing
the dilemma of the towers, remarked “if
they say anything at all ... it is that we
retreat into banality when the opportunity
comes for greatness.”
Richard G. Robbins, Department of
Health and Human Services, Public Health
Service, National Institutes of Health,
National Institute of Allergy and Infectious
Diseases, % Department of Entomology,
Museum Support Center, Smithsonian In-
stitution, Washington, D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 490-492
Book REVIEW
Outlines of Entomology. R. G. Davies
1988. Seventh Edition. Chapman and
Hall, London, New York; vii + 408 pp.
$37.50 Softcover, ISBN 0 412 26680 6;
$75.00 Hardcover, ISBN 0 412 26670 9.
This textbook is the newly-revised, con-
densed version of Imms’ General Textbook
of Entomology by Richards and Davies, and
replaces the sixth edition of ““Outlines.’’ The
seventh edition has been considerably ex-
panded, with new material on modes of life
in insects, a new chapter on the biology of
insect populations, and a new section on
injurious insects and pest management. The
book is designed to be an introduction to
the field of entomology for biological sci-
ence students and those in related agricul-
tural fields.
The introductory chapter briefly discuss-
es the general characteristics of insect struc-
ture, diversity, life histories, and habits. A
chapter on insect structure and function
deals with basic external and internal anat-
omy and morphology, insect flight, respi-
ration, circulation, and reproduction. The
following chapter treats development and
metamorphosis. A chapter on classification
and biology treats insects at the ordinal level.
A diagnosis is given for each order, followed
by a summary of the principal structural
features that are used in identification and
classification, habits, and habitats where
representatives are found. For larger orders,
major subdivisions and groups of families
are also discussed. A chapter on the evo-
lutionary relationships of insects discusses
the fossil record of insect groups, and sum-
marizes theories regarding relationships be-
tween orders or groups of orders, with em-
phasis on recent information. A chapter on
modes of life in insects deals with how in-
sects make a living, including the structural,
functional and behavioral adaptations to dif-
ferent environments and lifestyles, with sec-
tions on insects and plants, pollination, in-
sects and micro-organisms, predators,
parasitoids and parasites, aquatic insects,
mimicry and protective coloration, and so-
cial insects. This is followed by a chapter
on the biology of insect populations which
treats characteristics of population growth,
regulation, life tables, key factor analysis,
resource partitioning, predator-prey rela-
tionships, food webs and the trophic struc-
ture of communities, diversity, stability and
succession, and genetic diversity. A chapter
on the economic importance of insects gives
representative examples of the kinds of pest
insects, the nature of damage they cause,
methods of control by insecticides, biolog-
ical control, and integrated pest manage-
ment. A classified bibliography is subdivid-
ed by subject area, and includes references
that the reader can consult for additional
information. References up to 1987 are in-
cluded.
This textbook is an excellent introduction
to the field of entomology, and fulfills the
objective of the author to present the basics
of entomology, no mean task fora discipline
so large and diverse. The information pre-
sented is generally up-to-date. The text 1s
very well-written, readable, and the illus-
trations, with one or two exceptions, are
clearly labelled and easy to interpret, and
complement the text well. The classified
bibliography is especially useful, being cur-
rent, comprehensive, and providing an ex-
cellent beginning point for readers who wish
to pursue particular topics further. The clar-
ity of the text and illustrations make this
book suitable for readers with no special
scientific background, as well as for students
in the sciences.
The major criticism I have of this book
is the continued use of an archaic system of
higher classification. The author presents an
VOLUME 91, NUMBER 3
excellent discussion of the higher classifi-
cation of insect orders in the section on evo-
lutionary relationships, but justifies the use
of a long-outmoded system of ordinal clas-
sification, unwisely in my opinion, with the
statement that there is “substantial dis-
agreement” about some features of insect
phylogeny. In fact, there is substantial
agreement about many aspects of insect
phylogeny, and the classification used in a
modern textbook on entomology should re-
flect a “phylogenetic” classification when
there is a firm basis for it. While one might
quibble over the use of particular ordinal
names, I would prefer the use of Microco-
ryphia and Thysanura for the bristletails and
silverfish, respectively, with the names Ar-
chaeognatha and Zygentoma reserved for
the respective infraclasses to which these
orders are assigned. I also prefer the use of
Notoptera in preference to Grylloblattodea
for the mountain crawlers. The Isoptera
should be subsumed under the Dictyoptera,
along with the mantids and roaches, con-
sistent with current thinking, particularly in
view of the author’s statement that “the
members of this order [Isoptera] are struc-
turally very similar to the Blattaria.”” The
author acknowledges that the Neuroptera,
as treated in this text, is a “rather hetero-
geneous order... .”” Both the Megaloptera
(fishflies, dobsonflies and alderflies) and the
Raphidioptera (snakeflies) represent dis-
tinct lineages and should be treated as sep-
arate orders.
Inevitably, in a text of this breadth, there
will be minor problems of correct usage of
names, inadvertant misstatements, and
misleading generalizations. A sample of
these is given without intending to detract
from the general excellent quality of the text.
There are remarkably few typographical
errors in the text: Plannipennian for Plan-
ipennian (p. 65), Borellia for Borrelia (p.
315), Toxirhynchites for Toxorhynchites (p.
337), and hominovorax for hominivorax
were the only ones I found. Several old
names are still found in the text: Psi/opa =
49]
Helaeomyia (p. 2), Taeniorhynchus = Man-
sonia (p. 76), Culex fatigans = C. quinque-
fasciatus (p. 319), Reesimermis nielseni =
Romanomermis culicivorax (p. 337). The
use of Tahanus rusticus Linnaeus to illus-
trate a valid binomen (p. 116) is inappro-
priate since rusticus is now placed in Atylo-
tus. Use of another binomen, such as
Tabanus bovinus Linnaeus, to illustrate a
valid, original Linnean binomen would be
more satisfactory, with Atylotus rusticus
(Linnaeus) as an example of a species name
transferred from its original genus to
another. I would prefer the use of the term
binominal nomenclature instead of bino-
mial nomenclature (p. 116) to minimize
confusion with mathematical terminology.
There are a few minor errors of fact. Cy-
clorrhaphan Diptera have 3 preimaginal in-
stars, not 4 (p. 106), unless one includes the
puparium, but this is not clear in the text;
Grylloblattodea (Notoptera) are carnivo-
rous and predators, not omnivores (p. 132),
as clearly demonstrated by Pritchard and
Scholefield (1978); Stomoxys is inappro-
priately called the “biting house fly” (p. 165),
but more precisely the stable fly later in the
text; Sarcophaga is not a blow fly (p. 214),
but a flesh fly; visceral leishmaniasis occurs
in the New World as well as in the Old
World (p. 323).
A few generalizations by the author are
inappropriate or misleading to the reader.
Although many dipteran larvae pass through
3 or 4 instars (p. 166), most orthorrhaphan
Brachycera larvae have several additional
instars; the statement that “syrphids feed
largely on aphids” (p. 221) should refer more
appropriately to the Syrphinae; not only do
the males and females of Stomoxys feed on
blood (p. 230), so do both sexes of all Sto-
moxyinae (Haematobia, Haematobosca
etc.).
Occasionally, the author has failed to in-
corporate useful new information into the
text or uses curiously archaic terminology.
The discussion on the origin of wings in
insects (p. 25) is very superficial and in-
492
cludes none of the recent evidence on al-
ternative theories of the origin of wings in
insects, touting instead the thoroughly dis-
credited paranotal lobe theory. This 1s es-
pecially puzzling since the author includes
pertinent references by Kukalova-Peck, for
example, in the selected bibliography. Use
of the term “Protozoa” throughout the text
instead of Protista (or Protoctista) does not
reflect current usage of Kingdom-level clas-
sification in introductory biology texts or
the scientific literature. The treatment of
morphogenesis (p. 1 1 1-112) is seriously out-
of-date. No mention is made of the dra-
matic advances in understanding the mor-
phogenesis of Drosophila, with information
on pattern formation, the mosaic vs. regu-
lative models, importance of diffusible mor-
phogens and chemical gradients, progress
zones, homeoboxes, and the role of regu-
latory gene centers in specifying differentia-
tion of the developing embryo. The Order
Trichoptera has about 11,000-12,000 de-
scribed species, not “less than 3,000” (p.
175). The term “‘Siphunculata” (p. 230) for
the sucking lice (Anoplura) is archaic. Use
of “Culicinae” (p. 230 and elsewhere) to
refer to mosquitoes 1s archaic, since the Dix-
idae and Chaoboridae are generally consid-
ered separate families. The pathogens caus-
ing sleeping sickness in humans in Africa
are Trypanosoma brucei gambiense and T.
brucei rhodesiense. The brucei group 1s now
associated with human trypanosomiasis, not
nagana (p. 322). In addition to the blood-
sucking insects discussed (p. 230-232), the
curious blood-feeding Lepidoptera of the
genus Calyptra (Noctuidae) should be men-
tioned. A more precise, succinct definition
of pest management in the section on in-
tegrated pest management would be useful
to the uninitiated reader. The definition giv-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
en in Rabb and Guthrie (1970, p. 2-3) is an
excellent one.
Two general comments are in order. Some
of the common names of insects used in the
text may be unfamiliar to entomologists in
North America, who are more familiar with
the Entomological Society of America (ESA)
list of common names. Perhaps this is in-
evitable in a text written outside of North
America. The cost of the hardcover edition
of this text ($75.00) is far too high for an
introductory text, and far higher than the
prices of other introductory entomology
texts in hardcover available in North Amer-
ica ($39.00-$48.00). The quality of this text
is not sufficiently superior to others of its
genre to justify paying double the price of
others. The paperback edition, however, is
priced comparably ($37.50) with other in-
troductory texts.
In summary, this is an excellent intro-
duction to the field of entomology. It is well-
written, clearly illustrated, and highly rec-
ommended as a text for those interested in
a general introduction to insect biology,
classification, the importance of insects in
the natural environment, and the impor-
tance of insects to humans.
LITERATURE CITED
Pritchard, G. and P. Scholefield. 1978. Observations
on the food, feeding behaviour, and associated
sense organs of Grylloblatta campodeiformis
(Grylloblattodea). Can. Ent. 110: 205-212.
Rabb, R. L. and F. E. Guthrie, eds. 1970. Concepts
of Pest Management. North Carolina State Uni-
versity, Raleigh, 242 pp.
John F. Burger, Department of Entomol-
ogy, Nesmith Hall, University of New
Hampshire, Durham, New Hampshire
03824.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, pp. 493-495
Book REVIEW
Silent Spring Revisited. G. J. Marco, R.
M. Hollingworth and W. Durham [eds.].
American Chemical Society, Washing-
ton, D.C., 1987, xvili + 214 pp., $17.95.
That Rachel Carson’s book Silent Spring
was controversial is unrefutable. That all
the predictions and statements were correct
is questionable. That the book is important
is a fact. Silent Spring Revisited 1s based
upon a symposium organized to “‘address
the issues that Rachel Carson raised and to
focus on their pertinence for the past, pres-
ent and future.” The contributors provided
an excellent cross-section of opinions on all
sides of the issues—though the bias of the
authors usually was evident.
Shirley Briggs, of the Rachel Carson
Council, Inc., starts with a discussion of the
vision and legacy of Ms. Carson. Briggs dis-
cusses Carson’s motives, the reaction to Si-
lent Spring, and then goes on to the com-
prehensive view. She states that, “We at
Rachel Carson Council have yet to be shown
a valid example” [of inaccuracies or mis-
takes]. G. K. Kohn of Zoecon Corp., in
chapter 10 points out some of the inaccur-
acies, though recognizing Carson’s achieve-
ments. Kohn’s chapter is an excellent bal-
ance to the conflicting values of a pragmatic
approach to agricultural productivity and
the preservation of the environment. John
Moore of EPA reviews the evolution of gov-
ernment regulations. He states that, “the ex-
tensive requirements for current regulation
do not always provide clear insight as to
appropriate action. For example, the tech-
nical capability to routinely analyze in parts
per million, billion, trillion or quadrillion
clearly surpass the toxicologists’ and other
scientists’ ability to confidently interpret
human or environmental risk.’ Unfortu-
nately EPA gets blamed for not doing enough
by environmental zealots and for being over
zealous by the producers and users of pes-
ticides. It may be of interest to note that the
National Agricultural Chemical Associa-
tion estimates the cost and time of devel-
oping a pesticide, from the test tube to the
market, to be $40 million and 8 years. Most
of this is to fulfill EPA testing requirements.
C. F. Wilkinson of Cornell University
continues the discussion in a chapter on the
science and politics of pesticides. He very
succinctly states, ““For the last decade and
a half an emotional belief, often verging on
hysteria, has existed in the United States
that society is being not so slowly poisoned
by pesticides and other products of modern
chemical technology. Despite the facts that
we live longer and generally enjoy a much
better quality of life than at any other time
in our history, we have become a society
consumed with fear and obsessed by the
risks in our lives.”” Wilkinson then discusses
the regulatory decision-making process;
toxicology —a science and an art; toxicologi-
cal evaluations; assessment of acute toxic-
ity; carcinogenic risk assessment; science
or politics; scientific uncertainty and regu-
latory conservatism; and public perception
and acceptance of risk.
Chapters 4-7 assess the toxicity of pes-
ticides to aquatic organisms (by Nimo, Cop-
page, Pickering and Hansen of EPA), the
impact of pesticides on ground water con-
tamination (by Carsel and Smith of EPA)
and the impact of pesticides on bird pop-
ulations (by R. J. Hall of Fish and Wildlife
Service). To quote Nimo et al., “while rid-
ding the world of weeds, weevils and web-
worms/Be careful of fins, fur, and feathers.”
These chapters are primarily facts and fig-
ures on the use and effects of pesticides on
fish, water and birds, and are not contro-
versial in nature.
J. E. Davies and R. Doon in chapter 7
review the human health effects of pesti-
494
cides and discuss acute pesticide poisoning,
including suicides; cancer risk; residues; re-
sistance; and disposal. The beneficial as-
pects of pesticides in relation to public health
are neglected. Rosen and Grech present the
evaluation and impact of analytical chem-
istry of pesticides. This chapter is a straight-
forward discussion of the various assay
methods, from bioassay to gas chromatog-
raphy to mass spectrometry.
Virgil H. Freed of Oregon State Univer-
sity, in chapter 9, covers the global use of
pesticides and concerns. Freed points out
that despite the continuing and spreading
controversy over the use of pesticides, the
use of such chemicals has increased an-
nually by 4-5% on a global basis since the
publication of Silent Spring. Most of the
pesticides are used by developed nations.
He mentions a study in which pesticide poi-
soning was examined in a number of coun-
tries. He estimates a rate of 2.9-4.8 per
100,000 persons, with a mortality rate of
5.5%. This comes to 0.16-0.26 persons per
100,000. If this figure is compared to the
world wide morbidity and mortality for ma-
laria, I believe most developing nations
would consider the benefit far greater than
the risk.
G. K. Kohn presents the other side of the
coin in chapter 10, agriculture, pesticides
and the American chemical industry. He
discusses the demographic nature of Amer-
ican agriculture; the conflicting values; pes-
ticides produced before and after Si/ent
Spring, technology and farm income; spe-
cial problems of the Third World and use
of agrochemicals; and ends with a reassess-
ment of Si/ent Spring. He notes that his
rereading Si/ent Spring in no way altered
his original assessment that the volume is
important and would affect the publics’ at-
titude relative to the use of pesticides and
their impact on the environment.
David Pimentel of Cornell University, in
chapter 11 questions, “is Si/ent Spring be-
hind us?”. He starts with, “Fewer pesticide
problems during the past two decades” and
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
then follows with, ‘Increased pesticide
problems during the past two decades.” Ba-
sically he indicates that the amount of chlo-
rinated insecticide residues in the environ-
ment has declined since most of the
chlorinated insecticides have been banned.
However, pesticide production has in-
creased, and that some of the OP pesticides
may present a greater human risk. He then
goes on to discussing poisonings in humans,
animals and bees. His figures on pesticide
effects on crops seem to be at odds with
Kohn’s comments. Pimentel estimates a
negative impact on crops of about $70 mil-
lion annually. Kohn states that the addi-
tional wealth created by the technology to
be $9,125,000,000. Pimentel states that 37%
of all crops is lost annually to pests in spite
of the combined use of pesticidal and non
chemical controls. If he is correct, what
would it have been without the use of agro-
chemical technology? He concludes that
pesticides will continue to be effective pest
controls but the challenge now is to find
ways to use them judiciously to avoid many
of the environmental hazards and human
poisonings that exist today.
Marco et al. conclude the volume with an
overview of the symposium. They point out
that Si/ent Spring led society to evaluate the
new technologies in terms of risk versus
benefits. Whereas lower levels of pesticides
can now be detected, the true significance
of these small amounts of pesticide residues
is often quite uncertain. Toxicity evaluation
is a complex process with great uncertain-
ties, and it does not provide answers that
are nearly as precise as the analytical tech-
niques. The diverse opinions and opposing
views between manufacturer-user and en-
vironmentalist are clearly seen in the chap-
ters of this book. There are fewer farmers
and less labor available, yet more mouths
to feed, therefore there seems no likelihood
of completely eliminating chemical controls
in the near future. Rachel Carson was right
in many respects, and in a fewer respects
she was wrong. Nature, not just humans,
VOLUME 91, NUMBER 3
generates its share of carcinogens and other
poisons. Nature and humans both use
chemicals to their own advantage. The hu-
man life span is increasing and birds still
sing. What Rachel Carson accomplished was
to set in motion a philosophy of using all
495
tools in controlling pests, not relying exclu-
sively on chemicals.
Eugene J. Gerberg, /nsect Control & Re-
search, Inc., 1330 Dillon Heights Ave., Bal-
timore, Maryland 21228.
PROC. ENTOMOL. SOC. WASH.
91(3), 1989, p. 496
ENDANGERED SPECIES
On January 6, 1989, the U.S. Fish and Wildlife Service published an extensive ‘notice of review’ on United
States animal species including insects believed to be candidates for protection under provisions of the U.S.
Endangered Species Act, to be extinct, or to be previously considered names that are now known to be taxo-
nomically invalid or to represent taxa now thought to be in no danger of extinction. The Service is seeking
further information on these candidates or about other species that might be considered.
General comments or comments on species from more than one Service Region should be addressed to the
Director, U.S. Fish and Wildlife Service (FWE-SE), Washington, D.C. 20240. Specific comments on one or
more species from a specific Region should be addressed to the appropriate Regional Director listed below.
Copies of the notice may also be obtained from these addresses.
Region 1. CA, HI, ID, NV, OR, WA, Trust Territories.
Regional Director (FWE-SE)
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Regional Director (FWE)
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CONTENTS
(Continued from front cover)
STAINES, C. L. JR.—A new genus and species of Hispinae (Coleoptera: Chrysomelidae) from
Central America
TOGASHI, I.—Japanese sawflies of the family Blasticotomidae (Hymenoptera: Symphyta) ...
TRIPLEHORN, C. A.—A new species of Neobaphion Blaisdell, from Idaho (Coleoptera:
Tenebrionidae)
WARTHEN, J. D. JR.—Neem (Azadirachta indica A. Juss): Organisms affected and reference
list update
WEATHERBY, J. C., J. C. MOSER, R. J. GAGNE, and H. N. WALLACE—Biology of a pine
needle sheath midge, Contarinia acuta Gagné (Diptera: Cecidomyiidae), on Loblolly pine . .
WHEELER, A. G. JR.—Grypotes puncticollis (Homoptera: Cicadellidae), a Palearctic pine-feed-
ing leafhopper new to North America
ZAPPAROLI, M.—Notes on Pleurolithobius of Turkey (Chilopoda: Lithobiomorpha)
BOOK REVIEWS
BURGER, J. F.— Outlines of Entomology
GERBERG, E. J.—Silent Spring Revisited
ROBBINS, R. G.—The Fleas of the Pacific Northwest
ENDANGERED SPECIES NOTICE
VOE. 911 OCTOBER 1989 NO. 4
(ISSN 0013-8797)
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BOHART. R.M.—Review of Maricopodynerus (Hymenoptera: Eumenidae) ................
BOLDT, P. E.—Host specificity studies of Sto/as fuscata (Klug) (Coleoptera: Chrysomelidae) for
the biological control of Baccharis salicifolia (R.&P.) Pers. (Asteraceae) ................ 502
BUTLER, L.—Observations on Meganola spodia Franclemont (Lepidoptera: Noctuidae) with
AIdesenpuonl othe mairevanvan ii e4 Henk ae Na fetisnich seiise DE Kuosics dilsck be ste cede 615
BYERS, G. W.—Homologies in the wing venation of primitive Diptera and Mecoptera...... 497
CAVE, R. D. and M. J. GAYLOR—Longevity, fertility, and population growth statistics of
Telenomus reynoldsi (Hymenoptera: Scelionidae) .. 0.2... 0. cece eee eee ede e sees 588
DAVIS, D. R.—An exceptional fossil amber collection acquired by the Smithsonian Institution 545
GRISSELL, E. E.—Megastigmus floridanus (Hymenoptera: Torymidae), newly discovered in
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HEADRICK, D. and R. D. GOEDEN—Life history of Pteromalus coloradensis (Ashmead)
(Hymenoptera: Pteromalidae) a parasite of Paracantha gentilis Hering (Diptera: Tephritidae)
We CLES ISCICACADITU Ae he eis fees Gah Acedia eats WOE Y iG che keen bewt cones 594
HURD, L. E., P. E. MARINARI, and R. M. EISENBERG —Influence of temperature and
photoperiod on early developmental rate of Tenodera sinensis Saussure (Mantodea: Man-
HIGH) Rcm TM Mek APs ONE Oe et ee wee se, Pe OS aes oo Sl ge ee ee ee $29
KROMBEIN, K. V.—Systematic notes on some Bethylidae from Botswana: Pristocerinae (Hy-
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MACDONALD, J. F.— Review of Nearctic Metachela Coquillet, with description of a new species
(Pipteras Empididae: ‘Hemerodromimnae)} ..8 3.884%. 325.6% Se ah SLT a et 513
MANGUIN, S.—Sexual dimorphism in size of adults and puparia of Tetanocera ferruginea
Fallén (Diptera: Sciomyzidae) ....... Da pak bitte: AYR SAE Be gods BORIS o Recrctyetattes MARNE eye cote 523
SCHAUFF, M. E.—A new species of Horismenus (Hymenoptera: Eulophidae) parasitic on the
lesser cornstalk borer, Elasmopalpus lignosellus (Lepidoptera: Pyralidae) ............... 534
STRICKMAN, D. and J. PRATT—Redescription of Cx. corniger Theobald and elevation of
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(Continued on back cover)
THE
ENTOMOLOGICAL SOCIETY
OF WASHINGTON
ORGANIZED MARCH 12, 1884
OFFICERS FOR 1989
F. CHRISTIAN THOMPSON President NORMAN E. WooDLEY, Jreasurer
JEFFREY R. ALDRICH President-Elect WARREN E. STEINER, JR. Program Chairman
RICHARD G. RosBINs, Recording Secretary GEOFFREY B. WHITE, Membership Chairman
JOHN M. KINGSOLVER, Corresponding Secretary F. EUGENE Woop, Past President
HirAM G. LAREw, Editor
RosertT D. Gorpbon, Associate Editor
Publications Committee
REBECCA F. SURDICK T. J. HENRY GEORGE C. STEYSKAL
B. V. PETERSON
Honorary President
Curtis W. SABROSKY
Honorary Members
Louise M. RUSSELL ALAN STONE THEODORE L. BISSELL
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tution, Washington, D.C. 20560.
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STATEMENT OF OWNERSHIP
Title of Publication: Proceedings of the Entomological Society of Washington.
Frequency of Issue: Quarterly (January, April, July, October).
Location of Office of Publication, Business Office of Publisher and Owner: The Entomological Society of
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ington, D.C. 20560.
Editor: Hiram G. Larew, B-470, ARS, USDA, Beltsville, MD 20705.
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THIS PUBLICATION IS PRINTED ON ACID-FREE PAPER.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 497-501
HOMOLOGIES IN WING VENATION OF PRIMITIVE
DIPTERA AND MECOPTERA
GEORGE W. BYERS
Department of Entomology, The University of Kansas, Lawrence, Kansas 66045.
Abstract.—On the basis of evidence from tracheation of wing veins, distribution of
macrotrichia on veins, corrugation of the wing, and comparison of wings of Diptera to
those of Mecoptera, the media is four-branched and the anterior cubitus is unbranched.
Key Words:
Dipterists are concerned with the inter-
pretation of wing venation because they use
venational consistencies and differences for
recognition of taxa at all levels. According-
ly, it is awkward that conflicting views of
venational homology are currently in use.
In this paper, support is offered for one of
those interpretations.
J. H. Comstock and J. G. Needham pro-
duced the most widely accepted system of
nomenclature and homologies of wing veins
of insects (Comstock 1918). Familiar to most
entomologists, it need not be reviewed in
detail here. In diagramming the hypothet-
ical primitive or generalized insect wing,
Comstock and Needham noted that alter-
nating veins are elevated or convex (+), or
depressed, concave (—) with respect to the
plane of the wing. This corrugation, or flut-
ing, had been pointed out earlier by Adolf
(1879) and Redtenbacher (1886): the costa
is +, the subcosta strongly —, anterior
branch of the radius strongly +, the sector
(Rs) and its branches —, anterior media +
(but absent in extant orders of insects), pos-
terior media and its branches —, anterior
cubitus +, posterior cubitus —, and so on.
K. G. A. Hamilton (1971: 429) noted that
much of the fluting seen in wings of Pa-
leoptera is reduced in wings of Neoptera,
except in the costal area (C, Sc and R,) and
wing venation, Tipulidae, Tanyderidae, Mecoptera
along the cubitus. Nevertheless, the relative
elevation and depression of veins can in
most cases be readily determined.
The problem with which I am concerned
involves the branching of the posterior me-
dia (M) and the anterior cubitus (Cu,). Com-
stock and Needham determined that vein
M primitively was dichotomously four-
branched, M,, M;, M, and M,, all concave,
reaching the wing margin. They believed
that Cu was primitively two-branched rath-
er near its base, with the anterior cubitus
(Cu,) being strongly convex and the poste-
rior branch (Cu,) concave. Vein M,,, or
M, 1s, in their system, connected to Cu, by
the m-cu cross-vein. The anterior cubitus
may itself be divided into Cu,, and Cu,, (see
Snodgrass 1935: 216).
A problem arose for Comstock and Need-
ham with the observation that what ap-
peared to be the posterior branch of M was
moderately to strongly convex in Diptera.
Since all branches of M were supposed to
be concave, they initially said that this vein
is either an anterior branch of the anterior
cubitus (Cu,,) or Cu,, + My. Their eventual
conclusion, however, was that in Diptera
the media has only three branches and that
this convex vein is a branch of the cubitus;
that is, M, is always absent in wings of Dip-
tera.
498 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
\\ (« WA) (¢ uA)
Mees My
Vi
M Z j
DSSS SPO C See
Maya =< ~ Ms
2
1
Cu,
Figs. 1-2. Pseudolimnophila inornata (Osten Sacken). 1. Wing with venation labelled according to the
Alexander-Tillyard system, with branches of M and Cu also labelled (in parentheses) according to the Comstock-
Needham system. 2. Portion of venation showing distribution of macrotrichia on certain veins. Scale line = |
mm.
This interpretation was challenged only
eight years later, by R. J. Tillyard (1926),
who declared that M is four-branched in
flies and that Cu, is unbranched. Almost
simultaneously, C. P. Alexander had reached
the same conclusion, based on his study of
Tipulidae and other primitive Diptera (e.g.,
Tanyderidae). A student of Comstock at
Cornell University and at first a follower of
the Comstock system (from 1919 to 1925),
Alexander became convinced that the me-
dia has in Tipulidae four branches, as in the
hypothetical, generalized pattern, and that
what he had been calling the “anterior de-
flection of Cu,” in crane flies was really the
m-cu cross-vein.
The Alexander-Tillyard interpretation has
also been widely accepted, perhaps in part
because it holds that M in primitive Diptera
(that is, Diptera in which the venation is
not greatly reduced or modified) has the form
that Comstock and Needham claimed for
primitive insects generally. This view was
followed, for example, by D. H. Colless and
D. K. McAlpine (1970) in “The Insects of
Australia.” In contrast, the “Manual of
Nearctic Diptera” (J. F. McAlpine et al.
1981) adopted the Comstock-Needham in-
terpretation. Figure | illustrates these con-
flicting interpretations of M and Cu, in the
wing of a crane fly.
The problem is basically this, I think: Is
the vein (labelled m-cu in Fig. 1) connecting
what appears to be the posterior branch of
M with Cu, in fact a cross-vein, or is it the
basal part of a branch of Cu,? What kinds
of evidence can be obtained to support one
interpretation or the other?
Some similar problems have been solved
by examination of tracheal branching. In
Diptera, however, the veins are already
clearly established in the pupal wing (the
wing sheath) before tracheae enter the wing.
In crane flies of the genus Jipula (Fig. 3),
one trachea enters the radius and branches
at the arculus, one branch continuing along
the radius and through R, to the stigmal
area, the other proceeding along the cubitus
(Cu,). In several pupae of Tipula trivittata
Say examined (and in a few of other species),
the cubital trachea did not give off any
branch into the vein in question (which I
call m-cu). In some individuals, perhaps
preserved too soon, the trachea stopped be-
VOLUME 91, NUMBER 4
499
Figs. 3-4. Tipula trivittata Say. 3. Pupal wing (wing sheath) showing complete, slightly raised venation
(shaded) and tracheal branches chiefly in R, and Cu,. 4. Wing, indicating the convexity (+) or concavity (—) of
certain veins. Scale lines = 1 mm.
fore reaching m-cu; in others, it passed m-
cu without branching. In one specimen of
Tipula ignobilis , however, a short branch
entered m-cu for about one-fourth of the
length of that vein. I find this evidence in-
teresting but ambiguous.
A few morphologists and taxonomists
have observed that, in various taxa, macro-
trichia occur on certain longitudinal wing
veins but not on cross-veins. This contrast
is easily seen in Mecoptera, for example.
Numerous individuals of several genera of
Tipulidae (in all three subfamilies) were in-
spected with this in mind. While macro-
trichia usually occur on both M and its
branches and on Cu,, although often only
sparsely, they are almost never seen on m-cu.
One or two machrotrichia were found on
the cubital end of m-cu in a very few in-
dividuals. Pseudolimnophila illustrates well
the contrasting presence of macrotrichia on
M and Cu, and absence from m-cu (Fig. 2).
But macrotrichia are absent as well from
the basal, cross-vein-like portions of M,+,
and M,, so this evidence may not be very
convincing.
Homology of veins from one order to
another has been determined largely by the
previously mentioned corrugation, or flut-
ing. As a general observation, two main lon-
gitudinal veins—R, and Cu, —are ordinarily
strongly convex in virtually all neopterous
orders of insects. Such primitive flies as 77p-
ula (Tipulidae) and Protoplasa Tanyderi-
dae) show this condition (Figs. 4, 5). In both,
a strongly concave, darkened line closely
paralleling the convex Cu, but not reaching
the wing margin has been interpreted as Cu,
or the cubital fold, lying in the cubital fur-
row. Also in both, the vein that I suppose
is M, is clearly convex, while m-cu is much
less so and may be neither + nor —. But I
cannot accept the view of Comstock and
Needham that the convexity of “M,”’ must
be interpreted as this vein’s being a branch
of Cu,. My reason is this: In all crane flies
and tanyderids examined, vein R, or the
combined R,,; 1s moderately to strongly
convex, while the expectation is that it
should be concave. Comstock and Need-
ham, who held that the branches of the ra-
dial sector are concave, apparently took no
notice of this inconsistency. W. Hennig
(1969: 311,376) did, and suggested that the
apparent R, could include a remnant of the
convex anterior media. However, it is gen-
erally accepted, I think, that the anterior
media is absent in all modern orders of in-
sects. My interpretation of the convexity of
both R, and M, is that there is a structural
“need” for corrugation to give some relative
rigidity and strength to the respective parts
500 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 5-7.
M,
My
a
My,
Wing venation, showing the convexity (+) or concavity (—) of certain veins. 5. Protoplasa fitchii
(Osten Sacken) (Diptera:Tanyderidae). 6. Panorpa communis Linnaeus (Mecoptera:Panorpidae). 7. Nanno-
chorista neotropica Navas (Mecoptera: Nannochoristidae). Scale lines = 1 mm.
of the wing (which are otherwise rather
plane). I have no idea whether this is so;
but the point is that a convex M, is no more
remarkable than a convex R,, and no less
to be expected.
Occasionally, in local populations, one
finds numerous individuals that have the
branches of M variously dislocated (Byers
1961). Such aberrations appear to have a
genetic basis (cf. Laven 1957). While the
apparent M, 1s involved in these disloca-
tions, although less so than M, ;, and even
m-cu may be affected, the anterior cubitus
is not. This suggests that there are, to some
extent, different factors influencing the de-
velopment of M and Cu,.
Finally, there 1s the evidence from com-
parative anatomy, which is more convinc-
ing to me than any of the evidence already
cited. A survey of the orders of insects to
determine the most likely origin of the Dip-
tera leads us quickly to the Mecoptera. This
order, the fossil record of which goes back
to the lower Permian, displays the combi-
nation of characters most closely resem-
bling that found in primitive Diptera—
characters of general body structure, exter-
nal genitalia of both sexes, antennae,
mouthparts and wing venation. According-
ly, we may gain from examining the branch-
ing of the media and cubitus in Mecoptera
some understanding of equivalent venation
in Diptera.
Comstock and Needham thought that the
venation in Panorpa (Fig. 6), a common
genus of Mecoptera, agreed with their gen-
VOLUME 91, NUMBER 4
eralized plan, that is, Rs with four branches,
M with four and Cu separated basally into
Cu, and Cu,, but Cu, with no further branch-
ing. In the medial field, the venation of Pan-
orpa 1s Strikingly like that of the tanyderid
Protoplasa (compare Figs. 6 and 5), with
the same convexity and concavity of cor-
responding veins. Cu, in Tanyderidae, as in
Tipulidae, is only weakly developed as a
dark line in the cubital furrow behind Cu,.
Among the Mecoptera, Nannochorista
(Fig. 7) is most like the Diptera; the head
and mouthparts are not elongated as 1n most
Mecoptera, the mandibles are reduced in
the adult, and, in wing venation, M is fused
basally with Cu,. In this genus, as in Me-
coptera generally, the anterior cubitus is un-
branched. Moreover, in all Mecoptera the
veins R,; and M, are both distinctly convex,
as in the primitive Diptera.
From the evidence presented, I conclude
that in the Tipulidae (probably the most
primitive family of extant Diptera) and in
the Tanyderidae (which I consider also very
primitive flies, although their phylogenetic
position is still disputed) the media (M) typ-
ically has a dichotomous division, each
branch again dividing dichotomously, hence
four terminal branches. Further, I conclude
that the anterior cubitus (Cu,) in primitive
Diptera, at least, is not branched.
There is great variation in the branching
and the basal fusion of veins in the order
Diptera. I have not studied this in detail.
Nevertheless, I find the interpretation of ve-
nation based on the Tipulidae and Tany-
deridae, and compared to that of the Me-
coptera, as reasonable as any other for all
Diptera. An unbranched Cu, could be
thought of as one more character linking
Mecoptera and Diptera (as Antliophora of
Hennig, or Mecopterida of Boudreaux) and
differentiating this group (which also in-
cludes the Siphonaptera) from the Am-
phiesmenoptera, or Trichopterida.
501
ACKNOWLEDGMENTS
I thank Mr. Bryan N. Danforth for his
help in rearing pupae of Tipu/a and pre-
paring their wings for study and in verifying
the patterns of corrugation of veins in var-
ious genera. Charles D. Michener and Fenja
Brodo read an early version of this paper
and made useful suggestions.
LITERATURE CITED
Adolf, G. E. 1879. Uber Insektenfliigel. Nova Acta
Leop. Carol. 41: 215-291.
Boudreaux, H. B. 1979. Arthropod Phylogeny with
Special Reference to Insects. John Wiley & Sons,
New York. 320 pp.
Byers, G. W. 1961. The crane fly genus Dolichopeza
in North America. Univ. Kansas Sci. Bull. 62:
665-924.
Colless, D. H. and D. K. McAlpine. 1970. Diptera,
pp.656-740. Jn Waterhouse, D. F., et al., The In-
sects of Australia. Melbourne Univ. Press, Carl-
ton, Victoria. xii + 1029 pp.
Comstock, J. H. 1918. The Wings of Insects. Com-
stock Publishing Co., Ithaca, New York. 430 pp.
(based upon a series of papers by Comstock and
J. G. Needham, 1898-1899, on “The Wings of
Insects,” in the American Naturalist).
Hamilton, K. G. A. 1971. The insect wing, part I.
Origin and development of wings from notal lobes.
J. Kansas Entomol. Soc. 44:421-433.
Hennig, W. 1969. Die Stammesgeschichte der Insek-
ten. W. Kramer & Co., Frankfurt am Main. 436
pp.
Laven, H. 1957. Vererbung durch Kerngene und das
Problem der ausserkaryotischen Vererbung bei
Culex pipiens . 1. Kernvererbung. Zeitschr. Indukt.
Abstammungsund Vererbungslehre 88:443-447.
McAlpine, J. F. 1981. Morphology and terminolo-
gy—adults, pp. 9-63. Jn McAlpine, J. F., et al.,
Manual of Neartic Diptera. Vol. 1. Monogr. 27,
Res. Branch, Agriculture Canada. 674 pp.
Redtenbacher, J. 1886. Vergleichende Studien ber
das Fliigelgeader der Insekten. Ann. K. K. Natur.
Hofmus. 1: 153-232.
Snodgrass, R. E. 1935. Principles of Insect Mor-
phology. McGraw-Hill Book Co, New York. 667
pp.
Tillyard, R. J. 1926. The Insects of Australia and
New Zealand. Angus and Robertson, Sydney. 560
pp.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 502-508
HOST SPECIFICITY STUDIES OF STOLAS FUSCATA (KLUG)
(COLEOPTERA:CHRYSOMELIDAE) FOR THE BIOLOGICAL CONTROL OF
BACCHARIS SALICIFOLIA (R. & P.) PERS. (ASTERACEAE)
PAUL E. BOLDT
Grassland, Soil and Water Research Laboratory, Agricultural Research Service, USDA,
808 East Blackland Road, Temple, Texas 76502.
Abstract.—The leaf feeder Stolas (Anacassis) fuscata (Klug), from Argentina, was studied
in quarantine as a potential biological control candidate of seepwillow, Baccharis salicifolia
(R.&P.) Pers. in the United States. No-choice tests were conducted with first-instar larvae
on 34 plant species in 7 tribes of Asteraceae and 3 additional families. Pupation occurred
primarily on B. salicifolia and to a lesser degree on Gymnosperma glutinosum (Spreng.)
Less. and Aster subulatus Michx. No-choice tests were also conducted with adults on 23
plant species representing 5 tribes and 12 genera of Asteraceae. Leaf consumption and
longevity were greatest on B. salicifolia and A. subulatus, and oviposition occurred only
on these species. Stolas fuscata apparently has a high degree of host specificity. This insect
does not attack any economically important plant, and the risk of it feeding on any species
other than B. salicifolia is small.
Key Words: Insecta, Coleoptera, Chrysomelidae, Sto/as, Asteraceae, Baccharis, seep-
willow, host specificity, biological control
Seepwillow, Baccharis salicifolia (R.&P.)
Pers. (formerly = gl/utinosa Pers.) (Astera-
ceae) is an undesirable woody phreatophyte,
2 to 3 m in height, and has few, if any,
economic benefits. This shrub contributes
little to stream bank stability. Dense thick-
ets of seepwillow impede water flow and
contribute to channel migration, overflow
and increased sedimentation (Horton 1959,
Parker 1972). Seepwillow has little or no
value to wildlife except for the cover it pro-
vides and is not listed as a food source by
Martin et al. (1951) for any wildlife species
in the United States. No birds are closely
associated with this shrub (Engel-Wilson and
Ohmart 1978, Anderson and Ohmart 1984).
Stolas (Anacassis ) fuscata (Klug) is a po-
tential biological control agent for seepwil-
low. This beetle, a leaf feeder in both larval
and adult stages, occurs only in Argentina,
Brazil, Paraguay (Blackwelder 1946) and
Uruguay (Guido et al. 1956). Originally de-
scribed in the genus Cassida, fuscata was
later placed in Anacassis by Spaeth (1913).
Most recently Anacassis has been recog-
nized as a subgenus of the large neotropical
genus Stolas (Hincks 1952). This status is
maintained in Seeno and Wilcox (1982) and
followed in the present paper. Buzzi (1975)
and McFadyen (1987), working in South
America, continue to recognize Anacassis
as a valid genus separate from Stolas.
I conducted host specificity tests on S.
fuscata from Argentina as part of a project
to study the biological control of seepwil-
low. Tests by McFadyen (1987) indicated
VOLUME 91, NUMBER 4
that both Sto/as fuscata fuscata from Brazil
and S. fuscata unicolor (Burmeister) from
Argentina feed only on 3 species of Bac-
charis and the related Baccharidastium tri-
plenerve (Lessing) Cabrera.
The populations of S. fuscata used in these
tests are probably identical to S. fuscata uni-
color in Argentina and very similar to S.
fuscata fuscata in Brazil. The male and fe-
male genitalia of both subspecies were com-
pared by H. Cordo (pers. comm.). He found
no consistent differences in structure. My
specimens were identified as S. fuscata by
both Z. Buzzi (pers. comm.) and R. E. White
(pers. comm.), but neither identified a sub-
species. Possible differences between sub-
species in host feeding were reported by
McFadyen (1987). In 1979 several thou-
sand adults of S. Fuscata fuscata were re-
leased in Queensland, Australia, for the bi-
ological control of B. halimifolia. They
apparently fed on the plant but did not be-
come established in the field (McFadyen
1987). Stolas fuscata unicolor from Argen-
tina was also brought into Queensland but
did not survive in quarantine. The subspe-
cies name is not used in this manuscript for
lack of confirmation of its validity. Al-
though physiological differences may exist
between the populations, morphological dif-
ferences are not sufficiently unique to iden-
tify subspecies.
MATERIALS AND METHODS
Larvae and adults of S. fuscata were col-
lected on Baccharis salicifolia by H. Cordo
near Arroyito, Neuquen; Realico, La Pam-
pa; San Rafael, Mendoza; and Pedro Luro,
Buenos Aires, Argentina, during the months
of February and March from 1985 to 1987
and air-freighted to the Insect Quarantine
Facility, USDA-ARS, Temple, Texas. A
colony of this insect was maintained in
quarantine on excised leaves of potted B.
salicifolia or leaves periodically collected in
the field near Laredo, Texas.
All experiments were in the quarantine
facility at Temple, Texas. The room was
503
held at temperatures of 22 to 26° C, relative
humidity of 40 to 60% and a 12:12 h L:D
photoperiod under artificial light. Leaf con-
sumption was determined by placing a |
mm plastic grid over the leaf and counting
the squares or with a Li-Cor Model Li 3000
leaf area meter before and after feeding. Be-
cause an entire experiment could not be done
at the same time, there was a control treat-
ment of seepwillow each time any plant was
tested. Plants used in the tests were collected
in Texas and held out-of-doors except for
Baccharis pilularis DC., which was collect-
ed in California; those plants selected were
species closely related to seepwillow or
economically important. Plants were al-
lowed to flower when possible, and voucher
specimens were deposited in the permanent
collection of the Grassland, Soil and Water
Research Laboratory, Temple, Texas. In-
sect specimens were deposited in the Na-
tional Museum of Natural History, Wash-
ington, DC.
Larva host tests.—No-choice tests were
conducted by holding a single unfed, 0- to
48-hour old larva in a small (9.0 cm diam.)
petri dish on freshly excised leaves of a se-
lected plant species until death or pupation
occurred. Leaves were replaced every sec-
ond or third day. The amount of leaf con-
sumed, stage of development and number
of days survived were recorded. A replica-
tion consisted of 10 larvae per plant species.
Each of the 34 plant species was replicated
at least 3 times.
Adult host tests.—In the first of two adult
no-choice tests, a single unsexed, unfed, 3-
to 5-day-old adult was held in a small petri
dish on freshly excised leaves of each test
plant. Leaves were replaced every second or
third day, and the amount of leaf consumed
was recorded until death occurred. A rep-
lication consisted of | adult per plant species.
Each of the 15 plant species was replicated
10 times.
The second adult no-choice test was sim-
ilar except that 15, 2- to 6-day-old adults
were held on each plant species in a large
504 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Average feeding and longevity of larvae of S. fuscata on leaves of various plants.
Plant Species No. of Larvae Leaf Consumption (cm?) _ Longevity Time (days)*
ASTERACEAE
Astereae
Baccharis salicifolia (R. & P.) Pers. 110 52.6 + 12.4 NGS =9223)
Aster novae-angliae L. 30 0.1 + 0.1 4.0 + 0.3
A. subulatus Michx. 40 10.7 74:2 9.0 + 2.4
A. sp. (form Michaelmas) 30 0 4.0 + 0.9
Baccharis bigelovii Gray 40 0.6 + 0.7 5:3 26D
B. brachyphylla Gray 40 0 3330
B. halimifolia L. 40 0.6 + 0.4 5251+ 9 210)
B. neglecta Britt. 40 5:5) +.8.8 Tal 24)
B. pilularis DC. 30 O:3: 0:2 3:32 19.
B. pteronioides DC. 30 0.1 + 0.1 3.4 + 0.6
B. sarothroides Gray 30 0 32a Oe
Chrysothamnus nauseosus (Pall.) Britt. 30 OM 021 4.2+ 1.4
Ericameria austrotexana M. C. Johnst. 30 0 357/-E. 014
Grindelia lanceolata Nutt. 30 D4 323) Te Mipct Ae?)
Gutierrezia microcephala (DC.) Gray 30 0.2 + 0.4 5:2 see3all
Gymnosperma glutinosum (Spreng.) Less. 50 = 115.9 8:9FsE 75)
Haplopappus tenuisectus (Greene) Blake 30 0 3:4) ES
Isocoma coronopifolia (Gray) Greene 30 0 3.9 + 0.5
Solidago altissima L. 30 0.5 + 0.9 43) =F 166
Eupatorieae
Brickellia laciniata Gray 30 OF) 1081 Dia 2/0
Heliantheae
Helianthus annuus L. 40 OHs= OF 2.9 + 0.4
Rudbeckia hirta L. (form “‘gloriosa’’) 30 0 DED e
Viguiera dentata (Cav.) Spreng. 30 0) 325) E1016
Zinnia acerosa (DC.) Gray 30 Oo On 3570 10.5
Helenieae
Callistephus chinensis (L) Nees 30 0.2 + 0.4 Die 22
Gaillardia pulchella Foug. (form “‘grandiflora’’) 40 0 3.2.0 U7
Tagetes patula L. 40 0 Sele=ENO a],
Anthemideae
Artemisia filifolia Torr. 30 0 3:0°2- 1019.
Chrysanthemum cinerariifolium Vis. 30 0 352) == OW)
C. morifolium Ramat. 30 Of2'='0:3 2:6:=E10:5
Cynareae
Carduus macrocephalus Desfontaines 30 0 229221087,
Cichorieae
Lactuca sativa L. 30 0 3.0 + 0.4
ASCLEPIADACEAE
Asclepias viridis Walt. 30 0 Pra | aoa) ey
GRAMINAE
Zea mays L. 30 0 27 3403
*® Mean + standard deviation.
petri dish (14.1 cm diam). Leaf consump- except for Jsocoma wrightii (Gray) Rydb.
tion, survival time, the number of eggs laid, which was replicated twice and seepwillow
and percent hatched were recorded. Each of _ which was replicated 5 times. All adults were
the 10 plant species was replicated 3 times dissected after death to determine sex.
VOLUME 91, NUMBER 4
505
Table 2. Percentage of larvae of S. fuscata surviving on various test plants and number of days to pupation.
Larval survival
f Sth Instar Pupae
No. of
Test Plant Larvae No % No, % Days to Pupation*
Baccharis salicifolia 110 89 80.9 83 IS) [9:1 + 3:2
Aster subulatus 40 4 10.0 2 5.0 20:3 + 2.75
Baccharis neglecta 40 2 5.0 0 0
B. halimifolia 40 1 2.5 0 0
Callistephus chinensis 30 1 33 0 - 0
Gymnosperma glutinosum 50 8 16.0 6 12.0 Died V6
*Mean + standard deviation.
RESULTS
Larva hosts tests. — Significant feeding and
survival occurred on several species in the
tribe Astereae (Asteraceae) and one species,
Callistephus chinensis (L.) Nees, in the tribe
Helenieae (Table 1). More feeding occurred
on seepwillow than on either Gymnosperma
glutinosum (Spreng.) Less. or Aster subu-
latus Michx.; and only minor feeding oc-
curred on Baccharis neglecta Britt. and Gu-
tierrezia microcephala (DC.) Gray. Few
larvae on non-host plants, including the 5
other species of Baccharis, survived beyond
the second instar, although some larvae sur-
vived without feeding for up to 4 days.
Neonate larvae fed and developed to the
fifth instar only on the plants listed in Table
2. Normal pupation occurred for 75.5% of
the larvae restricted to seepwillow but only
for 12.0% of the larvae on G. glutinosum
and 5.0% of the larvae on 4. subulatus. Lar-
vae on other species died without pupating.
Adult host tests.—In the first adult no-
choice test, feeding occurred primarily on a
few plants in the tribe Astereae. Adults on
seepwillow consumed over 9 times more
foliage than those on /socoma coronopifolia
(Gray) Greene, the next most fed on plant
(Table 3). Mean longevity on seepwillow
was 26.1 days, but it was longer on J. co-
ronopifolia, G. glutinosum and G. micro-
cephala. Survival on Antennaria fallax
Greene was nearly as long as on seepwillow
although the adults apparently did not feed.
In the second adult no-choice test, sig-
nificantly greater longevity, feeding and ovi-
position were recorded on seepwillow than
on any other species in the test (Table 4).
The 39 females on seepwillow lived 34.3
days longer than the 19 females on Aster
subulatus Michx. and laid over four times
the number of eggs per female. The mean
percent egg hatch was 79.4% on seepwillow
as compared with 32.5% on 4. subulatus.
No eggs were laid on any other species.
DISCUSSION
Stolas fuscata appears to be a suitable bi-
ological control agent for seepwillow. Hosts
of S. fuscata are limited to a few plants in
the subtribe Baccharidinarum, as shown by
larval and adult multiple-choice tests on 52
plant species in 25 families in both Argen-
tina and Brazil (McFadyen 1987). My tests
of larvae on 34 additional plants and adults
on 23 plants in no-choice tests also indicate
a high probability that if S. fuscata were
released in the United States it would es-
tablish only on seepwillow.
The risk that S. fuscata might survive on
the perennial shrub G. g/utinosum is min-
imal. No eggs were laid, adults fed poorly
and only 12.5% of the larvae pupated on G.
glutinosum. This plant is more closely re-
lated to the genus Gutierrezia than to Bac-
charis and has few recorded benefits (Mar-
tinez 1959).
Eggs were laid by S. fuscata only on seep-
willow and A. subulatus. Calculations from
these tests, conducted under restrictions of
506
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 3. Feeding and longevity of adults of S. fuscata on leaves of various plants.
Amount
No. of | Amount Consumed Consumed/Day Longevity
Plant Species Adults (cm?)" (cm?)" (days)*
ASTERACEAE
Astereae
Baccharis salicifolia 10 95.1 + 73.9 33.0 + 19.0 26.1 + 17.9
B. neglecta Britt. 10 3:2) ==16.6 0.2 + 4.0 1723: 1016
B. halimifolia L. 10 Ost +310:2 —> 14.9 + 12.1
B. pilularis DC. 10 |e mee ep) OF AO 12.8 + 6.5
Ericameria austrotexana M. C. Johnst. 10 0 0 20.5 + 14.5
Grindelia squarrosa (Pursh) Dun. 10 3:07 729 On = 10 23:9) ES 1529
Gutierrezia microcephala (DC) Gray 10 NTE) Wee le 0.1 + 0.0 3226%-E wide
Gymnosperma glutinosum (Spreng.) Less. 10 Sd. + 1033 Oplg=s-240 S1OGES DE
Tsocoma coronopifolia (Gray) Greene 10 10:2" 116.6 0225053 38.2 + 19.6
Solidago altissima L. 10 0 -? 16.5 + 14.8
Anthemideae
Artemisia filifolia Torr. 10 Od! 22220 —> 16.0 + 13.0
Chrysanthemum morifolium Ramat. 10 JRO j=E92 —? PAT aI)
Cynareae
Centaurea macrocephala Pushk. 10 OSE O'S —? 19.9 + 14.6
Helenieae
Gaillardia pulchella Foug. 10 0 0) ldeSe-ee les
Inuleae
Antennaria fallax Greene 10 0) 0 21.0 + 16.9
“Mean + standard deviation.
»’ Amount consumed is less than 0.01 cm/day.
the quarantine, show that a hypothetical fe-
male fed leaves of seepwillow will lay 325
eggs which will result in 99.7 first generation
females (Table 5). However, one female and
offspring fed only on A. subulatus will pro-
duce less than | female. This difference is
the result of increased oviposition on seep-
willow and high mortality of eggs and larvae
on A. subulatus. Only 5 % of the neonate
larvae on A. subulatus survived to the pupal
stage.
Occasional feeding on A. subulatus, if it
does occur, should not be considered det-
rimental because this species is an annual
Table 4. Longevity and oviposition of S. fuscata on various species of Astereae in no-choice tests.
Longevity 2
No. of Leaf Consumption
adults Male Female per adult (cm?)* Eggs per Female*
Baccharis salicifolia TS 56:5. 30i1 » 169:9'-e 38.2) 59.4 .sEnli726 — 2+ 163.6
Aster subulatus Michx. 45; 44.9) += DIST 63526) 22-8..5 Silene Ou 2.0 + 64.8
A. ericoides L. 45 92 5.2 13 74,-655.9 0.4 + 0.4 0
A. novae-angliae 45 9.1 + 3.4 9.5 + 3.8 0:5 = 10:5 0
A. praealtus Poir. 452 2053) = 1BI0 29 18O 12a eS 0
A, spinosus Benth. 45 BSE 1S OM 15 OF 5081 0
A. texanus Burgess 45 7.6: = 246 13.4 + 8.6 OVSe=E O87 0
Ericameria laricifolia (Gray) Shinners 45 10.7 + 6.3 14.8 + 15.7 Ole 1051 0
Gymnosperma glutinosum (Spreng.) Less. 45 13.0 + 8.3 OOS 52. 0.2 03 0
Tsocoma wrightii (Gray) Rydb. 30 5.7 Ee 16 526) a= 2 O: = O11 0
* Means + standard deviation.
VOLUME 91, NUMBER 4
Table 5. Survival of S. fuscata when parent and offspring are fed B. salicifolia or A. subulatus.
B. salicifolia
Stage Sample size No. entering stage
Egg 39 325
Larva 110 258
Pupa 83 194.8
No. of females (51.2%) 99.7
native weed with no economic value. It is
abundant in ditches, margins of ponds,
streams, and poorly drained areas of the
United States from North Carolina to Cal-
ifornia (Correll and Johnston 1979). It is
widespread in Central America and is con-
sidered a weed in Australia and western Asia
(Faust and Strang 1983).
There are no Baccharis species listed in
the Federal Register of 1986 as being threat-
ened or endangered (Anon. 1986).
Biological control of seepwillow offers a
relatively inexpensive, environmentally-
compatible alternative to both mechanical
and chemical control. A comparison of the
climate where S. fuscata occurs in Argen-
tina with the climate associated with the
geographical distribution of seepwillow in
North America suggests that the insect may
adapt to the arid southwestern United States
and probably move into northern Mexico
(Walter et al. 1975). The release of S. fuscata
would represent one of the first deliberate
attempts to control a native plant with an
exotic insect in the continental United
States. The expected result is a reduction in
the density of this plant but not its elimi-
nation (Johnson 1984, Harris 1986).
ACKNOWLEDGMENTS
I thank Thomas Robbins, Lisa Bruggman
and Janie Cisneros of the Grassland, Soil
and Water Research Laboratory, USDA-
ARS, for their technical assistance; Richard
White, Systematic Entomology Laboratory,
USDA-ARS; and Zundir Buzzi, Universi-
dade Federal do Parana, Brazil, for identi-
fications of S. fuscata; and Hugo Cordo, Bi-
ological Control of Weeds Laboratory,
507
A. subulatus
% survival Sample size No. entering stage % survival
79.4 19 72. 32.5
ee) 40 23.4 5.0
= D 1.17 =
0.6
USDA-ARS, Huringham, Argentina, for
supplying S. fuscata for this study. Mention
ofa proprietary product does not constitute
an endorsement or recommendation for its
use by the USDA.
LITERATURE CITED
Anderson, B. W.andR. D. Ohmart. 1984. Vegetation
Management Study for the Enhancement of Wild-
life along the Lower Colorado River. Final Report.
U.S. Bur. Reclamation, Boulder City, NV.
Anonymous. 1986. Endangered and Threatened
Wildlife and Plants. SOCFR17.11 and 17.12. U.S.
Fish and Wildlife Service. Dept. Interior.
Blackwelder, R. 1946. Checklist of the Coleopterous
Insects of Mexico, Central America, the West In-
dies and South America. Bull. U.S. National Mu-
seum 185: 742-743.
Buzzi, Z. 1975. Contribuicao ao conhecimento da
biologia de Anacassis fuscata . Rev. Brasil. Biol.
35: 767-771.
Correll, D. and M. Johnston. 1979. Manual of the
Vascular Plants of Texas. pp.1558-1562. Univ.
Texas, Dallas.
Engel-Wilson, R. and R. Ohmart. 1978. Assessment
of Vegetation and Terrestrial Vertebrates along the
Rio Grande between Fort Quitman, Texas and
Haciendita, Texas. International Boundary and
Water Commission, El Paso, Tx.
Faust, W. and R. Strang. 1983. Important Weeds of
the World. p.88. Bayer, A. G., Leverkusen.
Guido, A. and A. Ruffinelli. 1956. Primer catalogo
de los parasitos y predatores encontiadoes en el
Uruguay. Fac. Agron. de Montevideo, Uruguay.
Bol. No. 32.
Harris, P. 1986. Environmental impact of agents used
for the biological control of weeds. Paper pre-
sented to the Annual Meeting, entomol. Soc. Amer.
Reno, NV. Dec. 1986. unpub. ms.
Hincks, W. 1952. The genera of the Cassidinae (Co-
leoptera: Chrysomelidae). Trans. Roy. Entomol.
Soc. London 103: 327-358.
Horton, J. 1959. The problem of phreatophytes. pp.
76-83. Symp. of Hannoversch-Munden I. Intern.
Assoc. Sci. Hydrology.
508
Johnson, H. 1984. Consequences of species intro-
ductions and removals on ecosystem functions—
Implications for applied ecology. Jn E. DelFosse
(ed.) Proc. VI Internat. Symp. Biol. Control of
Weeds, Vancouver.
Martin, A., H. Zim, and A. Nelson. 1951. American
Wildlife and Plants. McGraw-Hill, New York.
Martinez, A. 1959. Las plantas medicinales de Mex-
ico. 4a Ed. Batas, Mexico.
McFadyen, P. 1987. Host specificity and biology of
Lioplacis elliptica (Col.: Chrysomelidae) intro-
duced into Australia for the biological control of
Baccharis halimifolia (Compositae). Entomoph.
32:19-21.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Parker, K. 1972. An Illustrated Guide to Arizona
weeds. pp. 278-279. Univ. Ariz. Press, Tucson.
Seeno, T. and J. Wilcox. 1982. Leaf beetle genera
(Coleoptera: Chrysomelidae). Entomography 1: 1-
22110
Spaeth, F. 1913. Kritische Studien uber den Umfang
and die Begrenzung mehrer Cassiden—Gattungen
nebst Beschreibung neuer amerikan. Arten. Arch.
Naturg. Berlin. 79, Abt. A, Heft 6: 126-164.
Walter, H., E. Harnickell, and D. Mueller-Dombois.
1975. Climate Diagram Maps of the Individual
Continents and the Ecological Climate Regions of
the Earth. Springer-Verlag, New York.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 509-512
FINAL INSTAR LARVA OF THE EMBOLEMID WASP,
AMPULICOMORPHA CONFUSA (HYMENOPTERA)
RosBert A. WHARTON
Department of Entomology, Texas A&M University, College Station, Texas 77843.
Abstract.—The development of the embolemid wasp Ampulicomorpha confusa is sim-
ilar to that of dryinids. The embolemid larva is enclosed in a sac bulging from its homop-
teran host, and the sac is of larval origin. The final instar is described, and notes are
provided on the occurrence of embolemids in Texas.
Key Words:
Recent works on the Chrysidoidea, cul-
minating in that of Carpenter (1986), have
greatly clarified relationships within this
superfamily. The resulting classification
provides a convenient framework for the
observations recorded here on the Embo-
lemidae. Members of this family are rarely
collected, and what little 1s known about
their biology is limited to the observations
of Bridwell (1937, 1958). Buergis (1987)
provides a recent summary, including mor-
phological observations for the family as a
whole and collection records for European
species.
In 1982, I reared a single male of Am-
pulicomorpha confusa Ashmead. Unfortu-
nately, additional attempts to rear this
species have failed. The results presented
here add few new details to the observations
of Bridwell (1958), but do clarify some as-
pects of the biology of A. confusa.
MATERIALS AND METHODS
On February 13, 1982, I collected several
nymphs of Epiptera floridae (Walker) (Ho-
moptera, Achilidae) in San Jacinto County,
Texas. The habitat was identical to that de-
scribed by Bridwell (1958): beneath loose
bark of rotting conifer logs. Two of these
nymphs had sacs bulging laterally between
Embolemidae, Achilidae, larva
the second and third abdominal segments.
These two nymphs were placed in a separate
vial with pieces of bark and kept under ob-
servation for the next six weeks.
Members of two other wasp families
(Rhopalosomatidae and Dryinidae) devel-
op in sacs on their exopterygote hosts, and
representatives of these were used for com-
parison in the description of the embolemid
larva and its sac. Four crickets (Gryllidae)
parasitized by rhopalosomatids were col-
lected in College Station, Texas, in Septem-
ber of 1987 and 1988. Determined and un-
determined dryinids were also examined:
determined material consisted of host re-
mains of reared specimens in the genera An-
teon and Pseudogonatopus, undetermined
material included approximately 2 dozen
homopteran hosts containing dryinid larvae
in sacs bulging from their bodies. The latter
were from numerous localities and were all
preserved in 70% ethanol.
All hosts and parasitoids were identified
by the author, and voucher specimens have
been deposited in the Texas A&M Univer-
sity Collection as TAMU voucher #228.
RESULTS AND DISCUSSION
The embolemid sacs bulging from the
nymphal achilids were identical in appear-
510
Figs. 1-6.
lateral view, 100 =. 3, labrum, ventral view showing part of epipharyngeal region, 160 x. 4, left mandible, frontal
view, 400 x. 5, maxilla, ventral-lateral view, 160. 6, labium, ventral view, 160.
ance to those produced by the larvae of dryi-
nids on their homopteran hosts. These sacs
were in fact embolemid larval exuviae, as
evidenced by the rows of spiracles along the
sides of the sacs. As Carpenter (1986) has
noted, Bridwell’s (1958) description did not
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Head and mouth parts of final instar larva of Ampulicomorpha confusa. |, frontal view 100. 2,
clearly address the question of whether the
tissue composing the sacs was of host or
parasitoid origin. The observations here
confirm the parasitoid origin of the sac.
As in most dryinids I have examined, the
embolemid sac is more or less transversely
VOLUME 91, NUMBER 4
oriented to the body of the host as opposed
to the longitudinal orientation of the sacs
produced by rhopalosomatid larvae on their
gryllid hosts. In addition, the posterior por-
tion of the rhopalosomatid body is attached
in anchor-like fashion while the head re-
mains free. The rhopalosomatid larva lies
stretched out along the side of its host. Dryi-
nid and embolemid larvae have the head
more completely buried in the host’s body,
and are doubled up in the sac (and thus
cyphosomatic).
One embolemid larva emerged from its
host within two weeks and spun a cocoon
on the bark. The cocoon was identical to
that described by Bridwell (1958). On March
22nd of the same year, a male A. confusa
emerged from this cocoon. The remaining
embolemid larva was preserved in 70% eth-
anol after it emerged from its host. The brief
description given below is of this second
larva. The body was poorly preserved and
certain details are not evident.
The larva of Ampulicomorpha is readily
distinguished from that of Rhopalosoma
since the spinnerets of the latter are paired
and palpiform. Detailed comparisons will
be needed, however, before the larvae of
embolemids can be distinguished from those
of other chrysidoids. Evans (1987) provides
a synopsis of existing data on chrysidoid
larvae. The large, tridentate mandibles sep-
arate the larvae of 4. confusa from those of
known dryinids, but few of the latter have
been described. Some chrysidids and beth-
ylids also have tridentate mandibles, hence
A. confusa larvae key to these families in
couplet 50 of Evans (1987).
Chrysidoid larvae are insufficiently known
to permit placement of embolemids on the
basis of larval morphology alone. The de-
velopmental biology and host associations,
however, clearly confirm the sister group
relationship between embolemids and dry1-
nids, as outlined by Carpenter (1986).
I have collected adults of A. confusa in
Malaise traps from April through June in
central Texas. In the same traps, I have col-
511
lected male Embolemus from May through
October. The two genera are readily distin-
guished on the basis of the relatively larger,
more quadrate pronotum and weaker fore
wing venation of 4mpulicomorpha. The fe-
males of Embolemus, as far as is known,
are apterous.
Description of Larva.—3.5 mm long,
mostly white, mandibular teeth dark brown;
body segments and head capsule distinct.
Spiracles and atrium simple, without dis-
tinct peritreme, diameter of spiracle varying
from 0.024 mm on abdomen to 0.036 mm
on thorax. Integument largely smooth, not
obviously setose or spinose. Head (Figs. 1,
2) more or less prognathous, 0.60 mm wide,
0.62 mm long (length measured dorsally be-
tween apex of labrum and posterior margin
of head capsule), without evident. parietal
bands but with a median ecdysial line pos-
teriorly. Antenna small, round, not pro-
truding, bearing 3 minute sensilla. Clypeus
distinct distally, but frontal-clypeal suture
not visible; labral-clypeal suture below or
extending anteriorad base of mandibles. La-
brum (Fig. 3) dorsally with a pair of distinct
setae and a group of smaller sensilla along
anterior margin; epipharynx at least medi-
ally covered with minute sensilla. Mandi-
bles large, tridentate in frontal view (Fig. 4),
the teeth decreasing in length mesally. Max-
illae (Fig. 5) weakly bilobate, simple, with
large, protruding palpi, the latter with 4 mi-
nute sensilla; a galea was not observed. La-
bium (Fig. 6) with large median spinneret
and a pair of distinct palpi.
ACKNOWLEDGMENTS
I wish to thank J. Carpenter and A. Ras-
nitsyn for encouraging me to publish this
information; and L. Bradley for collecting
two of the rhopalosomatid larvae.
LITERATURE CITED
Bridwell, J.C. 1937. In: Minutes of the 478th regular
meeting of the Entomological Society of Wash-
ington, H. H. Richardson, recording secretary. Proc.
Entomol. Soc. Wash. 39: 14-15.
512 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1958. Biological notes on Ampulicomorpha Carpenter, J. M. 1986 Cladistics of the Chrysidoidea
confusa Ashmead and its fulgoroid host. Proc. (Hymenoptera). J. N.Y. Entomol. Soc. 94: 303-
Entomol. Soc. Wash. 60: 23-26. 330.
Buergis, H. 1987 Die Wespe Embolemus antenna- Evans, H. E. (Coordinator). 1987. Order Hymenop-
/is—ein seltener Fund aus Hessen. Nat. u. Mus. tera, p. 597-710. Jn Stehr, F. W. (ed.). Immature
117: 12-19. Insects. Kendall/Hunt, Dubuque.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 513-522
REVIEW OF NEARCTIC METACHELA COQUILLETT,
WITH DESCRIPTION OF A NEW SPECIES
(DIPTERA: EMPIDIDAE; HEMERODROMIINAE)
JOHN F. MACDONALD
Department of Entomology, Purdue University, W. Lafayette, Indiana 47907.
Abstract—The genus Metachela is reviewed for the Nearctic region. Generic diagnosis
and new diagnoses of two previously described species, Metachela albipes (Walker) and
M. collusor (Melander), are presented with discussion of intraspecific variation in male
terminalia. Metachela convexa n. sp. is described. A key to Nearctic males, illustrations
of male terminalia, and known distributions are provided.
Key Words:
Aquatic dance flies of the genus Mera-
chela Coquillett are small, slender, and pos-
sess strongly raptorial fore legs; wings lack
an anal lobe and thus are relatively slender.
The genus Metachela, therefore, is similar
in general morphology to other Hemero-
dromiinae genera, the Nearctic species of
which were treated by Melander (1902,
1928, 1947). The present study was facili-
tated by the relatively large numbers of
Metachela specimens added to North
American collections since Melander’s
(1947) revision. Generic diagnosis, diag-
noses of two previously described species,
description of a new species, known distri-
butions, and a key to Nearctic males are
provided.
Metachela species are uniform in general
morphology and coloration, and possess few
characters of taxonomic value. Male ter-
minalia are of little use in separating the
two widespread, common species, but are
distinct for a newly described species. In-
terpretation of male terminalia follows that
of Chvala (1983). The only other taxonomic
tool was related to the structure and vesti-
ture of the fore femora. Vestiture terminol-
ogy follows McAlpine et al. (1981). The ba-
Diptera, Empididae, Hemerodromiinae, Metachela, dance flies
sic form of macrotrichia is a seta (= a hollow,
articulated, epidermal outgrowth), with the
following descriptive terms applied to it: hair
(= long, weak seta); bristle (= long, strong
seta); and setula (= short, strong seta). In
addition to the various forms of macrotrich-
ia, Metachela specimens possess pollinosity
(= dense microtrichia) over much of the
body. No reliable characters were discov-
ered for females despite examination of sev-
eral hundred, and thus only collecting data
pertaining to males could be used to gen-
erate the distribution map.
The following institutions provided ma-
terial upon which this work is based: Amer-
ican Museum of Natural History, New York
(AMNH); Biosystematics Research Centre,
Ottawa (Canadian National Collection)
(CNC); California Academy of Sciences, San
Francisco (CAS); Cornell University, Ithaca
(CU); Florida State Collection of Arthro-
pods, Gainesville (FSCA); Illinois Natural
History Survey (INHS); United States Na-
tional Museum of Natural History, Wash-
ington D. C. (USNM); University of Cali-
fornia, Riverside (UCR); University of
Kansas, Lawrence (SNOW); University of
Minnesota, St. Paul (UMSP); Utah State
514
University, Logan (USU); and Washington
State University (JAMES). Specimens also
came from the author’s collection (MAC)
and the Purdue Entomological Research
Collection (PERC).
Genus Metachela Coquillett
Metachela Coquillett 1903: 253, 263. Type
species: Hemerodromia collusor Melan-
der (orig. des.).
Diagnosis.—The genus Metachela is dis-
tinguished from other genera of Hemero-
dromiinae by the combination of the fol-
lowing: relatively long, thick stylus of the
antennae, absence of crossvein bm-cu, and
existence of two veins (anterior one forked)
arising from apex of fused cell bm + dm.
Description. —Small (body length ca. 3.5
to 4.0 mm), slender, with strongly raptorial
fore legs. Nearctic species uniformly greyish
black with golden pollinosity on scutum,
white pollinosity elsewhere on thorax and
on head. Mouthparts and palps yellowish.
Legs concolorous yellow, except for dark-
ened apical one or two tarsomeres. Fore
femora and fore tibiae possess two median
rows of black setulae ventrally, with those
of fore femora flanked by row of light brown
bristles and with group of paler setae ba-
soventrally. Mid tibiae ventrally with short,
apical row of black setulae. Body setae yel-
lowish. Prominent setae of head including
pair of long ocellars and pair of long inner
verticals, with a pair of shorter, outer ver-
ticals detectable on some specimens. Prom-
inent thoracic setae including several short
setae on pronotum, pair of long notopleu-
rals, pair of long supraalars, and pair of long
scutellars apically, with very short pair im-
mediately lateral to scutellars. Male termi-
nalia composed of hypandrium, pair of gon-
ocoxites that each possess apical fringe of
setae and apicodorsal claw-like gonostylus,
and epandrium consisting of two, broad-
ened lobes that rest on top of gonocoxites.
Aedeagus arising from complex aedeagal
apodeme; relatively thick, with apex sur-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
rounded by several very small, sclerotized
plates. Segments eight and nine of female
abdomen forming exposed “‘ovipositor”’ that
is sclerotized and terminates in pair of slen-
der cerci.
Remarks. —Flies of the genus Metachela
closely resemble species of Chelifera, but
the difference in antenna stylus and stable
differences in wing venation support sepa-
rate generic status, a taxonomic treatment
proposed by Coquillett (1903) and followed
by Melander (1947). Metachela appears to
be a very small genus; in addition to the
Nearctic species treated in this paper, one
species occurs in central Europe (Frey 1956).
South American species placed in Meta-
chela by Collin (1933) and Smith (1962) are,
according to Smith (1962), not typical of
the genus, differing in several respects from
the type species; the taxonomic status of
these species has yet to be established.
Almost nothing is known about the bi-
ology of Metachela species, but adults have
been observed preying on adult Simultidae
(Peterson 1960). Most specimens in collec-
tions have been swept off riparian vegeta-
tion and taken in Malaise traps placed along
and across streams. Collecting data reveal
that adults appear by late June and continue
to be taken well into September. Larval and
pupal stages have not been described.
Nearctic species are either boreal or as-
sociated with western mountain streams,
with only one, M. al/bipes (Walker), extend-
ing into eastern North America.
Key TO NEARCTIC SPECIES OF
METACHELA COQUILLETT
(MALES ONLY)
1. Gonocoxites expanded laterally, strongly con-
vex in dorsal view; apex of epandrial lobes
turned inward as rounded process lined with
black setulae (Fig. 1)
ee bd .. Metachela convexa, new species
— Gonocoxites not expanded laterally, not
strongly convex in dorsal view; apex of epan-
drial lobes without in-turned, rounded process
lined with black setulae (see Figs. 4-7) ...... 2
Inner surface of fore femora with long setae;
length of basoventral setae of fore femora
to
VOLUME 91, NUMBER 4
Fig: 1.
hypandrium; aea = aedeagal apodeme). A = lateral view; only basal portion of aedeagal apodeme shown. B =
dorsal view; hypandrium removed. C = ventral view of hypandrium. Scale: 0.5 mm.
longer than greatest width of fore femur (Fig.
Denese ac eee Metachela albipes (Walker)
Inner surface of fore femora without long se-
tae; length of basoventral setae of fore femora
subequal to greatest width of fore femur (Fig.
3) ........Metachela collusor Melander
Metachela albipes (Walker)
Metachela albipes (Walker), 1849: 505
(Hemerodromia).
Diagnosis.— Male: Body length, includ-
ing terminalia, ca. 4.0 mm. Inner surface of
fore femora (Fig. 2) with long setae; fore
femora relatively slender, ca. 4 times longer
than greatest width, with group of baso-
ventral setae longer than greatest width of
fore femora. Male terminalia as in Fig. 4;
gonocoxites not expanded laterally; apical
fringe of gonocoxites with longest setae ca.
S15
Metachela convexa male terminalia (ep = epandrial lobe; gn = gonocoxite; gs = gonostylus; hy =
as long as epandrial lobes; apex of epandrial
lobes smoothly rounded, without black set-
ulae. Fema/e:indistinguishable from fe-
males of other Metachela species.
Type material.—Lectotype male (Smith
1971: 365), Canada, St. Martin’s Falls, Al-
bany River, Hudson’s Bay (G. Barnston la-
bel), deposited in British Museum (Natural
History). The specimen was examined by
K. V. G. Smith (June 1988), who reported
(pers. comm.) it to be in “very bad condi-
tion,” but “the fringe of long hairs on the
inner surface of the front femur are quite
easily seen.” The terminalia, mounted in
balsam and illustrated in Smith (1971), agree
with specimens that I have seen from Can-
ada and the United States.
Specimens examined.—164 males. Al-
berta: 2, Belly River, Aug.; 15, Wild Hay
R. at Hwy. 40, Aug. (CNC). British Colum-
516 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 2. Metachela albipes front femur, inner surface. Scale: 0.5 mm.
Fig. 3. Metachela collusor front femur, inner surface. Scale: 0.5 mm.
Fig. 4.
Metachela albipes male terminalia. A = lateral view; only basal portion of aedeagal apodeme shown.
B = dorsal view; hypandrium removed. C = ventral view of hypandrium. Scale: 0.5 mm.
bia: 1, Taylor Landing, Peace R., Aug.
(CAS); 3, Tyree, 27 mi. E. Prince Rupert,
June; 1, Lakelsch Bog nr. Terrace, June
(CNC). Manitoba: 1, Mosquito Pt., Chur-
chill R., Aug. (CNC); 4, 20 mi S. Churchill,
Aug. (UMSP). Quebec: 1, Brador Bay, July;
2, Great Whale R., July; 1, Indian House
L., Aug.; 4, Ft. Chino, Aug.—Sep. (CNC); 1,
Brador Bay, Aug. (FSCA). California: 1,
Carmel, July (AMNH); 32, Los Angeles Co.,
n. fork San Gabriel R., nr. Coldbrook Sta.,
June; 1, Modoc Co., Cedar Pass, Aug. (CAS);
4, Placer Co., Granite Flat cmpgr., Sep.
(MAC); 12, Mono Co., Lower Rock Cr.,
Sep. (UCR); 1, Lone Pine, July; 1, Sequoia
Nat. Pk., Aug. (SNOW); 1, Humboldt Co.,
Hydesville, Aug.; 1, Tulare Co., Camp Nel-
son, July; 1, Barton Flat, South Fork Camp,
Sep.; 1, Los Angeles Co., s. fork Santa Ana
R., Aug.; 2, Palm Springs, Nov. (USNM).
Montana: 2, Glacier Nat. Pk., Swiftcurrent
cmpgr., Aug. (CAS); 1, Thompson, Aug.
(USNM). New Hampshire: 1, “White Mts.”
(USNM). Oregon: 1, Grant Co., Beech Cr.
cmpgr., July; 1, Lane Co., Salt Cr. Falls
cmpgr., July; 1, Wasco Co., Warm Springs
Indiana Res., Beaver Cr., June (CAS); 1,
Baker Co., Big Cr., Aug. (JAMES); 1, Eagle
Cr., Aug.; 3, Humbug St. Pk., Aug. (USNM).
Washington: 1, Clallum Co., Tumbling
Rapids Rec. Area, July; 3, Grays Harbor
Co., Olympic Nat. Pk., Willaby cmpgr., July;
Fig. 5. | Metachela collusor male terminalia, predominant form. A = lateral view; only basal portion of aedeagal
apodeme shown. B = dorsal view; hypandrium removed. C = ventral view of hypandrium. Scale: 0.5 mm
Fig. 6. Metachela collusor male terminalia; digitate form. A = lateral view; only basal portion of aedeagal
apodeme shown. B = dorsal view; hypandrium shown. C = ventral view of hypandrium. Scale: 0.5 mm.
Fig. 7. Metachela collusor male terminalia; small form. A = lateral view; only basal portion of aedegal
apodeme shown. B = dorsal view; hypandrium removed. C = ventral view of hypandrium. Scale: 0.5 mm.
517
518 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 8. Distribution of Nearctic Metachela.
1, Jefferson Co., Olympic Nat. Pk., July
(CAS); 10, Asotin Co., Asotin Cr., Aug.; 1,
Jefferson Co., Hoh R., Cottonwood cmpgr.,
July; 40, Lewis Co., Rainbow Falls State
Pk., June-July (JAMES); 1, Yakima Co.,
e M.albipes
o M. collusor
3 M. convexa
Tieton R., 20 mi. W. Natches, Oct.;1, Yak-
ima Co., Oak Cr. canyon, 6 mi. W. Natches,
Oct. (MAC); 1, Adna, July; 1, Lilliwaup,
Aug. (USNM).
Remarks.—The vast majority of M. al-
VOLUME 91, NUMBER 4
bipes males are distinguished from other
Nearctic Metachela by the distinctive long
setae on the inner surface of the fore femora
and by the long setae basoventrally on the
fore femora (Fig. 2). The inner setae provide
the best diagnosis, although they tend to be
less prominent in some southern California
specimens; however, these specimens still
possess the long basal setae on the fore fem-
ora and thus are distinct from M. collusor.
Male terminalia of M. albipes (Fig. 4) and
M. collusor (Figs. 5-7) possess only subtle
differences, which are not taxonomically
useful because they are difficult to charac-
terize.
Five males collected in California, three
from Fresno Co., Bear Cr. (1- VIH-1979) and
two from Siskiyou Co., Klamath National
Forest., S. of Forks of Salmon R. (22-VII-
1980) (CAS), that agreed with M. albipes in
all other respects possessed terminalia that
were reduced in size, contained relatively
short setae in the apical fringe of the gon-
ocoxites, and had much more slender epan-
drial lobes. I consider these males as struc-
tural variants of albipes, based on the
existence of so few specimens, the occur-
rence of a similar pattern of variation in M.
collusor, and lack of additional evidence that
would suggest separate taxonomic Status.
Metachela albipes is the most widely dis-
tributed Nearctic species of Metachela, oc-
curring from eastern Canada and New
Hampshire to western Canada, and south
along the mountains of Montana, Washing-
ton, Oregon, and California (Fig. 8).
Metachela collusor (Melander)
Metachela collusor (Melander), 1902: 235
(Hemerodromia).
Diagnosis.— Male: Body length, includ-
ing terminalia, ca. 4.0 mm. Inner surface of
fore femora (Fig. 3) without long setae; fore
femora relatively stout, less than 4 times
longer than greatest width, with basoventral
setae subequal to greatest width of fore fem-
ora in most specimens. Male terminalia as
in Figs. 5—7; gonocoxites not expanded lat-
519
erally; apical fringe of gonocoxites with
longest setae ca. as long as epandrial lobes
in most specimens; apex of epandrial lobes
smoothly rounded, without black setulae.
Female: indistinguishable from females of
other Metachela species.
Type material.—LECTOTYPE male, la-
belled “Dubois, Wyo/7200 ft/ 1x 6. 95” here
designated from Melander’s cotype series
number 29180 (all in excellent condition).
The lectotype and two male and three fe-
male paralectotypes are deposited in the
United States Museum of Natural History
(USNM).
Other specimens examined.— 363 males.
Alberta: 11, Banff, Aug.; 4, Wild Hay R. at
Hwy. 40, June (CAS); 1, 20 mi. W. Calgary,
June; |, Jasper, July (CNC). British Colum-
bia: 1, Mt. Fitzwilliam nr. Lucerne, Aug; 5,
nr. Golden, Aug.; 10, Taylor Landing, Peace
R., Aug.; 1, Kootenay Nat. Pk, Dolly Var-
den Cr., July (CAS). Yukon Territory: 9, 14
mi. E. Dawson, July-Aug.; 1, Swift R.,
Rancheria, Aug.; 2, Whitehorse, Aug.
(AMNH); 1, Whitehorse, July (CNC); 2,
Whitehorse, Aug. (USNM). Alaska: 20, Tok
R., Glenn Hwy., Aug. (CAS); 2, Big Delta,
July; 15, King Salmon, Naknek R., July; 1,
Unalakleet, July (CNC). California: 55,
Fresno Co., Huntington L., Rancheria Cr.,
Aug. (MAC, PERC); 2, Lone Pine, July
(SNOW), 8, Alpine Co., Disaster Cr., July;
2, Alpine Co., Little Teton Cr., July; 1, Fres-
no Co., Bolsillo Cr. cmpgr., Aug.; 2, Fresno
Co., Edison L., Aug.; 1, Mono Co., Leavitt
Meadow, Aug.; 4, Tuolumne Co., Clark Fork
R., July (CAS); 1, Sierra Nat. For., Vee Lake,
Aug. (MAC). Colorado: 12, Boulder, Aug.;
1, Steamboat Springs, Aug.; 6, Chaffee Co,
Garfield, Aug. (CAS); 1, Estes Park, July; 1,
3 mi. SW. Idaho Springs, July; 2, Raymond,
St. Vrain Cr., Aug. (CNC); 1, Midland, Aug.
(INHS); 1, Boulder Co., Boulder Cr., Sep.;
15, Estes Park, July; 1, Florissant, Aug.; 1,
Rocky Mt. Nat. Pk., June; 2, N. St. Vrain
Cr., Sep (USNM). Idaho: 1, Latah Co., Big
Meadow Rec. area, June (JAMES); 5, Car-
ibou Co., Kendall Cr., June-July; 1, Frank-
lin Co., Aug. (USU). Montana: 1, Beaver
520
Cr., Aug.; 1, Ennis, Aug.; 1, Glacier Nat.
Pk., Lake McDonald, June; 1, Glacier Nat.
Pk., Logan Falls, July; 3, Glacier Nat. Pk.,
Second Medicine L., Aug. (USNM). Ne-
vada: 1, Washoe Co., Verdi, June (CAS).
Oregon: 4, Clackamas Co., Mt. Hood Nat.
For., Tollgate cmpgr., June; 28, Klamath
Co., 5 mi N. Chiloquin, June; 2, Wasco Co.,
Warm Springs Indiana Res., Beaver Cr.,
June (CAS); 1, Clack. Co., nr. Rhodendron,
June (JAMES); 2, Corvallis, June; 3, Hood
River, June (USNM). South Dakota: 19,
Lawrence Co., Spearfish Cr., June (USNM).
Utah: 9, Cache Nat. For., Wasatch Mts.,
July (USNM); 1, Cache Co., Logan Canyon,
June; 1, Jaub Co., Mt. Nebo, July—Aug.
(USU). Washington: 11, Mt. Ranier Nat.Pk.,
Fish Cr., Aug. (CAS); 1, Glenwood, Klick-
itat R., June (CNC); 3, Glenwood, Klickitat
R., June (CU); 15, Asotin Co., Asotin Cr.,
June; 2, Mt. Ranier Nat. Pk., Tahoma R..,
Aug.; 1, Yakima Co., Nile, June (JAMES);
24, Glenwood, Klickitat R., June-July; 3,
Husan, July; 1, Spokane, Aug. (USNM).
Wyoming: 1, Yellowstone Nat. Pk., Aug.
(UCR); 1, Canyon Camp, Aug.; 2, Indiana
Cr., July; 4, Madison R., Riverside, Aug.;
2, W. Craigs Pass, Aug. (USNM).
Remarks.— Absence of long setae on the
inner surface and existence of shorter setae
basoventrally on the fore femora of males
distinguish MV. collusor from M. albipes. The
latter trait is somewhat variable, with some
M. collusor males possessing basoventral
setae that are longer than the greatest width
of the fore femora. However, the fore fem-
ora of such specimens are relatively thick
(their length less than four times their great-
est width) and there are no long setae on the
inner surface.
Metachela collusor is a widespread west-
ern species, distributed from western Alaska
south into the Sierra Nevada Mountains of
central California and the central Rocky
Mountains (Fig. 8). Variation in pigmen-
tation of the fore coxae and in terminalia
structure exists throughout the range of this
species, as described below.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Variation in the pigmentation of the fore
coxae of males in several series of M. col-
lusor from Alaska, British Columbia, cen-
tral Colorado, and Yukon Territory resulted
in coloration ranging from entirely yellow
to entirely dark brown. Specimens with var-
ious forms of intermediate pigmentation of
the fore coxae were collected concurrently
and lacked consistent differences in other
characters, including details of male ter-
minalia.
The predominant terminalia structure of
M. collusor males is depicted in Fig. 5, but
variation in size and shape of the compo-
nent parts existed throughout the range. The
two extreme forms of this variation (Figs.
6, 7) are firstly, a hypandrium with some-
what digitate apical processes (Fig. 6) and
secondly, terminalia of substantially small-
er overall size with an accompanying re-
duction in the aedeagal apodeme and length
of setae in the gonocoxite fringe (Fig. 7).
The form of terminalia with two digitate
processes at the apex of the hypandrium
(Fig. 6) existed on 12 males from Naknek,
Alaska (CNC), four males from Caribou
County, Utah (USU), and five males from
Lawrence County, South Dakota (USNM).
The males from Alaska and South Dakota
were part of series of concurrently collected
males that included individuals with the
predominant form of terminalia (Fig. 5) and
a few specimens that could be placed with
either the digitate form or the predominant
form, depending on how they were viewed.
Macerated terminalia of one of these males,
which was difficult to place before dissec-
tion, revealed that only one side of the apex
of the hypandrium appeared “‘digitate’’; the
other side of the hypandrium resembled the
apex of the predominant form. The ap-
pearance of these two forms of M. collusor
terminalia stems, at least in part, from the
variable manner in which the hypandrium
collapses on killed specimens as they dry.
Another contributing factor to an appear-
ance of distinctiveness in structure is the
size of the hypandrium, with the digitate
VOLUME 91, NUMBER 4
form of hypandrium tending to be smaller
and more slender than the predominant
form.
The type of terminalia of an overall much
smaller size (Fig. 7) existed on 53 males
collected above Huntington Lake, Fresno
County, California (MAC), one male from
Alpine county and another from Tuolumne
County, California (CAS), two males from
Lone Pine, California (SNOW), six males
from Chaffee County, Colorado (CAS), two
males from near Raymond, Colorado
(CNC), three males from Cache County,
Utah (USU), and one male from near Lu-
cerne, British Columbia (CAS). The 53
males from above Huntington Lake (col-
lected ina Malaise trap set across a tributary
to Rancheria Creek) were part of a series,
collected concurrently, that included three
males with the predominant form of ter-
minalia and two males with terminalia that
were intermediate in terms of overall size,
size of the aedeagal apodeme, and length of
setae forming the gonocoxite fringe. I con-
sider males with this much smaller form of
terminalia (due largely to reduction in the
hypandrium) to be structural variants of M/.
collusor, based on the presence of inter-
mediates and the fact that other structural
variation in terminalia is common among
M. collusor males, as mentioned above.
Metachela convexa MacDonald,
New Species
Description. — Male: Body length, includ-
ing terminalia, ca. 3.8 mm. In general,
matching the generic description with re-
gard to coloration, vestiture, and venation.
Fore femora without fringe of long setae on
inner surface; relatively stout, less than 4
times longer than greatest width, with ba-
soventral setae about as long as greatest
width. Terminalia (Fig. 1) with large,
strongly convex gonocoxites that overlap
hypandrium below and epandrial lobes
above; apical fringe on gonocoxites with
longest setae shorter than epandrial lobes:
521
median margin of epandrial lobes expanded
into rounded, apicodorsal process lined with
black setulae. Female: unknown; none as-
sociated with concurrently collected males.
Type material.—HOLOTYPE male, top
label “U.S. A.: California:/Shasta County,
Honn/Creek Campground, /Hat Creek, 29
km. SE./Burney, 30-VII-1974” and bottom
label “Paul H. Arnaud, Jr./Calif. Acad. Sci.
Coll.”; macerated terminalia in glycerin mi-
crovial attached to pin. Holotype deposited
in the California Academy of Science Col-
lection (CAS). PARATY PES.—2 males. Cal-
ifornia: Shasta Co., Lassen Volcanic Nat.
Park, 2.4 km. W. Kings Cr., July (CAS).
Oregon: Klamath Co., 5 mi N. Chiloquin,
on Williamson R., June (CAS).
Remarks—The specific epithet refers to
the large, strongly convex gonocoxites that
are diagnostic for this species. Nothing is
known about the biology of M. convexa
which, based on only 3 specimens, appears
to be restricted to northern California and
southern Oregon (Fig. 8).
ACKNOWLEDGMENTS
I thank the following individuals for the
loan of material: P. H. Arnaud, Jr. (CAS);
R. W. Brooks (SNOW); P. J. Clausen
(UMSP); S. Frommer (UCR); W. J. Hanson
(USU); L. Knutson (USNM); J. K. Liebherr
(CU); R. T. Schuh (AMNH); H. J. Teskey
(CNC); W. J. Turner (JAMES); D. W. Webb
(INHS); and H. V. Weems (FSCA).
Sincere appreciation is extended to the
Dipterists in the Biosystematics Research
Centre, Ottawa, where I spent a sabbatic
leave in the fall of 1984. Special thanks are
given to H. J. Teskey for his generous as-
sistance during initial study, to K. V. G.
Smith for examining the type of Metachela
albipes, to J. Cumming and W. P. Mc-
Cafferty for critically reading the manu-
script, and to Arwin Provonsha for prepar-
ing the illustrations. This paper is Purdue
Agricultural Experiment Station journal No.
11,534.
522
LITERATURE CITED
Chvala, M. 1983. The Empidoidea (Diptera) of Fen-
noscandia and Denmark. II General Part. The
Families Hybotidae, Atelestidae and Microphor-
idae. Vol. 12. Fauna Entomol. Scand. 279 pp.
Collin, J. E. 1933. Diptera of Patagonia and South
Chile. Part 4. Empididae. Brit. Mus. (Nat. Hist.).
334 pp.
Coquillett, D. W. 1903. The genera of the dipterous
family Empididae, with notes and new species.
Proc. Entomol. Soc. Wash. 5: 245-272.
Frey, R. 1956. Jn Lindner, E. ed., Die Fliegen der
Palaearktischen Region, 28. Empididae, vol. 4,
part 4.
McAlpine, J. F. 1981. Morphology and terminolo-
gy—Adults. 2, pp. 9-63. In McAlpine, J. F. et al.,
eds., Manual of Nearctic Diptera. Res. Br., Agric.
Canada Monogr. 27 (1): 1-674.
Melander, A. L. 1902. A monograph of the North
American Empididae. Part I. Trans. Am. Ento-
mol. Soc. 28: 195-367; plates 5-9.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
_ 1928. Diptera, Fam. Empididae. Fascicle 185,
Pages 1-434, plates 1-8 in P. Wytsman, ed. Genera
Insectorum. Bruxelles.
1947. Synopsis of the Hemerodromiinae
(Diptera: Empididae). J. N.Y. Entomol. Soc. 39:
237—273.
Peterson, B. V. 1960. Notes on some natural enemies
of Utah black flies (Diptera: Simulidae). Can.
Entomol. 92: 266-274.
Smith, K. G. V. 1962. Studies on the Brazilian Em-
pididae (Diptera). Trans. Roy. Entomol. Soc. Lon-
don. 114: 195-266.
1971. A revision of Francis Walker’s types
of North American Empididae (Diptera). Bull. Brit.
Mus. Nat. Hist. Entomol. 26: 347-370.
Walker, F. 1849. List of the specimens of dipterous
insects in the collection of the British Museum,
vol. 3, pp. 485-687. London.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 523-528
SEXUAL DIMORPHISM IN SIZE OF ADULTS AND PUPARIA OF
TETANOCERA FERRUGINEA FALLEN (DIPTERA: SCIOMYZIDAE)
SYLVIE MANGUIN
Laboratoire de Biologie Animale, Faculté des Sciences, 33, rue L. Pasteur, 84000 Avi-
gnon, France.
Abstract. — Adults and puparia of Tetanocera ferruginea Fallén are sexually dimorphic.
In both stages, the size of the female is significantly greater than that of the male. Based
on a biometric study, puparial length demonstrated that the sex of the adults could be
estimated before emergence with an 80.4% precision. This information on sexual differ-
entiation is important for field studies of populations of sciomyzid flies.
Key Words:
paria
In general, numerous Diptera (Vargas
1968) and many other insects (Mellini 1973)
show sexual dimorphism in the size of adults
and immature stages. Usually, the male is
smaller than the female. For the puparia,
the determination or separation of sexes is
based on few parameters: average weight
(Féron and Serment 1963), color of the pu-
parium (e.g. Lucilia cuprina Wiedemann
(Whitten 1969)), examination after a non-
toxic chemical treatment (e.g., Musca au-
tumnalis De Geer (Tung et al. 1969)), ex-
ternal morphological characters (Kuitert
1975), or microscopic examination through
the puparium (Cunningham 1966).
Other studies have shown that for Dip-
tera, sexual dimorphism rarely appears
among the larvae (Atchley 1971) and never
in the eggs. However, dimorphism of larvae
and eggs does occur in Homoptera (Monti
1955) and Lepidoptera (Levesque 1963).
The determination of sex of the immature
stages of the Sciomyzidae is an important
factor in studies of the basic biology of these
flies. Sexual differentiation is particularly
important for field studies of population dy-
namics of these flies, or when an intensive
Sciomyzidae, Tetanocera ferruginea, sexual dimorphism, size, adults, pu-
rearing program is part of a biological con-
trol project.
From 1982 to 1987, samples of sciomyzid
species were collected periodically from an
aquatic habitat near Avignon, in southern
France. Studies revealed that there was a
distinct variation in the size of adults and
puparia of the species of flies, and particu-
larly of Tetanocera ferruginea Fallén, which
was the most common species in the scio-
myzid population of this habitat (Vala and
Manguin 1987).
MATERIALS AND METHODS
T. ferruginea has 3 or 4 annual genera-
tions (Vala and Haab 1984) and the flight
period extends from late March through
November. The species overwinters as dia-
pausing puparia (Berg et al. 1982).
The puparium is black, oval, measuring
7.3 to 7.7 mm long and 3.1 to 3.4 mm wide
(Fig. 1). The anterior part has the circular
line of dehiscence for adult emergence. The
raised posterior end has a vestigial respi-
ratory disc and vestigial lobes of the third-
instar larva. The spiracles are above the pu-
parium, which is an adaptation for flotation.
524 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
spiracle; 2, posterior spiracle.
The puparia examined were from labo-
ratory cultures of 7. ferruginea which were
held in incubators at 20°C with a photo-
period of LD 16:8. The larvae were reared
in petri dishes (17 cm diameter <x 5 cm
deep) with 5 mm of tap water, which con-
tained one sciomyzid larva and 10 snails
each of the species Physa acuta (Drapar-
naud) and Lymnaea palustris (Miiller).
These species of prey were readily attacked
by larvae of T. ferruginea and 20 prey per
unit were more than adequate for one larva
(Manguin et al. 1986, 1988).
All specimens were measured with a 1/50
mm Kanon gauge. The length of the adults
was measured from the frons to the termi-
nalia; the width was the distance between
the lateral margins across the median part
of thorax. For puparia, three parameters
were measured: length, width, and height.
All measurements were of the distance be-
tween the extremities along or across the
median part of the puparia. After measure-
Lateral view of the puparium of 7. ferruginea. A, female puparium; B, male puparium. 1, anterior
ments were taken the puparia were returned
to the incubator and held for emergence of
the adult to verify the sex of each specimen.
RESULTS
Sexual dimorphism of adults: Measure-
ments of the length and width of 30 speci-
mens of each sex produced a bimodal curve
for the distribution of size. (Fig. 2).
Males, the smaller sex, represented the
first peak of the curve, averaging 9.6 mm
in length (range 7.9 to 10.5 mm) and 1.9
mm in width (range 1.6 to 2.5 mm). The
larger females represented the second peak
with an average of 10.9 mm in length (range
9.9 to 11.9 mm) and 2.2 mm in width (1.7
to 2.8 mm).
These two parameters showed highly sig-
nificant differences between males and fe-
males (P <0.01). However, the measure-
ments of the sizes of males and females
produced an overlapping zone which was
narrower for length than for width.
VOLUME 91, NUMBER 4
@-@: Adults é
o—o : Adults :
Number =
X=9.640.2
5
4
3 e
2 e # e@
1 ® e
é 6
8.0 9.0 9.5 10.0 10.5 11.0 11.5 12.0 Length (mm)
Number
7
6
5
4
3
2
1
SS ee ee ee ee
5 2.0 2:5 Width (mm)
Fig. 2. Distributions of length and width (mm) of adults of 7. ferruginea. Males = broken line, solid circle;
females = solid line, clear circle; n = 30 specimens per sex.
the empty puparium, minus the cephalic
caps, again was measured for comparison
with the observed sex of the adult specimen.
Sexual dimorphism of puparia: We mea-
sured unemerged puparia for predicting the
sex of an adult. After an adult had emerged,
: .
+ °
x
Length oy Length yay Width
(mm) Tee oo (mm) Pe
‘i ys Re
/ A
/ x
8 P ee G a) a AQ
| °
6 / 6 } J:
| / x : x
22 a ak
MB 7 20 : ee
* / Nb
| et | ae
(
\o
3 2
T 7 T - T
3 4 Width 2 3 4 Height 2 3 a Height
(mm)
@ (mm) ©) (mm) ©
Fig. 3. Regressions between sizes and unemerged puparia of 7. ferrwginea. Males = broken line, solid circle;
females = solid line, clear circle; a, length/width; b, length/height; c, width/height.
526
Length ne
Length
Width 2 3
@ (mm)
Fig. 4.
® 5 ©
Regressions between sizes of puparia of 7. ferruginea. Males = broken line, solid circle; females =
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
4a Height 2 3
4 Height
(mm)
solid line, clear circle; a, length/width; b, length/height; c, width/height.
The results in Table | indicate that the
size of the female puparium 1s significantly
larger than that of the male (P < 0.01).
The size distribution was established by
correlating paired dimensions of unemerged
puparia (Fig. 3) and emerged puparia (Fig.
4). For males, the distribution of every mea-
surement was narrow, regardless of the
measurement considered. This was con-
firmed by the coefficient of variation which
varied within narrow limits, from 5.6 to 7.1
(Table 1). For females, the data clusters on
the graph were larger and defined a loose
type of dispersion. The coefficient of vari-
ation was significantly higher and ranged
from 5.9 to 8.5.
In superimposing graphs obtained for both
sexes (Figs. 3 and 4), the dispersion of the
Table 1.
data was unwedged, particularly when the
length of the puparium was considered (Figs.
3a,b and 4a,b). The width and height were
highly different between the sexes, but these
differences were not as important as those
determined for the length (Figs. 3c, 4c).
In general, the correlations obtained (Figs.
3, 4) were defined by equations in which the
range slope varied between 0.22 and 0.52
for the males and between 0.60 and 1.11
for the females. These slopes, steeper for the
females, revealed that the 3 parameters
(length, width and height) evolved in a si-
multaneous manner. However, for males,
the slopes were much more level and there
was not much difference between parame-
ters. Although the length increased, the
width and height varied only slightly.
Mean size (mm) in length, width, and height of male and female unemerged puparia and empty
puparia of 7. ferruginea (+standard deviation at 5% threshold). CV (%): coefficient of variation; n = number
of measured puparia.
Unemerged pupana
Emerged puparia
Length Width
n= 57
Males ¥ (mm) 730 OLLT 3.1 + 0.06
CV (%) 5.8 7.0
n= 48
Females (mm) Tf £0.14" 3:4-£50108
Xx
CV (%) 6.3 8.5
Height Length Width Height
n= 72
2.9 + 0.05 5:70:08 3:1 40:05 2:9:=0/05
Ta 5.6 6.8 7.0
n=7]
3.121007 6.2+ 0.09 3.4+0.06 3.1 + 0.06
1S 5.9 hel 8.2
VOLUME 91, NUMBER 4
The length of a puparium was more de-
finitive than either the width or height as a
character to recognize the sex.
Estimation of the sex of the puparium: We
measured the length of 240 unemerged pu-
paria and estimated that 143 puparia would
be male and 97 puparia would be female
(Table 2). After adult emergence these pre-
dictions were proven correct in 80.4% of
cases. The results were more reliable with
males, with only 9.8% of error, whereas
measurements of the females were 34% er-
roneous. This difference was explained by
the larger spread of measurements for the
female, shown by the coefficient of variation
(Table 1).
DISCUSSION
There is sexual dimorphism in sizes of
adults and puparia of 7. ferruginea. For
adults, the length and width of the females
was significantly greater than for males.
Among the puparia, the parameters of
length, width, and height increased propor-
tionally for the females. Consequently, the
shape of female puparia was more oval with
the dorsal side slightly convex. For the males
only the length of the puparia varied. The
width and height were relatively constant;
and in lateral view, the dorsal surface of the
puparium was more rectilinear.
The possibility of determining the sex of
the flies prior to emergence, with an 80.4%
accuracy, 1s important not only for popu-
lation studies of field collected material, but
also for purposes of release in biological
control programs. The sexual differentia-
tion permits an easy manipulation of inert
pupae rather than active living flies, and
eliminates the need to hold for adult emer-
gence if sampling only is required. An ad-
ditional benefit of this method would be its
use in the study of those parasites that
emerge from puparia, such as many species
of Ichneumonidae (Manguin 1987). By de-
termining the sex of the puparia, the distri-
bution of parasites emerging from male and
female hosts can be established. This was
527
Table 2. Estimation of the sex of puparia deter-
mined by the measurement of the length.
Number Puparia Percentage
Examined Accuracy
Males 143 90.2
Females 97 66.0
Total 240 80.4
not possible when the sex of the host was
based only on the adult specimen.
ACKNOWLEDGMENTS
I thank J. C. Vala (Faculté des Sciences,
Avignon, France) for his helpful sugges-
tions, J. J. Drea (Beneficial Insects Labo-
ratory, Agricultural Research Service,
USDA, Beltsville, Maryland) for his critical
review and his assistance in the translation
of the manuscript, L. Knutson (Biological
Control of Weeds, USDA, Rome, Italy) and
J. R. Coulson (Beneficial Insects Labora-
tory, Agricultural Research Service, USDA,
Beltsville, Maryland) for reviewing this pa-
per.
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Atchley, W. R. 1971. Components of sexual dimor-
phism in Chironomus larvae (Dipt. Chironomi-
dae). Am. Nat. 105: 455-466.
Berg, C. O., B. A. Foote, L. V. Knutson, J. K. Barnes,
S. L. Arnold, and K. Valley. 1982. Adaptive dif-
ferences in phenology in sciomyzid flies. Mem.
Entomol. Soc. Wash. 10: 15-36.
Cunningham, R. T. 1966. Sex identification of pupae
of three species of fruit flies (Diptera: Tephritidae).
Ann. Entomol. Soc. Am. 59(4): 864-865.
Féron, M. and M. M. Serment. 1963. Differenciation
sexuelle aux stades oeuf et pupe chez la mouche
méditerranéenne des fruits Ceratitis capitata Wied.
Rev. Pathol. Vég. Ent. Agr. France. 42(4): 239-
244.
Kuitert, L. C. 1975. Sexual dimorphism in Plecia
nearctica pupae (Diptera: Bibionidae). Fla. Ento-
mol. 58(3): 212.
Levesque, G. 1963. A technique for sexing fully de-
veloped embryos and early-instar larvae of gypsy
moth. U.S. Forest Service Res. Note NE-2, 3 pp.
Manguin, S. 1987. Dynamique des peuplements de
Sciomyzides et de Mollusques d’un biotope léni-
tique méditerranéen et etude expérimentale des
préférences de proie de TJetanocera ferruginea
528
(Diptera: Sciomyzidae). 298 pp. Thése doct., bio.
physio. organ. populat.; Univ. Sc. Tech. Langue-
doc, Montpellier.
Manguin, S., J. C. Vala, and J. M. Reidenbach. 1986.
Prédation de mollusques dulgaquicoles par les
larves malacophages de Tetanocera ferruginea
Fallén, 1820. Can. J. Zool. 64(12): 2832-2836.
. 1988. Action prédatrice des larves de Tetano-
cera ferruginea (Diptera: Sciomyzidae) dans des
systémes a plusieurs espéces de mollusques-proies.
Acta OEcologica, OEcol. Applic. 9 (3): 249-259.
Mellini, E. 1973. Sul riconoscimento esterno del ses-
so negli stadi preimmaginali degli Insetti. Boll.
Entomol. Bologna. 30: 265-287.
Monti, L. 1955. Ricerche etologiche su due Coccidi
Diaspini: Diaspis pentagona Targ. e Mytilococcus
ulmi L. nella regione romagnola. Boll. Entomol.
Bologna. 21: 141-165.
Tung, S. C., T. H. Cheng, and W. G. Yendol. 1969.
A method of clearing face fly puparia for sex de-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
termination without affecting survival and devel-
opment. J. Econ. Entomol. 62(6): 1412-1417.
Vala, J.C. and C. Haab. 1984. Etude expérimentale
du développement larvaire de Tetanocera ferru-
ginea Fallén, 1820. Influences de la température
et de la photopériode, diapause pupale, biomasse
alimentaire (Diptera: Sciomyzidae). Bull. Annls.
Soc. r. belge Ent. 120: 165-178.
Vala, J. C. and S. Manguin. 1987. Dynamique et
relations sciomyzides-mollusques d’un biotope
aquatique asséchable dans le sud de la France
(Diptera). Bull. Annls. Soc. r. belge Ent. 123: 153-
164.
Vargas, M. V. 1968. Sexual dimorphism of larvae
and pupae of Aedes aegypti L. Mosquito News
28(3): 374-379.
Whitten, M. J. 1969. Automated sexing of pupae and
its usefulness in control by sterile insects. J. Econ.
Entomol. 62: 272-273.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 529-533
INFLUENCE OF TEMPERATURE AND PHOTOPERIOD ON
EARLY DEVELOPMENTAL RATE OF
TENODERA SINENSIS SAUSSURE (MANTODEA: MANTIDAE)
L. E. Hurp, P. E. MARINARI, AND R. M. EISENBERG
Ecology Program, School of Life Sciences, University of Delaware, Newark, Delaware
19716.
Abstract. —Rate of development for the first two instars of the mantid Tenodera sinensis
(Saussure) increased with increasing temperature for cohorts maintained at 20°C, 25°C
and 32°C. An increase in incubation temperature from 20°C to 32°C decreased time in
instar by more than two thirds for photoperiods of 8L:16D, 16L:8D and 24L:0D. The
effect of photoperiod was less pronounced, partly because immobile mantid nymphs could
feed in the dark when nocturnally active prey came into contact with them. Total prey
consumed and biomass of nymphs at ecdysis did not differ among treatment groups. The
effect of temperature apparently was to increase feeding rate, thereby decreasing the time
required to consume the threshold amount of prey biomass necessary to trigger molting.
Key Words:
toperiod, predators
As with other univoltine insects inhab-
iting temperate regions, the life cycle of the
mantid Tenodera sinensis (Saussure) 1s con-
strained by seasonality: it must emerge early
enough in the spring to complete its life cycle
before the onset of freezing temperatures in
the fall, yet not so early that it is in danger
of late spring frosts. In addition to these
abiotic limits, prey are scarce early in the
spring and late in the summer and fall, so
that food limitation effectively narrows the
temporal window available for success in
terms of survival and fecundity (Eisenberg
et al. 1981, Hurd and Eisenberg 1984).
There are three potential disadvantages
to emergence early in the spring: 1) prey are
scarcer than later in the spring, 2) temper-
atures are cooler and the risk of freezing
from a late frost is significant, and 3) pho-
toperiods are shorter. Food shortages for
first instar 7. sinensis nymphs, decrease sur-
vivorship and retard development (Hurd
Mantodea, Mantidae, Tenodera sinensis, development, temperature, pho-
and Eisenberg 1984, Hurd and Rathet 1986).
The importance of temperature to insect de-
velopment in general has been well-docu-
mented (e.g. Tauber et al. 1986, Bowler and
Cossins 1987), which led us to ask how im-
portant this abiotic factor may be to growth
rate of 7. sinensis. Photoperiod, which in-
creases less dramatically than temperature
from early to late spring, might also be im-
portant to visual predators such as mantids
because it presumably determines the
amount of time available for prey capture.
Here we ask how temperature and photo-
period affect developmental rate in newly
hatched nymphs of 7. sinensis.
MATERIALS AND METHODS
Oothecae of 7. sinensis were collected
from New Castle County, Delaware, during
the winter of 1988 and kept in the labora-
tory at 10°C. These oothecae were subse-
quently incubated at 25°C to induce hatch-
530
ing. Upon hatching, nymphs were divided
among three treatment groups of 30 nymphs
each, which differed in hours of light vs.
dark per day: 8L:16D, 12L:12D, and 24L:
OD. The entire light:dark series was run at
three temperatures with three separate co-
horts: 20°C, 25°C, and 32°C. The broad range
of light and temperature was chosen to pro-
vide the greatest opportunity for these
abiotic factors to influence development.
Five additional nymphs were kept in total
darkness at 25°C and otherwise treated as
above, to serve as a check on whether any
feeding could take place in the dark.
Nymphs were housed individually 1n 130
ml glass vials fitted with plastic lids, which
were ventilated with fine mesh brass screen.
Cotton-tipped swabs were inserted through
holes in the caps and the tips were wetted
daily with distilled water. Mantid nymphs
were allowed to crawl into the vials upon
hatching, and were not handled thereafter
in order to avoid injury. All mantid nymphs
were provided prey ad libitum, in the form
of apterous Drosophila melanogaster (Mei-
gen). Numbers of flies consumed in whole
or in part were recorded, and all flies were
replaced daily.
The first 10 nymphs in each group to molt
were killed by freezing, measured for body
length (top of head to tip of abdomen), dried
at 60°C for 48 h and weighed. This exper-
iment was carried out for the first two stadia.
Data obtained in this experiment gener-
ally did not meet the homogeneity of vari-
ance criterion necessary for the application
of parametric ANOVA. Therefore the ef-
fects of photoperiod and temperature were
analyzed using nonparametric Kruskal-
Wallis tests (Sokal and Rohlf 1981).
RESULTS
Temperature had the most pronounced
effect on developmental rates for both in-
stars. Examination of the pattern of molting
among experimental cohorts revealed a
consistent decline in stadium length with
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
increasing temperature for all three photo-
periods (Table 1).
The effect of photoperiod was both less
dramatic and less consistent. Photoperiod
was not correlated with developmental rate
within temperature groupings for first in-
stars (Table 1). The only significant differ-
ence within a temperature group was a
slightly greater mean stadium length in
nymphs exposed to 16 hours of light at 25°C.
Photoperiod had a greater effect on devel-
opmental rate of second instars than firsts.
Stadium duration declined significantly be-
tween the 8L:16D and 16L:8D groups for
nymphs incubated at 20°C and 32°C, but
not at 25°C (Table 1).
Feeding rates of nymphs during both sta-
dia were also more consistently related to
temperature than to photoperiod. During
first stadium, feeding rate increased with in-
creasing temperature for all photoperiods
(Table 2), but was not affected by photo-
period. During second stadium, feeding rate
also increased significantly with increasing
temperature for all photoperiods (Table 2).
As with developmental rate (Table 1), feed-
ing rate increased significantly between the
8L:16D and 16L:8D groups of nymphs ex-
posed to 20°C and 32°C, but not 25°C.
Total flies consumed was not related either
to temperature or to photoperiod. Mean
number of flies eaten by first instars ranged
from 19-22 among the nine experimental
groups; for second instars the range was 35—
44 flies. There was also no significant dif-
ference in biomass attained by nymphs
among the various treatment groups. Dur-
ing first stadium, mean nymph biomass
stadium the range was 5.3-6.1 mg.
The five first instar nymphs kept in total
darkness (OL:24D) were able to feed and to
complete the instar. In fact there was no
significant difference between these nymphs
and those kept at 24L:0D, either in rate of
maturation or in feeding rate. Observations
of nymphs made in the dark under red light
indicated that these flies were active in the
VOLUME 91, NUMBER 4
Table 1.
531
Mean stadium length in days (+SE) for 7. sinensis nymphs. N = 10 in each cell. Asterisks indicate
P < 0.001 (Kruskal-Wallis test) for differences within rows and columns.
Temperature (°C)
20 25 32
First instar:
Light : dark regime 8L:16D 16.7 (0.21) 8.9 (0.10) 5.4 (0.16)*
16L:8D 17.3 (0.49) 9.4 (0.16) 5.0 (0.00)*
24L:0D 16.9 (0.10) 8.0 (0.00) 5.0 (0.00)*
ns as ns
Second instar:
Light : dark regime 8L:16D 27.2 (0.66) 9.4 (0.24) 6.4 (0.40)*
16L:8D 15.6 (0.60) 9.4 (0.51) 5.0 (0.31)*
24L:0D 16.2 (0.49) 9.4 (0.24) 5.0 (0.00)*
* ns *
dark, and were captured by stationary man-
tid nymphs when contact was made. There-
fore, T. sinensis nymphs are at least capable
of night feeding by tactile cues, even when
they cannot see prey. This, of course, does
not explain the significant effects of pho-
toperiod on second instar nymphs (see
above).
DISCUSSION
Our results suggest that temperature is
more important than photoperiod to de-
velopmental rates of the first two instars of
T. sinensis. The initial assumption that pho-
toperiod places a limit on time for prey cap-
ture was found to be incorrect, at least dur-
Table 2.
ing first stadium. However, we do not know
to what extent night feeding is a useful ad-
aptation, or simply a laboratory artifact of
confining nymphs to vials in which noctur-
nally active prey are abundant. Some am-
bush predators, such as thomisid spiders
which use flowers as feeding sites, routinely
feed at night as well as during the day (Morse
1981, R. Fritz pers. comm.). However,
mantid nymphs are not flower specialists,
which calls into question how often prey are
likely to blunder into them in the dark in
the absence of such an attractant.
Although temperature has been found to
have profound effects on body size and even
adult fecundity in other insects (e.g. Laugh-
Mean (+SE) feeding rates of mantid nymphs fed ad /ibitum, in terms of number of flies consumed/
nymph/day. Asterisks indicate P < 0.001 (Kruskal-Wallis test) for differences within rows and columns.
Temperature (°C)
20 25 32
First instar:
Light: dark regime 8L:16D
16L:8D
24L:0D
Second instar:
Light : dark regime 8L:16D
16L:8D
24L:0D
1.34 (0.34) 2.08 (0.09) 3.58 (0.14)*
1.32 (0.05) 1.92 (0.09) 4.43 (0.11)*
1.34 (0.08) 2.27 (0.09) 3.85 (0.11)*
ns ns ns
1.73 (0.08) 4.87 (0.04) 5.49 (0.56)*
2.88 (0.16) 3.74 (0.45) 8.16 (0.43)*
2.79 (0.14) 4.81 (0.37) 8.80 (0.34)*
* ns *
532
lin 1964), there was no relationship between
temperature and size at ecdysis for first and
second instar 7. sinensis. Additionally, there
was no relationship between temperature
and total prey consumed. Feeding rate,
however, consistently increased with in-
creasing temperature. Therefore, the effect
of increasing temperature was to decrease
the time necessary to consume a threshold
amount of prey biomass in order to trigger
molting.
These results raise a question about the
life history strategy of this species: Why does
egg hatch occur so early, at a risk of star-
vation and late frost, when hatching a few
weeks later would expose them to increased
temperature and prey density, both of which
facilitate faster development? For instance,
20°C and 25°C correspond roughly to day-
time temperatures during late April and late
May, respectively, in our area (Bair and
Ruffner 1985). This five degree increment
resulted in a 43% decrease (from 33 to 19
days) in the time it took to complete both
first and second stadia among mantid
nymphs fed ad /ibitum in our experiment
(16L:8D). Nymphs which have hatched at
the end of April in the field have taken up
to 45% longer to develop through second
instar than in this experiment (Rathet and
Hurd 1983, Hurd and Eisenberg 1984), but
field conditions entail variable tempera-
tures as well as food limitation.
Ifabiotic advantages per se do not entirely
account for early hatching in this species,
what biotic factor(s) might be responsible?
Possibilities include avoidance of predators
(or parasites), and reduction in competition.
Other mantid species, sympatric with 7. si-
nensis here and abroad, hatch later in the
spring (Rathet and Hurd 1983, Hurd 1988).
These differences in egg phenology could
mitigate interspecific competition by reduc-
ing size overlap among these species (Hurd
and Eisenberg 1989); mantids of different
sizes take different sized prey (Bartley 1983).
Furthermore, larger mantids have a greater
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
total range of prey sizes they can effectively
take than smaller ones (Bartley 1983). How-
ever, a disparity in size can also result in
predation of later hatching (smaller) nymphs
by those emerging earlier (Hurd 1988).
Therefore, early hatching may confer an ad-
vantage to 7. sinensis over other sympatric
mantid species, which offsets the apparent
abiotic disadvantages.
ACKNOWLEDGMENTS
This work was supported by a NSF Grant
BSR 8506181. This is contribution #122
from the Ecology Program, University of
Delaware.
LITERATURE CITED
Bair, F. E. and J. A. Ruffner. 1985. Weather of U.S.
Cities. Vol. 1. Gale Research Publishing Co., New
York.
Bartley, J. A. 1983. Prey selection and capture by the
Chinese mantid. Ph.D. Dissertation, Univ. Del-
aware, Newark.
Bowler, K. and A. R. Cossins. 1987. Temperature
Biology of Animals. Chapman and Hall, New York.
Eisenberg, R. M., L. E. Hurd, and J. A. Bartley. 1981.
Ecological consequences of food limitation for adult
mantids (7enodera sinensis Saussure). Am. Midl.
Nat. 106: 209-218.
Hurd, L. E. 1988. Consequences of divergent egg
phenology to predation and coexistence in two
sympatric, congeneric mantids (Orthoptera: Man-
tidae). Oecologia 76: 549-552.
Hurd, L. E. and R. M. Eisenberg. 1984. Experimental
density manipulations of the predator Tenodera
sinensis (Orthoptera: Mantidae) in an old-field
community. I. Mortality, development and dis-
persal of juvenile mantids. J. Anim. Ecol. 53: 269-
281.
1989. The temporal distribution of hatching
times in three sympatric mantids (Mantodea:
Mantidae) with implications for niche separation
and coexistence. Proc. Entomol. Soc. Wash. 91:
55-S8.
Hurd, L. E. and I. H. Rathet. 1986. Functional re-
sponse and success in juvenile mantids. Ecology
67: 163-167.
Laughlin, R. 1964. Biology and ecology of the garden
chafer, Phyllopertha horticula (L.). VIII: Temper-
ature and larval growth. Bull. Ent. Res. 54: 745-
759.
VOLUME 91, NUMBER 4
Morse, D. H. 1981. Prey capture by the crab spider
Misuma vatia (Clerck) (Thomisidae) on three
common native flowers. Am. Midl. Nat. 105: 358-
367.
Rathet, I. H. and L. E. Hurd. 1983. Ecological rela-
tionships among three co-occurring mantids, 7en-
odera sinensis (Saussure), 7. angustipennis (S.), and
533
Mantis religiosa (L.). Am. Midl. Nat. 110: 240-
248.
Sokal, R. R. and F. K. Rohlf. 1981. Biometry. W. H.
Freeman, New York.
Tauber, M. J., C. A. Tauber, and S. Masaki. 1986.
Seasonal adaptations of insects. Oxford Univ.
Press, New York.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 534-537
A NEW SPECIES OF HORISMENUS (HYMENOPTERA: EULOPHIDAE)
PARASITIC ON THE LESSER CORNSTALK BORER,
ELASMOPALPUS LIGNOSELLUS (LEPIDOPTERA: PYRALIDAE)
M. E. SCHAUFF
Systematic Entomology Laboratory, PSI, USDA, Agricultural Research Service, c/o
U.S. National Museum, NHB 168, Washington, D.C. 20560.
Abstract.— A new species of Horismenus (elineatus Schauff) (Eulophidae: Entedoninae)
is described and illustrated. This species is a primary gregarious endoparasitoid of the
lesser cornstalk borer, E/asmopalpus lignosellus (Zeller), and is native to Bolivia. This
species is being studied for possible introduction into the United States for control of the
borer.
Key Words:
The lesser cornstalk borer, E/asmopalpus
lignosellus (Zeller), is a pest of corn in the
southern United States because it damages
the lower part of the cornstalk (Metcalf et
al. 1962). During a search for natural ene-
mies in South America, a new species of
eulophid was discovered in Bolivia that was
attacking the larvae of the borer. This species
was submitted to the USDA’s Systematic
Entomology Laboratory for identification,
and the specimens were found to represent
an undescribed species in the genus Horis-
menus. | am naming this species so that
quarantine, importation, and work on the
biology can proceed.
Species of Horismenus can be differen-
tiated from other eulophids by the following
characters: scutellum with one pair of setae:
submarginal vein of forewing with 2 setae
(subfamily Entedoninae); median propo-
deum with a smooth or lightly sculptured
raised area bordered by sunken, usually
sculptured, areas (Fig. 4); scutellum gener-
ally with a nearly complete median longi-
biological control, Chalcidoidea, corn
tudinal suture; posterior edge of prepectus
interrupted (indented) by anterior margin
of mesosternum (Fig. 6); dorsellum
V-shaped (pointed) postero-medially; peti-
ole reticulate; second metasomal tergum
larger than remaining terga, often covering
half the length of the metasoma; stigmal
vein (Fig. 9) short, and without a distinct
petiole.
The genera most easily confused with
Horismenus are Paracrias and Edovum,
which also have similar patterns of carinae
on the propodeum. But, species of Paracrias
do not have the posterior edge of the pre-
pectus interrupted and the postero-median
dorsellum is not V-shaped, but straight
(Schauff 1985). Species of Edovum (Grissell
1981) can be distinguished by the presence
of an epicnemial carina on the anterior
mesosternum and the presence of longitu-
dinal striae on the petiole. In addition, Par-
acrias species are generally parasitic on wee-
vil larvae and the single known species of
Edovum is parasitic in the eggs of Chryso-
VOLUME 91, NUMBER 4
Figs. 1-7. Scanning electron micrographs of Horismenus elineatus n. sp. 1, Head of 2, frontal view. 2, ¢ Head
and antennae. 3, 2 Scutum and scutellum, dorsal view. 4, 2 Propodeum, dorsal view. 5, 2 Clypeus and lower
face, lateral view. 6, 2 Mesosoma, lateral view. 7, ° Metasoma, dorsal view. Scale line = 0.1 mm,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
aN
:
TAN
WAV A
SENG RAE
536
Lt
ro pa Si aa
Figs. 8-9. 8, 2 antenna. 9, 2 forewing. Scale line = 0.1 mm.
melidae. Horismenus is generally not as-
sociated with Coleoptera nor are they egg
parasites.
Horismenus elineatus Schauff,
New Species
Female.—Length 1.1-1.4 mm. Color
black except the following: apices of all fem-
ora, lateral surface of foretibiae, base of mid
and hindtibiae, first 3 tarsomeres of all legs,
light yellow to white; mid and hindtibiae
sometimes dark to light brown, fourth tar-
somere sometimes brown.
Head.— Antennae (Fig. 8) with three ring-
like anelli; funicular segments wider than
long; lower margin of clypeus protruding
out over mandibles (Fig. 5); entire face (in-
cluding intertorular area), frons, and vertex
lightly to distinctly alutaceous (Fig. 1).
Mesoscutum alutaceous; notauli distinct
posteriorly, fading anteriorly; scutellum
alutaceous except smooth at posterior edge,
more striate outside of the lateral sutures,
sculpture often slightly less strongly ex-
pressed than on scutum, but broadening
slightly behind the axillae with lateral
grooves present only in the expanded area
(Fig. 3); median longitudinal groove present
only in anterior 4 to '3 and often only dis-
cernible as a slight change in the pattern of
sculpture; lateral propodeum (Fig. 4) cov-
ered by alutaceous sculpture which is
strongest medially and posteriorly.
Metasoma approximately 1.2 x as long as
wide (excluding petiole), shorter than the
mesosoma; petiole 1.2 to 1.0 = as long as
wide, tergum 2 covering 2 to nearly entire
length of metasoma (posterior terga tend to
telescope inward in air dried specimens):
lightly alutaceous over dorsal and lateral
surface, expect near basal fovea where the
surface is smooth and shining (Fig. 7).
Male.—Length 1.1-1.2 mm. Similar to
the female except the following: antenna
(Fig. 2) with scape swollen medially, ventral
sensory ridge extending nearly its entire
length. Metasoma about as long as wide (see
below).
There is considerable variation in the ap-
pearance and relative length to width of the
metasoma in both males and females be-
cause of the way in which specimens are
preserved and mounted. When air dried, the
terminal segments of the metasoma tend to
telescope inward. In some males, nearly all
the segments past the second may be hidden
from view. Otherwise, very little morpho-
logical variation was evident in the speci-
mens available for study.
Using the key to North American species
VOLUME 91, NUMBER 4
of Horismenus (Burks 1971), this species
would key to /ixivorus (Crawford) because
it possesses dark tibiae combined with even
sculpturing of the scutellum. It is easily sep-
arated from that species by the lack of a
distinct longitudinal suture on the scutellum
(suture present for about 3/4 length of scu-
tellum in /ixivorus); the sculpturing of the
median propodeum (raised median area
smooth in /ixivorus); and the length of the
female metasoma (shorter than the head and
thorax in elineatus, longer than the head and
thorax in /ixivorus). There is no key to South
America Horismenus species. However, |
have examined representative specimens of
most of the described species from the Neo-
tropics and they share neither the darkened
femora and tibiae nor the almost complete
lack of a longitudinal scutellar suture.
Distribution.—Known only from Boliv-
ia.
Discussion.—The placement of this
species in Horismenus changes, to some de-
gree, the traditional limits of the genus. One
of the characters used to define Horismenus
has been the presence ofa more or less com-
plete longitudinal scutellar groove. But, my
study of the species of Horismenus and of
related genera has shown that not only is
the length and distinctness of the groove
variable in Horismenus, but that it also oc-
curs in other genera (e.g. Alachua in the New
World (Schauff and Boucek 1987) and Par-
zaommomyia from Australia (Boucek
1988)). Hence, the presence of this character
can no longer be considered a synapomor-
phy for species in the genus.
Biology.—This species 1s a primary gre-
garious endoparasite of the last three larval
instars of E. lignosellus. It pupates in the
dead body of the host (J. W. Smith, pers.
comm.). Whether this species will attack re-
lated lepidopterous hosts is being investi-
gated.
Types.— Holotype 2 on point, antennae
and forewing slidemounted: Bolivia, Santa
Cruz, CIMCA, 22-IX-1988. G. Pruitt.
T88048. Ex. Elasmopalpus lignosellus (Pyr-
537
alidae). Antennae and forewing slide
mounted. Deposited in the U.S. National
Museum of Natural History. Paratypes: 18
2 and | 4 with same data as holotype. 82 2
and 5 6 with data: Bolivia, 10-VI-1976. E.
Colque A. ex. Elasmopalpus lignosellus.
Paratypes are deposited in the USNM and
the following institutions: Texas A & M
Univ., College Station; Canadian National
Collection, Ottawa; British Museum (Nat-
ural History, London).
Etymology.—The species epithet e/inea-
tus is formed from “‘e”’ meaning not or with-
out and “linea” for line and refers to the
lack of a distinct longitudinal scutellar line
or groove.
ACKNOWLEDGMENTS
Iam grateful to J. W. Smith, Jr., M. Rose,
and J. B. Woolley for specimens from the
Texas A & M University and information
on the biology of the parasite. T. J. Henry,
E. E. Grissell, S. Heydon, J. B. Woolley and
J. LaSalle reviewed the manuscript and pro-
vided helpful comments. Linda Lawrence
(Systematic Entomology Lab., USDA) drew
the female wing and antennae.
LITERATURE CITED
Boucek, Z. 1988. Australasian Chalcidoidea (Hy-
menoptera): A biosystematic revision of genera of
fourteen families, with a reclassification of species.
C.A.B. International, Wallingford, UK. 832 pp.
Burks, B. D. 1971. The Nearctic species of Horis-
menus Walker (Hymenoptera: Eulophidae). Proc.
Entomol. Soc. Wash. 73: 68-83.
Grissell, E. E. 1981. Edovum puttleri, n. g., n. sp.
(Hymenoptera: Eulophidae) an egg parasite of the
Colorado potato beetle (Chrysomelidae). Proc.
Entomol. Soc. Wash. 83: 790-796.
Metcalf, C. L., W. P. Flint, and R. L. Metcalf. 1962.
Destructive and Useful Insects. Their habits and
Control. McGraw Hill Book Co., New York. 1087
pp.
Schauff, M. E. 1985. The new world genus Paracrias
Ashmead (Hymenoptera: Eulophidae). Proc.
Entomol. Soc. Wash. 87: 98-109.
Schauff, M. E. and A. Boucek. 1987. Alachua flori-
densis, a new genus and species of Entedoninae
(Hymenoptera: Eulophidae) parasitic on the Flor-
ida carpenter ant, Camponotus abdominalis (For-
micidae). Proc. Entomol. Soc. Wash. 89: 660-664.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 538-544
MEGALONOTUS SABULICOLA (HETEROPTERA: LYGAEIDAE),
AN IMMIGRANT SEED PREDATOR OF
CENTAUREA SPP. (ASTERACEAE):
DISTRIBUTION AND HABITS IN EASTERN NORTH AMERICA
A. G. WHEELER, JR.
Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn-
sylvania 17110.
Abstract.—The North American distribution of Megalonotus sabulicola (Thomson), a
European rhyparochromine lygaeid accidentally introduced to the Pacific Northwest and
East Coast, 1s reviewed. Delaware, New Jersey, Pennsylvania, Rhode Island, Virginia,
and West Virginia are listed as new state records, and additional localities are cited for
Maryland and New York. In the mid-Atlantic region this bug feeds mainly on fallen seeds
of spotted knapweed, Centaurea maculosa Lam. (Asteraceae), a noxious weed of western
rangelands and invasive plant in ruderal habitats of eastern North America. This bivoltine
seed predator could be considered a beneficial immigrant because it destroys spotted
knapweed seeds. Its effects on plant density, however, apparently are negligible. Characters
are provided that facilitate recognition of this Palearctic bug in the New World fauna.
Key Words: immigrant insect, distribution, eastern North America, seed predation, spot-
ted knapweed, Centaurea maculosa
Megalonotus sabulicola (Thomson) is a
widespread Old World lygaeid known in
Europe to range from Sweden and England
south to the Mediterranean countries, where
it seems particularly common, and east to
the European U.S.S.R; it also occurs in
northern Africa (Morocco) and in Turkey,
Syria, and Israel (Slater 1964). In Europe,
M. sabulicola was long considered a sub-
species of MM. chiragra (F.) and is so listed
in Slater’s (1964) world catalog. But, as
Southwood (1963) pointed out, the two
overlap in their range and sometimes even
in habitat; therefore, M. sabulicola cannot
be considered a subspecies of M. chiragra.
Southwood (1963) and Roubal (1965) de-
termined that these sympatric bugs do not
substantially intergrade and concluded that
they represent distinct species, although
morphological separation is not always
clearcut.
The habits of W/. sabulicola in Europe are
not well known. It has been collected from
the litter layer in various microhabitats and
apparently is most common in sandy areas
(Butler 1923, Southwood and Leston 1959).
Pfaler (1936) reported that adults overwin-
ter and that Finnish populations are bivol-
tine. Several workers have provided eco-
logical notes (Slater 1964 and references
therein), but detailed information on host
plants and biology is lacking for European
populations.
The first New World record of this rhy-
parochromine was from California (as Rhy-
parochromus chiragra californicus) (Van
Duzee 1928). Since then, M. sabulicola has
been recorded in the Pacific Northwest from
VOLUME 91, NUMBER 4
British Columbia, Oregon, and Washington
(Scudder 1960, 1961, Slater 1964).
Slater and Sweet (1958) first reported this
immigrant lygaeid from eastern North
America, listing it from three localities in
Connecticut. Sweet (1964) gave Massachu-
setts (Cape Cod) as a new state record and
additional Connecticut records. Inclusion
of District of Columbia, New York, and
Pennsylvania under the known eastern dis-
tribution of M. sabulicola (as M. chiragra)
in the world lygaeid catalog (Slater 1964)
was misleading. As Slater and Sweet (1958)
discussed, these records represent quaran-
tine interceptions at ports of entry rather
than established populations. The only east-
ern records subsequent to Slater’s (1964)
catalog are ones likely to be overlooked: Ith-
aca, New York, cited without collection data
(Pimentel and Wheeler 1973), and Talbot
Co., Maryland, based on one female (Hoe-
beke 1977).
Because M. sabulicola had been inter-
cepted at ports on produce originating in
Czechoslovakia or France and because this
species is less common in Great Britain than
M. chiragra, Sweet (1964) suggested that
continental Europe was the likely source of
established eastern U.S. populations. Slater
and Sweet (1958) postulated that 7. sabu-
licola was introduced to the Northeast with
imported plant material rather than with
ships’ ballast. A ballast origin of Pacific
Northwest populations, however, is prob-
able (Slater and Sweet 1958, Scudder 1961).
Herein, I give new distribution records of
M. sabulicola in the eastern states and dis-
cuss its association with spotted knapweed,
Centaurea maculosa Lam., a naturalized
weedy composite of railroad right-of-ways
and highway embankments. Characters al-
lowing this immigrant heteropteran to be
recognized in the Nearctic fauna are pro-
vided.
DISTRIBUTION IN EASTERN NORTH AMERICA
In addition to published records from
Connecticut, Maryland, Massachusetts, and
\
fe ae
Wee
Pee as x
a | _
‘ \
sou
A
Fig. 1. Known distribution of Megalonotus sabu-
licola in eastern North America. Open circles indicate
previously published records; closed circles represent
new records.
New York, the following records are avail-
able for M. sabulicola in the East (Fig. 1).
Except as noted, all collections were made
under Centaurea maculosa by the author
(AGW); collection data are cited for Pi-
mentel and Wheeler’s (1973) New York rec-
ord from Tompkins Co. Voucher specimens
have been deposited in the insect collections
of Cornell University (CUIC), Pennsylva-
nia Department of Agriculture (PDA), and
U.S. National Museum of Natural History
(USNM).
DELAWARE: New Castle Co., Newark,
15 Oct. 1987, AGW and T. J. Henry.
MARYLAND: Harford Co., 195 S, Aber-
deen Proving Grd. exit, 15 Oct. 1987, AGW
and T. J. Henry; Prince Georges Co., 195 §
nr. Laurel, 15 Oct. 1987, AGW and T. J.
Henry. NEW JERSEY: Atlantic Co., Rich-
land, 7 Oct. 1987; Cumberland Co., Vine-
land, 26-28 May 1979, E. R. Hoebeke. NEW
540
YORK: Chemung Co., Rt. 427 w. of Che-
mung, 19 Oct. 1987; Tioga Co., Rt. 79. 1
mi. e. of Richford, 29 Oct. 1987; Tompkins
Co., Savage Farm, Ithaca, 3 July, | Aug.
1966, on alfalfa. PENNSYLVANIA: Bed-
ford Co., Rt. 220, s. of Cessna, 3 May 1988;
Berks Co., 3 mi. e. of Bethel, 8 Oct. 1987;
Rt. 183 n. of Leinbachs and Rt. 198 s. of
Bernville, 22 Oct. 1987; Blair Co., Franks-
town, 17 Nov. 1987; Tipton, 17 Apr. 1988;
Bradford Co., Canton, 16 Oct. 1987; Cum-
berland Co., 181 N, 2 mi. s. of Rt. 114 nr.
New Kingstown; Dauphin Co., Early Mill
Rd., 4.5 mi. n. of Hershey, 10 Aug. 1987,
in house; Franklin Co., Rt. 75 s. of Metal;
PA Turnpike w. of Blue Mtn. tunnel and w.
of Rt. 997, 9 Oct. 1987; Fulton Co., Rt. 30
e. and Rt. 16 s.e. of McConnellsburg, 9 Oct.
1987; PA Turnpike, Sideling Hill Plaza, 3
May 1988; Huntingdon Co., Huntingdon,
17 Nov. 1987; Lackawanna Co., Rt. 9 (NE
Ext., PA Turnpike) nr. Scranton, 31 Mar.
1988: Lancaster Co., 1283 W at Rt. 772 nr.
Mount Joy, 29 Sept. 1987; Elizabethtown,
3 Aug. 1988, on Pinus mugo;, Lebanon Co.,
I81 N, nr. Rt. 934, 28 Sept. 1987; Rt. 934
nr. junc. Rt. 22 nr. Harper Tavern, 28 Sept.,
8 Oct. 1987 and 24 Mar., 17 & 24 May, 1
& 17 June 1988, AGW, J. E. Fetter, & J. F.
Stimmel; Luzerne Co., Hazleton, 29 Oct.
1987; Lycoming Co., Trout Run, 16 Oct.
1987; McKean Co., Bradford, 16 Aug. 1988;
Mifflin Co., 8 mi. e. of Mt. Union, 17 Nov.
1987; Perry Co., Rt. 322 nr. Millerstown,
19 Apr. 1988; Schuylkill Co., 181 N, .5 mi.
s. & | mi. n. of Pine Grove exit and s. of
Ravine, 8 Oct. 1987, AGW and J. F. Stim-
mel; Union Co., Rt. 15, 1.2 mi. s. of Lew-
isburg, 16 Oct. 1987; Wyoming Co., Tunk-
hannock, 29 Oct. 1987. RHODE ISLAND:
Kent Co., 1958, n. of Rt. 102 n.e. of Exeter,
3 Dec. 1987; VIRGINIA: Augusta Co.,
Staunton, 7 Apr. 1988; Roanoke Co., Rt.
220 n. of Boones Mill, 10 Apr. 1988; Rock-
bridge Co., Rt. 11, 2 mi. s. of Fairfield, 10
Apr. 1988; Shenandoah Co., Rt. 11, 1 mi.
n. of Edinburg, 26 Nov. 1987 and Rt. 11 s.
of Edinburg, 3 Dec. 1987; Wythe Co., junc.
I81S & 1778S nr. Ft. Chiswell, 22 Apr. 1988,
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
AGW & T. J. Henry. WEST VIRGINIA:
Berkeley Co., 1818 nr. Falling Waters, 26
Nov. 1987.
HABITAT AND Host PREFERENCES
Following the first Pennsylvania collec-
tion of M. sabulicola, a specimen taken in
my house in Dauphin Co. on 10 August
1987, a survey was initiated to learn more
about its distribution in the eastern states
and to determine its host plant and habitat
preferences. Slater and Sweet (1958) re-
ported a close association of this lygaeid
with cornflower or bachelor’s-button, Cen-
taurea cyanus L., in New England, and Sweet
(1964) observed the bugs using their beaks
to carry or drag fallen seeds to a sheltered
site for feeding. This introduced species ap-
pears to compete poorly with native rhy-
parochromines and in New England is near-
ly always restricted to C. cyanus colonies
growing in temporary habitats (Sweet 1964).
Because the European C. cyanus 1s not ex-
tensively naturalized in Pennslyvania
(Wherry et al. 1979), I began looking for /.
sabulicola under spotted knapweed, C. ma-
culosa,acommon European weed that grows
in ruderal situations similar to those in which
Slater and Sweet (1958) encountered C. cy-
anus and associated populations of M. sa-
bulicola, i.e. early-succession xeric sites.
Nymphs and adults were found initially
by looking under C. maculosa plants colo-
nizing shaly slopes along interstate high-
ways (181 in Lebanon Co. and 1283 in Lan-
caster Co., Pennsylvania). To allow more
sites to be surveyed, this rather time-con-
suming, scratch-and-search collecting
method (Slater and Baranowski 1978: 9) was
abandoned. Instead, a small shovel was used
to scoop soil (about 0.25 ft?) beneath spotted
knapweed into a plastic bag. In the labo-
ratory, specimens of M. sabulicola could be
obtained by processing soil samples in a
Berlese funnel, but its presence at a site could
be determined more easily by sorting soil
dumped into a white enamel tray. The bugs
typically became concentrated in the bot-
tom of the plastic bag and were collected
VOLUME 91, NUMBER 4
when the last batch of soil was placed in the
tray.
Populations were readily detected by this
method, and it was soon apparent that this
immigrant rhyparochromine is common in
eastern Pennslyvania. Fewer than five of the
nearly 30 soil samples taken were negative
for M. sabulicola. West of the Allegheny
mountains, however, populations were more
difficult to locate.
Megalonotus sabulicola was found con-
sistently under C. maculosa. Soil nearly al-
ways could be taken beneath pure colonies
owing to the plant’s allelopathic effects on
nearby vegetation (Fletcher and Renney
1963). Bugs were absent in the few soil sam-
ples taken under other composites, e.g. Eu-
patorium and Solidago spp. Nymphs and
adults were found not only under spotted
knapweed in sparse colonies on shaly high-
way embankments but under plants grow-
ing in other disturbed situations: in wet
clayey soil, in vacant lots and at the edge of
roads, in thick sod of pastures, in gravelly
soil at the edge of parking lots, in gravel pits
and stone quarries, and in fine sooty and
sandy ballast along railroads.
As Slater and Sweet (1958) and Sweet
(1964) noted, 7. sabulicola is restricted to
the litter layer near Centaurea plants. Dur-
ing the distribution survey, no adults were
taken by sweeping spotted knapweed.
Adults, probably dispersing individuals,
sometimes occur on other vegetation; for
example, they have been collected on straw-
berries (Scudder 1961) and corn (Hoebeke
1977). On two occasions at Ithaca, New
York, adults were collected on alfalfa, Med-
icago sativa L.; at Elizabethtown, Pennsyl-
vania, an adult was beaten from a small
Swiss mountain pine, Pinus mugo Turra.
During 1987-88, a population of . sa-
bulicola associated with spotted knapweed
growing at the edge of a gravel parking lot
along Rt. 934 near its junction with Rt. 22
(Lebanon Co., Pa.) was monitored more fre-
quently than other sites but only at irregular
intervals. Based on a composite of obser-
vations made in Pennsylvania and other
541
mid-Atlantic states, phenology conforms
generally with that reported for bivoltine
populations of the bug in New England
(Sweet 1964).
This lygaeid overwinters in the adult stage.
Overwintering individuals were found in
litter and soil taken beneath spotted knap-
weed from mid-October to early December
and during late March to early May. In
Pennsylvania, first and second instars oc-
curred in early June, with first generation
adults appearing by early July. Early instars
of a second generation were not collected,
but fourth and fifth instars were found dur-
ing mid-August. They were present until late
September and were found in southern New
Jersey with a few second and third instars
in early October.
At Canaan in Connecticut’s northern
highlands, Sweet (1964) found that over-
wintered adults oviposited from May to ear-
ly June, the tacklike eggs attached to litter
beneath hosts, sand, or plant fuzz. In early
June, Connecticut populations consisted
mainly of first and second instars; by late
June, a few fifth instars were present with
larger numbers of instars II-IV. First gen-
eration adults appeared during July, and first
instars of a second generation were ob-
served in late July. The two generations
overlapped considerably; some first gener-
ation females continued to oviposit into
August. Second generation adults entered a
reproductive diapause. By late September,
Sweet found that the Canaan population
consisted of 80% adults, 19% fifth instars,
and 1% fourth instars. By mid-October,
adults made up 95% of the population.
Compared to Canaan, phenology at Storrs
in eastern Connecticut was advanced by
about three weeks (Sweet 1964).
EFFECTS ON SPOTTED KNAPWEED
POPULATIONS
Centaurea maculosa is a biennial or short-
lived perennial (Watson and Renney 1974,
Maddox 1979) belonging to the composite
subfamily Cynaroideae, which includes the
well-known thistle genera Carduus and Cir-
542
sium. In some recent schemes of higher clas-
sification, Centaurea is placed in the tribe
Cardueae, subtribe Centaureinae (Dittrich
1977, Zwolfer 1988). Spotted knapweed is
thought to have been introduced to North
America in alfalfa seed that originated in
continental Europe or Asia Minor-Turk-
menistan (Maddox 1979). Although this in-
vasive plant is a pioneer species useful in
erosion control, it poses a serious threat to
western rangelands, where millions of acres
are infested (Maddox 1979). It also is pes-
tiferous in rangelands of British Columbia
(Strang et al. 1979) and is listed as a noxious
weed in Manitoba (Watson and Renney
1974). Well established in Ontario, Quebec,
and the Canadian maritime provinces
(Watson and Renney 1974), spotted knap-
weed generally infests the eastern United
States except for the Deep South (USDA
1971).
Since 1970, several seed-and root-feeding
insects have been evaluated and released for
their biocontrol potential against spotted
knapweed (Harris and Myers 1981, Miller
et al. 1988). Megalonotus sabulicola, an in-
advertent introduction, should be included
among the seed-destroying insects associ-
ated with C. maculosa. Sweet (1960, 1964)
discussed the ecological role of seed bugs
belonging to the large lygaeid subfamily
Rhyparochrominae. Seed destruction not
only results from the piercing of seed coats
but, according to Sweet (1964), probably
from subsequent invasion by pathogenic
fungi. Eyles (1964) also discussed seed-feed-
ing behavior of rhyparochromines, includ-
ing destruction of the embryo.
Habits of M. sabulicola have not been
studied in western North America, but the
identification of Centaurea cyanus seeds as
a preferred food source in New England
(Slater and Sweet 1958, Sweet 1964) gave
more meaning to the seemingly accidental
occurrence of the lygaeid in Oregon pea
fields. In reporting three specimens from the
Willamette Valley, Larson and Hinman
(1932) remarked that the crop was infested
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
by weeds, mainly cornflower, C. cvanus, and
Canada thistle, Cirsium arvense (L.) Scop.
The lygaeid probably was feeding on C. cy-
anus seeds in the litter layer of pea fields in
Oregon and may now be established in areas
of western rangeland infested with C. ma-
culosa.
In the East, /. sabulicola, as a seed feeder
closely associated with spotted knapweed,
should be considered beneficial. Prolific seed
production, however, is characteristic of C.
maculosa, with as many as 40,000 seeds/
m? capable of being produced (Watson and
Renney 1974). Some of the insects released
in biocontrol efforts against Centaurea spp.
significantly curtail seed production but have
failed to limit plant density (Harris and
Myers 1981, Miiller et al. 1988). Megalo-
notus sabulicola may also destroy large num-
bers of seeds, but its effects on plant repro-
duction must be minimal. Even in areas of
Pennsylvania where this accidentally intro-
duced seed predator has become abundant,
stands of spotted knapweed continue to
flourish.
RECOGNITION FEATURES
Megalonotus sabulicola (Fig. 2) may be
placed in the large subfamily Rhyparo-
chrominae by having the suture between ab-
dominal sterna 4 and 5 curving anteriorly,
not meeting the lateral margin. Adults are
hairy, medium-sized, usually macropterous
lygaeids with hemelytra a mottled brown-
testaceous, somewhat contrasting with the
dull piceous head, pronotum, and scutel-
lum. Adults can be additionally character-
ized as 3.80-5.40 mm long (New York and
Pennsylvania specimens), somewhat elon-
gate, broadest across posterior third of he-
melytra; head relatively short, dectivent,
rostrum extending to bases of mesocoxae;
pronotum trapeziform, punctate, separated
into anterior and raised posterior lobes, an-
terior margin lacking ringlike collar, lateral
margin distinctly carinate; scutellum punc-
tate, longer than wide; long, erect, bristlelike
setae on head, pronotum, and scutellum
VOLUME 91, NUMBER 4
Fig. 2. Megalonotus sabulicola, adult habitus; scale
bar = 1.0 mm.
(those on head and scutellum about 0.25
mm long) intermixed with finer, paler, more
appressed setae; forefemur black, incrassate
with prominent tooth ventrally on apical
third: meso- and metafemora black apical-
ly, yellowish basally; tibiae yellowish, with
stout dark spines.
The antennal and tibial characters South-
wood (1963) used to separate M. sabulicola
from M. chiragra—antennal segment II and
hind tibiae in sabulicola almost wholly yel-
low and antennal segment III with broad,
central, yellow band—do not always hold
true for eastern U.S. specimens. Sweet (1964)
noted that in certain New England popu-
lations antennae and hind tibiae showed the
543
dark chiragra-like color patterns. Speci-
mens (ca. 120) collected in this study, how-
ever, always have antennal segment II and
the hind tibiae yellow or pale yellowish
brown, whereas antennal segment III varies
from mostly yellow to entirely black.
Fifth-instar nymphs can be keyed in Sweet
and Slater (1961); nymphal characters dis-
tinguishing members of the tribe Megalono-
tini are discussed by Slater and Sweet (1961).
Slater and Sweet (1958) described instars
III-V.
ACKNOWLEDGMENTS
I am grateful to T. J. Henry (Systematic
Entomology Laboratory, USDA, c/o U.S.
National Museum of Natural History,
Washington, DC) for identifying MZ. sabu-
licola and for helpful comments on an early
draft of the manuscript, and to J. F. Stimmel
(BPI, PDA) for assistance in the field and
laboratory, the photograph used in Fig. 2,
and for helpful comments on the manu-
script.
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tera-Heteroptera. H. F. & G. Witherby, London.
682 pp.
Dittrich, M. 1977. Cynareae—systematic review, pp.
999-1015. In Heywood, V. H., Harborne, J. B.,
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of the Compositae. Vol. II. Academic Press, Lon-
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Eyles, A. C. 1964. Feeding habits of some Rhypar-
ochrominae (Heteroptera: Lygaeidae) with partic-
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R. Entomol. Soc. Lond. 116: 89-114.
Fletcher, R. A. and A. J. Renney. 1963. A growth
inhibitor found in Centaurea spp. Can. J. Plant
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Harris, P. and J. H. Myers. 1981. Centaurea diffusa
Lam. and C. maculosa Lam. s. lat., diffuse and
spotted knapweed (Compositae), pp. 127-137. Jn
Kelleher, J. S., and Hulme, M. A., eds. Biological
Control Programmes against Insects and Weeds
in Canada 1969-1980. Commonwealth Agric.
Bureaux, Slough, England.
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Larson, A. O.and F.G. Hinman. 1932. Insects found
on pea fields in the Willamette Valley, Oregon,
after harvest. J. Econ. Entomol. 25: 971-976.
Maddox, D. M. 1979. The knapweeds: Their eco-
nomics and biological control in the western states,
U.S.A. Rangelands 1: 139-141.
Miiller, H., D. Schroeder, and A. Gassmann. 1988.
Agapeta zoegana (L.) (Lepidoptera: Cochylidae),
a suitable prospect for biological control of spotted
and diffuse knapweed, Centaurea maculosa Mon-
net de la Marck and Centaurea diffusa Monnet de
la Marck (Compositae) in North America. Can.
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the world. Univ. of Connecticut, Storrs. 2 vols.
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PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 545-550
AN EXCEPTIONAL FOSSIL AMBER COLLECTION ACQUIRED
BY THE SMITHSONIAN INSTITUTION
DONALD R. DAvis
Department of Entomology, National Museum of Natural History, Smithsonian Insti-
tution, Washington, D.C. 20560.
Abstract.—One of the largest collections of fossil arthropods preserved in amber from
the Dominican Republic has been acquired by the Smithsonian Institution’s Department
of Entomology. The collection, known as the Brodzinsky/Lopez-Penha Collection, com-
prises over 5000 amber samples estimated between 20 and 30 million years old. Each
sample contains from | to over 65 specimens of arthropods representing 22 insect orders
as well as several arachnids and myriapods and some plant material.
Key Words:
The Department of Entomology of the
National Museum of Natural History, with
the support of the Smithsonian Institution’s
Collection Acquisition Program, has re-
cently acquired one of the largest collections
of fossil organisms 1n Dominican amber.
The collection was amassed by Jacob Brod-
zinsky and Marianela Lopez-Penha Brod-
zinsky of Santo Domingo, Dominican Re-
public. The Brodzinsky/Lopez-Penha
Collection comprises over 5000 amber
samples, with each piece containing from
one to approximately 65 fossilized organ-
isms (Figs. 3, 4). Although a tally has not
been completed, a conservative estimate
would place the number of insect inclusions
in this collection at well over 10,000.
For several millennia humans have been
fascinated by amber, not only by its beauty
but also for its use in amulets and medi-
cines. Its formation is still only partially
understood, although its general properties
are well known. Amber is a fossilized, highly
polymerized resin and, consequently, of
vegetable origin. It requires millions of years
to form, and exposure to the catalytic action
of salt water probably is needed to harden
amber, fossil insects, fossil arthropods
and polymerize it. It has a hardness varying
between | and 3, a specific gravity between
1 and 1.3, and a melting point ranging from
120°C to 400°C (usually 300°C to 400°C for
Dominican amber). These characteristics
distinguish amber from the more recent co-
pals and most of the plastic imitations
(Larsson 1978, Poinar 1985). Amber is not
a stable substance but, instead, can gradu-
ally oxidize, dry out, and crack once re-
moved from its natural ground or seabed
deposit and polished. Consequently, one of
the clues to preservation is storing the sam-
ples in containers with minimal exposure to
air but, as is true for pearls, with occasional
handling to provide a protective oil film.
Dominican amber is frequently clear and
honey colored although it can range in color
from nearly transparent to jet black through
yellow, red, blue, and green (Baroni-Urbani
and Saunders 1980). Amber deposits around
the world vary greatly in age, in degree of
fossilization, as well as in plant origin. In
the Dominican Republic this material 1s
found as secondary deposits in mid-Ter-
tiary sandstone marine silts which range be-
tween 20 to 30 million years in age. The
546 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
mamas |) GRRE
3
SROBELS RN’ RECTS oats
Figs. 1-5. Brodzinsky/Lopez-Penha Amber Collection. 1, Standard USNM drawer with trays containing
individual samples. 2, Detail of sample tray showing label data and samples enclosed in zip-lock plastic bags.
3, Sample 10747, containing approximately 50 specimens of mostly Dolichopodidae (4 mm). 4, Sample 10755,
containing approximately 65 specimens of Dolichopodidae (4 mm). 5, Sample containing a flower petal from
the tree (Hymenaea) and probable source of the original resin, and a mycetophilid fly. (Scale length in parentheses.)
VOLUME 91, NUMBER 4
primary plant source of Dominican amber
is believed to be Hymenaea, an extant genus
of Leguminosae currently represented by one
African species and about two dozen neo-
tropical species (Poinar 1985).
The scientific value of amber lies pri-
marily in the inclusions it often contains;
frequently these inclusions are insects. The
preservation of some specimens, in which
the object itself is often intact and is not
merely represented by an impression or
some mineral replacement, can be so perfect
to appear unreal considering their antiqui-
ty.This has prompted some scientists to lik-
en such samples as “windows to the past.”
The natural fixatives in some plant resins
have preserved tissues so well that section-
ing has revealed such minute anatomical
details as nuclei and other cellular inclu-
sions (Poinar and Hess 1982).
Amber from the Dominican Republic was
first reported by Christopher Columbus af-
ter his second voyage between September
1492 and March 1496 (Hale 1891). No fur-
ther mention of this material has been noted
until the early twentieth century. Sanderson
and Farr (1960) visited one of the major
Dominican deposits and reported on the di-
verse representation of fossil insects. Schlee
and Glockner (1978) lists 28 arthropod or-
ders represented in Dominican amber in
their general review of ‘“Bernstein.”” Only
two years later, this inventory was updated
by Baroni-Urbani and Saunders (1980).
Thus far the Dominican Republic is the only
locality in the West Indies where amber has
been found.
My first experience with Dominican am-
ber, other than examining occasional mu-
seum samples, occurred during a fieldtrip
in 1973. After visiting several amber shops
in Santo Domingo, I became duly im-
pressed at the amount of fossil material being
sold as jewelry and, for all purposes, lost to
science. It was apparent that a major source
of knowledge on the Tertiary insect fauna
was disappearing without adequate sam-
pling or documentation. Subsequently, I
547
routinely urged every amber dealer I occa-
sionally met to allow interested biologists
to examine and perhaps purchase the best
preserved fossils before these were sold on
the general market. In 1977 I had the good
fortune of meeting Mr. Jacob Brodzinsky
and his wife, who like so many previous
amber dealers, visited our entomology de-
partment with the desire to have their fossil
insects identified. The Brodzinsky’s were not
only sympathetic to my plea to allow ento-
mologists to examine their fossil material,
but they also began in earnest to amass a
sizeable collection of their own. A year later
Brodzinsky met Dr. Robert Woodruff of the
Florida Department of Agriculture, an ento-
mologist also interested in amber fossils.
Together with Brodzinsky, Woodruff began
to register this material as a means of per-
manently documenting the identification
and final deposition of the more valuable
specimens. This registry has now ap-
proached 12,000 samples. A major portion
of this material, all collected after 1977 and
containing some of the rarest and best pre-
served fossil specimens, was set aside as the
Brodzinsky/Lopez-Penha Collection.
Jacob Brodzinsky has estimated that ap-
proximately five percent of the amber sam-
ples discovered contain insect remains. This
estimate agrees with the overall inclusion
percentage others have expressed (Baroni-
Urbani and Saunders 1980). Brodzinsky also
has reported (in litt.) that on at least two
occasions he has encountered between 12
and 20 specimens for each pound of amber
examined. The overwhelming majority of
these inclusions either represent fragmen-
tary remains or are so poorly positioned that
identification, even to the family level, may
be impossible. As an example of the culling
process that must be accomplished, Brod-
zinsky estimates that he has examined ap-
proximately 140,000 amber samples con-
taining inclusions since 1977. From this
considerable material, he selected only the
finest examples for his personal collection
and the smaller collections which he sold
548 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
=
Figs. 6-11. Brodzinsky/Lopez-Penha Amber Collection. 6, Sample 9366, centipede (Geophilidae?) and For-
micidae (1 mm). 7, sample 9452, Archaeognatha (2 mm). 8, sample 8873, Gryllidae, Trigonidiinae, Annaxipha
sp. (2 mm). 9, Sample 5204, Strepsiptera (0.5 mm). 10, Sample 10939, Tipulidae, Trentepohlia sp. (2 mm). 11,
Sample 10668, Keroplatidae (Mycetophilidae, sensu latu). (Scale length in parentheses.)
VOLUME 91, NUMBER 4
piecemeal to various scientific institutions,
including the Smithsonian Institution. A
cursory inventory of the Brodzinsky/Lopez-
Penha Collection reveals the high degree of
his selection and desire to assemble a truly
representative sample. Included in the over
5000 samples are 22 of the 26 generally rec-
ognized insect orders. The four orders not
represented are Protura, Anoplura, Mallo-
phaga, and Mecoptera. Ants are among the
most common insects trapped by resin flows
and subsequently entombed in amber. Re-
alizing this, Brodinsky did not “load”’ his
collection with this one family (less than
300 samples contain Formicidae), but in-
stead he attempted to diversify the collec-
tion as much as possible. Among some of
the rarer specimens are: Embioptera (7 ex-
amples), Ephemeroptera (9), Neuroptera (6,
including one intact Hemerobiidae and one
Sialidae), Plecoptera (1), Siphonaptera (1),
Strepsiptera (3), Zoraptera (1), and many
rare Coleoptera, Diptera, Homoptera, Hy-
menoptera, Lepidoptera, and Trichoptera.
Other rare organisms include preserved
flowers (Fig. 5) of the tree (Hymenaea) be-
lieved to be the primary source of the am-
ber, centipedes (Fig. 6), millipeds, isopods,
pseudoscorpions, mites, and one small bird
feather. Among the more unusual samples
are such specimens as copulating pairs of
Diptera (Ceratopogonidae and Chironom-
idae) and a micropezid fly with a colony of
phoretic mites on the ovipositor. George
Poinar of the University of California, and
one of the appraisers of the collection, has
estimated (in litt.) that approximately 20%
of the collection consists of new species or
potential holotypic material. The absence
of some insect groups is also of some inter-
est. Although aphids are relatively common
in Baltic amber, none have yet turned up in
the large sample examined by Brodzinsky.
The Brodzinsky/Lopez-Penha Collection
is now stored in standard insect drawers (Fig.
1) within airtight steel cabinets. Each sam-
ple is enclosed within a small zip-locked
plastic bag which is then placed in individ-
549
ual trays arranged in numerical sequence
according to the Woodruff registry number
(Fig. 2). A permanent register number is
attached to the tray with a duplicate label
included inside the bag with the sample.
The fossil contents are currently in the pro-
cess of being inventoried and recorded in
dBase III for rapid access.
Future acquisition and research on Do-
minican amber could become more diffi-
cult. As recently reported (Anonymous
1988) the government of the Dominican
Republic has now taken more stringent steps
to regulate the export of amber fossils. De-
cree number 288-87 states that the amber
mines are government property under law.
Exportation of amber with plant or animal
fossils is prohibited unless accompanied by
official permission granted by the Museo
Nacional de Historia Natural, Santo Do-
mingo.
Jacob and Marianela Brodzinsky are still
very much involved with their Dominican
amber enterprise. It is hoped they will con-
tinue to discover new and exciting fossils
and bring these to the attention of interested
investigators for some time to come. For
their concern of this scientifically priceless
heritage, all biologists should be grateful.
ACKNOWLEDGMENTS
I wish to thank Jacob Brodzinsky and his
wife Marianela Lopez-Penha for their ded-
icated efforts over the years in assembling
an excellent representation of Dominican
amber fossils. Our Department is grateful
for the support of the Collection Acquisition
Program of the Smithsonian Institution
which provided funds for the purchase of
this collection. I am also thankful to George
Poinar, Jr. of the University of California,
Berkeley, John Tkach of Bozeman, Mon-
tana, and Robert Woodruff, Florida De-
partment of Agriculture and Consumer Ser-
vices for their helpful support. Assistance
with identifications was provided by Wayne
Mathis, Smithsonian Institution, David
Nickle, U.S. Department of Agriculture.
550
Luis Pereira, and Jon Gelhaus, Smithsonian
Postdoctoral Fellows. Acknowledgment is
especially due William Rowe of the Smith-
sonian’s Department of Entomology for his
excellent curation of the collection. The
photographs were taken by Victor Krantz
of the Smithsonian Photographic Labora-
tory, and the final draft of the manuscript
was typed by my secretary, Silver West.
LITERATURE CITED
Anonymous. 1988. Fossil amber from the Domini-
can Republic. ASC Newsletter, Association of Sys-
tematic Collections 16(1): 10.
Baroni-Urbani, S. and Saunders, J. B. 1980. The fau-
na of the Dominican Republic amber: The present
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
status of knowledge. Proceedings of the Ninth Ca-
ribbean Conference, Santo Domingo, pp. 213-223.
Hale, E.E. 1891. The Life of Christopher Columbus
from His Own Letters and Journals. G. L. Howe
& Co., Chicago.
Larsson, S. G. 1978. Baltic amber—a palaeobiolog-
ical study. Entomonograph 1-192.
Poinar, G. O., Jr. 1985. Glimpses ofa Tertiary forest
in amber. Pacific Horticulture 46(3): 39-41.
Poinar, G. O., Jr. and Hess, R. 1982. Ultrastructure
of 40-million-year-old insect tissue. Science 215:
1241-1242.
Sanderson, M. W. and Farr, T. H. 1960. Amber with
insect and plant inclusions from the Dominican
Republic. Science 131: 1313.
Schlee, D. and Glockner, W. 1978. Bernstein. Stutt-
garter Beitrage Naturkunde, Serie C 8: 1-72.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 551-574
REDESCRIPTION OF CX. CORNIGER THEOBALD AND
ELEVATION OF CULEX (CULEX) LACTATOR DYAR AND KNAB
FROM SYNONYMY BASED ON SPECIMENS FROM
CENTRAL AMERICA (DIPTERA: CULICIDAE)
DANIEL STRICKMAN AND JONATHAN PRATT
(DS) Walter Reed Biosystematics Unit, Department of Entomology, Walter Reed Army
Institute of Research. Current address: Department of Entomology, Armed Forces Re-
search Institute of Medical Sciences, APO San Francisco 96346-5000; (JP) Box 212, Spring
Arbor College, Spring Arbor, Michigan 49283.
Abstract. — Culex (Culex) lactator Dyar and Knab is resurrected from synonymy with
Culex (Culex) corniger Theobald on the basis of morphological differences in the adult,
male genitalia, and fourth-instar larva. Both species are redescribed and illustrated. Culex
lactator has been collected from Mexico to northern South America at elevations from
sea level to 1500 m. Most larval habitats were in direct contact with the ground, in contrast
to the container habitats of Cx. corniger.
Key Words: Diptera, Culicidae, Culex, lactator, corniger, coronator, restuans, taxonomy,
canonical variable, discriminant analysis, Central America
Culex (Culex) corniger Theobald is known
from the Caribbean and Mexico to Uruguay
(Knight and Stone 1977). It is easily rec-
ognized by the unusual form of the larva
and the ornamentation on the mesonotum
of the adult. We first saw evidence of two
forms of the species in Honduras during
extensive collecting in many parts of the
country. Study of this and other material in
the collections of the United States National
Museum resulted in the discovery of differ-
ences in all stages of two forms of this species
in Central America. One form resembled
Cx. corniger from its type locality in Para,
Brazil, and the other resembled the lecto-
type (Stone and Knight 1957) of the junior
synonym, Culex lactator Dyar and Knab
from Rincon Antonio, Oaxaca, Mexico. We
therefore elevate Cx. /actator from synon-
ymy to species status.
METHODS
First, detailed morphological examina-
tions were made of specimens from Mexico
to Costa Rica. Once these studies produced
clear means of identification, specimens
were examined from other parts of the Neo-
tropics to understand the distribution of Cx.
lactator. Except for material from the type
locality, no material of Cx. corniger from
outside of Central America was examined.
Morphological terminology is defined in
Harbach and Knight (1980) with modified
terms for the male genitalia taken from Har-
bach et al. (1983). Abbreviations used in
figures were taken from the same references.
Color designations were based on color
printing process, as outlined in Kueppers
(1982) and used previously for mosquitoes
by Strickman (1988). The system describes
colors as combinations of three of the fol-
552
lowing hues: Black (B), cyan (C; a deep sky
blue), magenta (M; a deep reddish purple),
and yellow (Y). Each component of the col-
or mix is expressed as a percentage of the
coverage of a white page (e.g. ByyMo Coo 1s
a sky blue color). To describe a color, an
area of the specimen and the appropriate
color chart were illuminated with light from
tungsten bulbs receiving five volts of elec-
tricity. The color on the specimen was then
matched by eye to the nearest color on the
chart, taking care to examine each color of
the chart surrounded by a neutral gray mask.
For light yellow, the color charts did not
offer enough resolution to describe color ob-
served by eye. In these cases, the color was
described as “yellowish.” The use of this
color system was not intended to replace
qualitative color descriptions; instead, 1t was
meant to provide objective measurements
of a species which would act as standards
for the authors’ perception of color.
Material examined was from the United
States National Museum. Abbreviations are
2 for female adult, ¢ for male adult, 6G for
male genitalia, P for pupa, and L for larva.
Canonical discriminant analyses of larvae
and pupae were performed in an effort to
use combinations of characters to provide
better separation than individual charac-
ters. The CANDISC procedure in SAS soft-
ware (SAS Institute Inc., Carey, NC) was
chosen for the analyses because of its sim-
plicity and availability. This procedure finds
linear combinations of quantitative char-
acters that best separate known classes.
Standardized coefficients enabled ranking
importance of characters regardless of nu-
merical size. Raw coefficients were used to
produce formulas involving characters that
were measured on a specimen. The sum of
products of characters and coefficients re-
sulted in a single value, the canonical vari-
able, used to separate species. The sensitiv-
ity and accuracy, as defined by Griner et al.
(1981), of this method ofidentification were
evaluated by calculating the canonical vari-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
able value for each specimen, then counting
the number of correct identifications.
TAXONOMY
Culex (Culex) corniger Theobald
Culex corniger Theobald, 1903: 173, Fig.
93 (Para, Brazil, male type, female); Belk-
in, 1968: 15 (lectotype designation).
Culex basilicus Dyar and Knab, 1906b: 169
(Arima, Trinidad, female).
Culex hassardii Grabham, 1906: 167 (New-
castle, Jamaica, male).
Culex subfuscus Theobald, 1907: 403 (Mo-
neague, Jamaica, male).
Culex lactator var. loquaculus Dyar and
Knab, 1909: 254 (Corozal, Panama Canal
Zone, female).
Culex rigidus Senevet and Abonnenc, 1939:
68 (Saut-Tigre, French Guiana, male).
Female (Figs. 2, 3-5).— Head: Proboscis
black with white band from 0.5 to 0.75
length; band incomplete on dorsal surface,
separated by as little as single row of dark
scales on either side of channel. Maxillary
palpus and clypeus entirely dark. Antennal
pedicel without scales, yellowish; first and
base of second flagellomere yellowish; re-
mainder black. Vertex with erect narrow
scales light medially, dark brown laterally,
but sometimes few brown ones; decumbent
scales yellowish and white, white on ocular
line. Broad white scales on postgena. Tho-
rax: Mesonotum as in Fig. 2; light scales
yellowish to almost white; background scales
dark brown (Boy Yoo.Mgo); bare areas laterad
of lateral and posterior scutal fossal and ac-
rostichal scales; bare area at posterior of
prescutellar area. Scutellar scales pure white.
Lateral of thorax similar to Cx. /actator (Fig.
18). Antepronotum with setae over most of
surface, narrow dingy white scales on ven-
tral portion. Postpronotum with setae on
posterodorsal margin, narrow pale scales
along dorsal margin. Proepimeron bare.
Proepisternum with numerous setae and
sparse to numerous small white scales. Low-
VOLUME 91, NUMBER 4
Figs. 1, 2.
553
Dorsal view of mesonotum and scutellum of females from Honduras (WRBU Acc. 1179). 1, Cx.
lactator (coll. HONC 241-100). 2, Cx. corniger (coll. HONC 230-108).
er mesokatepisternum with 1-6 setae and
group of dingy white, medium broad scales;
upper mesokatepisternum with row of setae
and group of dingy-white moderately broad
scales; prealar knob with numerous setae
and small, translucent scales; pigmentation
of integument between scale patches and
above middle scale patch conspicuously
darker than remainder of sclerite; hypostig-
mal and postspiracular areas bare, with
darkly pigmented integument. Mesanepi-
meron with up to 4 lower setae, 8 upper
setae, and anterior patch of moderately
broad, pure white scales; dark pigmentation
of integument as on mesokatepisternum.
Paratergite without scales. Pattern of pig-
mentation on integument of pleuron in 2
bands separated by white scale patches. Legs:
Dark scales similar in color to proboscis.
Forecoxa with small patch of light scales
dorsally, light ventrally and posteriorly; api-
cal fringe of light scales around segment.
Foretibia for most of its length dark dor-
sally, light ventrally; apical band of light
scales around segment. Foretarsus with bas-
al and apical light bands on tarsomeres |-
4, light scales on basal portion of tarsomere
5; bands sometimes very narrow. Midcoxa
with translucent scales, light integument.
Midfemur dark dorsally and anteriorly, light
ventrally and posteriorly; light scales sur-
round segment at base; dark scales surround
segment near apex; apical fringe of light
scales. Midtibia dark with light scales ven-
trally and apical band of light scales. Mid-
tarsus similar to foretarsus. Hindcoxa with
translucent scales and light integument.
Hindfemur dark dorsally; dark area pro-
gressively wider apically; dark scales sur-
round segment near apex, light scales sur-
round segment at base; border between light
and dark scales near apex roughly a right
angle; apical fringe of light scales. Hindtibia
dark with light scales ventrally and light api-
cal band 5-6 scales wide. Hindtarsus as in
Fig. 5. Wings: Scales on costa, subcosta, ra-
dius, and bases of other veins brown
(Boy Y¥;9My)) and overlapping. Abdomen:
Terga as in Figs. 3, 4; basal bands slightly
yellowish, whiter laterally; dark scales dark
brown (Boy Y+)M,4). Sterna V-VII each with
complete posterior band of dark scales.
Male adult (Figs. 6-8).— Head: Proboscis
dark brown with white band surrounding
false joint; band complete but on dorsal sur-
face dark scales intrude from apex. Maxil-
554 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 3-8. Adult Cx. corniger from Honduras (WRBU Acc. | 179: 2 HONC 230-14; 6 HONC 230-12). Scale
bars do not apply to magnified insets. 3, Female abdomen, dorsal view. 4, Female abdomen, lateral view. 5,
Female hindtarsus and apex of hindtibia, anterior view. 6, Male abdomen, dorsal view. 7, Male abdomen, lateral
view. 8, Male hindtarsus and apex of hindtibia, anterior view.
VOLUME 91, NUMBER 4
lary palpus 1.4 longer than proboscis; dark
brown dorsally with light scales on apical
0.5 of palpomere 5 and middle of palpo-
mere 3, narrow light bands at base of 5, base
and apex of 3 and 4; ventrally, bright white
patches of broad scales on base of 5, just
beyond middle and at base of 4; light scales
at middle and base of 3. Thorax: Light scales
usually not as extensively developed on
mesonotum as in female. Antepronotum
with longer, wider scales than on female.
Legs: Hindtarsus as in Fig. 8. Abdomen: As
in Figs. 6, 7; light scales more extensive than
in female.
Male genitalia (Figs. 9-13).—Ninth ter-
gum (Fig. 13): Shallowly cleft between se-
tose lobes; setae in 3-5 irregular rows, each
seta on a small tubercle; integument of lobes
aculeate. Proctiger (Fig. 12): Basal lateral
arm nearly constant in width to tip; base
with small mesal, rounded projection; ac-
etabulum prominent; cercal setae 1-5 (mode
= 3). Lateral plate (Fig. 10): Dorsal process
small or absent; lateral lobe prominent, aris-
ing from lateral ridge, curving posteriorly
along ventral margin, ending in concave ru-
gulose lobe; denticles directed dorsally,
tightly packed; individual denticles slightly
curved dorsally; ventral arm curved and
papery with mesal thickened ridge; dorsal
arm short, reaching to or slightly beyond
bases of denticles, with pointed or rounded
tip. Viewed dorsally in whole mount, lateral
lobe and ventral arm form 2 or 3 large lobes
with the denticles at their common base.
Gonocoxopodite (Fig. 9): Surface opposite
subapical lobe flattened; integument acu-
leate, subapical lobe protruded as a tab; seta
aa Slender rod, gently curved at tip, with
prominent sclerotized socket at base; seta b
stouter than a, curved and usually recurved
at tip, with prominent sclerotized socket at
base; seta c similar to seta a; seta d—f small
and hairlike; seta g on tubercle, foliform,
usually with weak striations; seta / finer than
a or b, socket much less developed; gono-
stylus with tip minutely divided, claw
trough-shaped.
555
Pupa (Figs. 14, 15; Table 1).—Cephalo-
thorax: Trumpet, median keel, scutum,
metanotum, median portions of metatho-
racic wing, sometimes dorsal portion of
mesothoracic wing darker than other areas
of cuticle; trumpet dark anteriorly. Trumpet
in shape of flattened cone. Abdomen: An-
teromedian edge of terga I, III, IV, and
sometimes terga II, V-VIII, sternum III, and
sometimes sterna II-VI darkly pigment-
ed; lighter pigmentation on median por-
tions of terga, sterna I-III, and sometimes
sterna IV-VII; pigmentation progressively
light posteriorly from III; surface of tergum
I reticulate at 200x; surfaces of terga, sterna
II-VIII, genital lobes minutely spiculate;
posterior of sternum II with fine spicules;
distal 0.5-0.67 of outer margin of paddle
minutely spiculate; spot of pigmentation on
anterior portion of buttress light, dark, or
absent; seta 2-P sometimes absent, very
small when present. Chaetotaxy: As in Ta-
ble 1; Figs. 14, 15.
Larva (Fig. 16, Tables 2, 5).— Head: More
pigmented around mouthparts, on lateralia
and collar; antenna slightly darker from seta
1 to tip; mean antennal tuft ratio (distance
from antennal base to seta 1-A divided by
antenna length) 0.51 (SD = 0.04, min =
0.43, max = 0.62, n = 34). Thorax: Integ-
ument coarsely spiculate, coarsest on me-
sothorax but clearly visible on all segments
at 40x; support plate of setae 9-12 with mi-
nute spines on prothorax, large spines on
meso- and metathorax. Abdomen: Spicu-
lation finer than on thorax; spicules not vis-
ible on all surfaces at 100 x; coarse spicules
on posterior edge of segment X progres-
sively finer ventrad. Siphon: Heavily pig-
mented on acus, base, spiracular apparatus,
spiracular apodeme, and around distal por-
tion of siphon; small projection sometimes
visible on basal side of ring formed by spi-
racular opening of spiracular apodeme;
mean spiracular apodeme ratio (length of
spiracular apodeme divided by dorsal length
of siphon) 0.45 (SD = 0.02, min = 0.42,
max = 0.50, n = 33); modal number of
556 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
lateral
mesal
mesal
y—
dorsal
dorsal
aM
dorsal 2
Figs. 9-15. Male genitalia and pupal exuviae of Cx. corniger. Drawn from Guatemalan specimen GUA 17-
11 (phallosome), Honduran specimen HONC 230-14, WRBU Acc. 1179 (dissected genitalia), and Nicaraguan
specimen NI 45-16 (pupa). 9, Gonocoxopodite, mesal view. 10, Lateral plate, lateral and mesal views. 11,
Phallosome, dorsal view. 12, Proctiger, dorsal view. 13, Ninth tergal lobes, dorsal view. 14, Pupal cephalothorax
(CT), anterior lateral quadrant. 15, Metathoracic wings and abdomen (P = paddle; each magnified inset is 0.05
mm across).
557
VOLUME 91, NUMBER 4
SSS
fal
nw
S-2
al
Larva of Cx. corniger, Nicaraguan specimen NI 45-1. Scale does not apply to enlargements of
Fig. 16.
dorsomentum, spiracular apodeme, pecten spine, or comb scale; each magnified inset of spiculation is 0.07 mm
across. (A = antenna; C = head; CS = comb scale, Dm = dorsomentum; M = mesothorax; P = prothorax;
PMPc = posterior median process; PS = pecten spine; S = siphon; SAd = spiracular apodeme; T = metathorax.)
558
Table 1.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Number of branches of pupal setae of Culex (Culex) corniger. Based on counts made on 10 specimens
from 10 collections in Mexico, Guatemala, El Salvador, Honduras, Nicaragua, and Costa Rica.
Abdominal Segments
Seta Paddle
No. cT I II Il IV Vv VI VII Vil IX
0) - — Is 1 1 1 21(0) 1 1 1 = _
1 1,2(1) 7-12(9) 7-12(10) 2-7(2) 2-5(2) 1-3(2) 1,21) 1,2()) - 1 125)
De ANG 22) M282 l 1 1 1 1 1 - = 1
3 l 2 2 2 3-5 (4) 14(1)) 1,2() 1 - - =
4 2-4(2) 2-5(4) 232)! <455)(5))425.3'@) 2-43), eZ) as 2l)y 210) - —
5 2-4(3) 2-5 (5) 12d) 1-3(@Q 1 1 1 1 — — -
6 1, 2(2) 1 1 1 1 1 1 2-5 (3) - — aa
7 2 1-3 (2) 1-4(2) 3-5(4) 24(2) 2-6(4) 1 1 —- = ~
8 1 ~ 1 2-4 (3) 2-4(3) 2-5(3) 2,3(3) 1-3(2) — - ~
9 1,,2(2) — 1=3:(1) 1 l 1 I 1 2,3(2) 47 (6) _ =
10 3-8 (4) _ = 15-21) a2) 1 1 ] - - -
11 2 — — l 1 12\(1) 1 1 _ - -
12 1, 2(2) — — = - — = = _ -
13 - — - ~ - - — — - - -
14 - — ~ 1 1 l 1 1 1, 25) — =
“ Range (mode).
siphon tufts 6 (min = 5, max = 6, n = 33);
other quantitative measurements in Table
5. Chaetotaxy: As in Table 2, Fig. 16.
Diagnosis. — Culex corniger can be distin-
guished from other members of the subge-
nus in Central America north of Panama by
the following features: Adult: Proboscis with
light band (though not always completely
surrounding proboscis); characteristic me-
sonotal pattern (Fig. 2); basal and apical light
bands on tarsomeres well defined and con-
tinuing beyond base of tarsus 3. In female,
basal light bands of abdominal terga II-VII
not connected laterally to lateral light spots
on at least one segment. Male genitalia:
Ninth tergum shallowly cleft with numerous
setae on lobes; lobes connected by thin strap
of cuticle; lateral plate of characteristic form
(Figs. 10, 11); dorsal process poorly devel-
oped; lateral lobe well developed; subapical
lobe with setae a, b, c, d—f, g (foliform), and
h present; gonostylus narrowed beyond bas-
al 0.3. Larva: Antenna not markedly ta-
pered in apical portion; tuft (seta 1-A) with
3 or fewer branches; linear combination of
characters (see Discussion) including (in or-
der of importance) saddle index, branches
of seta 4-C, siphon index, branches of seta
7-C, ratio of distance to seta la-S (from base
of siphon) to siphon length, number of pec-
ten teeth, and pecten row length index.
Remarks on types and original descrip-
tions.— We were able to examine type spec-
imens for three of the five remaining syn-
onyms for Cx. corniger. Culex basilicus Dyar
and Knab (from Trinidad, 1906b) (¢ lec-
totype designated by Stone and Knight
1957), Cx. hassardii Grabham (from Ja-
maica, 1906) (é lectotype designated by
Belkin et al. 1970) and Cx. Jactator var.
loquaculus Dyar and Knab (from Panama,
1909) (2 lectotype designated by Stone and
Knight 1957) all correspond to Cx. corniger
rather than Cx. /actator, based on characters
discussed in this paper.
The descriptions for the other two syn-
onyms of Cx. corniger indicate their equiv-
alence to that species. The description of
Cx. subfuscus Theobald (from Jamaica,
1907) includes an illustration of male gen-
italia showing a strongly developed seta c
on the subapical lobe, a character of Cx.
corniger. The male genitalia and other adult
characters of Culex rigidus Senevet and
Abonnenc (from French Guiana, 1939) also
correspond to those of Cx. corniger, as ac-
559
VOLUME 91, NUMBER 4
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PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
no 100 90
1 1
“AP
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ey
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Legend Grr. - a
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L109 Culex lactator lectotype f Bei 4
q
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Culex lactator collections e 5
—= y
asi ‘€
Countries from which x -
ae i
Culex corniger examined be,
us aus ao : Le
Published distribution of B q 3
Culex corniger, certain ———— aN >
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uncertain ———— a Noy
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é
Fig. 17.
knowledged by the authors when they later
synonymized their species (Senevet and
Abonnenc 1958).
We examined specimens of Cx. corniger
from its type locality, Para, Brazil. Adults
and male genitalia were similar to speci-
mens from Central America. In the original
description, Theobald (1903) mentioned
characteristics of Central American Cx. cor-
niger, including strong mesonotal pattern,
incomplete abdominal tergal banding on
some segments, and distinct hind tarsal
banding.
Distribution (Fig. 17).—According to
publications, the distribution of Cx. corni-
ger extends from Mexico to Uruguay, in-
cluding the Caribbean basin. Because our
paper is limited geographically to Central
America, we can only state with certainty
that Cx. corniger (as defined here) is present
in locations from which we have examined
specimens. The distribution of the species
in South America may include other species
in an unresolved complex (Belkin et al.
1970).
Geographical distribution of Cx. /actator and Cx. corniger in northern Latin America.
Collection records of specimens exam-
ined for this study indicated that the focal
distribution of larval Cx. corniger includes
treeholes, bamboo, and other container
habitats. Occasionally, the larvae were found
in ground pools. A number of authors have
reported these habitats as typical of Cx. cor-
niger, including Clark-Gil and Darsie (1983),
Belkin et al. (1970), Forattini (1965), and
Galindo et al. (1951).
Material examined.—99 2 75 664 8G 38P
36L from 55 collections. Mexico: Veracruz:
Cordoba, 900 m, 14-17 Jul 1964, E. Fisher
and D. Verity coll. no. MEX 34 and 41,
large treehole, 16 2 8 6 3 6G 2P 2L; 800 m,
28 Jul 1965, D. Schroeder coll. no. MEX
237, stream margin with rocks, 4 @ 2P 2L;
1000 m, 12 Jul 1965, C. L. Hogue coll. no
MEX 380, Heliconia flower bracts, 4 6 2
8G; 6 Jul 1970, D. and K. Schroeder coll.
no. MEX 524, treehole, 3 24 6 2 6G 2P 2L.
Oaxaca: Santa Lucrecia, 19 June 1905, F.
Knab coll. no. 262, old tank, 6 2 1 6 1 8G;
Rincon Antonio, 23 June 1905, F. Knab
coll. no. 272, trench, 4 2 2 6 1 6G; Tehuan-
VOLUME 91, NUMBER 4
tepec, 2 July 1905, F. Knab coll. no. 294,
tank, 5 2 1 4; Alomoloya, 21 Jul 1905, F.
Knab coll. no. 312, stream margin rock pool,
1 @ 241 8G. Campeche: Campeche, 5 m,
21 Jul 1970, D. and K. Schroeder coll. no.
MEX 591, rockhole, 1 2? 2 62 6G 2P IL; 5
Aug 1970, D. Schroeder coll. no. 602, large
rockhole, 22141 6G 2P 2L. Chiapas: Santo
Domingo, 815 m, 22 Aug 1987, Strickman,
Roberts and Wilkerson coll. no. MX 94,
WRBU Acc. 1250, can in cemetery, 12 2 4
61 8G 2P 2L. Guatemala: Chiquimula: 420
m, 17 Dec 1915, R. Morales, in house, | 2.
Guatemala: Guatemala City, 1500 m, 5 Jul
1964, T. J. and J. Zavortink coll. no. GUA
17, hole in cement, | éG. Retalhuleu:
Champerico, <5 m, 2 Jul 1964, V. P. Cow-
sill coll. no. GUA 21, ground pool, 1 4G;
San Sebastian, 300 m, 2 Jul 1964, V. P.
Cowsill and T. J. Zavortink coll. no. GUAK
29, cut bamboo, | 2 1 62 4G. Suchitepéquez:
Mazatenango, 380 m, 3 Jul 1964, V. P.
Cowsill coll. no. GUA 23, old truck axle, 2
2 16 6G 2P 2L. El Salvador: Sonsonate:
Izalco, 430 m, 6 Nov 1971, J. N. Belkin and
S. G. Breeland coll. no. SAL 53, treeholes,
12163 6G 2P 2L. Honduras: Atlantida:
Rio Macora, <5 m, 27 Jul 1985, N. Powers
coll. no. HONC 010, WRBU Acc. 1171,
stream pool, 92441 4G 2P 2L. Yoro: Rio
Jalegua, 690 m, 26 Aug 1986, D. Strickman
coll. no. HONC 176, WRBU Acc. 1179,
crumpled roofing paper, 3 25 6 1 6G 2P 2L.
La Paz: Marcala, 1260 m, 11 Sep 1986, D.
Strickman coll. no. 230, WRBU Acc. 1179
military foxhole, 4 24 46 1 6G 2P 2L. Nic-
aragua: Chinandega: Corinto, 29 May 1945,
1 6 1 6G. Zelaya: Blue Fields, <5 m, 12 Jul
1964, A. Quinonez coll. no. NI 40, ground
pool, 1 ¢1éG1P 1L; 13 Jul 1964, coll. no.
NI 45, ground pool amid tree roots, | 2 1
2 6G 2P 2L; 22 Jul 1964, coll. no. 59, tree-
hole, 1 2; 5 m, 25-26 Nov 1971, D. Schroe-
der coll. no. NIC 101, light trap, 1 2. Costa
Rica: Alajuela: Alajuela, 880 m, 18 Jul 1971,
D. W. Heinemann coll. no. CR 271, 1 @;
Turrucares, 650 m, 31 Jul 1971, S. J. Hei-
nemann coll. no. CR 304, rockholes, | 2 1
6 1 46G 1P 2L; 480 m, 31 Jul 1971, S. J.
561
Fig. 18. Lateral view of thorax of 2 Cx. lactator
(Honduran specimen HONC 241-100, WRBU Acc.
1179).
Heinemann and A. Berrios Arias coll. no.
CR 308, rockholes, 1 2 1 6 2 6G 2P; 3 km
E. of San Mateo, 260 m, 1 Nov 1971, D.
A. Schroeder coll. no. CR 496, 12142 64G
2P 2L; Ciruelas, 780 m, 4 Nov 1971, D. A.
Schroeder coll. no. CR 515, rockhole, | 2 1
6 2 6G 2P 2L. Cartago: Turrialba, 500 m,
23 Aug 1971, D. W. Heinemann coll. no.
CR 390, rockholes, | 6. Guanacaste: 10 km
NW of Liberia, 60 m, 6 Aug 1964, C. L.
Hogue coll. no. CR 187, rockhole, 1 2 1 4
2 6G 2P 2L; 23 Jun 1964, coll. no. CR 182,
resting on rock, | 2; Alajuelita, 1090 m, 23
Jul 1971, S. J. Heinemann coll. no. CR 284,
12161 4G; coll. no. CR 286, rockhole, 2
2; coll. no. CR 285, rockhole, 2 2: coll. no.
CR 288, rockhole, | 6G; Puerto Humo, 10
m, 20 Jun 1975, J. Hayes coll. no. CR 601,
tire, 1 2161 8G. Limon: Westfalia, <5 m,
4 Dec 1962, C. L. Hogue and W. A. Powder
coll. no. 72, 1 2; 8 km E. of Zent, 20 m, 17
Dec 1971, D. A. Schroeder coll. no. CR 564,
treehole, | 9; Limon, 10 m, 16 Jul 1975, J.
Hayes coll. no. CR 610, biting human, | °.
Puntarenas: Cerrillos, 1938, H. W. Kumm
coll. no. CRK 599, 1 6 1 6G; 7 km E. of
Palmar Norte, 40 m, C. L. Hogue coll. no.
CR 163, treehole, 1 2? 144 6G 2P 2L; 6 km
S. of San Vito, 30 Apr 1967, D. F. Veirs, 1
562
2; Las Loras nr. Puntarenas, 9 Sep 1905, F.
Knab coll. no. 334, ground pool, | 4, 1 4G.
San José: San José, 21 Sep 1905, F. Knab
coll. no. 343, barrel, 1 2: San Isidro del Gen-
eral, 750 m, 23 Jun 1964, C. L. Hogue coll.
no. CR 180, concrete pond, 1 2141 4G 2P
2L; Santa Ana, 860 m, 25 Jul 1971, A. Ber-
rios Arias and S. J. Heinemann coll. no. CR
293, broken bamboo, | 2 | ¢ 1 6G; Santa
Ana, 1938, H. W. Kumm coll. no. CRK
283, 12141 4G; 4kmE. of San Isidro de
Coronado, 1520 m, 30 Jul 1971, D. W. and
S. J. Heinemann coll. no. CR 300, ditch, 1
2. No location within country: 3 4G. Pan-
ama: Corozal, USNM Type No. 12050, 2,
lectotype of Culex lactator var. loquaculus.
Jamaica: Kingston, M. Grabham, 4, lecto-
type of Culex hassardii. Trinidad: F. W. Ur-
ich, USNM Type No. 10021, 2, lectotype
of Culex basilicus. Brazil: Para, Apr 1930,
N. C. Davis, 11 6 11 8G. Representative
specimens from Panama, Colombia, and
Venezuela were examined.
Culex (Culex) lactator
Dyar and Knab
Culex lactator Dyar and Knab, 1906a: 209,
Fig. 23 (Rincon Antonio, Oaxaca, Mex-
ico, larva). RESURRECTED FROM
SYNONYMY.
Culex lactator var. lactator Dyar and Knab,
1909: 254 (adult).
Culex corniger var. lactator Howard, Dyar,
and Knab, 1915: 240 (adult).
Culex corniger Theobald, Dyar, 1922: 23
(/actator in synonymy).
The following description lists characters
of Cx. lactator which are different from those
of Cx. corniger.
Female (Figs. 1, 18, 19-21).—Head: De-
cumbent scales of vertex yellowish, dingy
white on ocular line. Broad dingy white
scales on postgena. Thorax: Mesonotum as
in Fig. 1; background scales dark brown
(Boo Y 79 Moo) adjacent to light areas, on scutal
fossa, and along dorsocentral and acrosti-
chal lines; other background scales lighter
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
brown with gold reflection; lateral scutal
fossal light scales sometimes four rows wide
in anterior portion; or less numerous, caus-
ing break in pattern; posterior dorsocentral
light scale sometimes few, absent, or iso-
lated so they form spots or sometimes pat-
tern formed by light scales inconspicuous.
Scutellar scales narrowest on posterior me-
dian part of midlobe. Pleura as in Fig. 18.
Antepronotum with narrow yellowish scales
on ventral portion. Postpronotum with nar-
row golden scales along dorsal margin.
Proepisternum with sparse, broad, yellow-
ish scales. Legs: Foretarsus entirely dark.
Midtarsus dark with narrow, poorly defined
bands of slightly lighter scales at base and
apex of tarsomeres 1, 2, base of tarsomere
MT-3. Hindtibia with light apical band 3-
4 scales wide. Hindtarsus as in Fig. 21. Ab-
domen: Terga as in Figs. 19, 20; basal light
bands reach lateral spots on terga I-VII.
Sternum V covered with slightly yellowish
scales; VI and VII with posterolateral or
complete posterior bands of dark scales.
Male (Figs. 22—24).— Differences from fe-
male similar to those listed for Cx. corniger.
Male genitalia (Figs. 25—29).— Ninth ter-
gum (Fig. 29): Tergum deeply cleft between
lobes; setae in three or four irregular rows.
Proctiger (Fig. 28): Basal lateral arm usually
narrows slightly, then widens to rounded,
excavated tip; cercal setae 2-6 (mode = 3).
Gonocoxopodite (Fig. 25): Seta c only oc-
casionally present (one side of one speci-
men, both sides of another) as rod much
finer than seta c in Cx. corniger; gonostylus
with slightly lumpy outline.
Pupa (Figs. 30, 31; Table 3).—Abdomen:
Posterior of sternum II with stout spicules.
Chaetotaxy: As in Table 3; Figs. 30, 31.
Larva (Fig. 32; Tables 4, 5).— Head: An-
tenna gradually darker from seta | to tip;
mean antennal tuft ratio (distance from an-
tennal base to seta 1-A divided by antenna
length) 0.49 (SD = 0.04, min = 0.40, max
= 0.59, n = 104). Thorax: Spiculation
coarser on dorsal and sometimes ventral
surfaces of meso- and metathorax; spicu-
VOLUME 91, NUMBER 4 563
Figs. 19-24. Adult Cx. /actator from Honduras (WRBU Acc. 1179; 2 HONC 241-100; ¢ HONC 241-21).
Scale bars do not apply to magnified insets. 19, Female abdomen, dorsal view. 20, Female abdomen, lateral
view. 21, Female hindtarsus and apex of hindtibia, anterior view. 22, Male abdomen, dorsal view. 23, Male
abdomen, lateral view. 24, Male hindtarsus and apex of hindtibia, anterior view.
564 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
/ =
\ ‘
)y \
\
dorsal alls
Figs. 25-31. Male genitalia and pupal exuviae of Cx. lactator. Drawn from Mexican specimens MX 19-1
and 19-100, WRBU Acc. 1250 (genitalia) and Honduran specimens HONC 241-19 and 241-20, WRBU Acc.
1179 (pupa). 25, Gonocoxopodite, mesal view. 26, Lateral plate, lateral and mesal views. 27, Phallosome, dorsal
view. 28, Proctiger, dorsal view. 29, Ninth tergal lobes, dorsal view. 30, Pupal cephalothorax (CT), anterior
lateral quadrant. 31, Metathoracic wings and abdomen (P = paddle; each magnified inset is 0.05 mm across).
VOLUME 91, NUMBER 4
Table 3.
from 11 collections in Mexico and Honduras.
565
Number of branches of pupal setae of Culex (Culex) lactator. Based on counts made on 14 specimens
Abdominal Segments
Seta Paddle
No cT I Tl ir IV Vv VI VII Vill IX P
0) —_ - l 1 1 1 1 1 I, 2 (1) - —
1 1,2(1) 9-14(11) 6-14(10) 2-4(3) 2,3(2) 1-3(2) 1,2(1) 1,2) - I 1-3 (1)
2 2 22) 1,2 (1) 1 1 1 1 1 _ — 1,2 (1)
3 1 1, 2 (2) 2 2 3=9:(5)) 71,,2.(2)) 15 -24(2) 1 — -
4 2-4(2) 3-5(5) 2-5 (3) 3-6(4) 2-5(3) 2-4(3) 1-3(2) 1,2(1) 1,2) _ —
5 2-5 (3) 3-8 (3) 1,2(1) 1-3(2) l 1 l | _ - -
6 1-3 (2) l 1,2 (1) l 1 1 l 2-7 (4) - — -
7 2 2 1-3 (2) 3-5 (4) 2-5 (3) 2-5 (4) 1 1 — — _
8 1,2) _ _ 2-5 (3) 2-4(3) 2-4(3) 2-4(3) 1-3 (2) = = —
WI PACA) 1, 2 (2) 1 1 1 1 l 2-4 (2) 4-7 (5) = =
10 4-8 (5) - = 1.21) 15.2.2) 1 1 1 = = _
1 25.3.2) _ - 1 1 1 1 1.2 (1) — - -
12. 1,3(2) = = = = - _ — ~ = =
BO
ea
al
1 |
— ||
=|
“ Range (mode).
lation clearly visible on dorsal side of the
mesothorax at 40 x , elsewhere at 100 x. S7-
phon: Mean spiracular apodeme ratio (length
of spiracular apodeme divided by dorsal
length of siphon) 0.45 (SD = 0.035, min =
0.36, max = 0.56, n = 107); modal number
of siphon tufts 6 (min = 5, max = 7, n =
107); other quantitative larval measure-
ments in Table 5. Chaetotaxy: As in Table
4; Fig. 32.
Lectotype (Figs. 33-42).—Includes adult
male and associated genitalia, larval and pu-
pal exuviae on slides. Adu/t male: Glued to
point with right wing detached and glued
separately; right hindleg absent; dark scales
faded compared to recently collected spec-
imens (brown Boy Yoo Mgo On proboscis,
Bs YooM>, on hindfemur, Byy Yoo.Mgo on ab-
dominal terga); some scales of pleuron
rubbed off. Male genitalia: Includes VI-VIII,
cleared but undissected, gonostyli and sub-
apical lobes arranged for easier viewing.
Ninth tergum and proctiger have been
pushed downward (ventrally) and apically,
distorting their relationship with lateral
plates. In Fig. 34, ventral arms of lateral
plates appear mesad of lateral lobes. Larva:
Divided at abdominal segment VII, thorax
and abdomen twisted. Pupa: Good condi-
tion.
Diagnosis. — Culex /actator can be distin-
guished from other members of the subge-
nus in Central America north of Panama by
the following features. 4du/t: Proboscis with
light band (though not always completely
surrounding proboscis); mesonotal pattern
characteristic (Fig. 1); bands of tarsi narrow,
poorly defined and not extending beyond
base of tarsus 3. Basal light bands on ab-
dominal terga II-VII connected laterally to
lateral light spots. Male genitalia: Ninth ter-
gum deeply cleft with numerous setae on
lobes; lobes connected by broad strap of cu-
ticle; lateral plate of characteristic form,
dorsal process poorly developed; lateral lobe
well developed; subapical lobe with setae a,
b, d-f, g (foliform), and / present; gonostylus
narrowed beyond basal 0.3. Larva: Antenna
not markedly tapered in apical portion; tuft
(seta 1-A) with 3 or fewer branches; linear
combination of characters (see Discussion)
including (in order of importance) saddle
index, branches of seta 4-C, siphon index,
branches of seta 7-C, ratio of distance to
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
566
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VOLUME 91, NUMBER 4
567
Table 5. Comparisons of quantifiable larval characters used to distinguish Cu/ex corniger from Culex lactator.
The characters are listed in order of importance.
Mean + SD (n) and Range
Larval Character
Saddle index® 2.13 + 0.14 (106)
1.58-2.59
No. 4-C branches" 3 (97)
2-4
Siphon index: 1.71 + 0.21 (108)
1.18-2.48
No. 7-C branches? 6 (104)
4-9
la-S ratio® 0.21 + 0.05 (108)
0.14-0.60
No. pecten spines? 9 (108)
6-15
Pecten row index:
0.37-0.73
Cx. lactator
0.58 + 0.05 (107)
Cx. corniger
1.87 + 0.11 (33)
1.61-2.09
2 (31)
2-3
1.82 + 0.28 (33)
1.23-2.41
4 and 5 (33)
4-7
0.27 + 0.05 (33)
0.17-0.36
10 (33)
8-13
0.61 + 0.06 (33)
0.48-0.73
* Dorsal length of siphon divided by dorsal length of saddle.
» Mode (n) and range.
© Dorsal length of siphon divided by width of siphon at base.
4 Distance from base of siphon to first siphon tuft divided by dorsal length of siphon.
© Distance from dorsal base of siphon to last pecten spine divided by dorsal length of siphon.
seta la-S (from base of siphon) to siphon
length, number of pecten teeth, and pecten
row length index.
Distribution (Fig. 17).—We found spec-
imens of Cx. /actator in collections from
central Mexico to central Colombia. The
lack of specimens from Nicaragua and Cos-
ta Rica was unexpected, but more thorough
collecting in habitats suited to Cx. lactator
would probably find the species in those
countries. The species is apparently absent
from the Caribbean. Its distribution in South
America is uncertain because the corniger
group may include undescribed species
which resemble Cx. /actator. For example,
two male genitalia from Ecuador resemble
the genitalia of Cx. lactator, but because
they are not associated with adults, identi-
fication is uncertain.
Collections of larval Cx. /actator have
been made from sea level to 1500 m. The
species has usually been found in sunlit water
in contact with soil, such as ground pools,
stream margins, and lake margins. Most of
the sites had been fouled either by natural
vegetation or domestic activity. Only one
collection was taken from a container, a liter
can in a cemetery, but the can rested on the
ground and contained rotting vegetation.
Although the focal distribution of Cx. /ac-
tator is distinct from that of Cx. corniger,
these species occasionally share the same
larval habitat (Honduras: HONC 316,
WRBU Acc. 1221; Mexico: MEX 237).
Material examined.—Total 150 2 145 ¢
39 8G 163P 110L in 30 collections. Mexico:
Oaxaca: Rincon Antonio, lectotype, 23 Jun
1905, F. Knab coll. no. 2701, “Ditch con-
taining a small quantity of very thick and
foul water among bones and other rubbish,
and swarming with mosquito larvae and pu-
pae.”, 1 6 1 6G 1P IL, also in series 5 2 4
61 8G. Veracruz: Cordoba, 10 Jun 1905, F.
Knab coll. no. 258, 22 1 6 1 6G; 600 m, 26
Jul 1965, D. Schroeder and R. X. Schick
coll. no. MEX 235, margin artificial lake, 3
2142 68G 4P 9L; 1100 m, 28 Jul 1965, D.
Schroeder coll. no. MEX 237, stream mar-
gin, | 2 1P 1L; Orizaba, 1300 m, 11 Aug
1965, D. Schroeder and R. X. Schick coll.
568 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
\\M
WALL
ri i
1LOmm
] |! it
il
Fig. 32. Larva of Cx. /actator Honduran specimens HONC 241-19 and 241-20, WRBU Acc. 1179. Scale
does not apply to enlargements of dorsomentum, spiracular apodeme, pecten spine, or comb scale; each magnified
inset of spiculation is 0.07 mm across. (A = antenna; C = head; CS = comb scale; Dm = dorsomentum; M =
mesothorax; P = prothorax; PMPc = posterior median process; PS = pecten spine; S = siphon; SAd = spiracular
apodeme; T = metathorax.)
VOLUME 91, NUMBER 4 569
33
Figs. 33, 34. Male terminalia of Cx. lactator lectotype. 33, Eighth abdominal tergum. 34, Genitalia as they
appear on slide mount.
no. MEX 291, concrete pit, | ¢ 1 6G 1P. Wilkerson coll. no. MX 19, post hole, 8 2
Oaxaca: Matias Romero, 200 m, 1 Sep 1965, 244 8G 14P 6L; Tapachula, 100 m, 7 Sep
D. Schroeder coll. no. MEX 335,2 29463 1987, Strickman, Roberts, and Wilkerson
6G 12P 16L. Chiapas: El Gancho, sea level, coll. no. MX 176, can on ground, 1 21 4 1
15 Aug 1987, Strickman, Roberts, and 4G. Belize: Cayo: Central Farm, 70 m, 9-10
570
35
i | tl
Lf} init
S Nn |
0 05m
f ‘s
| / Cf,
0.1mm |
“ oy 40
Figs. 35-42.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
0.2mm
Details of pupal and larval exuviae of Cx. lactator lectotype. 35, Pupal trumpet. 36, Larval
antenna. 37, Terminal larval segments. 38, Dorsomentum. 39, Comb scale. 40, Setal support plates for setae 9-
12-M and 9-12-T. 41, Pecten spine. 42, Spiracular apodeme with base of posterior median process.
May 1967, D.S. Bertram coll. no. BH A128,
light trap, 1 ¢ 1 6G. Guatemala: Izabal: Ca-
yuga, | 6G. Honduras: Copan: nr. San Jose
de Copan, 12 Mar 1987, D. B. Francy,
WRBU Acc. 1254, light trap, 2 2 6 4G.
Cortés: Puerto Cortés, <5 m, 10 Aug 1967,
A. J. Adames coll. no. HON 63, ground
pool, 1 2 4P 1L. Atlantida: La Ceiba, 10 m,
18 Jul 1986, D. Strickman coll. no. HONC
92, WRBU Acc. 1179, roadside ditch, 1 2
IP 1L. Yoro: Camp Big Bear, 800 m, 29
Apr 1986, R. Johnson coll. no. HONC 316,
WRBU Acc. 1221, shower runoff, 25 ¢ 20
6 1 4G 44P 15L. Comayagua: Comayagua,
590 m, 20 Mar 1984, A. R. Gillogly coll.
no. HONK 16, WRBU Acc. 1076, stream
margin pool, 22 1 6 1 6G 2P 2L; Palmerola
Air Base, 630 m, 23 Nov 1983, J. J. Arnott
coll. no. HO 11-1-23, WRBU Acc. 1067,
road rut, 23 2 13 6 1P IL; 6 Jul 1985, N.
Powers coll. no. 002, WRBU Acc. 1171,
stream margin, 1P 1L; 6 Jul 1985, coll. no.
HONC 003, ditch, 2 2 1 6 5P 5L; 17 Jul
1985, coll. no. 006, ditch, 20 2 22 6 3 6G
42P 43L; 31 Jul 1986, D. Strickman coll.
no. HONC 117, WRBU Acc. 1179, shower
runoff, 1 2 2 6 1 6G; 31 Jul 1986, coll. no.
HONC 118, ditch, 2 ¢ 1 6 1 6G; 17 Sep
1986, coll. no. HONC 241, kitchen waste
water, 6 266 1 4G 12P 6L; Siguatepeque,
1100 m, 21 Jul 1986, D. Strickman coll. no.
HONC 94, WRBU Acc. 1179,221¢6164G
3P 3L. La Paz: Cerro Sosomico, 1200 m,
27 Jul 1986, D. Strickman coll. no. HONC
VOLUME 91, NUMBER 4
107, WRBU Acc. 1179, ground pool, | 9°.
Colon: Puerto Castilla, <5 m, 10 Aug 1964,
A. Quinonez coll. no. HO 21, ditch, 4 9 6
62 6G 8P. Gracias a Dios: Puerto Lempira,
<5m, 17 Jun 1986, D. Strickman coll. no.
HONC 27, WRBU Acc. 1179, ground pool,
1 2 1P. Panama: Canal Zone: Summit, 16
Aug 1923, Dyar and Shannon, 5 2 12 4 1
8G; 4 May 1935, coll. no. PAX 45, 6 2 15
é 1 8G. Colombia: Cundinamarca: Fusa-
gasuga, 1500 m, 3 Mar 1965, A. M. Alarcon
and E. Osorno-Mesa coll. no. COB 16,
ground pool, 3 2 3 6 1 6G. Venezuela: Ara-
gua: Turmero, 500 m, 5 Sep 1966, E. R.
Vasquez coll. no. VZ 32, ground pool, 21 2
21846G 8P IL.
DISCUSSION
Culex lactator and Cx. corniger are sim-
ilar species which are most easily separated
as adults. Culex corniger is generally darker
with greater contrast between light and dark
areas. The dorsum of the abdomen of fe-
male Cx. corniger has light bands at the base
of each segment that became progressively
less extensive posteriorly until, at segments
V and VI, the bands do not reach the lateral
spots. On some specimens only a few light
scales are present anywhere on the dorsum
of the abdomen. In contrast, females of Cx.
lactator have broader basal abdominal bands
which reach the lateral spots on all seg-
ments. There is a marked difference be-
tween the two species in the banding on the
hindtarsus. Culex corniger has distinct light
bands on the proximal tarsomeres and less
distinct bands on tarsomeres 4 and 5;
whereas, Cx. /actator has less distinct light
bands on the proximal tarsomeres and bands
either absent or reduced to a few scales on
tarsomeres 4 and 5.
In the male genitalia, Cx. corniger differs
from Cx. lactator in the consistent presence
ofa moderately stout third rod (seta c) prox-
imal to the leaf (seta g), a gonostylus that
narrows less markedly in its apical two
thirds, and a thinner, less deeply cleft con-
nection between the lobes of the ninth ter-
gum. The lateral plates of the two species
571
are very similar. The rare presence of seta
cin Cx. lactator (in 2 of 39 examined) places
some doubt on the diagnostic usefulness of
this character; however, in both cases seta
c was markedly weaker than in Cx. corniger.
The larvae of Cx. corniger and Cx. lac-
tator share the same distinctive form of head
and siphon. Seven characters enabled sep-
aration of some, but not all, of the speci-
mens examined (Table 5). Although the
saddle index could be used to identify many
of the specimens, separation of species was
improved by using combinations characters
(Table 6). Each character was multiplied by
a coefficient that weighted it according to
its importance in separating the species. The
sum of the products yielded the canonical
variable (CNV), the value of which deter-
mined the species identification. The for-
mula below was arranged so that the char-
acters appear in their order of importance
(i.e. the first character is the one which would
provide the best separation by itself, the sec-
ond character would separate the next larg-
est number of specimens, etc.). Use of fewer
characters simply requires elimination of
products beginning from the right side of
the formula.
CNV = 5.45(A) + 0.74(B) — 2.07(C)
+ 0.36(D) — 4.42(E)
— 0.15(F) — 3.50(G)
where:
A = saddle index (dorsal length of siphon/
dorsal length of saddle)
= branches of seta 4-C
= siphon index (dorsal length of siphon/
width of siphon at base)
= branches of seta 7-C
= base of siphon to insertion of seta la-
S/dorsal length of siphon
= number of pecten teeth
pecten row length index (dorsal base
of siphon to apical pecten tooth/dor-
sal length of siphon).
mo Aw
|
Q
I
Determination of specimens not included
in the original analysis supported the use of
the canonical variable. Both the lectotype
572 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 6. Canonical variable (CNV) values separating Culex /actator and Cx. corniger based on the use of
one, two, three, or seven characters. Use of 4-6 characters was no more accurate than use of three characters.
Specimens examined were from Central America.
No. of Parameters
Species Statistic 7 38 2 le
Cx. lactator CNV cutoff > 6.25 >9.25 >12.75 >11
Sensitivity’ 96% 93% 91% 84%
Mean CNV 8.11 10.28 13.79 11.61
CNV range 5.25-10.25 7.84-12.18 10.81-15.85 8.61-14.12
Cx. corniger CNV cutoff <6.25 <9.25 = 12:75 <11
Sensitivity 97% 93% 90% 88%
Mean CNV 4.98 8.04 11.78 10.19
CNV range 3.25-7.25 6.70-9.47 10.74-12.87 8.77-11.39
Both Accuracy® 96% 93% 90% 85%
“CNV = 5.45(saddle index) + 0.74(branches of seta 4-C) — 2.07(siphon index) + 0.36(branches of seta 7-C)
— 4.42(base of siphon to insertion of seta la-S/dorsal length of siphon) — 0.15(number of pecten
teeth) — 3.50(pecten row length index).
° CNV = 5.45(saddle index) + 0.74(branches of seta 4-C) — 2.07(siphon index).
© CNV = 5.45(saddle index) + 0.74(branches of seta 4-C).
4 CNV = 5.45(saddle index).
© Optimum value of CNV to accurately distinguish the most specimens of each species.
‘Sensitivity = percentage correctly identified out of total examined of that species.
* Accuracy = percentage of both species correctly identified out of total examined.
(CNV = 7.28) and specimen VZ 32-62 from
Venezuela (CN V=7.33) were well within the
range of CNV values (Table 6) for Cx. lac-
tator. Collection MEX 237 from Mexico in-
cluded adults of both Cx. corniger and Cx.
lactator. Associated larval exuviae yielded
canonical variable values that corresponded
to identifications of the adults: MEX 237-
41, CNV = 5.82; MEX 237-93, CNV =
5.29; MEX 237-95, CNV = 7.20.
Ata practical level, use of all seven char-
acters would rarely be necessary. The ninety
percent accuracy possible using a CNV cal-
culated from a combination of the saddle
index and seta 4-C (two parameters in Table
6) would probably be adequate for most
purposes, especially since usually only one
of the species would be collected from a
single site.
A similar discriminant analysis was per-
formed for pupae of the two species (152
specimens of Cx. /actator, 35 of Cx. cor-
niger). Although statistically significant (P
< 0.01) separation was achieved, accuracy
was only 83%, even using seven characters.
We do not consider this level of accuracy
sufficient for useful identification. The
stouter spiculation on the posterior of ster-
num II in Cx. /actator versus the finer spic-
ulation in Cx. corniger separates most
specimens, but it is difficult to evaluate
specimens without comparative material.
Of the other species in Central America
that have adults with banded hindtarsi, only
Cx. coronator Dyar and Knab and Cx. res-
tuans Theobald might be confused with Cx.
lactator or Cx. corniger. Culex coronator
may be identified by a weakly developed
mesonotal pattern and strongly developed
banding on the hindtarsus, particularly on
tarsomere 5. Culex restuans sometimes
closely resembles adults of Cx. /actator, but
the mesonotal pattern is weaker anteriorly
and the hindtarsal banding is wider and ex-
tends to the base of tarsomere 5 (Strickman
and Darsie 1988).
CONCLUSION
The wide distribution and morphological
variability of Cx. corniger led to the naming
VOLUME 91, NUMBER 4
of five species and one variety which were
later synonymized. We have found that one
synonym, Cx. lactator, represents a species
that is distinguishable in all stages but the
pupa from sympatric populations of Cx.
corniger. For this reason, we have elevated
Cx. lactator to species status.
The previous descriptive literature on Cx.
lactator was inadequate to identify the
species. Dyar and Knab’s (1906a) original
description of this species was part of a re-
vision of Culex larvae which did not include
Cx. corniger. Of the seven collections cited
by Dyar and Knab (1906a), two (from Cor-
doba and Rincon Antonio) were Cx. lac-
tator and five (from Santa Lucrecia, Te-
huantepec, Almoloya, Puntarenas, and San
Jose) were Cx. corniger. The various adult
forms were later organized into varieties,
culminating in the short key by Howard et
al. (1915). Dyar and Knab evidently never
associated Cx. /actator with its male geni-
talia, because they illustrated descriptions
of Cx. corniger with genitalia of Cx. lactator
in two publications (Howard et al. 1912,
Dyar 1928). Fortunately, certain identifi-
cation is possible since the male prepared
by Knab and selected by Stone and Knight
(1957) as the lectotype of Cx. /actator has
associated larval and pupal exuviae as well
as genitalia.
Subsequent treatments of Cx. corniger in
Central America gave little indication of a
separate form corresponding to Cx. /acta-
tor. Lane (1953) mentioned that seta c is
sometimes absent on the subapical lobe of
the male genitalia, but Forattini (1965) and
Bram (1967) did not describe this variation.
Clark-Gil and Darsie (1983) did not ex-
amine male genitalia and their key char-
acters for females and larvae fit both Cx.
corniger and Cx. lactator. Only Martinez
Palacios (1950) discussed the two forms,
stating that the form lacking seta c (1.e. Cx.
lactator) was the more common and that it
was widespread throughout the Neotropical
part of Mexico.
In spite of the possibility that the differ-
573
ences between Culex corniger and Cx. lac-
tator are environmentally induced (making
the two forms non-genetic ecophenotypes),
we believe that they are closely related, dis-
tinct species. The two collections of both
species from the same larval habitat support
this view. Separate species status may even-
tually require confirmation through cross-
ing, rearing in artificially adjusted habitats,
or chemical genetic studies. For the time
being, recognition of Cx. /actator as a sep-
arate species will facilitate accumulation of
data on this form.
ACKNOWLEDGMENTS
E. L. Peyton for helpful comments on tax-
onomy; Taina Litwak for preparation of the
figures; J. Mohatt and Medical Element,
JTF-Bravo, Palmerola Air Base, for orga-
nizational support in Honduras; J. Pecor for
technical assistance in the United States; R.
Koinzan for technical assistance in Hon-
duras; National Aeronautics and Space
Administration (NASA) contract W-16,306
to the Uniformed Services University of the
Health Sciences entitled Application of Re-
mote Sensing and Predictive Modeling to
Malaria Transmission and Vector Ecology
for funding of collections in Chiapas, Mex-
ico. Opinions and assertions contained
herein are the private views of the authors
and are not to be construed as official, nor
as reflecting the views of the supporting
agencies.
LITERATURE CITED
Belkin, J. N. 1968. The type specimens of New World
mosquitoes in European museums (Mosquito
Studies. IX). Contrib. Amer. Entomol. Inst. 3: 1-
69.
Belkin, J. N.,S. J. Heinemann, and W. A. Page. 1970.
The Culicidae of Jamaica (Mosquito Studies. X XI).
Contrib. Amer. Entomol. Inst. 6: 1-458.
Bram, R. A. 1967. Classification of Culex subgenus
Culex in the New World (Diptera: Culicidae). Proc.
U.S. Natl. Mus. 120. 122 pp.
Clark-Gil, S. and R. F. Darsie, Jr. 1983. The mos-
quitoes of Guatemala. Their identification, distri-
bution and bionomics. Mosq. Syst. 15: 151-284.
574
Dyar, H. G. 1922. The mosquitoes of the United
States. Proc. U.S. Natl. Mus. 62: 1-119.
. 1928. The mosquitoes of the Americas. Car-
negie Inst. Washington Publ. No. 387. 616 pp. +
123 pls.
Dyar, H. G. and F. Knab. 1906a. The larvae of Cu-
licidae classified as independent organisms. J. N.Y.
Entomol. Soc. 14: 170-230.
. 1906b. Notes on some American mosquitoes
with descriptions of new species. Proc. Biol. Soc.
of Washington 19: 159-172.
1909. Descriptions of some new species and
a new genus of American mosquitoes. Smithson-
ian Misc. Coll. 52: 253-266.
Forattini,O. P. 1965. Entomologica Medica. 20. Vol-
ume. Editora da Universidade de Sao Paulo. 506
pp.
Galindo, P., S. J. Carpenter, and H. Trapido. 1951.
Ecological observations on forest mosquitoes of
an endemic yellow fever area in Panama. Amer.
J. of Trop. Med. 31: 98-137.
Grabham, M. 1906. Notes on some mosquitoes from
Newcastle, Jamaica. Can. Entomol. 38: 167-173.
Griner, P. F., R. J. Mayewski, A. I. Mushlin, and P.
Greenland. 1981. Selection and interpretation of
diagnostic tests and procedures. Principles and ap-
plications. Ann. Intern. Med. 94: 557-600.
Harbach, R. E.and K. L. Knight. 1980. Taxonomists’
Glossary of Mosquito Anatomy. Plexus Publish-
ing, Inc., Mariton, N.J. x1 + 413 pp.
Harbach, R. E., E. L. Peyton, and W. L. Jakob. 1983.
Synonymy of Culex (Culex) oswaldoi with Culex
(Culex) maxi (Diptera, Culicidae). Mosq. Syst. 15:
310-317.
Howard, L. O., H. G. Dyar, and F. Knab. 1912. The
Mosquitoes of North and Central America and the
West Indies. Volume Two. Carnegie Inst. of
Washington. x + 150 pls.
1915. Mosquitoes of North and Central
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America and the West Indies. Volume Three. Car-
negie Inst. of Washington. 523 pp.
Knight, K. L. and A. Stone. 1977. A Catalog of the
Mosquitoes of the World (Diptera: Culicidae).
Second Edition. Thomas Say Foundation, Ento-
mol. Soc. Amer. 6. x1 + 611 pp.
Kueppers, H. 1982. Color Atlas. Barron’s, Wood-
bury, N.Y. 170 pp.
Lane, J. 1953. Neotropical Culicidae. Volume I. Uni-
versity of Sao Paulo, Brazil. 548 pp.
Martinez Palacios, A. 1950. Identificacion de los
mosquitos mexicanos del subgenero Culex (Dip-
tera: Culicidae) por la genitalia masculina. Rev.
Soc. Mex. Hist. Nat. 11: 183-189.
Senevet, G. and E. Abonnenc. 1939. Les moustiques
de la Guyane Francaise - IT. Le genre Culex. Arch.
Inst. Pasteur Alger. 12: 62-134.
. 1958. A propos de quelques Culex sud-amér-
icains. Arch. Inst. Pasteur Algér. 36: 343-350.
Stone, A. and K. L. Knight. 1957. Type specimens
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Wash. Acad. Sci. 47: 42-59.
Strickman, D. 1988. Redescription of the holotype
of Culex (Culex) peus Speiser and taxonomy of
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viously undetected presence of Culex restuans
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Theobald, F. V. 1903. A Monograph of the Culicidae
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PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 575-582
REVIEW OF MARICOPODYNERUS (HYMENOPTERA: EUMENIDAE)
RICHARD M. BOHART
Department of Entomology, University of California, Davis, California 95616.
Abstract.—The 16 known species of the genus include 7 described as new here. A key
to the species is given along with illustrations of important characters. Additional locality
records are included for previously described forms. New species are: arizonicus, Arizona,
New Mexico; differens, Arizona, Nevada, California; festivus, Texas, Arizona; /insleyi,
southern California; /issoides, Texas, New Mexico, Nueva Leon; optimus, New Mexico,
Arizona; sternalis, California.
Key Words:
H. L. Viereck described Maricopodynerus
(1908) as a subgenus of Odynerus for the
single new species, maricoporum. Bohart
(1948) elevated it to generic rank and added
2 new species. Then, in a revisionary effort,
Bohart (1950) added 4 more species. At that
time less than 150 specimens of the genus
were known. Now I have seen more than
400 and among them are 7 new species de-
scribed herein. This brings the total to 16,
all west of the 100th meridian in United
States and Mexico.
The generic characters were given by Bo-
hart (1950). Maricopodynerus is unique in
its greatly enlarged, posteriorly membra-
nous second tergum which is twice as long
as the second sternum, and can enclose the
remaining terga (Figs. 22-24). Specific char-
acters of most importance are clypeal shape,
punctation, and color pattern. Male geni-
talia are rather similar except in sericifrons
and arizonicus (Figs. 17, 18).
More than one species may occur in a
single locality; for example, sericifrons and
linsleyi at Palm Springs, California; rudiceps
and decorabilis at Boca, California; mari-
coporum and differens at Oak Creek Can-
yon, Arizona; and Jissoides and festivus in
wasps, Eumenidae, Maricopodynerus
Santa Elena Canyon of Big Bend Park, Tex-
as.
Type depositories indicated with the de-
scriptions are: Academy of Natural Sciences
(San Francisco), British Museum of Natural
History (London), Los Angeles County Mu-
seum (Los Angeles), University of Arizona
(Tucson), University of Idaho (Moscow),
Harvard University (Cambridge), Univer-
sity of California Bohart Museum (Davis),
University of Kansas Snow Museum (Law-
rence), U.S. National Museum (Washing-
ton), Oregon State University (Corvallis).
Abbreviations used are F-I etc., flagello-
mere; MOD, median ocellus diameter; PD,
puncture diameter; T-I etc., terga; S-I etc.,
sterna.
KEY TO THE SPECIES OF
MARICOPODYNERUS
1. T-II posterior ivory band deeply u-shaped
medially (Figs. 10, 12)
T-II posterior ivory band not deeply u-shaped
medially (as in Fig. 11) 3
Vertex swollen and partly polished behind
ocelli, forewing marginal cell darkened to-
ward apex (as in Fig. 20) arizonicus Bohart
Vertex not swollen, punctation close and
to
to
576
~
10.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
coarse; forewing marginal cell not darkened
toward apex (Figs 21). 2 .:..:.. optimus Bohart
. Forewing marginal cell not at all darkened
fowardiapexin eee GO petite teeter 4
Forewing marginal cell somewhat darkened
in apical third or fourth, especially in females
RR en oe Rice CoC Sore 7
. Tegula black and ivory or very dark red and
ivory; propodeum black posteriorly or mostly
so; male clypeus with black apical rim, female
clypeustallblack ..'2. c.caprasces rudiceps Bohart
Tegula light or bright red and ivory; propo-
deum red posteriorly; male clypeus without
black apical rim; female clypeus with consid-
erable red
. Forewing marginal cell unusually short and
stout (Fig. 19), male F-XI stout, less than twice
as long as thick in lateral profile, female clyp-
eus all red and black _ sericifrons Bohart
Forewing marginal cell more slender (shaped
about as in Fig. 20 or 21), male F-XI slender,
more than twice as long as thick in lateral
profile, female clypeus nearly always with some
ivory marks
. T-II with considerable red along i inner margin
of ivory, female pronotum mostly red dor-
sally, female postocular spot encircled with
red mats . chisosensis Bohart
T-II with little or no red along inner ivory
margin, female pronotum not mostly red dor-
sally, female postocular spot not encirlced with
red maricoporum (Viereck)
. T-II lateral spot isolated from marginal ivory
band (as in Fig. 22), male clypeus with free
apical margin black rimmed (as in Fig. 5),
female clypeus black with lateral ivory spot
(as in Fig. 1) or all black .... 8
T-II lateral spot attached to marginal ivory
band (as in Fig. 23), male clypeus with free
apical margin pale, female clypeus usually
partly red but never all black 10
T-I black except for posterior ivory band, T- i
punctation rather coarse (Fig. 14)
: . festivus Bohart
T-I mostly or all a except for ivory band,
T-II punctation unusually fine (as in Fig. 16)
.S- II mostly oral dark; femaléclypeusall black;
T-II punctures small, somewhat effaced by
microsculpture decorabilis Bohart
S-II mostly or all bright red; female clypeus
black with 2 white spots (Fig. 1); T-II punc-
tures small, but their outlines distinct (Fig.
16) eevee ten aes mete arate: sternalis Bohart
T-II punctation along midline in front of pos-
terior depression absent or practically so 11
T-II punctation distinct along midline 12
11. T-IL practically impunctate in midposterior
depression, female mandible not unusually
thick or bluntly curved toward apex .......
pulvipilus Bohart
T-II distinctly punctate in midposterior
depression, female mandible thick and bluntly
curved toward apex .... permandibularis Bohart
12. T-II punctation coarse, rather evenly distrib-
uted, most punctures | PD or less apart ...
differens Bohart
T-II punctation fine and/or unevenly distrib-
uted with most anteromedial punctures 2 or
S3PDiapartiias eet es Seo ee 13
13. T-II with anterior half mostly polished in me-
dian area, punctures sparse (Fig. 13)
T: II with numerous punctures in aeeheae area
(Fig. 15)
14. T-II punctures in lateral emargination of ivo-
ry band exceptionally coarse, some almost as
large as midocellus ............ lissoides Bohart
T-II punctures somewhat enlarged in emar-
gination of ivory band but much smaller than
midocellls: Vax. sictuesineee eae eelemcrneten 15
15. Clypeus with apical emargination moderately
shallow between sharp teeth (Figs. 2, 6); clyp-
eus of male lightly microsculptured but a little
shiny, that of female normally convex
POR COT oct cin orkicthc linsleyi Bohart
Clypeus imusually short, apical emargination
quite shallow (Figs. 3, 7); clypeus of male
densely microsculptured and dull, that of fe-
male unusually flat shannoni Bohart
Maricopodynerus arizonicus Bohart,
New SPECIES
Male holotype.—Length 7 mm. Black,
ivory, and reddish brown. Ivory are: clyp-
eus, dot above, ocular and postocular spots,
pronotum anterodistally, 2 spots on tegula,
2 spots on scutellum, metanotal band, large
upper mesopleural spot, distal spots on fore
and midfemora, tibiae outwardly, irregular
recurved band on T-I, markings on T-II (Fig.
24); posterolateral spot on S-II, reddish
brown are: flagellum inwardly, mandible,
pronotal edge posteriorly, legs mostly, cen-
tral spot on tegula, terminal tergal and ster-
nal segments; wings slightly dusky, forewing
marginal cell darkened in front and on distal
third. Pubescence pollinose, silvery on head
and thorax, fulvous on distal third. Pubes-
VOLUME 91, NUMBER 4 Se,
y
i 2. linsleyi . ee
|. sternalis y 3. shannoni 4. arizonicus
& :
; oo Ca
5. sternalis 7. shannoni
poe 6. linsleyi 8. arizonicus
9. sternalis
Loma)
11. linsleyi
10. optimus
a
[ A)
2 : 12. arizonicus
14. festivus
Oy eeu
S
——
17. sericifrons 18. arizonicus
16. sternalis
19. sericifrons 20. linsleyi 21. optimus
22. sternalis 23. linsleyi 24. arizonicus
Figs. 1-24. 1-4, clypeal pattern, females, x 30; 5-8, clypeal shape, males, = 30; 9-12, T-II pattern, posterior
view, males, 18; 13-16, punctation across T-II opposite lateral spot, females, x 18; 17, 18, flattened aedeagus
and cuspis, x60; 19-21, forewing marginal cell, females, 19, 20, x40, 21, 45; 22-24, T-I-II pattern, lateral
males, x 20. Suppled areas of Figs. 2, 3, 22, 23, red.
578
cence pollinose, silvery on head and thorax,
fulvous on T-II. Punctures moderate and
sparse on clypeus, moderately coarse and
close on frons, irregularly spaced across ver-
tex, moderate and somewhat spaced on no-
tum, pleuron and T-I, punctures of T-II
moderately coarse and rather evenly spaced
about a PD apart. Clypeal apex obtusely
emarginate between sharp teeth (Fig. 8);
F-XI small, not reaching base of IX; vertex
swollen, forming a low smooth transverse
swelling behind ocelli and a barely percep-
tible swelling near compound eye; cuspis
with a row of teeth along inner edge (Fig. 18).
Female.— About as in male except: length
8 mm, clypeus more coarsely punctate and
with a dark basal triangle (Fig. 4), reddish
brown of legs as far as tibiae replaced by
red, propodeum and T-I mostly red.
Holotype male, 5 mi w. Portal, Chirica-
hua Mts., Cochise Co., Arizona, VIII-7-58
(R. M. Bohart, DAVIS). Paratypes, 10
males, 21 females, topotypical, VIII-6 to
VIII-22 (R. M. Bohart, SAN FRANCISCO,
WASHINGTON, DAVIS). Other speci-
mens, not paratypes, VII-[X, 6 males, | fe-
male, Arizona: 5 mi w. Portal, Patagonia
and 6 mi se.; New Mexico: | 1 mi wsw. Car-
rizozo, Rockhound State Park (Luna Co.),
18 mi w. Guthrie.
Discussion.—The deeply divided black
markings posteriorly on T-II (Fig. 12), to-
gether with the partly polished vertex and
darkened apex of the forewing marginal cell,
characterize the species.
MARICOPODYNERUS CHISOSENSIS BOHART
Maricopodynerus chisosensis Bohart 1950:
20. Holotype male, Chisos Mts., Brewster
Co., Texas (WASHINGTON). The red col-
oration is usually extensive, bordering the
ivory postocular spot and ivory band of T-II.
Also, the female clypeus and pronotum are
mostly red. The species 1s known only from
Texas in Brewster Co. (type series), and 2
females in Terre Co. and Sanderson Co. (W.
F. Barr, MOSCOW).
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Maricopodynerus decorabilis Bohart
Maricopodynerus decorabilis Bohart 1950:
22. Holotype male, Leavitt Meadows, Mono
Co., California (SAN FRANCISCO).
Obviously similar to sternalis by its small
size, less rotund T-II, and free spot on T-II,
this species has the T-II punctures more
shallow and partly effaced by microsculp-
ture. Also, S-II is black instead of red. It is
known from southeast Oregon and eastern
California as far south as Inyo Co. Many of
the 24 specimens I have seen were taken at
or near Boca, Nevada Co., California on
Chrysothamnus in July.
Maricopodynerus differens Bohart,
New SPECIES
Male holotype.— Length 7.5 mm. Black,
ivory, and red. Ivory markings as described
for optimus except 2 tegular spots, a distal
one on midfemur, and less extensive T-II
spots posteriorly; red markings as in opti-
mus except propodeum laterally and pos-
teriorly; T-I basally; forewing marginal cell
darkened in apical half. Pubescence silvery,
inconspicuous. Punctation as described for
optimus, that of T-II as in Fig. 14. Clypeal
apex obtusely emarginate between angular
teeth, F-XI reaching base of F-IX, lateral
ivory margin of T-II moderately swollen,
cuspis without teeth.
Female.— About as in male: length 7.5-
8.5 mm; femora all red, propodeum all or
mostly red, T-I usually red except for pos-
terior ivory band; clypeus more coarsely
punctate, red and ivory or more often red,
black, and ivory.
Holotype male, Oak Creek Canyon, Co-
conino Co., Arizona, VI-11-40 (R. M. Bo-
hart, DAVIS). Paratypes (all DAVIS except
as noted), Arizona: III-VI, 5 males, 3 fe-
males, Maricopa Mts., 4 mis. Hoover Dam
(Mojave Co., MOSCOW), 4 mi n. White
River (TUCSON), Superior (TUCSON),
Sedona (TUCSON), Lake Mojave; Califor-
nia: 2 males, 9 females, IV-VIII, Surprise
Canyon and Big Pine Canyon (Inyo Co.),
VOLUME 91, NUMBER 4
near Bishop and Topaz Lake (Mono Co.,
DAVIS, SAN FRANCISCO); Nevada: 2
males, 4 females, IV-VIII, Nelson, 5 mi e.
Carson City, Mt. Montgomery, 2 mie. To-
nopah.
Discussion.—The coarse and rather
evenly distributed punctures of T-II, the
apically darkened forewing marginal cell,
and the absence ofa free spot on T-II, taken
together are differentiating.
Maricopodynerus festivus Bohart,
New SPECIES
Male holotype.—Length 5.5 mm. Black,
ivory and red. Ivory are: clypeus except black
apical rim, dot above, ocular and postocular
spots, irregular anterior pronotal margin,
scuttelar spots, metanotal band, anterior teg-
ular dot, mesopleural spot, forefemoral spot,
tibial streaks, posterior T-I band, T-II
markings (about as in Fig. 22), apicolateral
spot on S-II: red are: flagellum inwardly,
tegula mostly, legs mostly (as far as tibiae);
forewing marginal cell a little darkened dis-
tally. Pubescence silvery, inconspicuous.
Punctures moderate on shagreened clypeus,
moderate and close on frons and notum,
moderately coarse and close but evenly
spaced on T-I-II (as in Fig. 14). Clypeal apex
obtusely emarginate between sharp teeth.
Female.— About as in male except: length
6.5 mm, clypeus black with lateral ivory
spot, coarsely punctate, marginal wing cell
more darkened in distal third.
Holotype male, Santa Elena Canyon, Big
Bend National Park, Texas, Brewster Co.,
IV-9-65 (D. Bolinger, DAVIS). Paratypes,
3 females, [V-19-54 (R. H. Beamer, LAW-
RENCE, DAVIS); 1 female, Sabino Can-
yon, Santa Catalina Mts., Arizona, V-5-55
(F. G. Werner, TUCSON).
Discussion.— Although similar to differ-
ens in the marginal cell and punctation of
T-II, there are several points of difference.
In festivus the scape, propodeum, and basal
three-fourths of T-I are black. The clypeal
apex is more sharply toothed, and the fe-
579
male clypeus is black with 2 ivory spots.
Finally, the anterior ivory spot of T-II is
isolated from the lateral mark.
Maricopodynerus linsleyi Bohart,
New SPECIES
Male holotype.—Length 6.8 mm. Black
or dark reddish brown, ivory and red. Ivory
are: same areas given for arizonicus except
T-II spots attached laterally and posterior
spots smaller (Figs. 11, 23); red are: flagel-
lum inwardly, tegula medially, posterior
edge of pronotum, legs mostly, propodeum,
T-I except for ivory border; forewing nearly
clear except dusky distal third of marginal
cell. Pubescence as described for arizonicus.
Punctures of clypeus small and sparse, those
of frons, vertex, and notum moderately
coarse and close; those of mesopleuron well
separated, those of T-II moderate and about
a PD apart anteriorly but becoming close
beyond middle. Clypeal apex shallowly
emarginate between sharp teeth (Fig. 6),
F-XI moderately slender, nearly reaching
base of F-IX; T-II ivory margin not over-
hanging membranous edge laterally; cuspis
without teeth.
Female.— About as in male except: clyp-
eus more coarsely punctate and bimaculate
red and ivory (Fig. 2); anterior half of T-II
with punctures usually 1-3 PD apart.
Holotype male, Palm Springs, Riverside
Co., California VII-6-75, on Dalea spinosa
(E. G. and J. M. Linsley, DAVIS). Para-
types, 11 males, 30 females, topotypical, VI-
17 to VI-6 (E. G. and J. M. Linsley, SAN
FRANCISCO, DAVIS); 3 males, 7 females,
Corn Springs, Chuckwalla Mts., Riverside
Co., California (LOS ANGELES, DAVIS);
3 males, 17 females, Shaver’s Well, River-
side Co., California (LOS ANGELES, DA-
VIS). Other specimens, typical but not para-
types, California: Westmorland, Indio, Palm
Springs, Brawley, Winterhaven (DAVIS).
Discussion. — Features of the marginal cell
(Fig. 20), T-II markings (Figs. 11, 23), and
clypeus (Figs, 2, 6) are given in the key. T-II
580
is fairly well punctate but in the anterior
half there are considerable polished inter-
vals, but less so than in /issus (Fig. 13). Lat-
eral punctation of T-II separate /issoides and
linsleyi, which also has the clypeal apex more
sharply emarginate. This last feature also
distinguishes /ins/eyi and shannoni (Figs. 2,
3, 6, 7). The species is named for the emi-
nent Coleopterist and Hymenopterist, my
friend, E. Gorton Linsley.
Maricopodynerus lissoides Bohart,
New SPECIES
Male holotype.—Length 7 mm. Black,
ivory and red. Markings and pubescence as
described for /ins/eyi. Punctures of clypeus
small and sparse, those of frons, vertex and
notum moderate and slightly separated;
those of mesopleuron about 1.0-1.5 PD
apart; those of T-II coarse and 1.0-1.5 PD
apart on anterior half (Fig. 15), becoming
closer posteriorly; those in lateral emargi-
nation of ivory band quite coarse and close.
Clypeal apex weakly indented between blunt
teeth, F-XI small, not reaching base of T-IX,
ivory margin of T-II overhanging membra-
nous edge laterally, cuspis without teeth.
Female. — About as in male except: length
8 mm, clypeus bicolored red and ivory and
more coarsely punctate, apex nearly trun-
cate.
Holotype male, Santa Elena Canyon, Big
Bend National Park, Brewster Co., Texas,
VIII-25-67 (R. Gardner, C. Kovacic, DA-
VIS). Paratypes, | male, 17 females, topo-
typical, VIII-22-25-54 (R. M. Bohart, SAN
FRANCISCO, DAVIS, WASHINGTON);
5 males, Big Bend National Park, Texas,
April (R. M. Bohart, D. Bolinger, COR-
VALLIS, DAVIS). Other specimens, not
paratypes, 11 males, 2 females, Texas: 25
mis. Marathon, Stillwell Crossing and Study
Butte (Brewster Co.), Lajitas; New Mexico:
11 mi wsw. Carrizoza, Organ Mts., Pelon-
cillo Mts.; Nueva Leon: near Monterrey,
Mexico.
Discussion. — Differences from /ins/eyi are
given under that species, and in the key.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Maricopodynerus lissus Bohart
Maricopodynerus lissus Bohart 1950: 23.
Holotype male, Maricopa Mts., Arizona
(SAN FRANCISCO).
The much reduced punctation of T-II (Fig.
13) characterizes /issus, and the extensive
smooth areas are surpassed only by pulvi-
pilus. The 37 specimens I have seen came
from southwest New Mexico, Arizona, and
southeast California.
Maricopodynerus maricoporum (Viereck)
Odynerus (Maricopodynerus) maricopo-
rum Viereck 1908: 397. Holotype male, Oak
Creek Canyon, Coconino Co., Arizona
(LAWRENCE).
Odynerus chelonogastrus Cameron 1908:
202. Holotype female (no abdomen), Je-
rome, Yavapai Co., Arizona (LONDON).
New synonymy.
This generotype species is quite similar
to lissoides but the apically pigmented fore-
wing marginal cell of the latter is differen-
tiating. I have seen both types listed above
and about 80 other specimens from eastern
California (Inyo Co. to Imperial Co.), east-
ern and southern Nevada, Arizona, south-
ern New Mexico, southwestern Texas, and
southern Utah (Leeds).
Maricopodynerus optimus Bohart,
NEw SPECIES
Male holotype.—Length 7 mm. Black,
ivory, and red. Ivory are: clypeus, scape 1n
front, ocular and postocular spots, prono-
tum anterodorsally, tegular spot, 2 scutellar
spots, metanotal band, upper mesopleural
spot, distal spot on forefemur, tibiae out-
wardly, irregular recurved band on T-I,
markings on T-II (Fig. 10 and as in 23),
subapical band on S-II; red are: flagellum
weakly toward inner apex, mandible, pro-
notal edge posteriorly, tegula mostly, legs
mostly; forewing marginal cell not darkened
toward apex. Pubescence silvery, as long as
1 MOD on face and propodeum. Punctures
moderate and a PD apart on clypeus, coarse
VOLUME 91, NUMBER 4
and close on face and notum: punctures of
T-II coarse, rather evenly distributed, most-
ly less than | PD apart, unusually coarse in
lateral and posterior emarginations of ivory
band. Clypeal apex obtusely emarginate be-
tween sharp teeth, F-XI reaching apical two-
fifths of F-VIII, ivory margin of T-II over-
hanging membranous edge laterally.
Female.— About as in male except: length
8.5-9.0 mm, clypeus more coarsely punc-
tate, black with 2 ivory spots (as in Fig. 1);
pronotal band broken submedially, lateral
spot of T-II sometimes unattached.
Holotype male, Dripping Spring, Organ
Mts., Dona Ana Co., New Mexico (C.
Townsend, WASHINGTON). Paratypes,
female, New Mexico: topotypical (T. Cock-
erell, WASHINGTON); female, Texas: The
Basin, Big Bend National Park, Brewster
Co., X-4-56 (J. W. MacSwain, SAN FRAN-
CISCO); Arizona: 2 females, Lower Madera
Canyon, Santa Rita Mts., VI-I-61 (J. Be-
quaert, CAMBRIDGE, DAVIS).
Discussion.—The rather deeply divided
ivory markings posteriorly on T-II (Fig. 10)
are rivalled only by arizonicus which 1s eas-
ily separated by its somewhat bulging and
polished vertex, as well as the stained fore-
wing marginal cell.
Maricopodnerus permandibularis Bohart
Maricopodynerus permandibularis Bo-
hart 1948: 321. Holotype female, Tiburon
Island, Gulf of California, Mexico (SAN
FRANCISCO).
Known only from the type female, the
stout, blunt mandibles are diagnostic. The
punctation of T-II is similar to that of /issus.
MARICOPODYNERUS PULVIPILUS BOHART
Maricopodynerus pulvipilus Bohart 1948:
321. Holotype male, Tortuga Island, Gulf
of California, Mexico (SAN FRANCISCO).
In addition to the 6 specimens of the type
series from Baja California Sur, I have be-
fore me 7 males, 7 females collected by L.
D. French and E. O. Sugden 35 km s. Mu-
lege, Baja California Sur, IV-I-80 (DAVIS).
581
Also, there are 5 females (LOS ANGELES),
5 mi nw. San Ignacio, Baja California Sur,
IX; and 13 km nw. Rosarito, Baja Califor-
nia, IX. The practical absence of punctation
on T-II, including the posteromedial
depression, is characteristic.
Maricopodynerus rudiceps Bohart
Maricopodynerus rudiceps Bohart 1950:
21. Holotype male, Verdi, Nevada (SAN
FRANCISCO).
The moderate development of the T-II
ivory band posteriorly (about as in Fig. 11),
even coloration of the forewing marginal
cell, all-black female clypeus, and black-
rimmed male clypeus will differentiate rudi-
ceps from other species with close, coarse
T-II punctation. This species is rather widely
distributed in western United States from
southeastern Washington to central Mon-
tana, south to eastern California as far as
Mono Co., and in northern Arizona. I have
before me several hundred specimens col-
lected at Boca, Nevada Co., California, VII
(DAVIS).
Maricopodynerus sericifrons Bohart
Maricopodynerus sericifrons Bohart 1950:
19. Holotype male, 2 mi w. Edom, River-
side Co., California (SAN FRANCISCO).
This species can be distinguished by the
clarity and stout shape of the forewing mar-
ginal cell (Fig. 19), by the limited lateral
extent of the posterior membrane of T-II,
and by the spade-shaped aedeagus (Fig. 17).
I have seen about 80 specimens, mostly from
Riverside Co., California, but a few from
San Diego Co. (Borrego) and Inyo Co. (Lone
Pine). Out-of-state records are from Ari-
zona (Parker, Yuma, Wellton), and Sonora
(39 min. Puerto Penasco).
Maricopodynerus shannoni Bohart
Maricopodynerus shannoni Bohart 1950:
24. Holotype male, Stratford, Washington
(SAN FRANCISCO).
The short and nearly truncate clypeus
(Figs. 3, 7), which is unusually flat in the
582
female and densely microsculptured in the
male, are identifying characters.
Altogether, I have seen 18 specimens. Lo-
calities are Washington (Stratford, Lake
Paha), Idaho (6 mi nw. St. Anthony), Ne-
vada (3 mi w. Hazen, Yerington), and Cal-
ifornia (Siskiyou, Shasta, Mono, and Inyo
counties).
Maricopodynerus sternalis Bohart,
New SPECIES
Male holotype.—Length 5.5 mm. Black,
ivory, and red. Ivory are: clypeus except rim
(Fig. 5), scape in front, ocular and postoc-
ular spots, pronotum anterodorsally, scu-
tellar spots, metanotal band, tegular dots,
upper mesopleural spot, small tibial spots,
irregular recurved band on T-I, markings
on T-II (Figs. 9, 22); lateral dot on S-II; red
are: mandible apically, flagellum inside, teg-
ula mostly, legs as far as tarsi mostly, T-I
except posterior band, T-II basolateral spot,
S-II mostly; forewing marginal cell dark-
ened distally. Pubescence silvery, incon-
spicuous. Punctures of clypeus moderate, a
little spaced by dense microsculpture; punc-
tures of frons, vertex and notum moderate
and close; those of T-II small, distinct,
evenly distributed, spaced by about | PD
of microsculpture (Fig. 16). Clypeal apex
emarginate between sharp teeth (Fig. 5);
F-XI small, not reaching base of F-IX; lat-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
eral ivory margin of T-II hardly swollen;
cuspis without teeth.
Female.— About as in male except: length
6.5 mm, clypeus black with 2 ivory spots
(Fig. 1), pronotal band broken submedially,
tegula all red.
Holotype male, Townes Pass, Inyo Co.,
California, V-6-60 (A. S. Menke, DAVIS).
Paratypes, 2 females, California: 12 mi s.
Baker, IV-15-69 (E. E. Grissell, DAVIS);
Argus Mts. (Inyo Co.), V-22-37 (N. W. Fra-
zier, DAVIS).
Discussion.—The similar shape and
markings of T-II indicate a relationship to
decorabilis. However, in sternalis the mac-
ulate rather than all black clypeus (Fig. 1),
mostly red rather than black S-II, and finely
but distinctly punctate T-II, indicate a dif-
ferent species.
LITERATURE CITED
Bohart, R. M. 1948. Contributions toward a knowl-
edge of the insect fauna of Lower California, No.
9, Hymenoptera: Eumeninae. Proc. Calif. Acad.
Sci. (4) 24: 313-336.
. 1950. A synopsis of Maricopodynerus. Bull.
Brooklyn Ent. Soc. 40: 17-25.
Cameron, P. 1908. A contribution toward the knowl-
edge of the Odyneridae of the southwest of the
United States. Trans. Amer. Ent. Soc. 34: 195-
246.
Viereck, H. L. 1908. Notes and descriptions of Hy-
menoptera from the western United States. II.
Trans. Amer. Ent. Soc. 33: 381-409.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 583-587
DRUMMING BEHAVIORS OF THREE PENNSYLVANIA
STONEFLY (PLECOPTERA) SPECIES
DAvip D. ZEIGLER
Department of Biology, Southwest Texas State University, San Marcos, Texas 78666.
New address: Biology Department, Pembroke State University, Pembroke, North Carolina
28372.
Abstract. Drumming behaviors of three central Pennsylvania stonefly species were
described. Males of Tallaperla maria initially produced calls of a single scraping beat
which shifted to a two-pulse scrape/beat call as signal exchanges with females continued.
Agnetina capitata males produced a diphasic call that is the longest known in any stonefly
species. The signals of 7. maria and Prteronarcys biloba were compared to previous signal
descriptions from geographically separate populations, and some intraspecific divergence
of signal structure was found in these species.
Key Words:
perla, Pteronarcys, Agnetina
Drumming behavior in stoneflies is a sys-
tem of intersexual vibrational communi-
cation which aids in mate identification and
location. The vibrational pulses that make
up signals are typically produced when the
insects strike or scrape the substratum with
the posteroventral portion of the abdomen
(Rupprecht 1967, Maketon and Stewart
1984). Drumming behavior was first quan-
tified by Rupprecht in 1967. To date, nu-
merous workers have described the signals
of over 100 species (see Zeigler and Stewart
1987, Maketon and Stewart 1988 for liter-
ature citations). Continued descriptive work
of this nature contributes to systematics
(Zeigler and Stewart 1987) and to data banks
which may help answer other assorted evo-
lutionary questions, such as those raised by
the current hypotheses concerning sexual
selection’s role in the divergence of repro-
ductive behaviors (West-Eberhard 1984,
Searcy and Andersson 1986). To elucidate
Plecoptera, stoneflies, drumming behavior, reproductive behavior, Talla-
these and other evolutionary implications,
more species and various populations with-
in species must be studied. The present work
describes drumming in three North Amer-
ican stonefly species and suggests some di-
vergence of drumming between intraspecif-
ic populations.
MATERIALS AND METHODS
Virgin adults of two of the three species
were obtained by rearing pre-emergent
nymphs in styrofoam bait buckets. The
nymphs were collected as follows: Talla-
perla maria (Needham & Smith), Fisher
Run, Columbia Co., PA, 12-V-1987; Ptero-
narcys biloba Newman, Fishing Crk., Co-
lumbia Co., PA, 18-V-1987. The adults of
Agnetina capitata (Pictet) were collected in
early May 1987 along the banks of the Sus-
quehanna River at Danville, PA (Montour
Co.). The presence of numerous emergent
exuviae along the riverbank showed that A.
584
capitata nymphs were emerging from the
Susquehanna and not from any nearby feed-
er streams.
All adults were allowed to adjust to lab
temperature for at least one hour prior to
recording. Drumming exchanges were re-
corded in stereo (male and female on sep-
arate channels) on cassette tapes using a re-
cording setup similar to that described by
Zeigler and Stewart (1985). All recordings
of 7. maria and P. biloba were of adults one
to three days old. Signals were measured
and analyzed using a Tektronix 5111 stor-
age oscilloscope, and selected signals were
photographed with a Nikon FG SLR cam-
era.
RESULTS AND DISCUSSION
Tallaperla maria. —At 19-20°C males (n
= 7) produced calls of a single scraping beat.
Observations indicated that the scrapes were
produced by curling the abdomen antero-
ventrally to contact the substratum and then
straightening the abdomen posteriorly thus
scraping the abdomen tip across the sub-
stratum. The male’s body rocked slightly in
the posterior direction as the scrape was
produced. The scrape tone averaged 596 +
86 Hz with a duration of 47 + 10 ms as
measured on the oscilloscope screen (Fig.
1). However, scrape duration could have
been less, because the residual ringing was
impossible to distinguish from the scrape
itself on oscilloscope tracings. When un-
answered by a receptive female, male scrapes
(calls) usually occurred in series of three to
five calls. Within such a series, calls were
separated by 1379 + 175 ms.
Females (n = 5) produced answers con-
sisting of a single abdomen/substratum beat
which followed the male call after 178 + 21
ms. After one to three of these simple ex-
changes, the males added a response scrape
190 + 11 ms after the female’s answer, and
later exchanges evolved into four-part ex-
changes with a male call, female answer,
male response, and female “response” (Fig.
2). In one pair, the male response was a
simple beat rather than a scrape.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
In three of the five pairs, the male switched
to a different type of call in later exchanges
within an exchange series. This call con-
sisted of the initial scrape followed by a sim-
ple beat (Fig. 3). The scrape/beat interval
was 245 + 25 ms and the female answer
typically followed in 181 + 9 ms. In two of
the five pairs, scrape/beat calls initiated a
few three-part and even four-part exchanges
as described above, but the majority of later
exchanges consisted only of a male scrape/
beat call and a female answer. Females were
typically stationary between exchanges while
males searched when not signalling.
Maketon and Stewart (1988) describe five
“calls” from three males of another 7. ma-
ria population in southwestern Virginia.
They interpret the call as consisting of a
series of scrapes similar to those noted for
the Pennsylvania males when females were
not answering. It seems likely that these Vir-
ginia calls were really a call series similar to
those noted herein which would have bro-
ken down into single scrape calls if females
had answered. Single scrape calls are de-
scribed for 7. lobata and T. anna, and
another call series described in T. elisa
(Maketon and Stewart 1988). Since no male/
female exchanges were recorded for these
species, it is impossible at this time to say
if the scrape/beat calls produced by the
Pennsylvania males late in exchange series
are unique to this species or population. The
intercall intervals in the call series of the
Virginia population appear distinctly short-
er than those noted for the Pennsylvania
population (app. 450 ms vs. 1379 + 175
ms, respectively) even allowing for the 4°C
warmer recording temperatures for the Vir-
ginia population (see Zeigler and Stewart
1977 regarding drumming speed and tem-
perature).
Pteronarcys biloba. — At 20°C, males (n =
4) produced calls of 12.5 + 1.5 beats with
beat intervals which were close to constant
at 944 + 53 ms. Only two answers from
one female were recorded. These answers
contained 9.5 + 0.7 beats with relatively
constant intervals of 869 + 48 ms. These
; owe KA
re vy
“Try if ’ i ++
eee
Figs. 1-6. Oscillographs of drumming signals. 1, 7a//aperla maria, male scrape call at 20°C (20 ms)*. 2,
Tallaperla maria, four-part exchange at 20°C (200 ms)*. 3, Ta/laperla maria, male scrape/beat call at 20°C (200
ms)*. 4, Preronarcys biloba, overlapping male/female exchange at 20°C (Ist female beat is seen after 7th male
beat, some male and female beats overlap) (2000 ms)*. 5, Agnetina capitata, diphasic male call at 21°C (1000
ms)*. 6, Agnetina capitata, reinforcement event at 21°C (1000 ms)*. *Time indicated by white bar in upper right
comer
585
586
answers overlapped the male calls by five
to six beats (Fig. 4). The overlap of call and
answer is a relatively common occurrence
in the genus Pteronarcys (Zeigler and Stew-
art 1987).
Stewart et al. (1982) describe drumming
ina Tennessee population of P. biloba. They
report a male call of seven to eight beats
with beat intervals of 524 + 46 ms. Re-
cording temperature was not reported for
the Tennessee population, so there is no way
to guess whether a significant difference ex-
ists between the two populations in beat
spacing. The difference in the number of
beats per call seems significant (TN = 7.17
+ 0.8; PA = 12.5 + 1.5), but the Tennessee
data, taken from only one male, is insufh-
cient for drawing a conclusion.
Agnetina capitata.—At 21-22°C six of
seven males produced diphasic calls with
9.2 + 2.6 beats in the first phase (Fig. 5).
Within the first phase, beat intervals de-
creased from 256 + 12 to 203 + 22 ms.
The interphase interval was 123 + 19 ms.
The second phase consisted of 68.9 + 43.3
beats with intervals averaging 100 + 14 ms,
but typically starting and ending intervals
averaged | 10-120 ms while the central body
of this phase consisted of 80-90 ms inter-
vals. This diphasic call, which ranged up
over 150 beats, is the longest male call, in
terms of beat number, ever described in
stoneflies.
One male produced monophasic calls that
were indistinguishable from the first phase
of the other six males’ calls. This shortened
call could have resulted because, in the ini-
tial exchange, the female answered before
even this “first phase” was completed. Per-
haps the male simply aborted his second
phase due to the female’s ““‘premature” an-
swer. The female also answered “early” in
the second exchange, but not in the third
and fourth where the male also delivered
only a first phase type call. The longest calls
recorded from other males (197 and 170
beats) were unanswered calls. It appears that
males may stop signalling when the female
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
initiates her answer, otherwise delivering a
longer call.
Females (n = 3) produced long answers
of 185.4 + 36.9 beats with relatively con-
stant beat intervals of 82 + 3 ms. Answers
typically overlapped the last few beats of
the male call. In two of three pairs, ex-
changes continued beyond the call and an-
swer with what I am terming “reinforce-
ment events.” The male would overlap or
follow closely the end of the female’s answer
with 12.9 + 0.8 beats with intervals varying
widely around 100 ms. These male beats
would stimulate more female drumming in
groups of 29.8 + 11.4 beats with beat in-
tervals similar to those in her answer. Re-
inforcement events (Fig. 6) usually num-
bered three to four per exchange and
overlapped the previous event or followed
after a pause of less than two seconds.
In the third pair, the male produced a
signal of 121 + 12.2 beats which overlapped
or shortly followed the female’s answer and
terminated the exchange. Beat intervals in
this signal were similar to those in the sec-
ond phase of the call but with slightly longer
beat intervals starting and ending this series.
In two of the three pairs, males delivered a
few (less than five) irregularly spaced “‘re-
inforcement beats” during the last half of
the female’s answer. These beats were de-
livered on the move (i.e. males did not cease
their searching movements to produce these
beats). The females remained stationary
during and between exchanges. Males
searched when not signaling, especially dur-
ing the long female answers which could last
as long as 25 seconds.
Maketon and Stewart (1984) and Graham
(1983) report on drumming in populations
of Agnetina capitata in Oklahoma and Wis-
consin respectively. However, after a recent
genus revision by Stark (1986), both pre-
vious studies were found to be on 4. fla-
vescens (K. W. Stewart and S. W. Szczytko
personal communication). The present de-
scriptions, then, are the first for 4. capitata.
Males in both populations of 4. flavescens
VOLUME 91, NUMBER 4
produced a much shorter monophasic call.
Since monophasic calls are generally be-
lieved to be the plesiomorphic condition
(Zeigler and Stewart 1987), the extraordi-
nary length and diphasic nature of the 4.
capitata calls appear to be apomorphic
within the genus, although diphasic calls
have apparently arisen separately in other
genera and families (Zeigler and Stewart
1987). The reinforcement events noted
above may likewise be apomorphic addi-
tions since they were not seen in the Okla-
homa A. flavescens (no live male/female ex-
changes were recorded in the Wisconsin
population), but conversely they could rep-
resent a shorter ancestral exchange with the
initial diphasic call and long female answer
being the apomorphic additions.
ACKNOWLEDGMENTS
I would like to acknowledge the cooper-
ation of Dr. A. R. Mallard and the Com-
munication Disorders Program of South-
west Texas State University for allowing me
access to their Tektronix 5111 oscilloscope.
I also thank Dr. Bill P. Stark for checking
my species identifications.
LITERATURE CITED
Graham, E.A. 1983. Drumming communication and
pre-mating behavior of fourteen Nearctic stonefly
587
(Plecoptera) species. M.S. Thesis. University of
Wisconsin at Stevens Point. 79 pp.
Maketon, M. and K. W. Stewart. 1984. Further stud-
ies of the drumming behavior of North American
Perlidae (Plecoptera). Ann. Entomol. Soc. Am. 77:
770-778.
1988. Patterns and evolution of drumming
behavior in the stonefly families Perlidae and Pel-
toperlidae. Aquatic Insects 10: 77-98.
Rupprecht, R. 1967. Das Trommeln der Plecopteren.
Z. Vergl. Physiol. 59: 38-71.
Searcy, W. A. and M. Andersson. 1986. Sexual se-
lection and the evolution of song. Annu. Rev. Ecol.
Syst. 17: 507-533.
Stark, B. P. 1986. The Nearctic species of Agnetina
(Plecoptera: Perlidae). J. Kans. Entomol. Soc. 59:
437-445.
Stewart, K. W.,S. W. Szczytko, and B. P. Stark. 1982.
Drumming behavior of four species of North
American Pteronarcyidae (Plecoptera): Dialects in
Colorado and Alaska Pteronarcella badia. Ann.
Entomol. Soc. Am. 75: 530-533.
West-Eberhard, M. J. 1984. Sexual selection, com-
petitive communication and species-specific sig-
nals in insects, pp. 283-324. Jn Lewis, T., ed.,
Insect Communication. Academic Press, New
York.
Zeigler, D. D. and K. W. Stewart. 1977. Drumming
behavior of eleven Nearctic stonefly (Plecoptera)
species. Ann. Entomol. Soc. Am. 70: 495-505.
1985. Drumming behavior of five stonefly
(Plecoptera) species from central and western North
America. Ann. Entomol. Soc. Am. 78: 717-722.
1987. Behavioral characters with systematic
potential in stoneflies (Plecoptera). Proc. Entomol.
Soc. Wash. 89: 794-802.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 588-593
LONGEVITY, FERTILITY, AND POPULATION GROWTH STATISTICS OF
TELENOMUS REYNOLDSI (HYMENOPTERA: SCELIONIDAE)
RONALD D. CAVE AND MICHAEL J. GAYLOR
Department of Entomology and Alabama Agricultural Experiment Station, Auburn
University, Alabama 36849; Current address (RDC) is Dept. de Proteccion Vegetal, Es-
cuela Agricola Panamericana, Apartado 93, Tegucigalpa, Honduras, C.A.
Abstract. — Longevity and fertility of female Telenomus reynoldsi Gordh and Coker
parasitizing Geocoris punctipes (Say) eggs were determined at constant temperatures of
20, 25, 28 and 32°C. Females lived significantly longer at temperatures below 32°C, but
produced significantly more offspring at 28°C. Net reproductive rate was greatest at 25
and 28°C, whereas intrinsic rate of increase was highest at 28°C. Population growth
statistics are compared to those of G. pallens Stal.
Key Words:
trinsic rate of increase
Big-eyed bugs, Geocoris spp., are com-
mon predaceous lygaeids in many agroeco-
systems of the western and southern United
States. Nymphs and adults consume i1m-
mature stages of several crop pests (Cham-
plain and Sholdt 1967, Ables et al. 1978,
Lawrence and Watson 1979, Crocker and
Whitcomb 1980, McDaniel et al. 1981,
Ragsdale et al. 1981). For this reason, big-
eyed bugs may be an important component
of pest management programs.
The scelionid Telenomus reynoldsi Gordh
and Coker is a solitary endoparasitoid of
big-eyed bug eggs. Distribution of the para-
sitoid extends throughout the southern
United States from California to Florida.
Egg parasitism may range up to 65% in
southern California cotton (Coker 1973) and
Alabama cotton and soybean fields (Cave
and Gaylor 1988b). Thus, 7. reynoldsi may
limit the impact big-eyed bugs may have in
pest management programs.
Information concerning reproductive ca-
pabilities and longevity of 7. reynoldsi is
useful for evaluating the potential impact of
Geocoris pallens, Geocoris punctipes, sex ratio, net reproductive rate, in-
this parasitoid on big-eyed bug populations.
Nevertheless, little work has been done on
these aspects of the adult biology of the
parasitoid. Coker (1973) reported the fer-
tility of 10 females at 27°C ranged from 47
to 107 progeny per female. He also found
that adult longevity decreased with an in-
crease in temperature from 16 to 32°C and
increased when adults were provided with
honey or with honey + water.
This paper presents the results of a study
of longevity and fertility of 7 . reynoldsi
parasitizing G. punctipes (Say) at four con-
stant temperatures. Population growth po-
tential of the parasitoid is described by four
population growth statistics calculated for
each temperature. These statistics are com-
pared to those of another big-eyed bug, Geo-
coris pallens Stal (Tamaki and Weeks 1972),
since population growth statistics are not
available for G. punctipes.
METHODS AND MATERIALS
Parasitized G. punctipes eggs from a lab-
oratory colony (Cave et al. 1987) were ran-
VOLUME 91, NUMBER 4
Table 1.
589
Reproductive statistics of Telenomus reynoldsi at four constant temperatures.
Temperature
Vanable
20°C
255C 28°C
n
Adult female longevity (¥ + SE) Won
Mean number progeny produced (X + SE) 14.1 +
Sex ratio (M:F) 1.5:la
Net reproductive rate (29/2) 4.3
Generation time (days) 35:3
Intrinsic rate of increase (29/2/day) 0.042
20 20 20
0.6a 16.8 + 0.8a 17.0 + 0.8a 12.4 + 0.6b
2c) 39:0 =63.5b (Se 2 5:2a) AI Db
1.5:la 1.6:la 3.7:1lb
10.7 10.5 le?
21.0 16.1 13.1
0.117 0.152 0.043
Means within rows followed by the same letter are not significantly different (P > 0.05; Duncan’s multiple
range test and x? test of homogeneity of proportions).
domly assigned to controlled environment
chambers and reared under constant tem-
peratures of 20, 25, 28, or 32°C and a 14:
10 L:D photoperiod. On the morning of
eclosion, adult 7. reynoldsi females were
transferred individually to plastic cups (30
ml) containing 10-15 G. punctipes eggs <72
h old and returned to the temperature and
photoperiodic regimen in which they were
reared. Each female was accompanied by I-
2 males. A drop of honey-water was applied
to the inside of each cup as a food source.
Each female was checked daily for survival
and, if alive, transferred to a new cup con-
taining fresh hosts and food. Dead males
were replaced with live ones. A water-sat-
urated cotton ball was placed in the cup
containing the exposed eggs and the eggs
were incubated at 28°C in a rearing room.
After 5 days, parasitized eggs were counted,
separated from unparasitized eggs (Cave and
Gaylor 1988a), and held for adult emer-
gence. The sex of each emerging adult was
determined. Hosts with parasitoids which
failed to emerge were dissected and the sex
of the parasitoid was determined if possible.
Effects of temperature on female longev-
ity and fertility were analyzed using the GLM
procedure of the Statistical Analysis System
(SAS 1985) and means were separated with
Duncan’s multiple range test. The sex ratios
of offspring produced at the different tem-
peratures were separated using a x test of
homogeneity of proportions. Each adult fe-
male was considered a separate replicate.
A survivorship and fertility table was
constructed for wasps held at each of the
four experimental temperatures by deter-
mining for each day (x) the proportion of
surviving individuals (1,) and the mean
number of daughters per surviving female
(m,). The survivorship level at adult emer-
gence was set as equal to the survivorship
level previously determined for the end of
the immature stage (Cave and Gaylor
1988a). From the survivorship and fertility
tables, the following population growth sta-
tistics were calculated: Ro, the net repro-
ductive rate, is
> lm, (1)
for all age intervals; G, the mean generation
time, 1s
> lame Ry (
where x, = pivotal age, which includes the
mean developmental time of an immature
female at the particular temperature (Cave
and Gaylor 1988a); and r, the intrinsic rate
of increase, 1s determined by substituting
values for r into the equation
> 1.m,e-"? = | (3)
until equality is obtained.
2)
RESULTS AND DISCUSSION
Longevity.— Mean longevity of adult fe-
male 7. reynoldsi was significantly (F = 11.3;
df = 77; P < 0.05) shorter at 32°C than at
590
100
—~ 80
aS
|
a
z 60
5
: °
s
5
2
20
04 + r + oe +
4 8 12 16 20 24 28
ADULT AGE (DAYS)
Fig. 1. Age-specific survivorship of adult female
Telenomus reynoldsi at four constant temperatures.
the other three temperatures (Table 1).
Maximum adult longevity at 20, 25, 28, and
32°C was 22, 26, 23, and 17 days, respec-
tively (Fig. 1). Coker (1973) determined
mean longevity of adult females at 16, 21,
27 and) 32°@ tobe 34.6,, 23-1, 20:3, and
17.2 days, respectively.
Survivorship at 20, 25 and 32°C was 100%
until nearly one-half or more of the maxi-
mum adult age (Fig. 1). Survivorship at 28°
was 100% until only one-third of the max-
imum adult age. Survivorship decreased
steadily after these periods until the last fe-
male died, except at 25°C where two females
lived 5 and 6 days longer than the third
oldest female.
Fertility. — Mean number of progeny per
female was significantly (F = 21.8; df = 77;
P < 0.05) greater at 28°C than at the other
three temperatures (Table 1). Fertility at
20°C was significantly less than at 25, 28,
and 32°C. The most progeny produced by
a single female at 20, 25, 28, and 32°C was
22, 80, 99, and 65, respectively. Coker
(1973) reported an average of 79.1 offspring
per female at 27°C; the maximum for a sin-
gle female in his study was 107. More male
than female offspring were produced at each
temperature (Table 1). The sex ratio was
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
significantly (x? test of homogeneity of pro-
portions; P < 0.05) more skewed towards
males (79%) at 32°C than at the three cooler
temperatures. Since fertility at 32°C was not
significantly different from fertility at 25°C,
we concluded that the high proportion of
males produced at 32°C may have been due
to sperm inviability, which would have
caused unfertilized eggs to become males.
Also, different temperatures may unequally
affect mating activity, which ultimately af-
fects sex ratio. Sex ratios in the field are 1:
1 or slightly skewed towards males (Cave
and Gaylor 1988b). Contrary to our find-
ings, Coker (1973) found that female prog-
eny outnumbered males 3 to 2. The differ-
ence in sex ratio between our laboratory
findings and those encountered in the field
and those found by Coker may be influenced
by laboratory conditions, such as colony
density.
The fertility of 7. revnoldsi is comparable
to that of other species of Te/enomus. Y ear-
gan (1982) found that the mean number of
progeny/female Te/enomus podisi Ashmead
was ca. 40 at 21°C. Orr et al. (1986) reported
the fertility of Telenomus calvus Johnson
was ca. 22 offspring/female at 27°C and that
ca. 32 offspring/female were produced by
Telenomus cristatus Johnson. Conversely,
Schwartz and Gerling (1974) observed that
Telenomus remus Nixon produced ca. 165
offspring at 25°C, although females lived
only nine days. A species that parasitizes
hosts in dense patches, such as 7. remus
attacking egg masses of Spodoptera spp..,
might take advantage of high host abun-
dance by producing large numbers of prog-
eny at one time. However, species attacking
smaller egg masses (e.g. the pentatomid egg
parasitoids, 7. podisi, T. cristatus, and T.
clavus) or widely dispersed solitary hosts
(e.g. T. reynoldsi) might have lower fertility
if they expend more energy reserves for host
searching and less for egg production.
Daily production of progeny by 7. rey-
noldsi was highest during the first five days
of adult life at 25, 28, and 32°C (Fig. 2). At
VOLUME 91, NUMBER 4
20T
— T=Total
--- #=Males
--*-9=Females
NUMBER OF PROGENY / FEMALE
0 4 8 12 16 20 24 28
ADULT AGE (DAYS)
Fig. 2. Age-specific fertility of female Telenomus
reynoldsi at four constant temperatures.
28 and 32°C, fertility gradually declined af-
ter five days, but at 25°C it stabilized for
seven days after declining for seven days.
As reflected in the lifetime sex ratios, male
progeny outnumbered female progeny on
most days.
Most females oviposited within 24 h after
emerging from their host. At temperatures
= 25°C, ca. 96% oviposited during the first
day. At 20°C, ca. 63% of the females did
not oviposit on day 1, and three did not
oviposit during the first three days. Coker
(1973) determined the preoviposition pe-
riod of 7. reynoldsi to be less than 6 h at
271°,
Mean (+SE) number of days to comple-
tion of oviposition was 10.7 (+0.6), 12.8
(+0.9), 13.8 (+0.7), and 9.8 (+0.5) for 20,
591
26C
AGE-SPECIFIC lm,
) 4 8 12 16 20 24 28
ADULT AGE (DAYS)
Age-specific 1,m, values of female Te/eno-
Fig. 3.
mus reynoldsi at four constant temperatures.
25, 28, and 32°C, respectively. Maximum
oviposition period was 15, 25, 19, and 14
days, respectively. Some females lived for
several days after their last day of ovipo-
sition. Mean (+SE) postovipositional pe-
riod was 6.5 (+0.9), 3.9 (+0.5), 3.2 (+0.5),
and 2.6 (+0.4) days at 20, 25, 28, and 32°C,
respectively. Yeargan (1982) reported that
T. podisi also lived ca. 5 days after comple-
tion of oviposition.
Population growth statistics. — The repro-
ductive contribution of each adult female
age-class is represented by the product of
age-specific survivorship and age-specific
production of daughters (1,m,) as a function
of age (x) (Fig. 3). At 25 and 28°C, the curves
are similar, with the earliest ages contrib-
uting most (i.e. l,m, values highest). Ages
592
2-8 contributed the most at 20°C. At 32°C,
l,m, values were highest the first few days,
but were still as low as 25% of those at 25
and 28°C because of lower survivorship and
fewer female progeny at 32°C.
Net reproductive rates (R,) at 25 and 28°C
were similar and more than 2- and 6-fold
those at 20 and 32°C, respectively (Table 1).
Mean generation time (G) decreased with
increasing temperature. The mean genera-
tion time at 20°C was more than 2-fold that
at 28 or 32°C. The intrinsic rate of increase
(r) was highest at 28°C and lowest at 20 and
32°C. The intrinsic rates of increase convert
to doubling times (In 2/r) of 16.9, 5.9, 4.6,
and 16.1 days. Although immature parasit-
oids develop fastest at 32—33°C (Cave and
Gaylor 1988a), reduced survival of imma-
tures, shortened lifetime, and low produc-
tion of daughters at this temperature limit
the population growth potential.
Population growth statistics have been es-
timated for only two other species of Te-
lenomus. Hirose (1986) calculated an ry of
0.296 females/female/day at 30°C for Te-
lenomus dendrolimi Matsumura attacking
the eggs of the pine moth, Dendrolimus
spectabilis Butler. Orr et al. (1986) found
that the intrinsic rate of increase of the pho-
retic pentatomid egg parasitoid T. ca/vus at
27° was 0.149 females/female/day, which is
similar to that found for T. reynoldsi at 28°C.
Population growth statistics have not been
estimated for G. punctipes. However, these
statistics have been determined for G. pal-
lens and Geocoris bullatus (Say) (Tamaki
and Weeks 1972). Geocoris pallens is a
known host of 7. reynoldsi (Gordh and Cok-
er 1973), but parasitism of G. bullatus eggs
has not yet been demonstrated. The net re-
productive rate (26.8 females/female) and
generation time (59 days) of G. pallens are
more than twice those of 7. reynoldsi at 25°
and 28°C. The intrinsic rate of increase of
G. pallens was 0.056 females/female/day,
which converts to a doubling time of 12.3
days. These population growth statistics
suggest that, although the host produces
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
twice as many daughters as the parasitoid,
the parasitoid has greater population growth
potential due to its shorter generation time
and faster intrinsic rate of increase. How-
ever, as pointed out by Tamaki and Weeks
(1972), Geocoris species may have quite dif-
ferent population growth statistics with dif-
ferent diets and physical conditions, and
these conditions may at times favor the host.
ACKNOWLEDGMENTS
We extend our appreciation to Lacy
Hyche, Timothy P. Mack and G. W. An-
kersmit for their critical reviews of the
manuscript. This paper is approved for pub-
lication as Alabama Agricultural Experi-
ment Station Journal No. 17-881486P.
LITERATURE CITED
Ables, J. R., S. L. Jones, and D. W. McCommas, Jr.
1978. Response of selected predator species to
different densities of Aphis gossypii and Heliothis
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Cave, R. D. and M. J. Gaylor. 1988a. Influence of
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survival of Telenomus reynoldsi (Hymenoptera:
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1988b. Parasitism of Geocoris (Heteroptera:
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versity of California, Riverside.
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dae: Geocorinae). Environ. Entomol. 9: 508-513.
VOLUME 91, NUMBER 4
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prey relationship of Geocoris punctipes and He-
liothis virescens. Environ. Entomol. 8: 245-248.
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Reproductive biology and behavior of Telenomus
calvus (Hymenoptera: Scelionidae), a phoretic egg
parasitoid of Podisus maculiventris (Hemiptera:
Pentatomidae). Can. Entomol. 118: 1063-1072.
593
Ragsdale, D. W., A. D. Larson, and L. D. Newsom.
1981. Quantitative assessment of predators of
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75: 181-183.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 594-603
LIFE HISTORY OF PTEROMALUS COLORADENSIS (ASHMEAD)
(HYMENOPTERA: PTEROMALIDAE) A PARASITE OF
PARACANTHA GENTILIS HERING (DIPTERA: TEPHRITIDAE) IN
CIRSIUM THISTLE CAPITULA
Davip HEADRICK AND RICHARD D. GOEDEN
Department of Entomology, University of California, Riverside, California 92521.
Abstract. —The biology and behavior of Preromalus coloradensis, a solitary larval-pupal
parasite of Paracantha gentilis, a thistle-capitulum-infesting tephritid, are described. Pter-
omalus coloradensis, previously known from a single female collected in Colorado, is
reported herein from California. We describe larval and pupal development using labo-
ratory reared specimens, and oviposition, feeding, and reproduction from field and lab-
oratory observations. Scanning Electron Microscopy of adult mandibles was used to
determine the function of the dentition during emergence from the host puparium and
overwintered capitula. Morphometric analysis of individual host puparia and parasites
showed three distinct relationships between size and sex of the parasite to the size of the
host puparium, substantiating predetermination of sex by the ovipositing female. Court-
ship behavior and the side mounting of the female by the male during copulation have
not been previously described for males in Pteromalus. A description of the male is given.
Key Words:
courtship behavior
This study of Pteromalus coloradensis
(Ashmead) was undertaken while working
on the life history of its host, Paracantha
gentilis Hering, a stenophagous capitulum-
infesting tephritid on Cirsium thistle species
in western North America (Foote and Blanc
1963, Goeden and Ricker 1986a, b, 1987a,
b). Ashmead (1890) described Pteromalus
(= Rhopalicus) coloradensis from one fe-
male specimen taken at West Cliff, Colo-
rado; no rearing records were given. Rare 1s
the opportunity to describe the life history
of a parasite, but Pt. coloradensis lent itself
well to study, thus we were able to obtain
as complete a record of its biology and be-
havior as known for any other North Amer-
ican species in this family.
Insecta, Paracantha, Cirsium, Pteromalus, parasite, functional morphology,
MATERIALS AND METHODS
Cirsium californicum Gray capitula con-
taining host puparia were collected from Mill
Creek, San Bernardino National Forest, San
Bernardino Co., California, 16 km N.E. of
Mentone, and Cirsium proteanum J. T.
Howell capitula were collected from Saw-
mill Mt., Angeles National Forest, Los An-
geles Co., California, 25 km S.E. of Gor-
man, during the Spring and Summer of 1987
and 1988 (Headrick and Goeden 1990). The
capitula were dissected to remove the host
puparia which were then individually placed
in 60-ml clear plastic rearing vials. Newly
emerged adults of Pt. coloradensis were
maintained on a diet of honey which was
streaked onto the inside of the vials. Cou-
VOLUME 91, NUMBER 4
ples, consisting of virginal males and fe-
males, were placed together in a vial to ob-
serve courtship and copulatory behavior.
The larvae and pupae were placed onto
moistened filter paper in covered glass petri
dishes and held in darkened growth cham-
bers at 27°C to determine developmental
rates.
Specimens for Scanning Electron Micros-
copy were killed by freezing, then either air-
dried or fixed in osmium tetroxide for 24 h
(Sabatini et al. 1963), and finally placed on
stubs and sputter-coated with a gold-plati-
num alloy. The SEM used was a JOEL-JSM-
35C3 in the Department of Nematology,
University of California, Riverside. Speci-
mens were examined and micrographs pre-
pared at 15 kV accelerating voltage on Po-
laroid 55 P/N film. All micrograph negatives
are stored with Gordon Gordh, Department
of Entomology, University of California,
Riverside. Voucher specimens of Ptero-
malus coloradensis from this study are de-
posited in the research collection of RDG
and the United States National Museum of
Natural History for incorporation into the
Chalcidoidea collection.
RESULTS
Biology. —New California records. San
Bernardino Co.: Mill Creek, 16 km N.E. of
Mentone on State Hwy. 38, San Bernardino
Nat. Forest, 14 F, 16 M, ex. Pa. gentilis on
C. californicum 25-IV-87 to 14-VI-88 (D.
Headrick, UCR). Los Angeles Co.: Sawmill
Mt., 25 km S.E. of Gorman at Sawmill
campground, Angeles Nat. Forest, 10 F, 5
M, ex. Pa. gentilis on C. proteanum 29-VII-
87 (D. Headrick, UCR).
Immature stages.—Pteromalus colora-
densis is a solitary, larval-pupal endopar-
asitoid. The egg is laid within the host dur-
ing the larval stages, but the host continues
its development through pupariation. Dis-
sections revealed that parasite larvae com-
pletely consumed the pupa within the pu-
parium only after the completion of
pupariation. However, three (5%) of 60 pu-
595
paria taken from post-blossom capitula of
Cirsium proteanum contained parasite lar-
vae that fed on the late pupal stage of the
host, leaving an empty, unemerged host
pupa inside the puparium. Normally, how-
ever, the overwintered larva consumed the
entire contents of the host and used the hol-
lowed puparium for its own pupation.
The last instar larva pupated with its head
directed anteriorly inside the host puparium
beginning in February in both the field and
the laboratory. Pupal development lasted 3
to 4 days under insectary conditions. Four
partly opened host puparia containing last-
instar parasite larvae were held in darkened
growth chambers and observed every 12 h
during pupation. Within the first 24 h, the
larva developed into an opaque white pupa
resembling the size and shape of the adult
and which could be sexed (there were three
males and one female). After 36 h, the head
and mesosoma became iridescent green
while the metasoma distad of the propo-
deum remained white. After 60 h, the male
metasoma had completely turned an iri-
descent green; however, the female sterna
remained unpigmented and the gonopore
and unsclerotized ovipositor could be seen
rhythmically pumping ca. once per second.
The antennae and legs of individuals of both
sexes were observed moving within the con-
fines of the pupal integument. After 72 h,
the female’s first-four sterna distad from the
propodeum became an iridescent green, and
the ovipositor was sclerotized to a golden
color. The antennae and legs moved more
rapidly than previously was observed and
showed twisting movements, the mesosoma
swelled every 5 to 10 min. The males
emerged before 84 h had elapsed and the
female emerged later after 94 h.
Adult.— Host puparia (n = 82) dissected
from capitula of C. californicum and C. pro-
teanum were caged individually in venti-
lated rearing vials and held in the insectary
at constant conditions; 11 (13%) parasites
emerged within | month. From three of the
puparia obtained from C. californicum heads
596 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Fig. 1.
=
; aS
MPG ee.
(A) Emergence hole in over-wintered head of C. californicum. (B) Catacord. (C) Catacord in dissected
host puparium. (D) Inside surface of left mandible. (E) Outside surface of right mandible.
caged on 9-VI-1987, parasites first emerged
on 1|1-II-1988 after 8 months in the insec-
tary. Adults emerged between 7-II to 9-III-
1988, from overwintered heads collected on
3-II-1988 at the Mill Creek site. Emergence
from overwintered heads in the field began
on 12-IIIJ-1988. Males emerged first, and
after a week, females began to appear. Field
observations and laboratory rearings sug-
gested that a portion of the first generation
of the parasite population emerges in sum-
mer and parasitizes the last of the current
host generation. The remainder of the first
generation as well as this second generation
of parasites overwinters in puparia within
closed capitula and presumably emerges the
following spring.
Adults exit the host puparium by chewing
a small, circular hole near the anterior end,
but never at the apex. They exit the capit-
ulum by chewing a tunnel through the dried
floral tubes, pappus and bracts of the over-
VOLUME 91, NUMBER 4
wintered head (Fig. 1A). As the emerging
parasite adult progresses through the head,
the chewed material, here given the name
“catacord”’ (Fig. 1B), is systematically
packed into the empty puparium (Fig. 1C).
The mandibular structure was viewed
with the SEM to determine the method of
excision and cutting of the catacord. In this
genus there can be a difference in dentition
between left and right mandibles, e.g. in Pr.
coloradensis the right has four teeth and the
left has three teeth in both sexes (Fig. 1D,
1E). The dorsal tooth has a broad, chisel
shape and a sharp leading edge. The left
dorsal tooth 1s twice as broad as the right.
The middle teeth are rounded with sharp
leading edges. Both mandibles have a ven-
tral tooth which is elongate, conical and ta-
pers to a point apically. The emerging par-
asite faced two very different substrates
through which it had to chew. The first ob-
stacle was the puparium which was hard,
concave, and without much surface texture
(Headrick and Goeden 1990). The two ven-
tral teeth punctured the wall of the pupar-
ium and then were brought together to make
a cut. The parasite continued to cut a cir-
cular hole, until eventually the excised piece
of shell was freed, then pushed down into
the bottom of the puparium. The next ob-
stacle was the dried capitulum tissues con-
sisting of floral tubes and compact fibers of
pappus. As the parasite exited the pupar-
ium, it chewed through the capitulum at an
angle perpendicular to the fibers. By lifting
its head, the plane of the cutting teeth was
in a position to grasp and cut through the
fibers. The uniform width of the catacord
is thus defined by the limit of mandible ex-
tension and the tunnel diameter is a func-
tion of the degree of rotation of its head.
Orientation of exit holes in capitula lying
on the ground, remaining on upright plants
or in rearing vials was always the same re-
gardless of the positions of the capitula (Fig.
1A). Capitula stored in darkened rooms over
the winter and later dissected showed that
most parasites had tunneled within the head
597
2507
MF
PARASITE THORAX WIOTHimm)
=
20+ M a
M
P
ie) } tot =—- + +
oO 5 20 25
HOST PUPARIUM WIDTH (mm)
Fig. 2. The relationships of the parasite thorax width
at its widest point and the host puparium width at its
widest point. F, female; M, male; (central M-F, rep-
resents a shared data point).
in various directions but did not exit. Thus,
perception of increasing light intensity may
help guide successful emergence from a ca-
pitulum.
Three relationships between the maxi-
mum width of the parasite thorax and the
maximum width of the host puparium from
which the parasite emerged are diagrammed
(Fig. 2). There is a distinct separation be-
tween size and sex in the parasite, i.e. fe-
males are larger than males. There is a sep-
aration, with little overlap, between the sex
of the parasite and the host size. There 1s a
significant correlation (corr. coeff. = 0.926,
P = 0.0001) between parasite size and host
size. These relationships indicate that host
size, or available resource, is a significant
factor in determining the sex and size of the
parasite. We hypothesize evidence of selec-
tive oviposition by parasites based on host
sex, 1.e. female parasites will lay fertilized
(or female) eggs into female hosts, and un-
fertilized (or male) eggs into male hosts
(Crozier 1971, Slobodchikoffand Daly 1971,
Charnov 1982).
Behavior.—Hostfinding: Cirsium thistle
capitula that are infested do not shed achenes
and remain closed and overwinter on or near
the parent plant. The parasitized tephritid
puparia inside these heads are insulated by
the dried achenes and pappus against freez-
598
ing winter temperatures. These overwinter-
ing heads, from which adults of Pt. color-
adensis emerge in spring, are scattered
among the current season’s thistle plants,
thereby facilitating host finding by newly
emerged parasites.
Males and females were observed to fly
directly to and land on the capitula of this-
tles where they apparently searched for food,
mates, and oviposition sites. Usually, nei-
ther sex was observed on other parts of the
thistles, but occasionally males were ob-
served to land on the upper surfaces of
leaves. Adults also were observed resting on
the leaves of nearby plants other than this-
tles, but were not observed feeding, mating
or Ovipositing on these plants. The peak ac-
tivity period for Pt. coloradensis was from
1000 to 1500 h. They did not remain on the
thistles overnight, but were observed to fly
to them each morning.
Feeding: On three occasions individual
males and females were observed feeding
on the exudate from Paracantha gentilis
Oviposition wounds.
Oviposition: Females search the surface
of a head by antennating in an apparently
random manner. In selecting an oviposition
site, the female places her mouthparts on
the surface, moves away, then returns and
touches her mouthparts again to the same
site. The number of these return visits var-
ied from two to five. Then, positioning her-
self head-up on the capitulum over the site
just selected, she places the apex of her me-
tasoma on the outer surface of the bract,
planting the ovipositor tip so that she can
unhinge the ovipositor by stepping back-
wards. She continues stepping backwards
until the ovipositor is perpendicular to the
long axis of her body and to the outer surface
of the capitulum. This allows her to balance
upon the ovipositor and to pull herself
downward with her legs. Drilling begins with
a slight twist of the metasoma in a lateral
plane and a side-to-side rocking of the body.
By continuing to pull downward she pushes
the ovipositor though the bracts and finally
into the softer tissues of the capitulum.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Once the hole is drilled and the female
has the ovipositor inserted completely into
the head, she pushes with her legs in a slight
up-and-down motion, presumably search-
ing for a host larva. If none is found, she
then uses her legs to push up and away from
the capitulum surface to remove the ovi-
positor. Once removed, it is secured by step-
ping forward. Females were first observed
to oviposit in thistle capitula in late March.
They select young capitula that are fully ex-
posed, or at least free from surrounding
leaves. Apparently, Pt. coloradensis females
do not distinguish capitula infested by Pa.
gentilis, because early 1n the season they drill
into uninfested capitula. This indicates that
they failed to associate the oviposition
wounds of Pa. gentilis in capitula at which
the parasite adults fed with the presence of
host larvae. The time spent on a capitulum
by a female parasite varied from | min to
2.5 h. Females were observed to drill from
one to six holes during observation periods
that lasted a maximum of 2.5 h. The average
time spent in drilling holes was 8 + 2 (X +
SE) (range, 2-22, n = 13) min; the average
time between drillings was 8 + 2 (range, l-
15, n = 7) min. Dissections showed that if
no host was located inside a capitulum, or
only host eggs were present, no parasite egg
was deposited. Capitula dissected after ovi-
position showed that drill holes occurred in
rows along the vertical axis of the capitu-
lum, from about one-third the distance be-
tween the apex of the capitulum to the pe-
duncle. On three separate occasions females
were observed drilling into the undersides
of capitula near the peduncle. Five capitula
collected 3 weeks after the first observation
of parasite oviposition contained on aver-
age of 36 + 5 (range, 20-49) drill holes.
Courtship. — Laboratory: Two each, new-
ly emerged (<48 h old) virgin males and
females were caged as couples (one male and
one female) in separate screened plastic vials
for ca. 30 min observation, after which they
were separated. Courtship behavior began
an average of 15 + 4 (range. 5 to 30) min
after caging (n = 9). Males approached fe-
VOLUME 91, NUMBER 4
ee
lar 5 fe
Mo ((
SZ / ;
NY |
IN
A mse Xp
Fig. 3.
599
Ve
(A) Male initiating courtship: dual-directional arrows indicate wing fanning and lateral weaving. (B)
Antennation posture: dual-directional arrows indicate antennal movement. (C) Position of male and female in
copula viewed from both sides. Arrow shows depressed antennae in female. Male drawn with right pair of wings
removed.
males either laterally or head-on and stopped
when they were | cm or less apart, but not
touching. With his antennae extended, the
male began to sway his body from side to
side and fan his wings (Fig. 3A). Swaying
consisted of a lateral rocking motion, while
moving his body over his legs from one side
to the other.
Wing fanning as a form of auditory court-
ship communication has been described for
two other pteromalids, Pteromalus pupar-
jum L. and Nasonia vitripennis (Walker)
(Miller and Tsao 1974). Experimentation
has shown that males vibrate their wings at
a specific frequency and pulse which is then
followed by the male mounting the female.
Wing fanning by Pt. coloradensis males also
preceded their mounting of the females and
beginning antennation.
Antennation occurred while the male
stood on the female’s dorsum with his fore-
tarsi on the vertex of her head, his middle
tarsi in front of her tegulae, and his hind
tarsi grasping her axillae (Fig. 3B). The fe-
male drummed the upper surface of her fu-
nicle segments 1-3 against the closed man-
600
dibles of the male. This is in contrast to
the courtship behavior described for other
Pteromalidae, e.g. Nasonia and Euptero-
malus (cf. Barass 1960, van den Assem
1974), where males moved their open man-
dibles over the upper surface of the female’s
funicle segments. While the female of Pr.
coloradensis drummed her antennae on the
mouthparts of the male, he intermittently
stroked the lateral aspects of her antennae
with his funicular segments (Fig. 3B), as is
common in the Pteromalinae (van den As-
sem 1974). Single antennation episodes
lasted an average of 3 + 0.4 s (range, 1-5
s, n = 12). When the female was receptive,
only one such episode occurred before cop-
ulation and was of short duration, 1.e. ca. 2
s. If the female was unreceptive, yet still
confined with the male in the vial, anten-
nation occurred up to seven times during a
30-min period, including five times in a 10-
min period, each episode terminating with
the male dismounting. The intervals be-
tween male approaches varied from | to 30
min.
Mating.— Laboratory: If the female was
receptive after antennation, she lowered her
flagella and held them tightly against her
head, a motion that has been reported to be
the receptivity signal to the male (Barass
1960, van den Assem 1974), and simulta-
neously, raised the apical four segments of
her metasoma to expose the gonopore. The
male immediately moved backwards and
down her left side, bent his metasoma down
and under her metasoma, and placed the
apex on the gonopore (Fig. 3C). This cop-
ulatory position is unusual for Pteromalinae
males, which usually tend to copulate from
a position behind and underneath the me-
tasoma of the female (van den Assem 1974).
The female terminated the mating by relax-
ing her metasoma and “shaking-off’ the
male. Two matings lasted ca. 30 s each. No
post-copulatory behavior was observed. The
male did not try to remount the female, but
both actively moved inside the vial for the
duration of the observation time.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Field: One courtship and copulation se-
quence was observed on the apex of a C.
californicum terminal capitulum. The male
approached the female and began swaying
and wing fanning for ca. 2s. He then mount-
ed her for antennation as described above
for ca. 3 s. Copulation followed and lasted
about | min. Males were observed to ap-
proach, mount, antennate, and dismount
any female they contacted. On separate oc-
casions, three different males mounted and
antennated a single female as she was ovi-
positing.
DISCUSSION
Zwolfer (1988) discussed biogeography of
thistles and suggested that they crossed into
North America from the Palearctic prior to
the late Miocene via the Bering Strait. Few
of the associated Palearctic insect taxa fol-
lowed their Cirsium host plants, and few
insect taxa of true Nearctic origin use them
for host plants. The genus Paracantha, how-
ever, is an exception (Goeden and Ricker
1986b, 1987a, b, Zwoélfer 1988), being orig-
inally Nearctic. Pteromalus coloradensis
may have followed its host onto Cirsium,
having been pre-adapted, with an elongate
Ovipositor, to parasitizing Pa. gentilis in
other capitula. Such an adaptation may also
have resulted in the unique copulatory po-
sition of the male having to move to the
side of the metasoma of the female.
The thistle-insect complexes that have
been studied in the Palearctic (Zwolfer 1985,
1988) have phytophagous arthropod guilds
that are in turn attacked by guilds of para-
sites. In southern California, there is a pau-
city of phytophagous species on thistles
compared to the fauna of Europe (Zwolfer
1965, Goeden and Ricker 1986a, b, 1987a,
b), and parasite species attacking them also
are few. No other species of Pteromalus in
North America is known to attack tephritid
hosts (Krombein et al. 1978).
Information on host relationships for
Pteromalus in North America may not give
as clear an indication of generic relation-
VOLUME 91, NUMBER 4
ships as that of European species. Ptero-
malus coloradensis was placed in the genus
Habrocytus until Pteromalus and Habro-
cytus were synonymized by Boucek and
Graham (1978). In Graham’s (1969) dis-
cussion of the Pteromalidae of North-West
Europe, 67 Habrocytus species were includ-
ed, of which 35 (52%) had unknown biol-
ogies. Of the 32 species whose biologies were
known, 14 (43%) were parasites of Tephri-
tidae exclusively, and the other 18 species
parasitized various phytophagous Lepidop-
tera and Coleoptera. The Tephritidae hosts
listed in Graham (1969) are non-frugivo-
rous species infesting capitula of various As-
teraceae, mainly belonging to genera in the
tribe Cardueae (thistles and knapweeds), e.g.
Centaurea, Cirsium and Carduus. Two of
the Lepidoptera and Coleoptera hosts listed
were also associated with Cardueae. Of the
nine Pteromalus species included, none were
listed as parasites of Diptera. When Boucek
and Graham (1978) transferred Habrocytus
to Pteromalus, they invalidated the generic
significance of such host relationships. Data
on the biologies of a host and parasite are
never complete, and often not useful in tax-
onomy; however, when biological infor-
mation is available, such as that listed by
Graham (1969), it could help to clarify taxo-
nomic questions.
DESCRIPTION OF THE MALE
Unknown until this study, the male of Pt.
coloradensis is described here. Body irides-
cent blue-green, head and thorax coarsely
reticulate and tinged with gold, metasoma
smooth and tinged with copper. Antenna
mostly fuscous; scape, pedicel smooth and
testaceous; flagellum fuscous. Coxae con-
colorous with thorax; trochanters testa-
ceous; femora concolorous with coxae ex-
cept apices testaceous; tibiae completely
testaceous; tarsi testaceous proximally, fus-
cous distally. Tegulae testaceous. Wings
hyaline, venation testaceous.
Head (Fig. 4A).— Head as broad as meso-
scutum, coarsely reticulate, eyes separated
601
Fig. 4. (A) Anterior view of the male head. (B)
Posterior view of the male head. (C) Thoracic spiracle
of the male.
dorsally by 1.5 their length; malar space
nearly '2* eye height; clypeus striate, an-
terior margin cleft medially. Antenna in-
serted 3 above lower ocular line, scape
length ca. ¥%4 eye length, nearly reaching low-
er edge of median ocellus; combined length
of pedicel and flagellum 1.1 x head width;
pedicel width 1.5 x length, about 2 length
of first funicular segment; funicle segments
quadrate, stouter than pedicel; clavus length
not quite twice width, slightly longer than
combined length of preceding two funicular
602
segments; sensilla numerous, usually in one
row circumscribing funicle segment. Pos-
terior aspect of head with many longitudinal
furrows; setae sparse, a row of up to six setae
on either side of occipital foramen extend-
ing from the ventral edge of the posterior
tentorial bridge to the dorsal edge of the
maxillary insertion, occipital foramen tri-
angular, as wide as maxilla at insertion (Fig.
4B).
Pronotal collar sharply margined ante-
riorad. Propodeum width 1.8 _ scutellar
width; 3 = length; plica distinctly bordered
by fine reticulation; panels shiny with lon-
gitudinal coarse wrinkles; costula distinct as
a lateral band of reticulation between apical
foveae; median carina absent; nucha with
small wrinkles and a band of coarse reti-
culation at base. Spiracles elongate-oval set
at ca. 45° angle to midline, inner walls with
distinct reticulation (Fig. 4C).
Forewing with costal cell bare; speculum
open below, extending 2 way below mar-
ginal vein; stigmal vein with one row of
setae, and 1.1 x longer than marginal vein
and slightly longer than the postmarginal
vein.
Gaster elongate-oval, length 2.2 x width,
dorsally compressed and with a flat dorsal
surface, ventral plica; narrower than thorax;
basal tergum occupying ' of total length.
ACKNOWLEDGMENTS
We thank D. W. Ricker and M. Mora-
torio for technical assistance with this re-
search, E. E. Grissell (Systematic Entomol-
ogy Lab., ARS-USDA, Beltsville, Md.) for
the identification of specimens, T. S. Bel-
lows, G. Gordh, J. LaSalle and J. Pinto for
their reviews of earlier drafts of this manu-
script.
LITERATURE CITED
Ashmead, W. H. 1890. On the Hymenoptera of Col-
orado; descriptions of new species, notes, and a
list of the species found in the state. Bull. Colo.
Biol. Assoc. 1: 27-28, 45.
Assem, J. van den. 1974. Male courtship patterns
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
and female receptivity signal of Pteromalinae
(Hym., Pteromalidae), with a consideration of some
evolutionary trends and a comment on the taxo-
nomic position of Pachycrepoideus vindemiae.
Neth. J. Zool. 24: 253-278.
Barass, R. 1960. The courtship behavior of Mor-
moniella vitripennis Walk. (Hymenoptera: Pter-
omalidae). Behaviour 15: 185-209.
Boucek, Z.and M. W.R.de V.Graham. 1978. British
check-list of Chalcidoidea (Hymenoptera): Taxo-
nomic notes and additions. Entomol. Gaz. 29: 225-
235%
Charnov, E. L. 1982. Theory of Sex Allocation.
Princeton Univ. Press, Princeton, NJ.
Crozier, R. H. 1971. Heterozygosity and sex deter-
mination in haplodiploidy. Am. Nat. 105: 399-
412.
Foote, R. H. and F. L. Blanc. 1963. The Fruit Flies
or Tephritidae of California. Bull. Calif. Insect Surv.
7. 115 pp.
Goeden, R. D. and D. W. Ricker. 1986a. Phytoph-
agous insect faunas of two introduced Cirsium
thistles, C. ochrocentrum and C. vulgare, in south-
ern California. Ann. Entomol. Soc. Am. 79: 945-
952.
1986b. Phytophagous insect faunas of the
two most common native Cirsium thistles, C. cal-
ifornicum and C. proteanum in southern Califor-
nia. Ann. Entomol. Soc. Am. 79: 953-962.
1987a. Phytophagous insect faunas of the
native thistles, Cirsium brevistylum, Cirsium
congdonii, Cirsium occidentale, and Cirsium tio-
ganum, in southern California. Ann. Entomol. Soc.
Am. 80: 152-160.
1987b. Phytophagous insect faunas of native
Cirsium thistles, C. mohavense, C. neomexican-
um, and C. nidulum, in the Mojave Desert of
southern California. Ann. Entomol. Soc. Am. 80:
161-175.
Graham, M. W.R. de V. 1969. The Pteromalidae of
North-Western Europe (Hymenoptera: Chalci-
doidea). Bull. Brit. Mus. (Nat. Hist.) Entomol.
Suppl. 16. Pp. 1-908.
Headrick, D. and R. D. Goeden. 1990. Life history
of Paracantha gentilis Hering (Diptera: Tephriti-
dae). Ann. Entomol. Soc. Am. 83: [submitted].
Krombein, K. V. et al. [eds.]. 1970. Catalogue of
Hymenoptera in America north of Mexico, Vol.
1. Smithsonian Inst. Press, Wash., DC. 1979.
Miller, M. C and C. H. Tsao. 1974. Significance of
wing vibration in male Nasonia vitripennis (Hy-
menoptera: Pteromalidae) during courtship. Ann.
Entomol. Soc. Am. 67: 772-774.
Sabatini, D. D., K. Bensch, and R. J. Barrnett. 1963.
The preservation of cellular ultrastructure and en-
zymatic activity by aldehyde fixation. J. Cell Biol.
17: 19-58.
VOLUME 91, NUMBER 4
Slobodchikoff, C. N. and H. V. Daly. 1971. System-
atic and evolutionary implications of partheno-
genesis in the Hymenoptera. Am. Zool. 11: 273-
282.
Zwolfer, H. 1965. Preliminary list of phytophagous
insects attacking wild Cynareae (Compositae) in
Europe. Commonw. Inst. Biol. Control Tech. Bull.
6: 81-154.
603
. 1985. Insects and thistle heads: Resource uti-
lization and guild structure, pp. 407-416. Jn Del-
fosse, E. S., ed., Proc. VI Int. Symp. Biol. Cont.
Weeds, Vancouver, Canada. Agriculture Canada.
1988. Evolutionary and ecological relation-
ships of the insect fauna of thistles. Ann. Rev.
Entomol. 33: 103-122.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 604-610
MEGASTIGMUS FLORIDANUS (HYMENOPTERA: TORYMIDAE),
NEWLY DISCOVERED IN ILEX SEED (AQUIFOLIACEAE)
E. E. GRISSELL
Systematic Entomology Laboratory, PSI, Agricultural Research Service, USDA, c/o
U.S. National Museum, NHB 168, Washington, D. C. 20560.
Abstract. — Megastigmus floridanus Milliron was discovered to be phytophagous in seeds
of I/ex opaca and one of its hybrids with //ex cassine. This is the first report of a Mega-
stigmus associated with Aquifoliaceae. A brief review of phytophagous wasps associated
with //ex seed is given, and adults and larvae of the two New World seed feeders (Mega-
stigmus floridanus, Torymus rugglesi) are illustrated. Originally described from a single,
unreared specimen from Florida, M. floridanus was rediscovered there, where it was reared
from J/ex x attenuata. It is newly reported from Maryland from an unknown cultivar of
Tlex opaca. Here it was univoltine, overwintering as mature larvae in mature fruit. Adults
emerged in May and June.
Key Words:
Ilex ‘Foster #2’
In June of 1982, Dr. Douglass Miller
(USDA, ARS, Systematic Entomology Lab-
oratory) discovered in Beltsville, Maryland
a cultivar of J/ex opaca Aiton that was en-
gulfed in adult female wasps of the chalci-
doid genus Megastigmus Dalman. These
wasps were emerging from mature red fruit
that remained on the tree from the previous
season. Species of Megastigmus are known
to be seed-feeders, mainly on conifers and
rosaceous plants, but have never been re-
ported from //ex (Lessmann 1962), nor any
Aquifoliaceae.
I identified the wasp as Megastigmus flor-
idanus Milliron by comparison with the sin-
gle female holotype specimen collected in
Florida in 1949. Nothing was known of its
host or biology at the time of description,
and no additional specimens have been
found until this study. In 1987 I spent two
weeks in Florida looking for Megastigmus
Megastigmus floridanus, Torymus rugglesi, Ilex opaca, Ilex x attenuata,
floridanus under the auspices of The Holly
Society of America. In this paper infor-
mation is presented on the rediscovery of
M. floridanus in Florida and its discovery
in Maryland. Preliminary data is presented
on its biology as a phytophagous feeder in
the seeds of cultivars of [/ex opaca and nat-
ural hybrids of /. opaca with I. cassine Lin-
naeus. A synopsis of holly seed-feeding
wasps is given as well as methods to distin-
guish M. floridanus from Torymus rugglesi
Milliron, the only other New World holly
seed feeding wasp. The latter species is also
reported for the first time from a commer-
cial cultivar of //ex.
PHYTOPHAGOUS WASPS IN /LEX SEED
At a world level, wasp species known to
be phytophagous in //ex seed are confined
to the family Torymidae of the Chalcidoi-
dea. This is the family to which Megastig-
VOLUME 91, NUMBER 4
mus belongs. Although most torymid wasps
are parasitic on other insects, there are sev-
eral genera with phytophagous species.
Bootania Dalla Torre.—This Austral-
asian genus 1s closely related to Megastig-
mus and has two species in Japan which feed
in I/ex seed. Bootania japonica (Ashmead)
occurs in seeds of //ex serrata Thunberg and
I, purpurea Hasskarl (reported as shinensis
Sims, Kamijo 1962); Bootania hirsutum
(Kamyjo) occurs in the seeds of J/ex integra
Thunberg (Kamijo 1962, 1981). Records for
Japonica and hirstum prior to 1988 are un-
der the generic name Macrodasyceras Ka-
mijo which was recently synonymized un-
der Bootania by Boucek (1988).
Torymus Dalman.—Torymus rugglesi
(Figs. 2, 4) has been reared from the seeds
of Ilex opaca in Delaware, Maryland, and
Virginia (Milliron 1949). This species was
omitted from ‘Field Guide to Insect Pests
of Holly” by McComb (1986). During the
period of 16 August to 30 September 1988,
I discovered 7. rugglesi emerging in large
numbers from seeds of //ex ‘Foster #2’ in
Silver Spring, Maryland. This is the first
record of the species from a cultivated J/ex.
These specimens are in the collection of the
U.S. National Museum of Natural History.
Megastigmus. —Only Megastigmus flori-
danus Milliron (Figs. 1, 3) is known from
Ilex. Taxonomically M. floridanus may be
distinguished from 7. rugglesi, the only oth-
er phytophagous wasp with which it might
be confused, by several characteristics.
Megastigmus floridanus (Fig. 1) is entirely
yellow, has an ovipositor that is about half
the body length, and has the stigmal vein
of the wing enlarged. Torymus rugglesi (Fig.
2) is metallic green dorsally and yellow on
the remainder of the body, has an ovipositor
that is slightly longer than the body, and has
the stigmal vein barely indicated. The ma-
ture larvae of both species may be identified
most easily based on setae. Megastigmus
floridanus (Fig. 3) has setae confined to the
first three body segments (excluding the
head), whereas 7. rugglesi (Fig. 4) has setae
605
over most of the body. Setae in both species
are white or clear and can be overlooked.
MATERIALS AND METHODS
Field.—Over a five year period (1983-
1988), observations were made at the USDA
Beltsville Agricultural Research Center-
West, Beltsville, Maryland, during the
months of April through July. Initially two
trees of J/ex opaca (unknown cultivar, det.
T. R. Dudley) were found to be infested, but
other trees within a one mile vicinity were
swept and examined visually on a yearly
basis for the presence of adult wasps. Be-
cause adults were abundant and fruit was
readily available for dissection, I made no
rearings from fresh fruit at Beltsville.
The search for Florida populations of MV.
floridanus centered upon Osceola County
because the only known specimen of the
wasp (the holotype) was collected there on
7 August 1929. Because the wasp I found
in Maryland was on holly, I searched for
habitats that could support holly species in
general and J. opaca in particular. No spe-
cific locality was given within Osceola
County, so I surveyed 23 localities over a
nine day period from | to 9 August 1987.
All hollies encountered were sampled with
a sweep net for the presence of free-living,
adult wasps. Both red (mature) and green
(immature) fruit were collected when avail-
able and stored in paper bags that were
placed in plastic bags to prevent excessive
dehydration. Emergence was checked daily
and all wasps were collected and killed in
alcohol. Collections were also made about
100 miles north in the Ocala National For-
est, Florida (Marion and Lake Counties)
where nine sites were sampled over a three
day period. Voucher herbarium specimens
were taken from each tree from which a fruit
collection was made and were determined
to species by Dr. T. R. Dudley, U.S. Na-
tional Arboretum, Washington, D.C. These
specimens are now in the herbarium at the
Arboretum. Reared wasp specimens were
606 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LL MG Fh Ewes
2 \ x
=
Figs. 1, 2. Adult females (scale = 0.5 mm). 1, Megastigmus floridanus. 2, Torymus rugglesi, ovipositor
extended upward in drawing, but normally horizontal or down-curved.
VOLUME 91, NUMBER 4
determined by me and are in the U.S. Na-
tional Museum, Washington, D.C.
Laboratory.—Fruit for dissection was
collected at Beltsville, Maryland from a sin-
gle tree of Z/ex opaca (one of the two men-
tioned above). This tree was part of a foun-
dation planting and had been in place a
number of years based upon the height of
the tree (ca. 12 ft.). Because the presence of
wasp larvae could not be determined by the
external appearance of fruit, and because
wasp larvae were difficult to find at the be-
ginning of the study in 1983, fruit was first
examined with x-ray equipment. After sev-
eral years, as populations increased, larvae
could be found in almost every fruit and
dissections were made at random. Dissec-
tions were made the day after collection and
larvae were killed 1n boiling water and stored
in alcohol for future study. On one occasion,
withered fruit from previous years’ crops
was taken from under the tree to determine
if live wasps might be present in them.
RESULTS OF FIELD SURVEY
Florida. — During the Florida field survey
I swept the following holly species: [/ex am-
bigua (Michaux) Torrey, //ex = attenuata
Ashe (/lex cassine x Ilex opaca), Ilex bus-
wellii Small, Ilex cassine, and Ilex glabra
Gray. No specimens of //ex opaca were
found. Only one free-living, adult female of
M. floridanus was collected, and this was on
Tlex x attenuata.
Fruit collections were made as follows
(mature/immature): J. ambigua 0/1200, J.
x attenuata 612/3970, I. bushwelli 0/100,
I. cassine 0/16,500. Of these only J. atten-
uata produced wasp adults: 3 from mature
(red) fruit and 116 from immature (green)
fruit. These wasps emerged from fruit col-
lected in Osceola County at Three Lakes
Wildlife Management Area (25 mi. S. St.
Cloud) and Prairie Lakes (29 mi. S. St.
Cloud). Wasp emergence occurred from 7
to 26 August 1988.
Ilex x attenuata and Ilex cassine were
the only hollies of those listed above that
607
£7?
Poy
PEA
Aut py ta
ici emee le sie
fem
Poe!
oN el lV
S A \ \ B
4 °° hey!
Figs. 3, 4. Larvae of //ex seed feeding Torymidae
(A, side view, B, ventral view; scale = 0.5 mm). 3,
Megastigmus floridanus, overwintering larva. 4, To-
rymus rugglesi, mature larva.
exhibited all stages of floral development on
the same tree at the same time. That 1s,
every stage from bud and flower to imma-
ture green fruit and mature red fruit was
available as a resource at one time.
Maryland.— During the period of study,
adult, free-living Megastigmus floridanus
were collected only in the months of May
and June. Adults were seen exiting from
mature, red fruit still attached to the tree.
During the first four years, wasps were col-
lected only on two trees which occurred
within 100 feet of each other. Although oth-
er nearby female trees were swept (from 100
yards to a mile away), no wasps were found.
In 1987, adult wasps were found for the first
time on four trees about 100 yards to the
east. Five trees 100 yards to the west had
608
no wasps. The tree first found to be infested
with M. floridanus in 1982 still supported
a large population of wasps in 1988.
The trees at Beltsville bloomed only in
May and June. At that time, the previous
season’s mature red fruit remained on the
tree, but there was no green fruit. After June
there were no buds or flowers present and
the developing fruit matured synchronous-
ly, reddening by early fall.
RESULTS OF LABORATORY DISSECTIONS
Larvae of M. floridanus were found to be
solitary endophytes within an //ex seed. Each
Ilex opaca fruit produced four seeds. Among
28 infested fruit the most common number
of larvae per fruit was 2 (65%), but 1 (14%),
3 (14%), and 4 (7%) larvae were also found.
In randomly selected fruit sampled during
July, August, and October from a heavily
infested tree, 80 to 100 percent (avg. 93%)
of the fruit (n = 30) was infested. On a per
seed basis (n = 120), 38 to 63 percent (avg.
50%) were infested, 25 to 40 percent (avg.
34%) were distorted (indicating possible
ovipositor probing and perhaps egg laying),
and 12 to 23 percent (avg. 16%) appeared
to be sound. There was no observable dif-
ference in appearance between infested and
non-infested seed, and there appeared to be
no obvious external affect on the growth of
the holly fruit. A small, clean, circular emer-
gence hole in the mature red fruit was the
only evidence that Megastigmus floridanus
had been present.
In 1987, I dissected 40 fruits (160 seeds)
on 11 June and the same number again on
9 July and found no evidence of eggs or
larvae. Larvae were first detected on 20 July
and averaged 0.6 mm in length (range 0.4
to 0.9 mm, n=14). As the overall seed in-
festation rate was 38 percent at this time, I
probably overlooked the eggs and larvae on
11 June and 9 July. By 26 August the larvae
averaged 2.4 mm in length (range 1.8 to 2.7
mm, n=17) and by | October they averaged
2.3 mm (range 1.8 to 2.9 mm, n=21). Ran-
dom dissections of fruit in early January
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
yielded larvae which averaged 2.2 mm in
length (range 2.1 to 2.5 mm, n=13).
A single, incidental collection was made
of old, blackened fruit from beneath a tree
on 15 May. Forty seeds were dissected and
contained three live adult females. This fruit
could have fallen prematurely from the cur-
rent year’s crop or could have been on the
ground for several years.
DISCUSSION
Milliron (1949) outlined a general life his-
tory for Megastigmus in which the female
adult emerged in spring and immediately
oviposited into immature host seeds; one
larva developed per seed, attained full
growth by late summer, and remained dor-
mant over winter; pupation occurred in
spring and adults emerged with the onset of
new seed initiation. I believe that Mega-
stigmus floridanus follows this develop-
mental phenology at least in the Maryland
population. Adult females were present in
May and June, and small larvae were first
detected in late July. Larval development
was completed at least by late August after
which time larval length remained approx-
imately the same through January when dis-
sections were no longer made. I believe that
I overlooked egg and early larval growth in
June and early July and found only mature
larvae after the August dissections. Because
mature larvae were present in January I as-
sume that this is the overwintering phase.
Ilex opaca in Beltsville, Maryland had a
definite seasonal phenology, with one crop
of buds developing synchronically, then
flowers, and then fruit. Megastigmus flori-
danus appeared to track this phenology and
produced one generation per year. In Flor-
ida populations of J/ex x attenuata, all stages
of floral and fruit development occurred si-
multaneously in August. Thus there was less
synchronized fruit development than on
Maryland trees and more overlap of avail-
able resources. It is possible that 7. flori-
danus is multivoltine in Florida, but this
was not determined.
VOLUME 91, NUMBER 4
In Maryland the discovery of adult fe-
males in fallen fruit of unknown age indi-
cated the possibility of a small portion of
the population diapausing for one or more
years. Diapause 1s known to occur in Mega-
stigmus, and Milliron (1949) cited examples
of several species which have retarded
emergences of up to two years. This is
thought to be a method of ensuring a con-
tinuing wasp population in case there should
be a year of poor or bad seed production.
It would be interesting to determine what
percent ofa year’s wasp population remains
in diapause and how long such dormants
might survive in fallen fruit on the ground.
(I have observed phytophagous wasps of the
genus Euryfoma emerging from seeds of
Rhamnus crocea Nuttall over a four year
period in the laboratory.)
It seemed surprising, at first, that Mega-
stigmus floridanus was found in seeds of
both //lex = attenuata in Florida and J. opa-
cain Maryland. Ilex x attenuata, however,
is a naturally occurring hybrid of J. opaca
and J. cassine (T. R. Dudley, personal com-
munication) so that the host trees are pre-
sumably closely related. What is more sur-
prising, however, 1s that none of the seeds
of pure J. cassine yielded adult wasps in
spite of the large number of collected fruits
(16,500). It may be that J. opaca and its
hybrids are the only hosts suitable for M.
floridanus development.
The known distribution of //ex opaca is
Massachusetts south to Florida, west to
Pennsylvania and Texas (Eisenbeiss and
Dudley 1973). Megastigmus floridanus
might be expected to have the same distri-
bution as its host tree, but additional sur-
veying within this potential range will be
necessary to determine this.
It is odd that only recently has an appar-
ently common and widespread seed-feeding
wasp been discovered on a common and
widespread plant such as American holly.
In Great Britain, records of the cultivation
of introduced American holly extend back
as far as 1744, and over 1,000 worldwide
609
selections have been given cultivar names
in the last 40 years alone (Eisenbeiss and
Dudley 1973). It would be interesting to
discover the natural geographic and host
range of Megastigmus floridanus and to de-
termine what impact, if any, this seed-feed-
ing wasp might have on its host trees.
ACKNOWLEDGMENTS
I thank Douglass R. Miller, USDA, ARS,
Systematic Entomology Laboratory, Belts-
ville, MD, for finding and collecting Mega-
stigmus floridanus. For a grant in support
of my fieldwork in Florida and for the pub-
lication of this paper I thank the Research
and Development Committee of The Holly
Society of America, Inc., Barton M. Bauers,
Sr., Chairman. Additionally, I thank Charles
W. McComb, Insect Population Monitor-
ing, Bena, Virginia, for suggesting that the
Holly Society might be interested in my
work. For help in several aspects of my Flor-
ida field work I thank Jim Smith (retired
Osceola County Agent) and Eleanor C.
Foerste, Osceola County Extension Service,
St. Cloud, Florida. I am grateful to T. R.
Dudley, U. S. National Arboretum, Wash-
ington, D.C., for the identification of //ex
specimens. I also thank Robert J. Amox and
the Nicolet XRD Corporation, Fremont,
California for use of their x-ray instrumen-
tation to determine the presence of wasp
larvae. Finally, for reviewing this paper and
offering numerous useful comments, I thank
Steve Heydon, Department of Entomology,
Smithsonian Institution, Washington, D.C.,
Manya Stoetzel and Paul Marsh, Systematic
Entomology Laboratory, Washington, D.C.,
and Charles McComb.
LITERATURE CITED
Boucek, Z. 1988. Australasian Chalcidoidea (Hy-
menoptera). C.A.B. International, Wallingford,
Oxon, UK. 832 pp.
Eisenbeiss, G. K. and T. R. Dudley. 1973. Interna-
tional checklist of cultivated //ex. Part 1. Ilex opa-
ca. Natl. Arbor. Contr. No. 3. 85 pp.
Kamio, K. 1962. A revision of the species of the
610
Megastigminae occurring in Japan (Hymenoptera:
Chalcidoidea). Insecta Matsumurana 25: 18-40.
. 1981. Description of the male and other notes
of Macrodasyceras hirsutum (Hymenoptera:To-
rymidae). Akitu 38: 1-4.
Lessmann, von D. 1962. Ubersicht der bisher be-
kannten Megastigmus-Arten und ihrer Wirts-
pflanzen. Z. Angew. Entomol. 50: 233-238.
McComb, C. W. 1986. A field guide to insect pests
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
of holly. Holly Society of America, Baltimore, MD.
v + 122 pp.
Milliron, H. E. 1949. Taxonomic and biological in-
vestigations in the genus Megastigmus. Am. Mid.
Nat. 41: 257-420.
. 1959. Anew species of Torymus reared from
the seeds of American holly (Hymenoptera:To-
rymidae). Ann. Entomol. Soc. Am. 52: 52.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 611-614
APTILOTUS MARTINI, A NEW SPECIES OF THE APTILOTUS BECKERI
GROUP (DIPTERA: SPHAEROCERIDAE) FROM
CAVES IN THE CANARY ISLANDS
TERRY A. WHEELER AND S. A. MARSHALL
Department of Environmental Biology, University of Guelph, Guelph, Ontario, Canada
N1G 2WI1.
Abstract. —Aptilotus martini n.sp., a brachypterous sphaerocerid with strongly reduced
eyes, 1s described from caves on La Palma, Canary Islands. The taxonomic status of the
A. beckeri group within the genus Apti/otus is briefly discussed.
Key Words:
The sphaerocerid genus Aptilotus Mik
comprises 18 species of apterous, brachyp-
terous, and macropterous flies, widely dis-
tributed in the Holarctic region. Six species
from the Canary Islands constitute the Ap-
tilotus beckeri species group (see Papp and
Rohacek 1981, Rohacek and Papp 1983—
as Paralimosina beckeri group). This paper
provides a description of a new cave-in-
habiting species of the Aptilotus beckeri
group and summarizes the taxonomic his-
tory of the group.
Aptilotus martini Wheeler and Marshall,
NEw SPECIES
(Fics. 1-9)
Description.— Body length 2.1-3.3 mm.
Color dark brown, pruinose; trochanters,
distal ends of tibia, and tarsomeres light
brown. Interfrontal bristles in three pairs,
middle pair much longer and cruciate, an-
terior pair shortest. Face carinate. Arista as
long as thorax, with long pubescence. Eyes
with indistinct margins, facets flattened,
pruinose. Eye height 1.0-1.2 times genal
height. Katepisternum with posterodorsal
bristle strong. Two pairs of postsutural dor-
socentral bristles, posterior pair longer. Pre-
sutural acrostichal setulae in ten rows. Scu-
Sphaeroceridae, Aptilotus, Aptilotus martini, taxonomy
tellum 1.5 times wider than long, apical
scutellar bristles 1.5—2 times scutellar length.
Mid tibia with one long and one short
anterodorsal and one posterodorsal bristle
in basal third, one anterodorsal, one dorsal,
and one posterodorsal bristle in distal third,
one apicoventral bristle. Wing reduced, ex-
tending only to posterior margin of second
abdominal tergite, wing membrane light
brown. Costa ending at junction with R4+ 5.
R4+5 straight, with R2 +3 diverging at close
to 90° angle. M, CuA1, and r-m present, dm-
cu absent. Halter without knob, reduced to
small yellow stub.
Male abdomen.—Preabdomen heavily
sclerotized; densely setose except sternite |
and anterior half of syntergite 1 +2. Sternite
5 (Fig. 4) with bifid, strongly deflexed pos-
teromedial tab. Tab lightly sclerotized with
darker, roughly X-shaped region. Single
transverse row of 8-10 stout spines anterior
to tab, with group of 3-5 longer bristles at
each end of row. Lateral regions of sternite
with scattered long setae. Synsternite 6+7
simple, dextral lobe narrow, extending over
posteromedial tab of sternite 5. Sternite 8
free, not fused with synsternite or epan-
drium. Epandrium uniformly setose, bris-
tles longer along posterior margin (Fig. 1).
pa
iy cae te te? aa
! Vb aed Piece 54 wee
aan ea pu! Hy phurc ea See sats
helt Vey ThA iy EU
Sn rte RSLS Nhe i f
Sia SAS eae Se Pe BAC cil MNCL]
i
1.
Figs. 1-9. Aptilotus martini new species. |, Male genitalia (left lateral view); CE, cercus; EP, epandrium; HP,
hypandrium; SS, surstylus; scale bar = 0.2 mm. 2, Aedeagal complex (left lateral view); AA, aedaegal apodeme;
BP, basiphallus; DP, distiphallus; EA, ejaculatory apodeme; PM, paramere; same scale as Fig. 1. 3, Surstylus
(left lateral view); scale bar = 0.1 mm. 4, Male fifth sternite (ventral view); scale bar = 0.2 mm. 5, Wing (dorsal
view); scale bar = 0.2 mm. 6, Female spectacles-shaped sclerite; scale bar = 0.1 mm. 7, Female terminalia
(dorsal view); CE, cercus; EPT, epiproct; T8, tergite 8; scale bar = 0.3 mm. 8, Female terminalia (ventral view);
HPT, hypoproct; S8, sternite 8; same scale as Fig. 7. 9, Spermathecae.
612
VOLUME 91, NUMBER 4
Cercus simple, fused with epandrium, with
ca 5 bristles, basal bristle longer than any
epandrial bristle. Sternite 10 (interpara-
meral sclerite) small but well sclerotized,
arms narrow, medial part visible as a nar-
row dark strip between cerci. Surstylus with
a setose posterolateral outer part and a long,
anteriorly dark-pointed inner part bearing
two large, flattened spur-like setae; anterior
spur-like seta with a broad basal lobe and
tapered apex (Fig. 3). Basiphallus simple,
carinate. Distiphallus with a narrow collar-
like base, expanding to a broad, membra-
nous, laterally spinulose part then tapering
to a distal part with a large, Y-shaped dorsal
sclerite, lateral lobes bearing long flat spi-
nules, and distal ventral part covered with
small spinules. Paramere simple, slightly
sinuate, truncate (Fig. 2).
Female abdomen.—Sternite | well-de-
veloped, bare except posterior margin: syn-
tergite 1+2 similar in size to tergite 3, bare
on anterior half; tergites and sternites 2-5
densely setose. Tergites and sternites 6 and
7 short, with only posterior bristles. Tergite
and sternite 8 paler than preceding sclerites;
tergite 8 tripartite, median part bare; lateral
parts setulose and setose except along bare
anterior margin. Sternite 8 setulose and se-
tose on posterior half only, posteromedial
margin with a row of 4 tuberculate bristles.
Hypoproct setose and setulose along pos-
terior margin only (Fig. 8). Epiproct broad,
setulose except along anterior margin (Fig.
7). Cerci short, blunt, with short, stout api-
cal and preapical dorsal bristles. Internal
vaginal sclerotization (Spectacles-shaped
sclerite) hyaline, with large, narrow rings
(Fig. 6). Spermathecae dark, spherical, with
distinct darker outpocketings: sclerotized
parts of ducts short (Fig. 9).
Types.— Holotype (¢): CANARY IS-
LANDS. La Palma: Cueva El Diablo,
24.vii.1986, J. L. Martin. Paratypes: CA-
NARY ISLANDS. La Palma: Cueva E1
Diablo, 24.vii.1986, J. L. Martin (5 4,14
2); Cueva del Rincon, 22.viii.1986, J. L.
Martin (22 6,12 9); Cueva Arrebolas,
613
28.vill.1986, J. L. Martin (1 4,1 ?); Cueva
Todoque, 18.viii.1986, J. L. Martin (5 4,1
2); Cueva Franceses, 2.1x.1986, J. L. Martin
(2 2).
Holotype deposited in the Biosystematics
Research Centre, Ottawa, Canada. Para-
types deposited in the Biosystematics Re-
search Centre, the University of Guelph col-
lection, the British Museum (Natural
History), the Museo Insular de Ciencias
Naturales de Tenerife, and the Departa-
mento de Zoologia de la Universidad de La
Laguna (Tenerife, Canary Is.).
Etymology.—This species is named for
Jose L. Martin, who collected the type se-
ries.
Comments.—Aptilotus martini is easily
distinguished from all other species in the
beckeri group by its flattened eyes, with in-
distinct margins. Other distinguishing char-
acters include the shape and degree of scler-
otization of the posteromedial tab of the
male fifth sternite, and the shape of the male
surstyli and parameres.
DISCUSSION
Papp and Rohaéek (1981) described four
brachypterous species of Canary Islands
sphaerocerids closely related to the mac-
ropterous species Limosina beckeri Duda.
These four species, along with L. beckeri,
were tentatively assigned by the authors to
the genus Paralimosina Papp as P. anap-
tera, P. beckeri, P. franzi, P. gomerensis, and
P. pilifemorata. Rohaéek (1983), recogniz-
ing that the P. beckeri group is morpholog-
ically distinct from other Paralimosina
species, erected the subgenus Paralimosina
(Canarisina) for the above five species. Al-
most simultaneously, Marshall (1983) not-
ed that the P. beckeri group shared a number
of synapomorphies with the apterous Eu-
ropean species Aptilotus paradoxus Mik, and
transferred the five members of the beckeri
group to the genus Apzti/otus.
An additional species of the P. beckeri
group, P. avolans, was described by Roha-
614
éek and Papp (1983), who suggested that P.
avolans is most closely related to beckeri.
Subsequently, these authors transferred P.
avolans to Aptilotus and considered the sub-
genus Canarisina a junior synonym of Ap-
tilotus (Rohaéek and Papp 1988). For the
present, pending a complete revision of Ap-
tilotus, we prefer to treat the beckeri group
as a species group, without subgeneric sta-
tus.
ACKNOWLEDGMENTS
We would like to thank Dr. Pedro Oromi
and Jose L. Martin, Universidad de la La-
guna, Tenerife, Canary Islands, who pro-
vided us with the type series of A. martini.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
LITERATURE CITED
Marshall, S. A. 1983. A revision of the genus Aptilo-
tus Mik in North America (Diptera, Sphaerocer-
idae). Can. J. Zool. 61: 1910-1924.
Papp, L. and J. Rohaéek. 1981. New species of the
Paralimosina beckeri-group from the Canary Is.
(Diptera: Sphaeroceridae). Folia Entomol. Hung.
42: 143-154.
Rohaéek, J. 1983. A monograph and re-classification
of the previous genus Limosina Macquart (Dip-
tera, Sphaeroceridae) of Europe. Part II. Beitr.
Entomol. Berlin 33: 3-195.
Rohaéek, J.,and L. Papp. 1983. Another new species
of the Paralimosina beckeri-group from the Ca-
nary Islands (Diptera: Sphaeroceridae). Folia
Entomol. Hung. 44: 147-152.
. 1988. A review of the genus Paralimosina L.
Papp (Diptera, Sphaeroceridae), with descriptions
of ten new species. Annls. Hist.-Nat. Mus. Natn.
Hung. 80: 105-143.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 615-619
OBSERVATIONS ON MEGANOLA SPODIA FRANCLEMONT
(LEPIDOPTERA: NOCTUIDAE) WITH A DESCRIPTION
OF THE MATURE LARVA
LINDA BUTLER
Division of Plant & Soil Sciences, P.O. Box 6108, West Virginia University, Morgan-
town, West Virginia 26506-6108.
Abstract.—In 1985, Franclemont gave the name Meganola spodia to a species whose
genitalia he had illustrated in 1960 but had left unnamed. The only mention of the larva
of this species was in the 1960 paper in which a brief description of coloration was given
along with the implication that Dyar’s (1899) description of the M. phylla (= minuscula)
larva was actually of MW. spodia. Meganola larvae collected in West Virginia were reared
to adults which are clearly M. spodia. The larvae are distinctly different from those
previously described. Herein, I describe the last instar larva, cocoon construction and
field observations for adults and larvae of M. spodia.
Key Words:
West Virginia
In 1983 during a study of an outbreak of
spring defoliating geometrids, I collected
partially mature larvae of an unknown No-
linae (Noctuidae) on oak in two eastern
counties of West Virginia. Genitalia of adults
reared from these larvae matched those il-
lustrated for Franclemont’s (1960) un-
named Meganola. Subsequently, this species
was named Meganola spodia Franclemont
(1985).
Franclemont (1960) included the follow-
ing description of the larva of WZ. spodia:
“dorsum pale cream color, mottled; with a
dark brown subdorsal stripe, joined by
transverse bands on A4 and 8; venter pale:
hair pale, that on i11 very long; head whitish,
mottled with brown on the lobes, jaws
black.” Apparently, Franclemont believed
Dyar’s (1899) larval description of . phyl-
la Dyar (= minuscula Zell.) to be a descrip-
tion of M. spodia. Larvae which I reared in
Noctuidae (Nolinae), Meganola spodia, oak defoliation, larval description,
this study were a rich green in color: dorsal
stripes were yellow and red; no transverse
bands were present and longest body setae
were dark brown. I do not consider Dyar’s
previous description of M. phylla (minis-
cula) to be that of M. spodia.
This paper includes field observations of
larvae, a description of the last instar larva,
cocoon construction and blacklight trap rec-
ords for adults in West Virginia. Voucher
specimens are in the West Virginia Uni-
versity Arthropod Collection, Morgantown.
BIOLOGICAL OBSERVATIONS
Meganola spodia larvae were collected at
Cacapon State Park, Morgan Co., WV on
chestnut oak (Quercus prinus L.) on 3 May
1983 soon after leaf expansion had begun.
The larvae appeared to be about half mature
and were yellowish to pale green with a yel-
616
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 1-3.
of last instar larva (4.6 x).
low mid-dorsal stripe. I collected additional
larvae on the same host species at this lo-
cation on 17 May. One larva was taken from
this site 24 May on black oak (Q. velutina
Lam.). Larvae were collected from post oak
(Q. stellata Wang.) 13, 18, and 24 May on
Elkhorn Mountain near Dorcas (Grant
County) WV. A total of 20 larvae were col-
lected.
Dyar (1892) described Meganola (as Nola)
Meganola spodia. |, larvae and damage on oak in field (1.3 x). 2, cocoons (4.1 x). 3, dorsal view
minuscula larvae as hiding “by day in dry
curled leaves that adhere to the twigs or in
some other place of concealment on the
branch.” The larvae I observed during this
study (Fig. 1) fed diurnally and unconcealed
which resulted in noticeable marginal and
hole feeding damage on host foliage.
Field collected larvae were maintained in
the laboratory on the host species on which
they were collected. All larvae produced co-
VOLUME 91, NUMBER 4
coons by 29 May and adults emerged be-
tween 29 May and 18 June.
It appears as though M. spodia produces
one generation per year and overwinters as
half mature larvae.
M. spodia adults have been recorded from
New Jersey, New York, Virginia, North
Carolina and Kentucky with collection dates
ranging from 18 June to 16 July (Francle-
mont 1985). West Virginia records of adults
at blacklight traps are as follows: Boone Co.
(Danville) 15 June 1979, R. Swope; Fayette
Co. (Babcock State Park) 11 July 1979, L.
Butler; Grant Co. (Dorcas-Elkhorn) 7 July
1980, L. Butler; Lincoln Co. (Big Ugly Pub-
lic Hunting Area) 28 June 1979, L. Butler;
McDowell Co. (Panther State Forest) 23
June 1979, L. Butler; Monongalia Co.
(Triune) 14 July 1979, L. Butler; Morgan
Co. (Cacapon State Park) 17 July 1979; and
Wyoming Co. (Twin Falls State Park) 3 July
1980, L. Butler. Seventeen adults from
blacklight traps are in the West Virginia
University Arthropod Collection.
COoCcOON CONSTRUCTION
Stehr (1987) described Nolidae cocoons
as being well formed, boat shaped with an
anterior ridge forming a valve-like slit.
Packard (1884) described Nola ovilla Grt.
cocoons as boat shaped, oval, and cylindri-
cal attached to leaves and spun with silk
and bits of leaves. Additional observations
on N. ovilla (as Lebena ovilla) were ‘‘the
larva builds up two parallel walls and unites
them at the top. Cocoon elliptical, flat at
base, size 7 x 2'’2 mm. The anterior end is
a little higher and more pointed than the
posterior’ (Dyar 1894). N. triquetrana
(Fitch) (as N. sexmaculata Grt. and N. tri-
notata Walker) constructs the cocoon on a
piece of wood from bits of bark laid together
like bricks (Dyar 1890, 1891). The cocoon
of Nola minna Butler (as Nola hyemalis
Stretch) was described as “not strong, com-
posed entirely of silk, and not firmly fas-
tened to a support. It is elliptical, opaque
617
white” (Dyar 1891). Meganola minuscula
(Zeller) (as N. phylla Dyar) cocoons were
described as triangular and constructed of
little pieces of bark (Dyar 1899). The co-
coon of M. minuscula (as Nola minuscula
Zeller) was described as “elliptical, opaque,
sordid white, composed of white silk, quite
tough and intermingled sparsely with larval
hairs. Dimensions 8 x 4mm” (Dyar 1892).
Cocoon construction by M. spodia larvae
is detailed. The larva lies along the twig
facing what will ultimately be the front of
the cocoon and lays down a thin silk mat
on the twig, lies over the mat and begins
plucking small bits of twig bark (thin trans-
parent cortex flakes and darker, bark
chunks), and incorporating them into the
silk, working alternately on left and right
sides of the twig, thus producing flanges on
each side of the twig. The larva stretches
forward to pluck bark from in front of the
mat and curls its body into a “C” over the
edge of the mat to pluck bark from the bot-
tom of the twig below the mat. Larvae stretch
farther on smaller diameter twigs to reach
enough construction material.
When the sides of the mat are each about
4 mm wide; the larva backs up, plucks bark
from the twig above the mat, and applies
silk and bark to produce cupping of the two
sides. When each side of the mat reaches
about 5 mm wide, the larva completes the
front of each side by adding anteriorly ex-
tending points where each side of the mat
attaches to the twig and at their outermost
edges to produce flaps. The left flap is ex-
tended about 1.5 mm beyond the right. Just
prior to closure, the larva begins inverting
its body, alternately examining each end of
the cocoon and adding bark or silk to weak
spots. Closure begins at the back of the co-
coon at twig level as the larva pulls the two
sides together, connects them with silk and
continues to close the top of the cocoon from
posterior to anterior. For front closure, the
larva generally adds silk to the left flap,
grasps the flap with the mandibles, pulls it
toward the right flap and attaches it; flap
Figs. 4-8.
tal view. 5, left mandible, oral surface. 6, prothorax
and mesothorax, lateral view. 7, abdominal segment
1, lateral view. 8, abdominal segments 6-10, lateral
view. Figs. 6-8 anterior to left. Scale lines = 0.5 mm.
Meganola spodia. 4, head capsule, fron-
closure generally progresses from bottom to
top.
While cocoons were generally constructed
of silk and twig bark, one larva which pro-
duced a cocoon on post oak incorporated
oak bud scales into the cocoon. One cocoon
on a leaf petiole consisted of petiole epi-
dermis and leaf pubescence held together
with silk.
The average cocoon construction time for
those which I timed was 7 h, 42 min (range
6 h, 40 min to 8 h, 50 min).
Cocoons (n = 15) averaged 12.7 mm in
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
length (range 11.8-13.2 mm), 3.54 mm in
height (range 3.2-4.0 mm) and were pro-
duced on twigs which averaged 2.5 mm in
diameter (range 1.8—3.2) (Fig. 2).
DESCRIPTION OF LAST INSTAR LARVA
Live last instar larvae medium green;
“knobs” of verrucae reddish; narrow red
dorsal stripe, continuous between segments
and slightly widened at middle of each seg-
ment, bordered by yellow stripes edged lat-
erally in red (Fig. 3).
The length of five mature ethanol pre-
served larvae ranged from 14.0 to 17.5 mm.
Head (Fig. 4): Pale tan with darker yel-
low-tan freckles. Greatest head width 1.78
mm. Adfrons weakly defined; frontoclypeus
slightly less than one-half the distance to
epicranial notch. Stemmata I-4 evenly
spaced; 6 close to 4; 5 at posterolateral edge
of antennal socket. All head setae long and
fine. P2 and P1 closely adjacent with P2
above and closer to epicranial suture. L1
distant from and slightly below P1 and di-
rectly above A2, which 1s slightly below A1;
A3 slightly above stemma 3, 02 near lower
edge of stemma | and O01 slightly above
stemma 4. AFI at junction of frontal and
coronal sutures and directly above Fl. Cl
and C2 closely adjacent and at membranous
fronto-clypeal junction. Mandibles (Fig. 5)
with four outer teeth; strong tooth on inner
surface; two well-separated outer setae pres-
ent.
Thorax (Fig. 6): Ventral gland absent.
Most primary setae obscured by tufts of ir-
regularly lengthened secondary setae on
verrucae. Tl shield weakly developed. D1
verrucae low, lying along anterior edge of
Tl, expanded transversely and almost
meeting at dorsal midline; all setae brown
tipped. D2 setae large, prominent, closely
adjacent at midline. L verruca slightly
knobbed, protuberant; setae of irregular
lengths, longest uniformly brown, inter-
mediate ones all pale and shortest setae
brown tipped. SV verruca very protuberant,
all setae pale. T1 spiracle elliptical, yellow-
VOLUME 91, NUMBER 4
ish brown with darker brown peritremes; T 1
spiracle 2x size of those of Al—-A7.
On T2, D1 verruca low, rounded; setae
of irregular lengths, none very long, most
brown tipped. SD and L verrucae very fleshy,
protuberant, slightly knobbed; two long,
brown whip like setae on SD verruca, other
setae shorter and entirely brown or brown
tipped; one long, brown whip like seta on
L verruca, other setae in decreasing lengths
all pale, all brown, and brown tipped. SV
verruca small, but very protuberant; setae
irregular, all pale. T3 setal arrangement sim-
ilar to that of T2, but SD and L setae not
large and whip like.
Abdomen (Figs. 7, 8): On Al-8 all D ver-
rucae low, rounded, all SD and L large, pro-
tuberant and slightly knobbed. L setae very
irregular in length, most longer setae pale.
One brown whip like seta on L verruca of
A8. SV verrucae small, sparsely haired, all
setae pale. A8 spiracle 2 the size of spi-
racles on Al-A7. On A9 (Fig. 8), D and SD
well developed, fleshy, protuberant, each
with two prominent, brown, whip like setae.
L verruca smaller with one prominent brown
seta. SV greatly reduced, sparsely haired.
A10 (Fig. 8) with most setae primary. Most
prominent feature is anal fork formed by
four long, stout intercrossing setae. Cro-
chets all uniordinal mesoseries; A4 and AS
with 21-22 crochets, A6 with 20-21 and
A10 with 22-23.
ACKNOWLEDGMENTS
I thank Vicki Kondo for laboratory as-
sistance and J. E. Weaver, J. W. Amrine
and J. E. Hall for review of the manuscript.
This paper is published with the approval
of the West Virginia Agricultural and For-
estry Experiment Station as Scientific Ar-
ticle No. 2127. The research was supported
with funds appropriated under the Hatch
Act.
LITERATURE CITED
Dyar, H.G. 1890. Description of certain lepidopter-
ous larvae. Insect Life 3: 61-62.
1891. Notes on Bombycid larvae I. Psyche
6: 110.
1892. The larva of Nola miniscula. Psyche
6: 248-249.
1894. Notes on Bombycid larvae. Psyche 7:
137-138.
. 1899. On the larvae of North American Noli-
dae with descriptions of a new species. Can. Ento-
mol. 31: 61-64.
Franclemont, J. G. 1960. Nolidae, pp. 50-55. In
Forbes, W. T. M. Lepidoptera of New York and
Neighboring States, Part IV Agaristidae through
Nymphalidae Including Butterflies. Cornell Univ.
Agric. Exp. Sta. Memoir 371.
1985. A new species of Meganola Dyar from
Eastern North America (Lepidoptera: Noctuidae:
Nolinae) Proc. Entomol. Soc. Wash. 87: 871-874.
Packard, A. S. 1884. The transformation of Nola.
Amer. Nat. 18: 726-727.
Stehr, F. W. 1987. Nolidae, pp. 548-549. Jn Stehr,
F. W., ed., Immature Insects. Kendall/Hunt Pub.
Co., Dubuque, Iowa.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 620-631
SYSTEMATIC NOTES ON SOME BETHYLIDAE FROM BOTSWANA:
PRISTOCERINAE (HYMENOPTERA: ACULEATA)
KARL V. KROMBEIN
Department of Entomology, NHB Stop 105, Smithsonian Institution, Washington, DC
20560.
Abstract. —Apenesia forchhammeri, new species, is described from both sexes from
Botswana. Apenesia punctulata is proposed as a replacement name for A. punctata Kieffer,
1904, from Cameroon, preoccupied in Apenesia by Epyris punctatus Cameron, 1888.
Males and the previously unknown females of Pristocera rhodesiae Turner, 1928, and
Prosapenesia lacteipennis Kieffer, 1910, are described from Botswana.
Key Words:
Per Forchhammer, Serowe, Botswana, has
operated several Malaise traps for the
Smithsonian from 1986 to the present time,
collecting all kinds of flying insects. The traps
were placed in a vegetation type known as
Acacia nigrescens/Combretum apiculatum
tree savanna, at an altitude ca 1500 m, with
an average annual rainfall ranging from 214
to 721 mm over a period of 10 years.
Forchhammer’s collecting has yielded
many interesting Hymenoptera, three of the
more unusual of which are described below.
All species of the bethylid subfamily Pris-
tocerinae have small wingless females. Ap-
terous female bethylids do not enter malaise
traps by their own volition. The respective
males are larger, fully winged forms with
large multidentate mandibles that are well
adapted to grasping the small female and
carrying her during a mating flight. So it is,
rarely of course, that one of these mating
flights occasionally terminates in a trap.
The following abbreviations, mostly as
used by Evans (1963), are employed in the
descriptions that follow:
LH—length of head from middle of clyp-
eal margin to midpoint of vertex;
Hymenoptera, Bethylidae, Apenesia, Pristocera, Prosapenesia
WH-—greatest width of head including
eyes;
WF—width of front (i.e. least interocular
distance);
HE—height of eye measured in lateral
view;
EV —distance from top of eye to crest of
vertex in lateral view;
WOT — width of ocellar triangle including
posterior ocelli;
OOL—ocello-ocular line, least distance
between posterior ocellus and inner eye
margin;
LT—length of thorax, collar excluded,
from anterior margin of pronotal disk to
posterior end of propodeum;
propodeal formula (females only)—an-
terior width: narrowest width (usually at
spiracles): greatest posterior width.
Apenesia forchhammeri Krombein,
NEw SPECIES
Figs. 1-13
Kieffer (1904, 1910b) described six Af-
rican species from females, none of which
is similar to forchhammeri. Kieffer’s species
are either glossy and impunctate, or sparsely
VOLUME 91, NUMBER 4 621
Figs. 1-8. Female Apenesia forchhammeri Krombein, paratype. 1, Dorsum of head, 80x; 2, head, lateral,
80x; 3, eye, 535%; 4, palpi, 870 x; 5, mandible, 270 x; 6, terminal segment, maxillary palpus, 1140 x; 7, dorsum
of thorax, 80; 8, apex of propodeum and base of abdomen, 80.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
1.0mm
13b
Figs. 9-13. Male Apenesia forchhammeri Krombein, paratype. 9, Dorsum of head, 35 x: 10, mandible. 100~:
11, dorsum of thorax, 35 x; 12, propodeum and base of abdomen, 35x: 13, genitalia (a), and (b) sixth sternum
and subgenital plate, dorsal aspect at left, ventral at night.
VOLUME 91, NUMBER 4
punctate and delicately alutaceous. The fe-
male of forchhammeri is quite densely
punctate on the head, more moderately so
on the thorax, and the integument of the
head and thorax is strongly alutaceous.
Holotype female.—Length 2.7 mm, LH
0.6 mm, LT 0.9 mm. Head and thorax pre-
dominantly ferruginous, narrowly infuscat-
ed above coxae, apex of mandible dark, an-
tenna light red except flagellum testaceous
beneath, legs pale testaceous, abdominal
petiole black, rest of abdomen light red ex-
cept large infuscated blotch on first tergum
and narrow infuscated band at base of sec-
ond and third terga. Vestiture pale, short,
erect and sparse.
Head (Figs. 1-2 of paratype) dull. strongly
alutaceous, moderately densely punctate,
many of punctures separated from each oth-
er by diameter of a puncture or less, LH
1.4 WH,; labial palpi 2-segmented, max-
illary palpi 3-segmented (Fig. 4), terminal
segment of latter (Fig. 6) with apical seta
1.3 x as long as segment; mandible (Fig. 5)
quadridentate; clypeus narrow, apical mar-
gin truncate medially, midline strongly car-
inate; eye (Fig. 3) dark, ovate, small, with
5-6 facets; scape about three times as long
as thick; flagellum about 2.7 as long as
scape, slightly clavate toward apex.
Thoracic dorsum (Fig. 7) strongly aluta-
ceous, more sparsely punctate than head;
pronotal disk 1.3 x as long as apical width;
mesonotum narrower than pronotum, about
0.7 as long as wide; mesopleuron with
moderately broad dorsal surface rounding
abruptly to broad lateral surface; propo-
deum 1.75 as long as maximum width,
minimum width 0.81 x maximum width,
propodeal formula 23:21:26; forefemur
2.6 as long as broad; mid tibia with weak
spines, hind tibia with setae only.
Abdomen shining, not alutaceous; petiole
(Fig. 8) composed of sternum only, stout
and short, about 1.6 x as wide as long.
Male.—Length 3.9-5.3 mm, forewing 2.6-
3.7 mm. Body shining except propodeum
rather dull from sculpturing; head black,
623
thorax black except pronotal collar some-
times light red, abdomen black, rarely
brown, last two segments light red; antenna,
mandible except apex, tegula, trochanters
and tarsi light red, coxae black to dark
brown, latter two pairs rarely light red, fem-
ora dark to light brown, occasionally red-
dish beneath, tibiae brown to light red.
Wings clear, stigma dark brown, veins light
brown to testaceous.
Head (Fig. 9) slightly longer than broad,
WH 0.83-0.95 x LH; mandible (Fig. 10) ro-
bust, quinquedentate; clypeus with median
apical margin broadly truncate, central ca-
rina rounded in profile, evanescent just be-
fore apical margin; WF 0.61-0.64 x WH and
1.22-1.47 x HE; front with relatively small,
shallow, dimpled punctures separated from
one another by 1.0-1.5 x diameter of punc-
ture; ocelli not enlarged, OOL 1.08-1.33 x
WOT, front angle about 90°; first four an-
tennal segments in a ratio approximately
20:7:11:9, third segment 1.75—2.00 x as long
as wide, pubescence very short, suberect,
glittering.
Thorax (Fig. | 1); pronotal disk with strong
anterior carina, posterior half shallowly de-
pressed, surface sculpture variable, ranging
from sparsely punctate, more densely so lat-
erally on anterior half in smaller specimens
to rather densely punctate on anterior half
in largest specimens, depressed posterior half
with closer, small punctures except apical
rim; notauli and parapsidal lines well de-
veloped, surface of scutum variably punc-
tate, relatively sparsely so in smaller spec-
imens and more densely in larger; scutellum
anteriorly with transverse groove, with
sparse punctures concentrated along ante-
rior groove and lateral margins of central
raised disk; metanotum with a small, shal-
low setose pit in middle, areas laterad of
this divided into a series of deeper, larger
foveae separated by longitudinal carinae;
propodeal disk 0.94= as long as wide,
rounding apically into posterior surface (Fig.
12), posterior carina lacking, lateral carina
weak, basal triangle irregularly rugose, areas
624
laterad of triangle irregularly rugulose; pos-
terior surface almost smooth in smallest
specimens, finely, closely, arcuately carin-
ulate in larger specimens; tarsal claws bi-
dentate apically, without an additional bas-
al tooth; forewing with costa extending
beyond stigma about 0.6 x length of stigma.
Abdomen with short petiole (Fig. 12)
composed of both tergum and sternum;
subgenital plate and sixth sternum (Fig. | 3b),
the former deeply and narrowly divided al-
most to base, very broad, lateral third weak-
ly sclerotized and doubled over to form a
pocket opening anteriorly on ventral sur-
face, sixth sternum with apical margin
broadly, shallowly emarginate; genitalia (Fig.
13a) with aedeagus moderately stout, the
parts closely consolidated.
Variation.— The two female paratypes are
2.5 and 2.6 mm long, and are very similar
to the holotype in all details. One topotypic
male, collected Dec 1987, is not included
in the type series. Its measurements fall
within the ratios listed above, and it agrees
with other males in essential details of punc-
tation, genitalia and subgenital plate, but
the antennae and last two abdominal seg-
ments are testacous, the mandibles and tarsi
are ivory, and the wing venation is white.
Type series. — Holotype: 2, BOTSWANA,
Serowe, Farmer’s Brigade, malaise trap, Dec
1987, Per Forchhammer. Paratypes: 3 2, 18
3, same label data except Feb (2 4), Dec (2)
1986, and Jan (6), May (8), Jul (4), Oct (2,
4), Nov (2, 6), Dec (11 4) 1987. [All USNM.]
A pair of paratypes will be deposited in the
British Museum (Natural History).
Etymology. —The species is named for Per
Forchhammer who has greatly enriched the
national collection by his collecting in Bo-
tswana.
Apenesia punctulata Krombein,
New NAME
Apenesia punctata Kieffer, 1904: 366 (2;
“Afrique occ., Mt. Camerun, 800-1200
m;” unique holotype in Genoa). — Kieffer,
1908: 25 (listed). — Kieffer, 1914: 393 (re-
described in German). Preoccupied by
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Epyris punctatus Cameron (1888: 174-
175), transferred to Apenesia by Evans
(1963: 280).
Remarks.—The coloration of punctulata
is somewhat similar to that of forchham-
meri, but the mandibles are bidentate in
females of the former species rather than
quadridentate, the pronotum is twice as long
as the mesonotum rather than thrice, and
the head and thorax are smooth and sparse-
ly punctate rather than strongly alutaceous
and rather densely punctate.
Etymology.—The name is the Latin di-
minutive of punctata.
Pristocera rhodesiae Turner
Figs. 14-21
Pristocera oriphila rhodesiae Turner, 1928:
142 (6; Sawmills, Southern Rhodesia; ho-
lotype now in South African Museum,
Capetown).
Pristocera rhodesiae Turner, Benoit, 1963:
50-51, fig. 81 (holotype redescribed).
Female.— Length 4.9 mm, LH 0.85 mm,
LT 1.40 mm, abdomen almost 4 its great-
est width. Body entirely light red, except
mandible with teeth and outer and inner
margins castaneous, eye black.
Head shining, not at all alutaceous, in
dorsal (Fig. 14) and lateral (Fig. 15) views;
mandible (Fig. 17) with three teeth; labial
palpi 3-segmented, maxillary palpi 6-seg-
mented; clypeus with median carina not
arched, ending in a small, rounded median
projection on apical margin on either side
of which the margin has a weak rounded
lobe with two tiny, short, rounded projec-
tions; LH 1.32 x WH, sides narrowing very
slightly nearly to posterior margin where
they converge more strongly to straight ver-
tex, punctures on front small and shallow,
quite sparse medially but more crowded an-
teriorly and laterally; eye (e, Fig. 15) small,
rounded, slightly longer than high, with
about eight facets, covered by a flat lens;
antenna thickening gradually toward apex,
segment 11 about 0.75 = as long as wide.
Thorax shining, not at all alutaceous, in
VOLUME 91, NUMBER 4
625
Figs. 14-17.
16, dorsum of thorax, 55 =; 17, mandible, 135.
dorsal view (Fig. 16); pronotal disk abruptly
declivous to collar, 2.13 = as long as wide,
with very sparse, shallow, small punctures;
mesonotum 0.75 x as long as wide, with a
few small lateral punctures; propodeum
2.44 as long as maximum width, 12.2 x
as long as minimum width, with a few small
punctures near lateral margin beyond con-
striction, propodeal formula 20:5:25:; fore-
femur flattened, 2.18 x as long as wide; mid
tbia with about 20 moderately stout spines.
Abdomen not petiolate, shining, with
sparse, subrecumbent setae laterally and
Female Pristocera rhodesiae Turner. 14, Dorsum of head, 55 =; 15, head, lateral (e = eye), 55 x;
apically on the segments, terga 1-5 also with
very tiny micropunctures separated from
each other by several times diameter of
puncture.
Male.— Length 5.1—7.5 mm, forewing 3.5-
5.0 mm. Black and shining; mandible ivory
to light red, except outer and inner margins
and teeth dark red; tegula and basal seg-
ments of legs dark brown; seventh abdom-
inal segment and occasionally apex of sixth,
tibiae and tarsi light red. Wings subhyaline,
veins light brown, stigma dark brown. Ves-
titure pale, glittering, suberect and moder-
626 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
7
g
x
N
‘
’
'
/
j
1.0mm
19, mandible, 50 x; 20, dorsum
21a
Figs. 18-21. Male Pristocera rhodesiae Turner. 18, Dorsum of head, 30 x;
of thorax, 30 x; 21, genitalia (a), and (b) sixth sternum and subgenital plate, dorsal aspect at left, ventral at right.
VOLUME 91, NUMBER 4
ate on head and thorax, decumbent and
sparse on sides and apices of abdominal seg-
ments; eyes bare.
Head (Fig. 18) about as wide as long, WH
0.98-1.07 x LH; mandible (Fig. 19) robust,
quinquedentate; clypeus with median cari-
na moderately arched in profile, apical mar-
gin broadly rounded; WF 1.59-1.67* HE
and 0.63-0.67 x WH; front with relatively
small, dimpled punctures that are some-
what sparser in smaller specimens; ocelli
not enlarged, OOL 0.70-1.00 x WOT, front
angle about 90°; first four antennal segments
in a ratio ranging from 13:3:7:7 to 17:4:11:
9, third segment 1.58—2.00 x as long as wide.
Thoracic dorsum (Fig. 20); pronotal disk
transversely carinate anteriorly, posterior
third depressed, anterior part with shallow
dimpled punctures that are denser toward
side; scutum with larger shallow punctures
that are denser on area between notauli and
parapsidal lines; scutellum anteriorly with
transverse groove, disk with small scattered
punctures; metanotum with a small median
pit anteriorly, area behind pit with fine,
dense piliferous punctures, laterad of pit a
larger fovea; propodeal disk about 0.8 = as
long as wide, rounding apically into poste-
rior surface, medially with a weak carina
reaching almost to apex, basally without a
well-defined triangular area, discal surface
with moderately close, irregular, transverse
rugulae that are sparser in smaller speci-
mens, lateral carina weak; posterior pro-
podeal surface lacking median carina, with
fine, close, transverse, arcuate rugulae that
are sparse or evanescent in smaller speci-
mens; tarsal claw bidentate at apex, inner
tooth subparallel to outer, but much shorter
and truncate apically, base of claw with small
tooth; costa of forewing not extending be-
yond stigma.
Abdomen not petiolate; subgenital plate
and sixth sternum (Fig. 21b), the former
deeply and narrowly divided almost to base,
lobes broadly rounded at apex, sixth ster-
num with a weak rounded median lobe on
apical margin; genitalia (Fig. 21a) with ae-
627
deagus relatively slender, the parts closely
consolidated.
Specimens examined.—2, 10 6, BOT-
SWANA, Serowe, Farmer’s Brigade, ma-
laise trap, Per Forchhammer, Sep (2, 5 4)
and Nov (é) 1986, Jan (3 6) and Dec 1987
(3).
Prosapenesia Kieffer, 1910
Benoit (1981) established the tribe Usa-
kosiini for Prosapenesia, Usakosia Kieffer,
1914, and Neusakosia Benoit, 1981. The
tribe was based essentially on the confor-
mation of the seventh abdominal sternum
(subgenital plate) of the male (Fig. 29b),
which has the subtriangular section between
the apical lateral lobes angled abruptly up-
ward, 1.e. toward the seventh tergum, as
contrasted to the broadly and slightly con-
vex surface of that sclerite in other Pristo-
cerinae.
The fortuitous circumstance of a wingless
female being carried into a Malaise trap dur-
ing her mating flight provides an opportu-
nity to describe the female of Prosapenesia.
It shares many characters including aptery
with the relatively few known pristocerine
females. The eyes are entirely lacking,
whereas other pristocerine females have
several to many well differentiated facets.
Evans noted (1964: 63) that the eyes of
Pseudisobrachium are reduced to a single
facet that is sometimes indistinct. The only
other differentiating character of female
Prosapenesia is that the median clypeal ca-
rina terminates at a narrow, truncate, thick-
ened apical lobe (see L, Fig. 22). These two
characters are too trivial to justify retention
of the tribe Usakosiini.
Prosapenesia lacteipennis Kieffer
Figs. 22-29
Prosapenesia lacteipennis Kieffer, 19 10a: 43
(6; Namibia; unique holotype in Ber-
lin).— Kieffer, 1914: 424 (6; rede-
scribed).— Benoit, 1981: 835-836, figs. 1
a—c (6; Namibia; redescribed).
Female.—Length 3.0 mm, LH 0.6 mm,
628 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 22-24. Female Prosapenesia lacteipennis Kieffer. 22, Frontal aspect of head (L = clypeal lobe), 220;
23, dorsum of head, 100 =; 24, dorsum of thorax, 100.
VOLUME 91, NUMBER 4
LT 1.2 mm. Head, mesopleuron and ex-
treme base of abdomen very dark brown;
dorsum of thorax and prosternum beneath
medium brown; sides of pronotum and pro-
sternum ferruginous; mandible except apex,
clypeus, antennae, legs and abdomen light
red. Vestiture pale, erect to suberect, sparse
and short.
Head (Fig. 23) slightly longer than wide,
WH 0.95 LH, moderately alutaceous,
rather shiny, with small punctures most of
which are separated from each other by 2-
3 times diameter of a puncture; mandible
(Fig. 22) bidentate, lower tooth more acute
and longer: clypeus narrow, with median
longitudinal carina that is straight in profile
and terminates at a narrow, truncate, thick-
ened apical lobe (L, Fig. 22), lower margin
of truncation slightly and shallowly emar-
ginate; eyes completely lacking; antenna 1 3-
segmented, scape relatively slender, 4.2 x as
long as wide, flagellum 2.3 x as long as scape,
not enlarged toward apex, segments 6-10
equally wide and 0.82 as long as wide.
Thoracic dorsum (Fig. 24) moderately
alutaceous, punctures smaller and sparser
than on head; pronotal disk slightly longer
than apical width, not carinate anteriorly,
abruptly declivous to collar; mesonotum at
base narrower than pronotum, sides con-
verging strongly toward apex, |.16 = as long
as basal width; mesopleuron with large dor-
sal area rounding gradually to lateral sur-
face; propodeum more delicately aluta-
ceous, 1.95 as long as greatest width,
minimum width 0.35 maximum width,
propodeal formula 9:7:20; forefemur rather
slender, 3.0 as long as wide; mid tibia
strongly spinose, hind tibia with setae only.
Abdomen not petiolate, shiny not aluta-
ceous, about 2.6 as long as wide.
Male. —Length 3.7-6.5 mm, forewing 2.5-
4.4 mm. Smallest specimens with integu-
ment mostly dark brown except legs lighter
brown, mandible except teeth and upper and
lower margins and clypeus light red; largest
specimens with integument predominantly
light red except apex and margins of man-
629
dible and extreme base of abdomen dark
brown, scutum, metanotum, propodeum,
mesosternum and lateral blotches on mid-
dle of second to seventh abdominal seg-
ments light reddish brown. Wings with a
slight milky cast, stigma dark to light brown,
veins light brown to testaceous. Vestiture
sparse, pale, short, suberect.
Head (Fig. 25) slightly broader than long,
WH 1.12-1.17= LH, posterior margin
straight in smaller specimens, angularly in-
dented in larger specimens, front in middle
with bilobed process projecting above and
between antennal insertions, weakly to
strongly grooved along midline nearly to an-
terior ocellus; mandible (Fig. 26) robust,
quinquedentate; clypeus narrow, apical
margin broadly and shallowly arched in-
ward on median third, surface slightly con-
cave; WF 1.54-2.13 x HE and 0.61-0.66 x
WH, with small, scattered punctures that
are quite shallow in smaller specimens, be-
coming denser and deeper in larger speci-
mens; eye slightly protuberant, EV 1.23-
1.79 HE, ocular setae lacking; ocelli not
enlarged, placed well forward on head, OOL
0.70-0.86 x WOT, front angle of triangle
about 110°; first four antennal segments in
a ratio ranging from 18:6:8:9 to 36:9:19:19,
third antennal segment 1.50-2.38 « as long
as wide; pubescence very short, dense and
erect:
Thorax (Figs. 27, 28); pronotal disk
abruptly declivous to collar, anteriorly with
transverse, erect lamella that is narrowed
toward middle, disk with scattered fine
punctures, posterior half depressed; scutum
with fine scattered punctures, notauli and
parapsidal lines well developed; scutellum
with anterior transverse groove and a few
fine punctures; metanotum with small, shal-
low pit in middle, laterally with small fo-
veae; propodeal dorsum rounding into pos-
terior and lateral surfaces, median and lateral
carinae weak, anteriorly usually with weak,
close, longitudinal rugulae beyond which
surface with close transverse rugulae; pos-
terior surface with close arcuate rugulae; lat-
630 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Figs. 25-29. Male Prosapenesia lacteipennis Kieffer. 25, Dorsum of head, 40x; 26, mandible, 85x; 27,
dorsum of thorax, 40 x; 28, lateral aspect of thorax, 40 x; 29, genitalia (a), and subgenital plate (b), dorsal aspect
at left, ventral at right.
VOLUME 91, NUMBER 4
eral surface with more separated, sinuous
rugulae.
Abdomen not petiolate; seventh tergum
broadly rounded at apex; subgenital plate
(Fig. 29b) bilobate at apex, subtriangular
section between lobes folded inward toward
tergum; genitalia (Fig. 29a) with aedeagus
massive, parts closely consolidated, para-
mere subtruncate to slightly rounded at apex,
deeply concave ventrally in middle for re-
ception of digitus and apex of cuspis.
Variation. — The forewing length is a more
reliable measure of size than total body
length owing to the variable degree of tele-
scoping of the posterior abdominal seg-
ments. When the series of /acteipennis males
is arranged in order of increasing forewing
length, two significant differences are ap-
parent. The smaller specimens, 3.3 mm or
less in forewing length, include the darkest
specimens, and the larger, 3.4 mm or longer,
have gradually increasing amounts of light
reddish brown to red integument.
The shape of the head also changes with
increasing size. The posterior margin of the
head is straight in the smallest specimens
and the median, bilobate frontal process
above and between the antennal insertions
projects rather weakly. At about 3.0 mm
forewing length the posterior margin of the
head becomes angularly indented in the
middle and the frontal process projects more
strongly (Fig. 25), both of these characters
becoming more accentuated with increasing
size.
Specimens examined.—?, 19 6, BOT-
SWANA, Serowe, Farmer’s Brigade, mal-
aise trap, Per Forchhammer, Sep (¢) and
Nov (2, 10 4) 1986, Jan (2), Feb (2 4), Apr
(4 6) and Nov (é) 1987.
631
ACKNOWLEDGMENTS
I am grateful to Beth Norden, Depart-
ment of Entomology, for skillful prepara-
tion of specimens for SEM study, and to
Susann Braden, Scanning Electron Micro-
scope Laboratory, for making the micro-
graphs from uncoated specimens. I thank
George Venable, Department of Entomol-
ogy, for making the line drawings, and for
mounting the plates.
LITERATURE CITED
Benoit, P. L. G. 1963. Monographie des Bethylidae
d’Afrique Noire (Hymenoptera). I, Sous-famille
Pristocerinae: Tribu Dicrogeniini; Tribu Pristo-
cerini, Gen. Pristocera Klug. Ann. Mus. Roy. Afr.
Centr., Sci. 119: 1-95.
1981. Bethylidae africains (Hymenoptera).
La Tribu Usakosiini. Rev. Zool. Afr. 95: 833-842.
Cameron, P. 1888. Descriptions of twenty-three new
species of Hymenoptera. Mem. Manch. Lit. & Phil.
Soc. (4) 1: 159-183.
Evans, H. E. 1963. A revision of the genus Apenesia
in the Americas (Hymenoptera, Bethylidae). Bull.
Mus. Compar. Zool. 130: 249-359.
. 1964. Asynopsis of the American Bethylidae
(Hymenoptera, Aculeata). Bull. Mus. Compar.
Zool. 132: 1-222.
Kieffer, J.-J. 1904. Description de nouveaux Dryi-
ninae et Bethylinae du Musée de Génes. Ann. Mus.
Civ. Stor. Nat. Genova 41: 351-412.
1908. Hymenoptera, Family
Genera Insectorum 76: 1-50.
1910a. Description de nouveaux Béthylides
(Hymeén.). Ann. Soc. Ent. France. 79: 31-56.
1910b. Serphidae, Cynipidae, Chalcididae,
Evaniidae und Stephanidae aus Aquatorialafrika.
Wiss. Ergeb. Deut. Zent.-Afr.-Exped. 1907-1908.
3, Zool. I: 91-119.
1914. Bethylidae. Das Tierreich. 41: 1-595.
Turner, R. E. 1928. New Hymenoptera of the Family
Bethylidae. Ann. & Mag. Nat. Hist. (10) 1: 129-
152.
Bethylidae.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 632-633
NORTH AMERICAN SPECIES OF STILBOPS
(HYMENOPTERA: ICHNEUMONIDAE)
HENRY K. TOWNES
American Entomological Institute, 3005 SW 56th Avenue, Gainesville, Florida 32608
Abstract.—The American species of the ichneumonid genus Sti/bops are described:
Stilbops latibasis, new species, from eastern North America and S. mexicana, new species,
from Mexico. The other species of the genus occur in Eurasia, Japan, Taiwan, and the
Philippines.
Key Words: wasps, Stilbopinae, parasite
There are two species of Sti/bops in North
America, both undescribed. This genus is
the only North American representative of
the subfamily Stilbopinae. A key identifying
the genus and a habitus figure are in Townes,
1970. Mem. Amer. Ent. Inst. 13, pages 5,
7, and 196.
Genus Silbops
Stilbops Foerster, 1869. Verh. Naturh. Ver.
Rheinlande 25: 163. Type: Pimpla vetula
Gravenhorst. Designated by Ashmead,
1900.
Aphanoroptrum Foerster, 1869. Verh. Na-
turh. Ver. Rheinlande 25: 168. Type:
(Lissonota rugicornis Gravenhorst) = ab-
dominalis Gravenhorst. Included by
Thomson, 1877.
Aphanoroptra Thomson, 1877. Opuscula
entomologica 8:736. Emendation.
Aphanorrhoptrum Dalla Torre, 1901. Ca-
talogus hymenopterorum... 3: 528.
Emendation.
Eritrachynus Schmiedeknecht, 1913. Opus-
cula ichneumonologica 5: 2709. Type:
Eritrachynus asper Schmiedeknecht. In-
cluded by Schmiedeknecht, thidem, p.
2724.
The genus Sti/bops is mostly Holarctic.
There are five described species in Europe
(Hinz 1981), a moderate number in Russia,
and ten undescribed species in other re-
gions. Five of the undescribed species are
from Japan, two from Taiwan, one from the
Philippines, one from Mexico, and one from
eastern North America. The latter two are
described below. The Stilbopinae were first
recognized as a separate subfamily by Short
(1957).
Two of the European species are known
to parasitize Adelidae (Lepidoptera), ovi-
positing into adelid eggs and emerging from
cocoons of the mature larvae (Hinz 1981).
Adelidae are presumed to be the hosts of
the entire genus.
Key TO THE NORTH AMERICAN
SPECIES OF S'TILBOPS
1. Underside of antenna stramineous or light
brown. Tergites 6-9 black in both sexes. Ter-
gites 2 and 3 of female mostly to entirely fer-
ruginous, black in male. Northeastern North
America. 1. /atibasis, new species
. Underside of antenna black. Tergites 6-9 black
in male, fulvous in female. Tergites 2 and 3
black in both sexes. Mexico. 2. mexicana, new
species
to
1. Stilbops latibasis, NEW SPECIES
Front wing 3.4 to 3.8 mm long. Body
moderately stout. Cheek about 0.1 as long
as basal width of mandible. Genal carina
VOLUME 91, NUMBER 4
weak, gradually curved inward to reach oral
carina about 1.1 as far above base of man-
dible as basal width of mandible. Clypeus
with basal 0.4 convex, the apical 0.6 weakly
concave. Flagellum about 0.68 as long as
front wing, with 17 segments, the last few
segments enlarged and the last segment quite
large. Epomia rather weak, evanescent dor-
sad. Mesopleurum polished, with moderate
sized punctures that are separated by about
0.8 their diameter. Propodeal areolation
complete and strong. Second lateral area of
propodeum polished, with moderate sized
punctures that are separated by about 0.8
their diameter. Areolet small, obliquely tr-
angular, sometimes absent. First tergite
stout, 1.4 as long as wide in male, 1.2 as
long as wide in female, its median dorsal
carinae strong on basal 0.7 and dorsolateral
carina sharp from base to apex. Second ter-
gite 0.88 as long as wide in male, 0.68 as
long as wide in female, with a preapical shal-
low transverse impression, its surface mod-
erately mat, with moderately large punc-
tures that are separated by about 0.5 their
diameter. Ovipositor sheath about 0.58 as
long as hind tibia.
Black. Mandible yellow. Palpi and under
side of antenna stramineous or light brown.
Tegula, front and middle coxae, and all tro-
chanters pale yellow, the coxae and hind
trochanters usually stained with fulvous
dorsally. Legs beyond trochanters light or
pale fulvous, the last segment of front and
middle tarsi light brown. Hind tibia weakly
infuscate apicad, and hind tarsus fulvous
brown. Wings faintly infuscate. Male ab-
domen black with apical margin of tergites
2 and 3 fulvous and apical margin of tergite
4 obscurely fulvous. Female abdomen with
first tergite, black with apical 0.3 to 0.4 often
ferruginous. Tergites 2-4 or 2-5 ferruginous
or sometimes partly infuscate, and tergites
5-8 or 5-9 blackish.
Type: Female, collected under trees on a
moist bank at edge of a marsh, E. S. George
633
Reserve, near Pinckney, Mich., June 24,
1956, H. Townes (AE]I).
Paratypes: Male, Crawford Notch Road,
6 miles south of Gorham, N.H., May 27,
1965, G. S. Walley (CNC). Female, Ring-
wood (near Ithaca), N.Y., June 7, 1951, C.
Dasch (Dasch collection). Female, summit
of King Mt., 1150 ft., Old Chelsea, Que.,
July 12, 1965, Malaise trap (CNC). Female,
Plummers Island, Md., June 6, 1911, H. S.
Barber (USNM).
2. Stilbops mexicana, NEw SPECIES
Front wing 3.8 to 4.3 mm long. Structure
similar to that of S. /atibasis except as fol-
lows: Propodeal carinae in female weak,
evanescent. Areolet present, narrowly trun-
cate above. First tergite 1.9 as long as wide
in male, 1.3 as long as wide in female, its
median dorsal carinae reaching a little past
spiracle in male, reaching 0.8 the distance
to spiracle in female.
Black, including under side of antenna.
Mandible, palpi, tegula, and front and mid-
dle coxae and trochanters pale yellow, the
middle coxae of female tinged with fulvous
basad. Front and middle femora and tibiae
fulvous, their tarsi brown. Hind coxa black.
Hind trochanters and tibiae of male dark
brown. Hind femur and tarsus of male
blackish. Hind leg of female beyond coxa
brownish fulvous, its tarsus brown. Tergites
6-9 of female fulvous, of male black.
Type: Female, kilometer 185.5 on Tux-
tepec to Oxaca road, Mexico, 2200 m, Oct.
22, 1962, H. & M. Townes (AEI).
Paratypes: 2 males, same data as type
(AED).
LITERATURE CITED
Hinz, R. 1981. Die europaischen Arten der Gattung
Stlbops Foérster (Hymenoptera, Ichneumonidae).
Nachrichtenblatt Bayerischen Ent. 30: 60-64.
Short, J. R. T., 1957. On the final instar larva of
Stilbops (Aphanoroptrum) abdominale (Grav.)
Hymenoptera: Ichneumonidae, Proc. R. Ent. Soc.
London (B) 26: 175-176.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, p. 634
NOTE
First Record of Thripobius semiluteus
(Hymenoptera: Eulophidae) from the New World
Thripobius semiluteus Boucek (Hyme-
noptera: Eulophidae) was described from
Africa (S40 Tomé) and India (Mysore)
(Boucek 1976, Entomophaga, 21: 401-414),
and it has subsequently been recorded from
Australia (New South Wales) (Boucek 1988,
Australasian Chalcidoidea (Hymenoptera),
CAB International, 832 pp.). As a result of
ongoing studies on the biological control of
greenhouse thrips, Heliothrips haemorrhoi-
dalis(Bouché), by one of the authors
(McMurtry), 7. semi/uteus was recently col-
lected in Brazil. This represents the first New
World record for this parasite. Collection
data is: BRAZIL, Minas Gerais, Lavras,
12.v.1988, J. A. McMurtry, ex. Heliothrips
sp. on Croton sp. (Euphorbiaceae) and av-
ocado (Persea americana: Lauraceae). This
species was imported into the University of
California, Riverside Quarantine facility
(S&R# 88-1 1-1). It is being propagated in
the laboratory on H. haemorrhoidalis on
citrus and investigated as a possible biolog-
ical control agent against this thrips in av-
ocado and citrus orchards.
T. semiluteus is known as a parasite of
Thripidae in the subfamily Panchaetothri-
pinae. Recorded hosts are Brachyurothrips
anomalus Bagnall, Panchaetothrips indicus
Bagnall, and H. haemorrhoidalis; known
host plants for this parasite are Hibiscus,
Croton, Liquid Amber, and Valencia or-
anges (Boucek 1976, loc. cit; 1979, loc. cit.).
Avocado represents a previously unrecord-
ed host plant for the parasite.
Parasite identification was done by La-
Salle. Specimens are in the collection at UC
Riverside, and exemplars have been placed
in the U.S. National Museum, Canadian
National Collection, and British Museum
(Natural History).
John LaSalle* and James A. McMurtry,
Department of Entomology, University of
California, Riverside, California 92521.
*Current address: CAB International, In-
stitute of Entomology, 56 Queen’s Gate,
London, SW7 SJR.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 635-636
NOTE
Hymenoptera Associated with a California Population
of the Russian Thistle Biological Control Agent
Coleophora klimeschiella Toll
(Lepidoptera: Coleophoridae)
Coleophora klimeschiella Toll, whose lar-
vae are casebearers which feed on the foliage
of the weed pest Russian thistle (Sa/sola
australis R. Brown), is native to Turkey and
probably the Soviet Union (Khan and Bal-
och, 1976. Entomophaga 21: 425-428). This
moth was imported into U.S. quarantine
from Pakistan in 1975 and tested for host
specificity. In 1977, it was released in Cal-
ifornia to control Russian thistle (Hawks
and Mayfield, 1978. Environ. Entomol. 7(2):
257-261).
Several years later at a C. klimeschiella
release site two miles northwest of Coalinga
(Fresno County) California, several species
of native Hymenoptera were discovered to
be parasitizing the C. klimeschiella case-
bearers. The C. kAlimeschiella casebearers
were gathered, the wasps reared, identified,
and the rates of parasitism established.
Coleophora klimeschiella casebearers were
collected for rearing on six dates (Table 1).
By rearing C. Alimeschiella individually in
gelatin capsules and dissecting their cases
after parasitoids emerged, parasitization
rates and the C. klimeshciella stage killed
were established.
Five species of Hymenoptera were reared
from C. klimeschiella: Macroneura n.sp.
Walker (Eupelmidae) from larvae and pu-
pae, Agathis gibbosa (Say) (Braconidae) from
larvae and pupae, Spilochalcis side (Walker)
from pupae, Spilochalcis torvina (Cresson)
from pupae (both Chalcididae), and Cato-
laccus aeneoviridis (Girault) (Pteromali-
dae). The stage which C. aeneoviridis killed
was not determined, but it is recorded from
many Lepidoptera and Coleoptera and as a
secondary parasitoid of braconid, ichneu-
monid, and bethylid wasps (Burks, In
Krombein et al., 1979, Catalog of Hyme-
noptera in America North of Mexico, Vol.
I (Parasitica), Smith. Inst. Press: 806).
Hence, this wasp is possibly a primary para-
sitoid of C. klimeschiella or a secondary
parasitoid of 4. gibbosa.
Four additional wasp species were reared
in sleeve cages containing Russian thistle
and C. klimeschiella casebearers: Halti-
chella rhyacionia Gahan (Chalcididae),
Epipteromalus n.sp. Ashmead, Norbanus
perplexus Ashmead, and Pachyneuron sp.
Walker (all Pteromalidae). These species
were possible parasitoids of C. klimeschiel-
la or from undetected hosts on or in the
Russian thistle. A female specimen of H.
rhyacionia (Coalinga, 2-X-1985, H. Misthe,
reared from Coleophora klimeschiella in
Salsola australis, em. 31-X-1985) in the
University of California Riverside collec-
tion supports the former association. These
records of a known central and eastern
United States species (Burks 1979, ibid.:
860) in California 1s a significant range ex-
tension for this species. Norbanus perplexus
is reported from and in this case possibly
parasitized Coleophora parthenica Meyr-
ick, a stem burrowing Russian thistle feeder
introduced from the Mediterranian region
(Goeden, Ricker, and Muller, 1978. Envi-
ron. Entomol. 7(2): 294-296). This is the
first rearing information for the genus Epip-
teromalus. Solenopsis xyloni McCook (For-
micidae) was observed preying on C. k/i-
meschiella casebearers. In three of five
observations, two to four ants attacked a
single C. klimeschiella casebearer.
In all, 1523 C. klimeschiella and 453
636 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Table 1. Parasitism of Coleophora klimeschiella (CK) casebearers near Coalinga, California, 1982-1984.
Acronyms for hymenopterous parasitoids are AGGI—Agathis gibbosa (Braconidae); CAAE—Catolaccus aeneo-
viridis (Pteromalidae); MASP—Macroneura n. sp. (Eupelmidae); SPSI—Spilochalcis side (Chalcididae); and
SPTO-— Spilochalcis torvina (Chalcididae).
% Parasitism
No. CK Cases % CK
Coll. Date w/Emerg. Emerg MASP AGGI CAAE SPSI SPTO
V-26-82 587 79 20 <1 <1 0 <1
VI-10-82 177 52 47 <1 0 0 0
VI-23-82 374 90 7 <1 0 1 l
VILI-3-82 250 97 2 0 (0) <1 <1
VI-2-83 151 95 <1 4 0 0 0
V-8-84 437 56 5) 38 <1 0 0
Total 1976 77 13 9 <1 <i <1
parasitoids were reared to adult emergence
(Table 1). No insects emerged from many
cases. Parasitism (based on emergence) av-
eraged about 22 percent but ranged from 3
to 48 percent among collection dates (Table
1). Macroneura n.sp. and Agathis gibbosa
were most numerous; the other parasitoids
were rarely reared.
The wasps Macroneura sp., Cerambyco-
bius sp. (both Eupelmidae), Hockeria n.sp.
(Chalcididae), Catolaccus aeneoviridis (Gi-
rault) (Pteromalidae), an unidentified ich-
neumonid, and a coleopterous predator
Phyllobaenus atriplexus (Forster) (Cleridae)
were reared in a field colony of C. klimes-
chiella at Indio, California (Goeden et al.
1978). The Hockeria specimens belong to
a widely distributed, polyphagus species
currently in manuscript (Halstead, in prep.).
With the exception of C. aeneoviridis and
possibly Macroneura sp., the parasitiods that
I reared are not recorded from C. klimes-
chiella. However, some of these species are
known parasitoids of other Coleophora
species (Burks 1979, ibid; Carlson, In
Krombein et al., 1979. Catalog of Hyme-
noptera in America North of Mexico, Vol.
I, Smith. Inst. Press: 315-740). The Macro-
neura and P. atriplexus reared by Goeden
et al. (1978) are also known enemies of cec-
idomyid (Diptera) gall-makers on Atriplex
spp., a species which commonly inhabits
areas with Russian thistle, and hence was
thought to be the source of those insects.
Additional host or plant associations for the
C. klimeschiella parasitoids at the Coalinga
site are unknown.
The adaptation of native parasitoids was
possibly a reason for the poor establishment
and ineffectiveness of C. klimeschiella as a
biological control agent in southern Cali-
fornia (Goeden et al. 1978). The same ap-
pears to be occurring at the Coalinga site.
I thank E.E. Grissell and S.R. Shaw, both
Systematic Entomology Laboratory, ARS-
USDA; G. Gibson, Department of Ento-
mology, University of Alberta, Edmonton;
G.J. Couch, Department of Entomology,
University of Massachusetts, Amherst; and
M.S. Wasbauer, Analysis and Identifica-
tion, California Department of Food and
Agriculture (CDFA), Sacramento, for iden-
tifying the parasitoids and predators. I thank
also D.J. Burdick, California State Univer-
sity Fresno, and B.E. Valentine, Kings Riv-
er Conservation District, Fresno, California
for comments on this paper and B. Villegas,
T.E. Esser, and G. Luna, Biological Control
Services Program (BCSP), CDFA, Sacra-
mento, for field assistance and editorial
comments. I thank also the BCSP-CDFA,
Sacramento for permission to publish this
information. The author was employed by
the BCSP during the study.
Jeffrey A. Halstead, 1/0 W. Barstow #112,
Fresno, California 93704.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 637-639
NOTE
Taxonomic Status of Mantispa sayi, Mantispa fuscicornis,
and Mantispa uhleri (Neuroptera: Mantispidae)
The North American mantispines Man-
tispa sayi Banks (1897. Trans. Am. Ento-
mol. Soc. 24: 21-31), Mantispa fuscicornis
Banks (1911. Trans. Am. Entomol. Soc. 37:
335-366), and Mantispa uhleri Banks (1943.
Psyche 50: 74-81) are currently considered
sibling species (Macleod, in Hughes-
Schrader, 1979. Chromosoma 75: 1-17),
even though recent investigators have found
them to be essentially indistinguishable
both karyotypically and morphologically
(Hughes-Scrader 1979), (Redborg, 1982. J.
Arachnol. 10:92—93). There have been ex-
tensive biological studies on M. whleri(Red-
borg and MacLeod. 1985. Ill. Biol. Monogr.
53, 130 pp.) and natural history observa-
tions on M. fuscicornis (Gilbert and Rayor.
1983. J. Kans. Entomol. Soc. 56: 578-580),
(Rice, 1986. J. Kans. Entomol. Soc. 59: 121-
126), although no significant biological dif-
ferences were noted. All three species re-
portedly were reared in the laboratory, but
only data on M uhleri was published (Red-
borg and MacLeod 1985). I recently ex-
amined the type specimens of these species
in the Museum of Comparative Zoology
(MCZ), Harvard University, as part of an
ongoing revision of the New World Man-
tispinae. A reevaluation of the taxonomic
status of these species was prompted by the
examination of these types and of numerous
other specimens from Arizona, Arkansas,
Connecticut, Florida, Georgia, Illinois,
Kansas, Maryland, Minnesota, Mississippi,
Missouri, Nebraska, New Mexico, Ohio,
Oklahoma, South Carolina, Texas, Utah,
and Mexico.
The original description of M. sayi was
based on one specimen from Texas and two
from Lake Worth, Florida, coll. Mrs. Slos-
son. The only type specimen of M. sayi in
the MCZ is a male with the following ver-
batim label data: “Type,” “Brazos Co.,
Tex.,’’ “Collection N. Banks,’’ ‘‘Type
10767,” “type Mantispa sayi Bks.”’ Man-
tispa fuscicornis was described from two
specimens collected in Florida, one from
Kissimmee and another from Lake Worth,
coll. Mrs. A. T. Slosson. Both type speci-
mens of M. Fuscicornis are present in the
MCZ. There is a male with the label data
“Kissimmee Fla,” “Collection N. Banks,”
“M.C.Z. Type 10769,” “type Mantispa fus-
cicornis Bks,” and a female with the label
data “type,” “L. Worth, Fla,” “Collection
N. Banks,” “M.C.Z. Type 10769.” The ab-
sence of the other two M. sayi syntypes and
the similarity of their collection data with
those of the female syntype of M. fuscicornis
suggest that this specimen was originally one
of the type specimens of M. sayi. While the
lack of a 10767 type label of the M. fusci-
cornis female might argue against such a
conclusion, the numbers currently assigned
to the type specimens of both MM. sayi and
M. fuscicornis were most likely assigned well
after the original descriptions of both species.
All specimens of a given type series at the
MCZ are assigned identical type numbers,
but prior to 1920 Banks did not designate
type numbers in his species description of
mantispids. The type numbers (10767,
10769) assigned to the two species in ques-
tion are very close to the type numbers
(10758-10762, 10770) currently assigned to
the type specimens of species described by
Banks in 1913 (Trans. Am. Entomol. Soc.
39: 201-242), as well as to the type numbers
(10774-10778) designated by Banks in his
descriptions of mantispid species in 1920
(Bull. Mus. Comp. Zool. 64: 299-362).
Therefore, the specimen from Lake Worth
would not be expected to bear a M. sayi
type number even if it were part of that type
638
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
0.5 mm
igs, 172%
Mantispa sayi. Color patterns on heads of male specimens collected on 20 July 1968 at Corpus
Christi Lake State Park, San Patricio Co., Texas. Anterior view.
series. Because of the uncertain status of the
specimen from Lake Worth, I hereby des-
ignate the syntype male of M. sayi from
Brazos Co., Texas, as the lectotype of M.
sayi and the missing specimens from Lake
Worth, Florida, as paralectotypes. In ad-
dition, the male of M. fuscicornis from Kis-
simmee, Florida, 1s hereby designated the
lectotype of M. fuscicornis and the female
from Lake Worth, Florida, is designated as
the paralectotype.
Banks (1911) did not include any char-
acters to distinguish M. fuscicornis from M.
sayi, and noted only that M. fuscicornis was
similar to M. sayi in general structure. Red-
borg (1982. J. Arachnol. 10: 92-93) sub-
sequently used facial markings to separate
M. fuscicornis from both M. sayi and M.
uhleri, stating that only M. fuscicornis pos-
sessed a broad, inverted Y-shaped line on
the frons, with the forks of the Y looping
under the antennal sockets. I have found
that the face markings of M. sayi vary con-
siderably (Figs. 1, 2) and that the range of
variation encompasses what I assume are
the facial markings attributed both to M.
fuscicornis (Fig. 1) and to the other two
species. (Fig. 2). Examinations of genitalic
and other morphological characters of the
type specimens and of specimens from the
aforementioned locations has revealed no
basis for the continued separation of these
species. Therefore, I hereby consider Man-
tispa fuscicornis Banks to be a junior syn-
onym of Mantispa sayi Banks (NEW SYN-
ONYMY).
The holotype female of WZ. uhleri is lo-
cated in the MCZ along with four female
paratypes. In distinguishing this species from
the preceding two, Banks (1943) referred to
its almost entirely black abdomen and to
the upward extension of the dark face mark
to the vertex. I have found that the dark
brown to black markings on abdominal seg-
ments of M. sayi females are often more
extensive than those of males, causing fe-
male abdomens to look darker. Redborg and
MacLeod (1985) also noted this dimor-
phism for M. whleri. Therefore, the color
differences noted by Banks could be attrib-
uted to this sexual dimorphism if Banks was
comparing the female types of M. uhleri to
the male types of M. sayiand M. fuscicornis.
Also, the abdomen of the holotype of M.
uhleri is discolored, darkened, partially ad-
hered to the left hind wing, and missing both
the third abdominal tergite and all segments
posterior to the fifth. Therefore, the poor
condition of the holotype may have con-
tributed to Banks’ diagnosis if the present
condition of the holotype is essentially un-
changed from 1943. The difference in facial
markings mentioned by Banks has been dis-
cussed above. In addition, I examined first
VOLUME 91, NUMBER 4
instars obtained from M. sayi females col-
lected in Texas and could find no differences
between these specimens and the descrip-
tion of M. uwhleri first instars provided by
Redborg and MacLeod (1985). Therefore,
for essentially the same reasons given earlier
when designating MM. fuscicornis a junior
synonym of M. sayi, I consider Mantispa
uhleri Banks to be a junior synonym of
Mantispa sayi Banks (NEW SYNONY-
MY).
The synonomy of both M. fuscicornis and
M. uhleri with M. sayi results in the bio-
logical information on the two former
species being combined and attributed to
the latter. The clarification of the status of
these names at this time facilitates future
studies on the ecological characteristics of
M. sayiand allows meaningful comparisons
between life-history characteristics of this
species and other mantispines.
I am grateful to the following persons for
the opportunity to examine specimens un-
der their care: Robert W. Brooks (Snow En-
tomological Museum, University of Kan-
sas), Philip J. Clausen (University of
639
Minnesota), Lewis L. Deitz (North Carolina
State University), Herbert L. Dozier (Clem-
son University), David K. Faulkner (San
Diego Natural History Museum), Oliver S.
Flint, Jr. (National Museum of Natural His-
tory), Norman D. Penny (California Acad-
emy of Sciences), Marlin E. Rice (lowa State
University), Terence L. Schiefer (Mississip-
pi Entomological Museum), Scott R. Shaw
(MCZ), Cecil L. Smith (University of Geor-
gia), Lionel A. Stange (Florida State Collec-
tion of Arthropods), and Charles A. Triple-
horn (The Ohio State University). I also
thank Peter H. Adler and John C. Morse
(Clemson University) and one anonymous
reviewer for their helpful comments on this
manuscript. This study was funded in part
by an Ernst Mayr Grant from the MCZ.
This 1s Technical Contribution No. 2923 of
the South Carolina Agricultural Experiment
Station, Clemson University.
Kevin M. Hoffman, Department of Ento-
mology, Clemson University, Clemson,
South Carolina 29634-0365.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, p. 640
NOTE
Black Walnut, Juglans nigra: A New Host Record for
Colaspis favosa (Coleoptera: Chrysomelidae)
Colaspis favosa Say is a small, metallic-
green, eumolpine chrysomelid that 1s wide-
spread in the southeastern United States
(Blake. 1977. Proc. Entomol. Soc. Wash.
79: 209). Adults of C. favosa have been as-
sociated with several species of woody plants
including apple (Malus spp.), silverleaf scurf-
pea [Psoralea argophylla (Walt.) Cory.]
(Douglas. 1929. J. Kansas Entomol. Soc. 2:
10), wax myrtle (Myrica sp.) (Kirk. 1970.
South Carolina Agric. Exp. Stn. Tech. Bull.
1038, Part 2. p. 88; Chapin. 1979. Coleopts.
Bull. 33: 450), azalea (Rhododendron sp.)
(Loding. 1900. Geol. Surv. of Alabama
Monogr. 11: 1), blackberry (Rubus spp.), live
oak (Quercus virginiana Mill.) and pine (Pi-
nus spp.) (Kirk. op. cit.). Except for Loding’s
(op. cit.) mention of C. favosa as a pest of
azaleas, it is not clear whether other host
records for this insect represent 1) instances
in which C. favosa actually fed upon the
plant listed or 2) incidental captures on a
non-food plant (e.g. pine is a suspect record
given the preponderance of angiosperm rec-
ords for this apparently oligophagous in-
sect).
During early June 1988, I collected adults
of C. favosa from black walnut (Juglans ni-
gra L.) trees growing in a small grove in
Blacksburg, Montgomery County, Virginia.
I observed beetles feeding on black walnut
foliage both on the tree and when confined
in petri dishes with fresh leaflets. Feeding
was restricted to leaflet margins and damage
to trees was negligible since beetles were
present in low numbers. This is the first
record of C. favosa feeding upon foliage of
black walnut. Host plant and insect voucher
specimens have been deposited in the her-
barium and insect museum at Virginia
Polytechnic Institute and State University,
Blacksburg.
Reported hosts of C. favosa (e.g. Myrica
sp. and Rhododendron sp.) were absent from
the vicinity of the black walnut grove from
which I collected beetles (the grove is sur-
rounded by several kilometers of agricul-
tural land), suggesting that the use of black
walnut by this species was not just a tem-
porary shift from a preferred primary host
to an adjacent secondary host. Rather it is
possible that black walnut may be a signif-
icant but overlooked host plant for C. favosa
in the southern Appalachian Mountains,
particularly since plants of the genus Myrica
[apparently important coastal plain hosts
(Chapin op. cit.)] are absent from this region
(Harvill et al. 1986. Atlas of the Virginia
Flora. p. 97). Interestingly, the Myricaceae
and the Juglandaceae are placed into the
order Juglandales by some authors (Benson.
1959. Plant Classification. p. 316) suggest-
ing a close relationship between these two
plant families.
I thank Dr. M. C. Thomas of the Florida
Department of Agriculture, Florida State
Collection of Arthropods, Gainesville, Flor-
ida, for identifying Colaspis favosa. K. Wil-
liams and an anonymous reviewer provided
useful comments on the manuscript.
Charles E. Williams, Department of Bi-
ology, Washington and Lee University, Lex-
ington, Virginia 24450.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 641-644
Book REVIEW
The Guild Handbook of Scientific Illus-
tration. Edited by Elaine R. S. Hodges,
with Lawrence B. Isham, Marsha E. Jes-
sup, and G. Robert Lewis. Van Nostrand
Reinhold. 1989, xv + 575 pp., 631 black-
and-white and halftone images, 36 color
images, 3 charts. Library of Congress Cat-
alogue Card Number (LCCCN) 88-10599,
International Standard Book Number
(ISBN) 0-442-23681-6, hardbound, 8!
x 11°’ matcoated stock, $85.95 US.
After approximately 14 years since its in-
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Illustration evolved into the most ency-
clopedic volume on scientific illustration
available. The text is in a 3-column format
and is written in an easily readable style
throughout, a characteristic not found in
most multi-authored volumes, regardless of
subject. The numerous black-and-white,
halftone, and color images represent not only
“fine art,” but conceptualize information
found in the text. The appendix includes
sources for most tools and equipment men-
tioned in the text, available degree pro-
grams, professional organizations, and
sources of business information. A bibli-
ography arranged by chapter rounds out this
comprehensive work.
The Guild Handbook of Scientific Illus-
tration emphasizes the fact that scientific
illustration is more than the rendering of
attractive images, and that scientific illus-
trators must possess (1) knowledge of their
subjects and of the conventions within the
area of science for which they work, (2) abil-
ity to distinguish between important details
and those that are anomalous, (3) command
of basic rendering skills so that the most
appropriate technique can be chosen for a
specific subject to ensure optimal reproduc-
tive results, and (4) knowledge of current
copyright laws and general business opera-
tions and practices. The volume is divided
into five parts: Basics, Rendering Tech-
niques, Subject Matter, Beyond Basics, and
The Business of Scientific Illustration. Each
part details one of the knowledge and skill
prerequisites necessary for all scientific 1l-
lustrators.
Part 1, Basics (Chapters 1-4), discusses
the roles and responsibilities of the scientific
illustrator and the scientist/client. Protocols
from the initial meeting, through the de-
tailed preliminary drawing and final ren-
dering, to storage of originals are examined
in detail. The relationship between scientific
illustrator and scientist/client is described
as dynamic, and vital to the scientific ac-
curacy and aesthetics of the final product.
The artist relies on the client for the subject
matter and information about what must be
represented, while the client relies on the
skills of the artist to convey the character-
istics that are diagnostic. Recommenda-
tions on tools and materials that scientific
illustrators use from day-to-day are exam-
ined with insights on surfaces, media, draw-
ing tools, mounting materials, measuring
devices, and general drawing procedures.
Because scientific illustration is important
as a permanent record, archival consider-
ations pertaining to the durability and per-
manence of surfaces and media, as well as
storage equipment, are discussed in detail.
These features are not treated in other ref-
erences I examined (see References below).
The section closes with an illustrative pre-
sentation on different types of lighting and
tips on how to depict shadows, transpar-
ency, iridescence, and highlights.
Part 2, Rendering Techniques (Chapters
5-11), introduces the reader to various kinds
of rendering techniques using line-and-
ink, pencil, coquille board, carbon dust,
watercolor-and-wash, gouache-and-acrylics,
scratchboard, and airbrush. Each technique
642
is presented in a step-by-step illustrative
fashion and followed by a discussion of the
technique. Included in this section are vari-
ations and modifications of basic rendering
techniques derived from the experiences of
the authors. In addition, materials, media,
and tools used for each technique are
presented along with the author’s encour-
agement to experiment with a variety of
mixed-media techniques, e.g. wet-with-wet,
dry-with-dry, and wet-with-dry media on
various surfaces, because only through ex-
perimentation are innovations made.
Part 3, Subject Matter (Chapters 12-22),
introduces the reader to the general aspects
of classification and morphology of selected
taxa, and to current methods and tech-
niques in illustrating plants, fossils, inver-
tebrates, fishes, amphibians and reptiles,
birds, mammals, wildlife illustration, an-
thropological artifacts, and medical sub-
jects. Examples demonstrating the method
of triangulation for rendering fish and re-
constructing archaeological ruins, the use of
coordinate measurements for rendering an-
imal skulls, and the reconstruction of fossil
taxa are illustrated and presented in great
detail. In addition, methods for rendering
plants from hebarium specimens, and the
use of potsherds to estimate the physical
dimensions and original painted surface
patterns of ancient pottery are discussed.
Optical and measuring devices, drafting aids,
specimen-handling tools and props, special
containers, wetting agents and preserva-
tives, and miscellaneous materials used to
handle specimens characteristic to the var-
ious subdisciplines of natural history are also
discussed. As some study objects may be
represented by only one specimen, the care
and handling of such objects is an important
consideration in scientific illustration.
The last two parts of The Guild Handbook
of Scientific Illustration (Parts 4, 5) discuss
pertinent background information for the
scientific illustrator and the scientist. Except
for topics in Chapters 24, 26, and 27, which
are thoroughly covered by Anderson et al.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(1988), other texts I examined either su-
perficially treat or omit such information
(see References below). The remainder of
the review thus summarizes each chapter in
parts 4 and 5.
Part 4, Beyond Basics (Chapters 23-27),
discusses a potpourri of information from
rendering charts, diagrams, and geograph-
ical maps to giving practical advice on the
use and mechanics of the microscope as well
as tips on copy photography and the print-
ing process.
Chapter 23 discusses the function, use,
and proper maintenance of the microscope.
Because most subject areas within scientific
illustration require the use of compound and
stereomicroscopes, it is essential for the il-
lustrator and scientist to understand how to
make the best use of these instruments. For
example, relationships between the focal
point, focal length, and optical axis are dis-
cussed according to the properties of posi-
tive and negative lenses. Resolution is ex-
plained in conjunction with the numerical
aperture of objectives, eyepieces, and sub-
stage condenser unit. In addition, illumi-
nation for both compound and stereoscopic
microscopes is discussed.
Chapter 24 explores the use of diagram-
matic illustration for the presentation of sci-
entific results by the use of charts and dia-
grams. Information on various instruments,
surfaces, and media are presented. Line and
bar charts, scattergrams, pie graphs, stacked
bar charts, tables and diagrams are defined
and illustrated, each according to its most
appropriate use(s). In addition to various
lettering techniques, conventional drafting
techniques and applications to computer-
graphics methods are compared.
Chapter 25 introduces cartography to the
scientific illustrator. A list of academic
courses is given as an indication of educa-
tional experience needed to become a well-
rounded cartographer. Similarly, instru-
ments and materials (surfaces, media, and
tools) used in cartography are listed. Car-
tographical rendering techniques are pre-
VOLUME 91, NUMBER 4
sented with respect to the understanding and
use of projections and various kinds of map
rendering techniques. Figure-ground maps,
visual-hierarchy maps, areal maps, and var-
ious relief representation maps are illus-
trated and discussed. The importance of the
kind, use, and placement of lettering, point
symbol designs, and line symbol designs are
emphasized.
Chapter 26 discusses the three basic ele-
ments of copy photography, 1.e. alignment,
composition, and exposure. The procedure
of photographic copying involves illumi-
nation, preliminary alignment, determina-
tion of exposure, and finally taking the pic-
ture. Specific photographic applications
related to linework, continuous tone, and
color reproductions are discussed. Addi-
tional considerations and special materials
used for archival purposes also are consid-
ered. Finally, evaluation of reproductions
with respect to excessive or inadequate con-
trast, loss of sharpness and detail, uneven
lighting, and discrete blemishes are exam-
ined.
Chapter 27 discusses the printing process
and how line and halftone reproductions are
made. These considerations are important
because original illustrations not only need
to be tailored to the dimensions of the jour-
nal or book they will be printed in, but also
to the printing process used by the publish-
ing company. The complementation of let-
tering with the original and use of labelled
overlays also are discussed. The chapter ends
with a comparison of spot color reproduc-
tion for line illustration and process color
reproduction for tone illustration.
Part 5, The Business of Scientific Illustra-
tion (Chapters 28-30), discusses the legal
rights and general business practices of the
scientific illustrator. Topics of importance
include reproduction and ownership rights
to specific works, contracts, and general du-
ties for the proper and efficient operation of
a freelance business.
Chapter 28 introduces the reader to the
new copyright law of | January 1978, and
643
how it applies to scientific illustrators. This
chapter emphasizes the importance of copy-
right registration necessary for particular
types of work. By law, the owner of the
copyright has the exclusive right to repro-
duce, sell and distribute the work, prepare
derivative works, and perform the work
publicly. All or part of the copyright priv-
ileges are negotiable items between the 11-
lustrator and the client. Finally, the copy-
right law is discussed as it pertains to
infringements and “‘fair use”’ practices.
Chapter 29 stresses the importance of
written contracts. These documents vary 1n
complexity depending on the nature of the
work to be done, and the artist-client rela-
tionship. Consequently, contracts generally
possess several of the following terms of
agreement: an accurate description of ser-
vices, fees, deadlines and payment sched-
ules, copyright, ownership of artwork, cred-
itlines and signatures, alterations, additional
artwork, provisions for termination of art-
work, and incapacitation or death of illus-
trator.
Chapter 30 introduces the reader to the
“real world” of freelance business. Strate-
gies for surveying geographical areas in
which contracts can be established and tips
for estimating potential markets are pre-
sented. Types of portfolios and the presen-
tation of art work to prospective clients are
considered. Studio and office management
duties and responsibilities are outlined ac-
cording to the daily routine of storing and
filing of business records and supplies,
maintaining ledger books, as well as keeping
track of deficits. Other aspects of freelance
businesses, such as the option of incorpo-
ration, home office tax considerations, use
of employees, insurance and retirement
plans are discussed.
In conclusion, The Guild Handbook of
Scientific Illustration gives the serious stu-
dent of scientific illustration an introspec-
tive view of the basic knowledge, skills, and
drive necessary to be successful within the
field. Its holistic approach and detailed con-
644
ceptual descriptions surpass the informa-
tion content and instructive qualities of any
other volume previously written on the sub-
ject (see References below). And in my opin-
ion, these factors by far justify the price. I
give this volume my highest recommen-
dation as a reference for all professional sci-
entific illustrators, students interested in sci-
entific illustration, and all scientists who
consider “‘fine art” as important a tool in
science as their computer algorithms.
References
Comparative statements in this review are
based upon examination of the following
current works on scientific illustration:
Anderson, J. et al., 1988. I//ustrating Sci-
ence— Standards for Publication. Council of
Biological Editors, Bethesda, Maryland, x
+ 297 pp., 112 black-and-white and half-
tone images, 10 color images.
Barlowe, D. and S. Barlowe. 1982. I/lus-
trating Nature—How to Paint and Draw
Plants and Animals. The Viking Press, 111
pp., 176 black-and-white and halftone 1m-
ages, 29 color plates.
Downey, C. and J.L. Kelly. 1982. Biolog-
ical Illustration—Techniques and Illustra-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
tion. The Iowa University Press, vil + 126
pp., 74 black-and-white and halftone im-
ages.
Jastrzebski, Z.T. 1985. Scientific Illustra-
tion—A Guide for the Beginning Artist.
Prentice-Hall Inc., iv + 319 pp., 129 black-
and-white and halftone text images, 11
within appendices, 10 color plates.
Leslie, C.W. 1980. Nature Drawing—A
Tool for Learning. Prentice Hall Press, xvii
+ 206 pp., 257 black-and-white and half-
tone images.
Wood, P. 1982. Scientific Illustration—A
Guide to Biological, Zoological, and Medi-
cal Rendering Techniques, Design, Printing
and Display. Van Nostrand Reinhold, 148
pp., 214 black-and-white and halftone im-
ages, 34 color images.
Zweifel, F.W. 1961. A Handbook of Sci-
entific Illustration. The University of Chi-
cago Press, xv + 132 pp., 66 black-and-
white and halftone images.
David Adamski, Department of Ento-
mology, National Museum of Natural His-
tory, NHB-127, Smithsonian Institution,
Washington, D.C. 20560.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 645-648
Book REVIEW
Alms for the Spore
Insect-Fungus Interactions. Edited by N.
Wilding, N. M. Collins, P. M. Hammond,
and J. F. Webber. 14th Symposium of the
Royal Entomological Society of London
in collaboration with the British Myco-
logical Society. Academic Press, London.
xv1 + 344 pp. $33.00 U.S.
The associations of insects and fungi are
numerous and diverse, but until recently
have received little attention. There have
been few efforts to either compile references
to an extensive and scattered literature or
to promote communications between the
expansive fields of entomology and mycol-
ogy. As co-editor, with mycologist Meredith
Blackwell, of one such effort (Fungus- Insect
Relationships, 1984)— that complemented
an earlier volume on symbioses edited by
L. R. Batra (1979)— I would disagree with
this volume’s jacket which reads in part
“The invited papers given at this meeting
provide the first summary of this fascinating
field...’ But in the product of the sym-
posium is found much to agree with. The
book complements these earlier contribu-
tions in many useful ways.
The volume consists of ten chapters and
a substantive appendix. Four chapters deal
explicitly with associations between insect
and fungus taxa (Coleoptera-fungi by Law-
rence; Macrotermitinae-7ermitomyces by
Wood and Thomas; leaf-cutting ants-fungi
by Cherrett, Powell and Stradling; and bark
and ambrosia beetle-fungi by Beaver. The
remaining six are focused on one or more
basic or applied aspects of ecology.
Lawrence’s chapter on mycophagy in the
Coleoptera is a masterful contribution. The
descriptive ecological catagories proposed
for the modes of feeding in beetles will help
in the general categorization of the habits
of fungus-associated beetles, and will be
found applicable to taxa outside the Co-
leoptera. Treatment of marginal substrates
is important because many groups of beetles
can only be ecologically characterized by
their associations with such habitats, and
not with particular fungus hosts (see, e.g.
Crowson 1984). Plasmodial feeding in
Agathidium has recently been confirmed in
a second subgenus (Wheeler 1987), adding
to the suspicion that such habits may be
more widespread. While Lawrence suggests
that it is too soon to generalize about co-
evolution of beetles and fungi, he does pro-
vide some fascinating correlations with
structural features, notably mouthparts.
Lawrence’s suggestion that mycophagy or
saprophagy was the ancestral habit in the
Coleoptera is not unequivocal. It does agree
with another leading coleopterist’s ideas on
the subject (Crowson 1981), and is perhaps
as good an estimate as is today possible. I
would only point out the following. The
groundplan for Archostemata is not firmly
established and if this suborder is used as
an outgroup for the other three, no conclu-
sion about habits is yet unambiguous. While
rhysodids are a fascinating departure for the
Adephaga into mycophagous habits, it is
doubtful that these are anything but a sec-
ondary adaptation to such habits and can
not be unequivocally argued as a ground-
plan for the adephagans. Myxophagan
mouthparts may well be independently
evolved for their peculiar aquatic habits,
and can not yet be tied overtly to anything
resembling a mycophagous life style. In the
Polyphaga, within which mycophagous
habits are common and widespread, insuf-
ficient knowledge of relationships exists to
say with any certainty what the ancestral
habits were. Further, some taxa suggested
to be possible outgroups for the Coleoptera
646
(e.g. Megaloptera) are predatory. I think that
Lawrence has done the best job possible to-
day in summarizing the breadth of associ-
ations in the order, and he has paved the
way for the more phylogenetically detailed
studies that are badly needed.
This chapter was only one of two excel-
lent contributions by Lawrence. The appen-
dix, co-authored with Peter Hammond,
alone is worth the price of the volume. This
is the first overview of fungus-insect asso-
ciations in a concise, tractable form. It is
fully as authoritative as I have come to ex-
pect from either of its authors. In my view,
the only shortcoming of the appendix (and
one that pervades the entire volume), is the
use of a letter-and-number codon system for
taxonomic references (cf. Tables I and II of
appendix). I did not find any cases where
the system led to mistakes in the text, but
these might be difficult to detect. A similar
system was used by Crowson (1981), with
numerous resultant problems in the text
(Wheeler 1982). Most of those errors I at-
tribute to the publisher failing to have the
manuscript thoroughly reviewed. Academic
Press (the same publisher) seems to have
done an outstanding job in the case of Wild-
ing et al. My problems with such systems
are simple. First, when errors are made, they
are not easily spotted. Second, the reader is
forced to refer to the table for most taxo-
nomic citations including those she is al-
ready familiar with. And last, mycologists
and entomologists are cushioned from ex-
posure to the “other” nomenclature that will
ultimately lead to familiarity. These short-
comings are, at least from a publisher’s view,
offset by the cost savings of not printing long
and repetitious scientific names. I am less
convinced of the benefits accrued to readers.
Hanski provides an informative charac-
terization of the fungus as an insect habitat,
and suggests that several general ecological
questions can be addressed using this sys-
tem. While polyphagy might be explainable
in the terms suggested by Hanski (strong
selection for polyphagy or low-cost of such
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
diverse habits), historical momentum may
be of importance in the ultimate answer. It
may be that in many kinds of insects po-
lyphagous ancestors beget polyphagous de-
scendant species and that this phylogenetic
(historical) component is stronger than either
selection for polyphagy or for lower energy
costs for any one of these taxa.
Several authors discuss mutualistic as-
sociations, including those shared by ter-
mites and fungi (Wood and Thomas in
Chapter 3), leaf-cutting ants and fungi
(Cherrett, Powell and Stradling, Chapter 4),
bark- and ambrosia-beetles and fungi (Bea-
ver, Chapter 5; Berryman, Chapter 6). These
chapters provide excellent introductions to
several mutualistic fungus-insect relation-
ships, but some fall to a greater or lesser
degree into ecological just-so stories. Per-
haps the least constrained Neo-Darwinistic
yarn was that about the evolution of am-
brosia beetles by Beaver (p. 136):
“It seems probable that this evolutionary
process occurred in tropical rain forests,
in which the high temperatures and hu-
midity are particularly favourable for
fungal growth, and where associations be-
tween beetles and fungi would frequently
have occurred. Atkinson and Equihua-
Martinez (1986) suggest that the utiliza-
tion of fungi as a primary source of food
may have arisen from competition be-
tween beetles and fungi for the same sub-
strate. Such competition might frequently
have occurred in different beetle-fungus
associations, leading to the multiple evo-
lution of the ambrosia habit.”
Evolution of ambrosia habits need not have
taken place first in tropical forests. Law-
rence (Chapter 1) has suggested that fungus
associations are taxonomically widespread,
including many beetles and fungi in the tem-
perates. An examination of similar habits
in Lymexylidae, for which there exists a
cladogram (Wheeler 1986), leaves the pic-
ture equivocal. The Hylecoetinae, sister-
group to the other subfamilies, is today only
North Temperate in distribution while the
VOLUME 91, NUMBER 4
common ancestor of Melittomminae plus
Lymexylinae is hypothesized to have been
Pantropical. Where did such habits evolve?
A model of a /Hylecoetus-like ancestral
species, living in temperate forests, evolving
its mutualistic association with fungi is no
less tenable than a Pantropical model. For
the ambrosia beetles, no such cladogram yet
exists. Any attempt to interpret the evolu-
tion of ecological strategies will be flawed
outside of the context of a credible cladistic
model for evolutionary history.
Several chapters address applied aspects
of fungus-insect interactions, including the
dissemination of tree pathogens (Webber
and Gibbs, Chapter 7) in an excellent over-
view of modes of transmission and several
case studies; the role of an Jps beetle and
Urocerus woodwasp in the transmission of
disease in larches (Redfern, Chapter 8); a
very good introduction to mycopathogens
of insects in epigeal and aerial habitats by
H. C. Evans (Chapter 9); and a similarly
good introduction to mycopathogens of soil
insects by Keller and Zimmermann (Chap-
ter 10).
The overall emphasis of the volume is on
ecology, with comparatively less attention
given to evolution. Where discussions about
evolution were given, they did not reflect
recent advances in analytical taxonomy. As
one measure, I did not find a single citation
to a paper dealing with systematic theory.
No major emphasis is necessarily appro-
priate to this book, but many conclusions
reached in it are dependent upon an accu-
rate picture of evolutionary pattern and that
picture depends on the application of cred-
ible analytical methods.
In general I would observe that the issue
of the separation of pattern and process (e.g.
Eldredge and Cracraft, 1980) has not yet
reached studies of fungus-insect interac-
tions. Such criticism does not apply to the
chapters in Wilding et al. uniformly, but it
is widespread enough to merit mention. The
abundant, complex interactions of insects
and fungi represent the end products of a
647
great deal of evolution. Until these products
are tied together in an historical framework
we shall never make sense out of them. Some
authors, Lawrence for example, have rec-
ognized this (“Clearly there is a need for
further rigorous cladistic analyses of both
beetles and fungi,” p. 19) and are leading
the way toward more robust theoretical
frameworks within which to interpret this
wondrous diversity.
This volume is an important contribution
to the ecology and evolution of fungus-in-
sect associations and to the application of
this knowledge to systems of applied inter-
est. It is exceptionally well written, edited,
and produced and is a credit to the authors,
editors, and sponsoring societies. Some of
my criticisms point to better interpretive
and synthetic contexts that are available for
looking at comparative data. Others reflect
the analytical compromises likely to be
found in the face of the enormous numbers
of insects and fungi that interact with one
another and the small number of scientists
studying them. These we may assume will
be corrected in time. This volume repre-
sents an important compilation of current
knowledge in this vast field. Several contri-
butions, including the appendix, will be es-
sential referential material for mycologists
and entomologists alike, and the others uni-
formly suggest productive lines of addition-
al research. Science progresses through the
collective efforts of her practicioners. Those
of us interested in fungus-insect relation-
ships have just been given an important
stepping stone, and fungi and their insect
associates are at last beginning to receive
the attention they deserve.
Quentin D. Wheeler, Department of
Entomology and L. H. Bailey Hortorium,
Cornell University, Ithaca, New York 14853.
LITERATURE CITED
Batra, L. R. (Ed.). 1979. Insect-Fungus Symbiosis,
Nutrition, Mutualism, and Commensalism.
Montclair, N.J. Allanheld, Osman.
648
Crowson, R. A. 1981. The Biology of the Coleoptera.
London, Academic Press.
1984. The associations of Coleoptera with
Ascomycetes, pp. 256-285 Jn Wheeler, Q. and
Blackwell, M., eds., Fungus-Insect Relationships.
New York, Columbia University Press.
Eldredge, N. and J. Cracraft. 1980. Phylogenetic Pat-
terns and the Evolutionary Process. New York,
Columbia University Press.
Wheeler, Q. D. 1982. The biology of beetles. Syst.
Zool. 31: 342-345.
. 1986. Revision of the genera of Lymexylidae
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
(Coleoptera: Cucujiformia). Bull. Amer. Mus. Nat.
Hist. 183: 113-210.
1987. A new species of Agathidium associ-
ated with an “epimycetic” slime mold plasmo-
dium on Pleurotus fungi (Coleoptera: Leiodidae—
Myxomycetes: Physarales— Basidiomycetes: Tri-
cholomataceae). Coleopt. Bull. 41: 395-403.
Wheeler, Q. D. and M. Blackwell (Eds.). 1984. Fun-
gus-Insect Relationships: Perspectives in Ecology
and Evolution. New York, Columbia University
Press.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 649-650
Book REVIEW
Catalog of the Heteroptera, or True
Bugs, of Canada and the Continental
United States. T. L. Henry and R. C.
Froeschner, editors. 1988. E. J. Brill. xix,
958 p.
Noctuidae. R. W. Poole. 1989. E. J. Brill
(Lepidopterorum Catalogus, new series,
Fasc. 118 (Pt. xi, 500 p.; Pt. 2: 501-1015
p.; Pt. 3: 1015-1314 p.).
Both obtainable in the United States
and Canada from E. J. Brill (U.S.A.), Inc.,
P.O. Box 467, Kinderhook, New York
12106, at $58.50 for the Heteroptera cat-
alog and $250 for the Noctuid catalog (3
vols.), plus in both cases USP shipping
charges.
These two catalogs of large and econom-
ically important insect groups are very dif-
ferent in nature and content.
The Henry and Froeschner catalog of
Heteroptera is a catalog in the traditional
senses of the term. All species known to
occur in its stated range, viz., 3834 species,
are listed with full literature references,
summary distribution, and synonymy. Each
family is preceded by a general discussion
and one to several habitus figures. The world
classification 1s outlined and the place there-
in of the regional species is given. In short,
it is virtually all that could be expected of
a regional catalog and would serve well as
a model for such. All that I miss in it 1s
specification of the nature of data given in
the references, whether there are figures (ha-
bitus, genitalia, etc.), sexes described, de-
scriptions of immature stages, mention of
host plants, etc.
The catalog of noctuid moths is very dif-
ferent from the foregoing. It is more than a
checklist, inasmuch as complete reference,
type data, and some other items are includ-
ed, but it is otherwise narrowly minimal.
The fact that it lists for the first time all
species of Noctuidae of the world, approx-
imately 38,000 through 1980, and that the
group is of great economic importance, have
both limited its scope and enhanced its val-
ue.
The 3 volumes consist of 2 volumes of
text and one of bibliography and index. The
arrangement is entirely alphabetical by gen-
era and species. Suprageneric classification
is indicated only by a subfamily name fol-
lowing the generic name, these being basi-
cally according to the Nye classification. It
is also stated in the introduction that a new
classification as well as lists of genera in-
cluded under each suprageneric category is
in preparation. Subspecies are cited under
species aS synonyms because the author
“does not subscribe to the subspecies con-
Cepts
Only the original form of each species
name is cited, regardless of gender of any
genus with which it may presently be com-
bined. I have inveighed against this practice
in a previous review in this journal in 1984
(Vol. 86, pages 328-329), stating then that
I thought the practice was confined to a small
number of systematists. I still think so and
now believe that the number is even less
than I thought then. Ifa user of this catalog
wishes to follow the rules of the Interna-
tional Code of Zoological Nomenclature in
regard to gender concord, he will receive no
help therefrom. In the genus Plagiomimicus
(masculine) 37 species names are listed, only
3 of them in masculine form, 7 in neuter
form because they were originally proposed
in Stibadium, and at least 7 in feminine
form.
After each species name (in semibold type)
are cited the author’s name and the name
of the genus in which the species was orig-
inally proposed, both of these latter in the
same regular Roman type without paren-
theses, punctuation, or in distinctive type-
650
face. This seems to me to be carrying econ-
omy a little too far.
Synonymy is complete and references to
type data, figures, larval descriptions, and
food plants are cited. No locality records
except that of the type are cited. References
to generic transfers, except for new ones here
made, are not given; for example, it is not
indicated where or when the transfers of
Stibadium species to Plagiomimicus were
made. The author states that the many new
combinations were “based on my own re-
search but more commonly they are the
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
combinations or synonymies in the collec-
tions of the U. S. National Museum of Nat-
ural History and British Museum of Natural
History that have never been published but
are based on the curation of past system-
atists, often of unknown identity.”
George C. Steyskal, Cooperating Scien-
tist, Systematic Entomology Laboratory,
Agricultural Research Service, U.S. Depart-
ment of Agriculture, c/o U.S. National Mu-
seum of Natural History, Washington, D.C.
20560.
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 651-658
SocieTY MEETINGS
944th Regular Meeting—December 1, 1988
The 944th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President Gene Wood in the
Naturalist Center, National Museum of
Natural History, at 8 p.m. on December 1,
1988. Twenty-three members and 8 guests
were present. Minutes of the November
meeting were read and approved with one
emendation.
Membership Chairman G. B. White read
the name of the following applicant for
membership: Rick McMullan, German-
town, Maryland. Chairman White also re-
ported that the Executive Committee had
unanimously elected Frank W. Fisk of De
Land, Florida, to Emeritus status. Dr. Fisk
has been a member of our Society since
1968; he retired from Ohio State University
in 1976.
Annual reports were given by the out-
going President, Membership Chairman,
Corresponding Secretary, Treasurer (read
by F. C. Thompson), Custodian, and Pro-
gram Chairman. President Wood warmly
thanked all Society officers for setting a stan-
dard of job performance that is the envy of
other entomological organizations.
President Wood and D. M. Anderson
again presented the slate of nominees for
officers in 1989, which this time included
Jeffrey R. Aldrich as candidate for Presi-
dent-Elect. Dr. Wood then called for addi-
tional nominations from the floor, of which
there were none. A motion was made and
seconded that the slate be accepted as pre-
sented. The motion was approved by accla-
mation.
President Wood read a letter from Manya
B. Stoetzel, President of the American As-
sociation for Zoological Nomenclature
(AAZN), an organization formed in 1983 to
(1) raise money in the United States for sup-
port of the International Commission on
Zoological Nomenclature (ICZN) and (2)
provide a liaison between the American sys-
tematic community and the ICZN. Estab-
lished in 1895, the ICZN is the only world-
wide authority that considers problems of
zoological nomenclature and that is em-
powered to rule on such problems. Amer-
ican systematists generate 25% of the ICZN’s
workload, and 35% of all cases deal with
the class Insecta. All entomologists, regard-
less of specialization, must know the correct
scientific names of the insects with which
they are working. Yet, in recent years the
finances of the ICZN have been increasingly
strained by rising costs and reduced bud-
gets. In her letter, Dr. Stoetzel asks that the
Entomological Society of Washington be-
come a supporting member of the AAZN
and that it continue its membership on a
yearly basis for the foreseeable future. In-
dividual ESW members are also encouraged
to join, pledging whatever amount they can
afford (generally not less than $20 per an-
num). Dr. Wood urged the membership to
seriously consider this request and an-
nounced that he would forward Dr. Stoet-
zel’s letter to incoming President F. Chris-
tian Thompson for a decision at the next
meeting of the Executive Committee.
Corresponding Secretary J. M. Kingsol-
ver displayed a series of remarkable scan-
ning electron photomicrographs showing
apparent stridulatory structures on 3 species
of bruchid beetles. In Acanthoscelides chi-
ricahuae (Fall) (western United States), the
scraper is a ridge on the mesal margin of
the hind femur, and the file is a radiating
set of fine ridges on the face of the hind coxa.
In Amblycerus eustrophoides (Schaeffer)
(Florida), the scraper is a series of ridged
setal bases on the mesal face of the hind
femur and the file is a transversely striate
652
ridge on the ventral border of the metepi-
sternum. In Amblycerus n. sp. (Mexico), the
scraper is a striated tooth on the ventral
margin of the hind femur, and the file is a
ridge similar to that of 4. eustrophoides but
nearer the middle of the metepisternum.
Program Chairman W. E. Steiner, Jr. and
Jil M. Swearingen exhibited a beautiful
hammer dulcimer built with genuine worm-
eaten American chestnut by Nicholas Blan-
ton of Shepherdstown, West Virginia, and
incorporating lifelike models of several ar-
thropods, including one of the Locust Borer,
Megacyllene robiniae (Forster), over the res-
onance hole.
R. G. Robbins displayed a copy of a new
reference for acarologists, Provisional Atlas
of the Ticks (Ixodoidea) of the British Isles,
by Kieran P. Martyn, Biological Records
Centre, Institute of Terrestrial Ecology,
Monks Wood Experimental Station, Hun-
tingdon.
President Wood announced that although
Gary Larson will not be made a Fellow of
the Entomological Society of America at this
year’s National Conference in Louisville, a
sizable collection of cartoons evidencing this
artist’s uncanny insight has been assembled
as a gift on behalf of the entire entomolog-
ical community. In Dr. Wood’s words,
“Gary Larson has probably done more to
encourage insect study than anything since
yellow fever!”
With around of applause, President Wood
welcomed the return of J. R. Aldrich, our
Society’s Program Chairman in 1983 and
1984 and our President-Elect for 1989. Dr.
Aldrich thanked the Society for its support.
He then noted with sorrow the untimely
passing of his colleague Elton Warren Her-
bert, Jr. (November 5, 1943—November 16,
1988), a research entomologist at the Belts-
ville Agricultural Research Center from
1966 to 1969 and again from 1972 until his
death. Dr. Herbert is remembered for his
studies of the nutritional requirements of
honeybees and for his research on methods
to control bee diseases and mites parasitic
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
on bees. Specifically, he worked to develop
a substitute for pollen and to formulate
chemicals that block the molting hormones
of Varroa jacobsoni Oudemans without
affecting honeybee physiology. Memorial
contributions may be made to the Dr. Elton
W. Herbert, Jr. Scholarship Fund in care of
the Entomological Society of America, 9301
Annapolis Road, Lanham, Maryland 20706.
The speaker for the evening was James
M. Hill, Ecologist, Maryland National Cap-
ital Park and Planning Commission, whose
talk was entitled “Life History Studies of
Cicindela dorsalis, an Endangered Tiger
Beetle.” With the ongoing degradation of
beachfront in Calvert County and other
Maryland jurisdictions, several species and
subspecies of tiger beetles have been locally
extirpated. Beach-inhabiting species require
untrammeled sand in which to complete
their development. The onslaught of com-
mercial and residential construction, off-the-
road vehicles, and sheer human numbers
has rendered much of the Atlantic coast un-
inhabitable for both tiger beetles and their
prey. In an attempt to conserve these at-
tractive insects, Mr. Hill has conducted life
history studies focusing on population dy-
namics, territoriality, food preferences,
thermoregulation, parasites and predators.
His presentation was illustrated with superb
close-up photographs taken by Nate Erwin
of this Society and with slides of distribu-
tion maps prepared by Barry C. Knisley of
Randolph-Macon College, Ashland, Virgin-
la.
President Wood congratulated Mignon
Davis and Don Anderson for their unstint-
ing loyalty to this Society. Mignon has been
bringing and preparing refreshments for
many years, while Don has set up chairs
and minded the slide projector for longer
than even he can remember! Such members
are the linchpins of our organization. With
these kind words, Dr. Wood handed the
gavel to incoming President Chris Thomp-
son, who adjourned the meeting at 9:20 p.m.
Refreshments (including some normally re-
VOLUME 91, NUMBER 4
served for lightly sclerotized specimens) fol-
lowed, together with holiday entertain-
ment a la dulcimer.
Richard G. Robbins, Recording Secretary
945th Regular Meeting—January 5, 1989
The 945th Regular Meeting of the Ento-
mological Society of Washington was gav-
eled to order by President F. Christian
Thompson in the Naturalist Center, Na-
tional Museum of Natural History, at 8:05
p.m. on January 5, 1989. Thirty-two mem-
bers and 23 guests were present. Minutes of
the December meeting were read and ap-
proved.
Membership Chairman G. B. White read
the names of the following applicants for
membership: Robert Carleton Brown, Mor-
ristown, Tennessee; Candy Feller, Depart-
ment of Entomology, Smithsonian Institu-
tion; Thomas O. Robbins, USDA, ARS,
Grassland, Soil and Water Research Lab-
oratory, Temple, Texas; and William P.
Weaver, Jr., Panorama City, California.
Chairman White also reported that the Ex-
ecutive Committee had unanimously elect-
ed the following members to Emeritus sta-
tus: Lauren D. Anderson, University of
California at Riverside, who has been with
us since 1944; and George R. Manglitz,
University of Nebraska, Lincoln, a member
since 1956.
President Thompson discussed the high-
lights of the Executive Committee meeting
held earlier in the day. With evident plea-
sure, he described the Committee’s decision
to throw down the gauntlet by donating $750
to the American Association for Zoological
Nomenclature (see previous minutes). Oth-
er priority items on this year’s agenda in-
clude selection of an Associate Editor, to
succeed Hiram Larew in 1990; sale at re-
duced rates of our vast back stock of Pro-
ceedings and Memoirs; and a survey of the
interests and expectations of the entire ESW
653
membership. T. E. Wallenmaier asked
whether the Committee had agreed to grant
other entomological organizations brief ac-
cess to the Proceedings. Dr. Thompson re-
plied that this matter remains under dis-
cussion.
Dr. Wallenmaier described his Christmas
sojourn in Alabama, where he encountered
the cattail toothpick grasshopper, Leptysma
marginicollis (Serville), an unusually slen-
der acridid that cryptically clings to grasses,
sedges and rushes bordering ponds and wa-
tercourses. According to the orthopterist
Jacques R. Helfer, this species ranges from
Maryland west to Kansas and Nebraska and
thence south to the Gulf Coast. Its quick
flight and agile dodging behavior make it a
difficult insect to see, let alone capture, but
collectors have learned to beat wetland
vegetation, thereby knocking the grasshop-
pers into the water, where they are easily
retrieved. A pinned specimen was exhibited
to the audience.
M. B. Stoetzel displayed a notebook con-
taining copies of numerous cartoons drawn
in honor of Gary Larson, who was sent the
originals after they were shown at a special
exhibition during last year’s National Con-
ference of the Entomological Society of
America, in Louisville, Kentucky.
The speakers for the evening were Dan
Polhemus and Warren E. Steiner, Jr., both
of the Department of Entomology, Smith-
sonian Institution. Their talk, entitled
“Natural History of Madagascar, with Em-
phasis on Entomological Studies Past and
Present,” was a delicious romp over the
length and breadth of the world’s fourth
largest island (after Greenland, New Guinea,
and Borneo), which at 587,041 square ki-
lometers is somewhat larger than France.
Climatically, Madagascar 1s divided into two
main areas: a narrow humid belt running
along the east coast and separated from the
rest of the island by the steep eastern slopes
of the central mountains; and a series of
western slopes and plains that are charac-
terized by a distinct dry season. In separate
654
but complementary expeditions, Dan and
Warren sampled the insect fauna of Mad-
agascar’s multifarious biotopes, including
the rain forests of the east coast, the remnant
deciduous woods of the Central Highlands,
the broad savannas of the west, and the
thorny scrub of the xeric southwest. Ac-
cording to Dr. Vincent Razafimahatratra,
an entomologist and professor at the Uni-
versity of Madagascar, fully 90% of the is-
land’s insect species remain undescribed. A
tiny sample of this diversity was exhibited
in several insect drawers crammed with
moths, butterflies, beetles, and other insects
that well evoke their mysterious homeland.
Each speaker also chronicled his travels with
scores of photographs, some so striking that
listeners could almost taste the island’s red
lateritic dust or hear its groves of windswept
bamboo. Warren warmly thanked Patrick
Daniels, Claire Kremen, and Patricia
Wright, all of Duke University, for their
logistical support and unfailing camaraderie
throughout his expedition.
Visitors were introduced and the meeting
was adjourned at 9:25 p.m. Refreshments
followed.
Richard G. Robbins, Recording Secretary
946th Regular Meeting— February 2, 1989
The 946th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President F. Christian Thomp-
son in the Naturalist Center, National Mu-
seum of Natural History, at 8:07 p.m. on
February 2, 1989. Twenty-two members and
two guests were present. Minutes of the Jan-
uary meeting were read and approved with
one correction.
President Thompson announced that the
Executive Committee had selected Robert
D. Gordon, Research Entomologist, Sys-
tematic Entomology Laboratory, USDA, as
the new Associate Editor of our Proceed-
ings. Dr. Gordon will succeed Hiram Larew
as Editor at the end of this year.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
Membership Chairman G. B. White read
the name of the following applicant for
membership: Jon K. Gelhaus, Postdoctoral
Fellow, Department of Entomology, Smith-
sonian Institution.
Custodian Anne M. Wieber warmly
thanked Herb Jacobi, Trevor Lubbert, and
Geoff White for taking time from their busy
weekend schedules to help her organize and
spruce up the Society’s vast backlog of Pro-
ceedings and Memoirs.
W. E. Bickley displayed a new book for
culicidologists, The Natural History of Lar-
val Mosquito Habitats, by Marshall Laird,
published in September of 1988 by Aca-
demic Press, 555 pages, ISBN 0-12-434005-
9, $135.00/cloth.
President-Elect J. R. Aldrich exhibited a
postcard on which the sender had affixed a
colorful National Wildlife Federation stamp
featuring the southern green stink bug, Ne-
zara viridula (Linnaeus) (Hemiptera: Pen-
tatomidae). Dr. Aldrich is gratified by pub-
lic recognition of this species because he has
been investigating its physiology for several
years.
The speaker for the evening was Past
President Gene Wood, whose talk was en-
titled “Insect Short Stories,” some reminis-
cences and anecdotes of an extension ento-
mologist. Drawing on a lifetime of
experience in urban entomology, Dr. Wood
recalled his adventures with head lice (Pe-
diculus humanus capitis de Geer) on school
children, body lice (P. h. humanus Lin-
naeus) on vagabonds, and German cock-
roaches (Blattella germanica (Linnaeus))—
not to be confused with the brown-banded
cockroach, Supella longipalpa (Fabricius),
which, alas, has lost its lovely alliterative
epithet supellectilium (Serville)—in restau-
rants and apartments throughout the mid-
Atlantic region. He also related the results
of his research on termite colonies, includ-
ing observations on behavior, rearing meth-
ods, and regulation via growth inhibitors.
There appear to be at least six components
in effective urban pest management: fre-
quent surveillance; establishment of pest
VOLUME 91, NUMBER 4
thresholds; accurate and detailed documen-
tation; public education (including regular
liaison); enlistment of alternative control
strategies; and proper pesticide usage. These
factors were illustrated with photographs
taken during actual control operations.
President Thompson thanked Dr. Wood
and, amid much applause, presented him
with a certificate of appreciation in recog-
nition of his service to our Society over the
past year.
Mignon Davis thanked Dr. Bickley for
offering to tote in the cider this evening and
called for additional beverage volunteers.
On that refreshing note, the meeting was
adjourned at 9:20 p.m.
Richard G. Robbins, Recording Secretary
947th Regular Meeting— March 2, 1989
The 947th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President F. Christian Thomp-
son in the Naturalist Center, National Mu-
seum of Natural History, at 8:08 p.m. on
March 2, 1989. Twenty-five members and
16 guests were present. Minutes of the Feb-
ruary meeting were read and approved.
Membership Chairman G. B. White read
the names of the following applicants for
membership: Prem Bhatt, Baltimore,
Maryland: Richard Duffield, Department of
Biology, Howard University, Washington,
D.C.; Kenneth S. Frazer, Department of Bi-
ology, University of Alabama, Tuscaloosa;
Alan R. Olsen, Altadena, California; and J.
David Warthen, Jr., USDA, ARS, PSI,
ICEL, Beltsville, Maryland.
Lance A. Durden exhibited some local
pedilid beetles that are characteristically at-
tracted to members of an unrelated coleop-
terous family, the Meloidae. While collect-
ing in a section of Patuxent River Park,
Prince George’s County, Maryland, Lance
obtained a single Pedilus terminalis (Say)
that had attached itself to a living speci-
men of Meloe angusticollis Say. During a
655
field trip of 5 or 6 hours, the pedilid re-
mained firmly joined to the meloid’s dor-
sum but, perhaps predictably, it detached
while being readied for photography. In Be-
thesda, Montgomery County, Maryland,
four P. terminalis were found clinging to a
dead specimen of Lytta aenea Say, dem-
onstrating that this peculiar heterospecific
attraction does not require a live meloid.
Apparently, pedilids are drawn to the can-
tharidin in meloid hemolymph. Canthari-
din is a potent toxin that repels potential
predators; it thus seems odd that pedilids,
together with their relatives the anthicids,
are sometimes attracted to this substance.
Dr. Durden thanked T. J. Spilman for his
assistance in identifying these beetles and
for providing valuable life history infor-
mation. An animated discussion ensued,
during which T. J. Henry, T. J. Spilman, A.
G. Wheeler, Jr., and J. R. Aldrich noted that
cantharidin has been used by entomologists
(among others) to attract at least two
subfamilies of Miridae (Hemiptera) as well
as ceratopogonid midges (Diptera: Nema-
tocera).
R. F. Whitcomb exhibited a male speci-
men of an undescribed leafhopper (Ho-
moptera: Cicadellidae: Limotettix n. sp.) that
feeds on spike-rushes (Cyperaceae: Eleoch-
aris sp.) associated with serpentine soils. The
specimen was collected on June 20, 1970 in
Beltsville, Prince George’s County, Mary-
land, but appears to have been subsequently
extirpated there. It still occurs at Soldiers
Delight, in western Baltimore County.
President Thompson displayed a singu-
larly interesting book that should not be
overlooked by entomologists with a yen for
travel. A Birder’s Guide to Japan, by Jane
Washburn Robinson, 1987, Ibis Publishing
Company, Santa Monica, California, is
crammed with advice on how to conduct
field work—and oneself—in a land that is
the antithesis of East Coast America. Topics
include when to go, what to bring, finding
your way about, safety, making arrange-
ments, and getting help. The insightful dis-
cussion of good manners might be profit-
656
ably perused even by stay-at-homes. Scores
of detailed maps together with summaries
of useful kanji should prove invaluable to
entomologists working in rural Japan. As
for the Occident, the President also exhib-
ited a flyer from the British Biology Cura-
tors’ Group exhorting young naturalists to
““beetle-down” to their neighborhood mu-
seum.
The speaker for the evening was A. G.
Wheeler, Jr., whose talk was entitled “Some
Insects of Eastern Serpentine Barrens.” Ser-
pentine rocks or ultramafics, i.e. igneous
rocks containing high concentrations of fer-
romagnesian minerals but a low comple-
ment of silica, yield soils that support an
impoverished but distinctive xeromorphic
flora characterized by endemism, disjunct
ranges, ecotypic differentiation, and nu-
merous morphological variants. To survive
on serpentine exposures, plants must tol-
erate low nutrient levels, high concentra-
tions of toxic heavy metals (nickel, chro-
mium, cobalt), and unfavorable calcium/
magnesium quotients that are equally del-
eterious to animals. No group of “‘serpen-
tine animals” has received more attention
than the insects, which have evolved nu-
merous endemic or quasi-endemic species
attuned to the biotic and edaphic strictures
of their environment. Among the species
discussed this evening were chrysomelid
beetles of the genus Diabrotica, the lygaeid
bug Blissus breviasculus Barber, the tingid
Hesperotingis antennata Parshley, and a
suite of striking myrmecomorph mirids:
Schaffneria davisi (Knight), S. pilophoroides
(Knight), and S. schaffneri Knight. Dr.
Wheeler’s presentation was illustrated with
slides prepared by Tom Smith and Jim
Stimmel. Two recent comprehensive texts
were also exhibited: Floras of the Serpentine
Formations in Eastern North America, with
Descriptions of Geomorphology and Min-
eralogy of the Formations, by Clyde F. Reed,
1986, Contributions of Reed Herbarium,
No. XXX, Baltimore, Maryland; and Ser-
pentine and Its Vegetation, A Multidisciplin-
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
ary Approach, by Robert Richard Brooks,
1987, Dioscorides Press, Portland, Oregon.
W. E. Bickley announced the availability
of a new book, Nymphs of North American
Stonefly Genera (Plecoptera), by Kenneth
W. Stewart and Bill P. Stark, with illustra-
tions by Jean Stanger. This work, volume
12 in the Thomas Say Foundation series, is
billed as the definitive sequel to P. W. Claas-
sen’s 1931 monograph, Plecoptera Nymphs
of America (North of Mexico), and may be
purchased by members of the Entomolog-
ical Society of America for $65.00 (non-
members $108.00). However, Dr. Bickley
perceptively notes that genera do not have
nymphs.
As one of its centennial activities, the En-
tomological Society of America 1s proposing
that the United States adopt a national in-
sect. Recently, their members selected the
monarch butterfly, Danaus plexippus (Lin-
naeus), as a candidate for this distinction.
President Thompson distributed some
“campaign literature” in the form of a col-
orful ESA brochure describing the mon-
arch’s life history and recent collaborative
efforts to preserve its overwintering grounds
in California and Mexico. Organizations and
individuals who would like to help sponsor
this project are invited to contact Douglas
W.S. Sutherland, ESA National Insect Sub-
committee, 9301 Annapolis Road, Lan-
ham, Maryland 20706-3115, or call (301)
731-4535.
T. J. Spilman called the membership’s
attention to a temporary exhibit of wooden
insect sculptures in the Insect Zoo, National
Museum of Natural History. Carved from
fine woods by Patrick Bremer, these ana-
tomically exact likenesses average three to
four feet in length and were featured in the
December 1988 issue of Smithsonian mag-
azine.
Among our numerous visitors this eve-
ning were four members of the Maryland
Forest, Park and Wildlife Service— Fraser
D. Bishop, Gene Cooley, Katharine Mc-
Carthy, and Frank Ryan—who discussed the
VOLUME 91, NUMBER 4
distribution and biota of their state’s ser-
pentine soils.
President Thompson thanked Rich Rob-
bins and Ed Saugstad for bringing refresh-
ments, wet and dry, toward which he ad-
journed the meeting at 9:25 p.m.
Richard G. Robbins, Recording Secretary
948th Regular Meeting—April 6, 1989
The 948th Regular Meeting of the Ento-
mological Society of Washington was called
to order by President F. Christian Thomp-
son in the Naturalist Center, National Mu-
seum of Natural History, at 8:07 p.m. on
April 6, 1989. Nineteen members and five
guests were present. Minutes of the March
meeting were read and approved with one
emendation.
Membership Chairman G. B. White read
the names of the following applicants for
membership, all from Maryland: R. Mark
Beach, Hanover; Michael B. Dimok, Co-
lumbia; Mary Anne Erickson, Baltimore;
Deborah Hahn Fay, College Park; and Mark
J. Rothschild, Salisbury.
Past President Gene Wood and Don H.
Messersmith circulated a draft of a letter
inviting other entomological organizations
to publish summaries of their minutes or
notices of their activities in our Proceedings.
The letter reads as follows: “The Entomo-
logical Society of Washington is inviting
your society to participate in a new coop-
erative project. We believe that greater co-
operation in the dissemination of infor-
mation among the many entomological
organizations in North America could be
useful to all of them. Therefore we would
like to invite you to periodically submit
items of general interest to our Editor for
inclusion in our quarterly Proceedings of the
Entomological Society of Washington. A
short paragraph or two would suffice be-
cause space is limited. You could include
your important society news such as when
657
and where you meet, elections of officers,
projects and activities, field trips or other
special events, news about members (hon-
ors received, promotions, retirements,
deaths), speakers at meetings, etc. Our Ed-
itor would modify your submissions to fit
the space we will reserve for these news
items. There will be no charge for this ser-
vice. Please let us know if you wish to par-
ticipate in this activity.”” There were no 1m-
mediate comments from the membership.
Corresponding Secretary J. M. Kingsol-
ver displayed a valuable new reference for
curators and conservators. 4 Guide to Mu-
seum Pest Control, edited by Lynda A.
Zycherman and John Richard Schrock, 29
November 1988, $36.00/paper, ISBN
0-942924-14-2, is published jointly by the
Foundation of the American Institute for
Conservation of Historic and Artistic Works
and the Association of Systematics Collec-
tions. This book provides expert guidance
on pest identification and pesticide use in
the context of current federal statutes and
museum policies. Sections of special inter-
est to entomologists include “Illustrated
Guide to Common Insect Pests in Mu-
seums,” by J.M. Kingsolver (drawings by
Candy Feller); “Cockroaches in the Mu-
seum Environment,” by W.J. Bell; “Biology
and Control of Wood-infesting Coleop-
tera,” by W.H. Robinson; “Trapping Tech-
niques for Dermestid and Anobiid Beetles,”
by W.E. Burkholder and J.K. Phillips; and
“List of Insect Pests by Material or Appar-
ent Damage,” revised by J.R. Schrock. The
work concludes with a partially annotated
bibliography on pest control in museums,
compiled by Karen Preslock, Branch Li-
brarian at the Smithsonian’s Museum Sup-
port Center in Suitland, Maryland.
Edd Barrows exhibited a bag of bag-
worms, Thyridopteryx ephemeraeformis
(Haworth) (Lepidoptera: Psychidae), which
he had collected and reared during the past
winter. Few investigators have bothered to
conduct laboratory research on this com-
mon moth, but Dr. Barrows has found that
658
eggs from Washington-area females collect-
ed in winter will hatch in about one month
if held at 80°F (27°C).
R.G. Robbins exhibited a set of colorful
commemorative stamps from Mozambique
belonging to one of this evening’s guests,
Ralph P. Eckerlin, Northern Virginia Com-
munity College, Annandale Campus. These
stamps feature paintings of some of Africa’s
most beautiful tick species: Amblyomma
eburneum Gerstacker, A. hebraeum Koch,
A. pomposum Déonitz, A. theilerae Santos
Dias (= A. hebraeum), Dermacentor cir-
cumguttatus Neumann, and D. rhinocerinus
(Denny). To the best of the Recording Sec-
retary’s knowledge, no other nation has of-
fered its citizens the opportunity to lick a
tick. Dr. Eckerlin will attempt to obtain ad-
ditional sets for entomophilatelists.
President Thompson circulated a bro-
chure announcing the Second International
Congress of Dipterology, to be held in Brati-
slava, Czechoslovakia, August 27-Septem-
ber 1, 1990. This Congress will be organized
by the Slovak Academy of Sciences, Come-
nius University, and the Slovak Entomo-
logical Society. Participants are invited to
join in dipterological field trips, tours of
Prague, a journey to the Tatra Mountains
(the highest range of the Carpathians), and
even a visit to a Czechoslovakian spa.
The speaker for the evening was Charles
Griswold, Postdoctoral Fellow, Depart-
ment of Entomology, Smithsonian Institu-
tion, whose talk was entitled ‘Natural His-
tory of Southern African Social Spiders.”
Social spiders are a pantropical assemblage
of many families. In Africa, several species
live along watercourses, where they capture
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
emerging aquatic insects, but others are
found in savannas and some occur even in
deserts. All share one nest for retreat and
construct one large sheet or tangle web for
prey capture. No orb weavers or running
species are known. Group capture strata-
gems may enable these spiders to subdue
larger prey, but Dr. Griswold’s research in-
dicates that the chief benefit of cooperation
is a greater diversity of prey. Tactile and
chemosensory stimuli probably help spiders
distinguish prey from conspecifics. Old prey,
vegetative debris, and silk are incorporated
into the nest, which consists of numerous
interconnected tubes with openings to the
outside. Egg sacs are communally guarded,
and newly emerged young are fed regurgi-
tated food by females, which may also
“adopt” any orphaned progeny. Young spi-
ders do not assist in catching prey until they
have reached their third instar. The ratio of
males to females is about 1:10, with females
founding new colonies either as groups or
individuals. All these remarkable behaviors
were captured with stunning close-up pho-
tographs taken by the speaker’s wife, Teresa
Meikle-Griswold.
J. R. Aldrich reminded the membership
that this year’s banquet will be held at the
National Museum of Natural History, with
dinner in the Associates’ Court.
President Thompson thanked Doug
Sutherland, who in turn thanked his daugh-
ter, for making this evening’s cookies. Vis-
itors were introduced and the meeting was
adjourned at 9:18 p.m.
Richard G. Robbins, Recording Secretary
PROC. ENTOMOL. SOC. WASH.
91(4), 1989, pp. 659-664
PROCEEDINGS
of the
ENTOMOLOGICAL SOCIETY
OF
WASHINGTON
Volume 91
OFFICERS FOR THE YEAR 1989
President F. Christian Thompson
President-Elect
Recording Secretary
Corresponding Secretary
Treasurer
Program Chairman
Membership Chairman
Custodian
Editor
Past-President
Published by The Society
WASHINGTON, D.C.
1989
Jeffrey R. Aldrich
Richard G. Robbins
John M. Kingsolver
Norman E. Woodley
Warren E. Steiner, Jr.
Geoffrey B. White
Anne M. Wieber
Hiram G. Larew
F. Eugene Wood
TABLE OF CONTENTS, VOLUME 91
ARTICLES
ADAMSKI, D.—See ASQUITH, A.
APPEL, A. G. and R. C. SPONSLER—Water and temperature relations of the primitive
xylophagous cockroach Cryptocercus punctulatus Scudder (Dictyoptera: Cryptocercidae)
ASQUITH, A. and D. ADAMSKI—Description of the predaceous larva of Pseudogaurax
signatus (Loew) (Diptera: Chloropidae) .........................
BAKER, G. T. and M. M. ELLSBURY— Morphology of the mouth parts nda antenna Vof the
larva of the clover stem borer, Languria mozardi Latreille (Coleoptera: Languriidae)
BAKER, G. T.—See ELLSBURY, M. M.
BARROWS, E. M.—See LAGOY, P. K.
BECKER, V. O.—See MILLER, S. E.
660 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
BOECKLEN, R. P.—See FERNANDES, G. W.
BOHART, R. M.—Review of Maricopodynerus (Hymenoptera: Eumenidae) .............. 575
BOLDT, P. E.—Host specificity studies of Stolas fuscata (Klug) (Coleoptera: Chrysomelidae)
for the biological control of Baccharis salicifolia (R.&P.) Pers. (Asteraceae) ............. 502
BOLDT, P. E.—See GAGNE, R. J.
BURKE, H. R.—See CLARK, W. E.
BUTLER, L.—Observations on Meganola spodia Franclemont (Lepidoptera: Noctuidae) with
andescription ofthe mmature dlarvalerrnns ca) tee nacsectetes ce saeeecreey-tae evecare en eee etter 615
BYERS, G. W.—Homologies in the wing venation of primitive Diptera and Mecoptera ... 497
CASTRO, A. G.—See FERNANDES, G. W.
CAVE, R. D. and M. J. GAYLOR—Longevity, fertility, and population growth statistics of
Telenomusyreynoldasi (Aymenoptera:Scelionidae)) =i ncschos occas 22 ars ewact to soe a ee 588
CHAN, K. L.—See LEE, K. M.
CLARK, W. E. and H. R. BURKE—Revision of the ornatus species group of the genus
Anthonomus’ Germar (Coleoptera: Curculionidae). 3 o.5 <6,o2 = cc.2,01c.e cag) ra = es Sieve cree ee 88
COLLINS, M. S.—See NICKLE, D. A.
DAVIS, D. R.—An exceptional fossil amber collection acquired by the Smithsonian Institu-
ELON ee cn sne eee Nee ee pare es BE vos She NDT ee oy Ame ete ce eB Pe RE fee ea eherercc sea eidyeyerts onsite ls, SreheuSaigele eeWs 545
DIETRICH, C. H.—Surface deuietinine of the abdominal erica ne Membracidae and
other Auchenorshyncha’ (Homoptera) err cecrsserss. ciate cre sionensheis siete o.chsrecert.s exereietacdlaraiereiersisisiece = 143
EISENBERG, R. M.—See HURD, L. E.
ELLSBURY, M. M. and G. T. BAKER—Biological and morphological aspects of the egg stage
and oviposition of Languria mozardi (Coleoptera: Languriidae) ........................ 9
ELLSBURY, M. M.—See BAKER, G. T.
FERNANDES, G. W., W. J. BOECKLEN, R. P. MARTINS and A. G. CASTRO —Ants
associated with a coleopterous leaf-bud gall on Yy/opia aromatica (Annonaceae) ......... 81
FORATTINI, O. S. and M. A.M. SALLUM—Redescription of Culex (Melanoconion) delpontei
Duret, 1968 and Cx. (Mel.) pereyrai Duret, 1967, from southern Brazil ................. 473
GAGNE, R. J. and P. E. BOLDT—A new species of Neolasioptera (Diptera: Cecidomyiidae)
from Baccharis (Asteraceae) in southern United States and the Dominican Republic ..... 169
GAGNE, R. J.—See WEATHERBY, J. C.
GAYLOR, M. J.—See CAVE, R. D.
GOEDEN, R. D.—Host plants of Neaspilota in California (Diptera: Tephritidae) .......... 164
GOEDEN, R. D.—See HEADRICK, D.
GRISSELL, E. E.—Megastigmus floridanus (Hymenoptera: Torymidae), newly discovered in
Tlexsseed\(Aquifoltaceae) gncia5 < siics.caireinedtttva ana are al Gkeate catest bes trek eeeteere ee 604
HANSSON, C.—New World of Holcopelte aa BALE (Hymenoptera: Eulophidae), ath
descriptions of two new species A 0 a an Ti Re nt ee See RST Oe Cele He 59
HEADRICK, D. and R. D. GOEDEN—Life history of Pteromalus coloradensis (Ashmead)
(Hymenoptera: Pteromalidae) a parasite of Paracantha gentilis Hering (Diptera: Tephritidae)
INNGIFSIU 7 thistle: Capita esmyecejeceatm = sya cc. otaras oc Ne acl Aes sees catcumsiohs oak sant eres 594
HELLENTHAL, R. A. and R. D. PRICE— Geampduee us (Mallophaga: Trichodectidae) from
the Texas and desert pocket gophers (Rodentia: Geomyidae) ................00..00000.. 1
HENRY, C. S.—The unique purring song of Chrysoperla comanche (Banks), a green lacewing
of the rufilbaris species group (Neuroptera: Chrysopidae) ............0.00.0000 000 ee eee 133
HEYDON, S. L.—A review of the world species of Notoglyptus masi (Hymenoptera: Pter-
omalidae) TA SRE en hn) eS a ee SSO ete eee OS eR AP Oe 112
HURD, L. E. and R. M. EISENBERG—A mid-summer comparison of sizes and growth rates
among nymphs of three sympatric mantids (Mantodea: Mantidae) in two old-field habitats 51
HURD, L. E. and R. M. EISENBERG—Temporal distribution of hatching times in three
sympatric mantids (Mantodea: Mantidae) with implications for niche separation and co-
EXIStENGe) 221... Senne a. deapasie )SaEa fey Bud auavaysr Macnee shot tece ce DRE APD Se accep sae eae 55
VOLUME 91, NUMBER 4 661
HURD, L. E., P. E. MARINARI, and R. M. EISENBERG—Influence of temperature and
photoperiod on early developmental rate of Tenodera sinensis Saussure (Mantodea: Man-
[BCC E: YS J Pere te eee eRe cd tN Coe RCTS ee EER EP ene IEE 529
HUSBAND, R. W.—Two new species of Coccipolipus (Acari: Podapolipidae) parasites of
Chilocorus spp. (Coccinellidae) from Vera Cruz and Morelos, Mexico and Florida and Wis-
CODSID ES CAC aero ore nes Renee toes are cone aaa acteee tarot te venan’ asec nego eveesls Cray Meine a eT vera 429
IRWIN, M. E.—See WEBB, D. W.
JACKSON, D.—See ROCK, E. A.
KELLEY, R. W.—New species of micro-caddisflies (Trichoptera: Hydroptilidae) from New
Caledonia Vianvatikan ebiy ty sarc reuun sails caste veetees Aas as seeps Seite er te nae ane 190
KIRCHNER, R. F.—See YOUNG, D. C.
KNISLEY, C. B., D. L. REEVES, and G. T. STEPHENS—Behavior and development of the
wasp Prerombrus rufiventris hyalinatus Krombein (Hymenoptera: Tiphiidae), a parasite of
larval tigeribeetles:(GoleopterariGicindelidae)) ..5 .. 10.2 nsec ocntte es oscesuis eevee see sau. 179
KONDRATIEFF, B. C.—See YOUNG, D. C.
KROMBEIN, K. V.—Systematic notes on some Bethylidae from Botswana: Pristocerinae (Hy-
MEDOD elas ACULCata) ieamune lagers cena safle shee asc atenaraversi ial eta ata eveoeesst a tuahala dats ci cystetens ue eee 620
LAGOY, P. K. and E. M. BARROWS-—Larval-sex and host-species effects on location of
attachment sites of last-instar bagworms, Thyridopteryx ephemeraeformis (Lepidoptera: Psy-
(2 01 (0 F: Y=) Jel oer RE PAR PPS eae ee to DP Se 468
LASALLE, J.—Notes on the genus Zagrammosoma (Hymenoptera: Eulophidae) with descrip-
Lomo frame wISDCClESS cacicam aca Heme crt tees ine ase oya areata oe Santas sigeettas Ete 230
LEE, K. M., W. W. WIRTH, and K. L. CHAN—A new species of Dasyhelea midge reared
from drains in Singapore (Diptera: Ceratopogonidae) ............00.000 000000 eee ee 452
LEIDY, N. A. and H. H. NEUNZIG—Taxonomic study of the larvae of six eastern North
American Dioryctria (Lepidoptera: Pyralidae: Phycitinae) ..............000.00..000000.. 325
LEIDY, N. A.—See NEUNZIG, H. H.
MacDONALD, J. F.—Review of Nearctic Metachela Coquillet, with description of a new
species (Diptera: Empididae; Hemerodromiinae) ..............0. 0000000 cee eee ee eee 513
MacKAY, W. P. and S. B. VINSON—Two new ants of the genus Solenopsis (Diplorhoptrum)
from eastern’ Texas (Hymenoptera: Formicidae) .. 22.5.2 .02css.cscecee ec eec eee senses 75}
MANGUIN, S.—Sexual dimorphism in size of adults and puparia of Tetanocera ferruginea
Fallén (Diptera: Sclomyzidae) ....... Pe iretnc dee Sem oon elo Bern rea * 523
MARINARI, P. E.—See HURD, L. E.
MARSH, P. S.—Notes on the genus Hybrizon in North America (Hymenoptera: Paxylom-
Wate ae) Mee. ae eesti vices tea esis ralata a, so ce grace, eae oke outs 9 wad ce Beer lous mein eeGe.to em hereto a= bradiore ds 29
MARSHALL, S. A.—See WHEELER, T. i
MARTINS, R. P.—See FERNANDES, G. W.
MILLER, S. E. and V. O. BECKER—North American moths described by L. A. G. Bosc
D’Antic (Lepidoptera: Noctuidae, Pyralidae) ........ 0.2.6 eee 22
MILLER, T. D.—First Nearctic record of the genus Nordlanderia (Hymenoptera: Eucoilidae),
with descriptions of two new species ........0..00.0 0000 eee Bone Acc seeveet fend : 158
MOSER, J. C.—See WEATHERBY, J. C.
MUNROE, E.—See SHAFFER, J. C.
NAKAHARA, S.—A new species of A/eurotulus (Homoptera: Aleyrodidae) ............... 436
NEUNZIG, H. H. and N. A. LEIDY—A new species of Dioryctria (Lepidoptera: Pyralidae:
Phycitinae) from the southeastern United States .......................... 321
NEUNZIG, H. H.—New records of Phycitinae from Bolivia melden new species of Peagus
(Lepidoptera: Pyralidae) . haat pirceke 355
NEUNZIG, H. H.—See LEIDY, N. A.
NICKLE, D. A. and M. S. COLLINS—Key to Kalotermitidae of eastern United States with a
new Neotermes from Florida (Isoptera) : ; Ane Mae aN acide ouimiee n eavetone LOO.
PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
PAOLETTI, M. G. and B. R. STINNER—Two new terrestrial Isopoda (Oniscidea) from cor-
allineicaysiof Venezuela's Caribbeani coast) +--+ 22. 2es ores se eeiecs se telecon
POLHEMUS, J. T. and P. J. SPANGLER—A new species of Rheumatobates Bergroth from
Ecuador and distribution of the genus (Heteroptera: Gerridae) .........................
PRATT, J.—See STRICKMAN, D.
PRICE, R. D.—See HELLENTHAL, R. A.
PURRINGTON, F. F. and D. H. STINNER—Wasp parasites of the burdock seed moth,
Metzneria lappella Zeller (Gelechiidae): New host record for Bracon mellitor Say (Hyme-
noptera: BraGonidae) ae tresses ses sous slseistas rotor eats seve cele ereremeloie miele atetsieclatels
REEVES, D. L.—See KNISLEY, C. B.
ROCK, E. A. and D. JACKSON — Naturally occurring host sites for xylophilic Cecidomytidae
(Diptera) A Seer sara trsrete teeta evar stelsvavadetarere ornate cts Sucrauan trey eve arane sav camara are orf ala: elabemenay ater Ace tMa
ROTH, L. M.—The cockroach genus Margattea Shelford, with a new species from the Krakatau
Islands, and redescriptions of several species from the Indo-Pacific region (Dictyoptera:
Blattaria) Blattellid ae) Wyseter: cic ac eee cos SE ernest Es eS he ee
ROTH, L. M.—Siiferia, a new ovoviviparous cockroach genus (Blattellidae) and the evolution
ofovovivipanitysiniBlattana:(Dictyoptera)) tert creer tees 12) wots Rtciceede vein tetee et siateh eetomets
SALLUM, M. A. M.—See FORATTINI, O. S.
SCHAUFF, M. E.—A new species of Horismenus (Hymenoptera: Eulophidae) parasitic on the
lesser cornstalk borer, E/asmopalpus lignosellus (Lepidoptera: Pyralidae) ...............
SCHWARTZ, M. D.—Polymerus castilleja, anew mirine plant bug from California and Oregon,
with remarks on generic characters of the genus Po/ymerus Hahn (Heteroptera: Miridae) . .
SHAFFER, J. C.—Two new phycitine moths of the genus Coenochroa (Lepidoptera: Pyralidae)
AKOMMMBLAZI ener cress eakeche Rts ene Sos tLe tee ee eee Me merge ore ee te ete
SHAFFER, J. C.—Review of Goya Ragonot and description of a new species, G. simulata
irom Parana, Brazil (Lepidoptera: PyralidaesReominae)! = aan-. vec teenie sees «ee eile
SHAFFER, J. C. and E. MUNROE—Type material of four African species of Notarcha Meyrick,
with designations of lectotypes and changes in synonymy (Lepidoptera: Crambidae: Pryaus-
THOS) Peg eee A re ey he Te ot PR Se NE CeO ENE CORE Ace iM CASS oo Re Bl ic
SHAFFER, J. C. and E. MUNROE—Type material of two African species of Herpetogramma
and one of Pleuroptya (Lepidoptera: Crambidae: Pyraustinae) .....................0....
SPANGLER, P. J.—A new species of Neotropical water bug, Paravelia biae, from Brazil
(HeteropteraavVielitdae)) © «2.2%. cicusts et ovctaed chatna 4 attccracatanemercbss Nav ontatche eae Come remo atie ee ere one
SPANGLER, P. J.—See POLHEMUS, J. T.
SPONSLER, R. C.—See APPEL, A. G.
STAINES, C. L., Jr.—A new genus and species of Hispinae (Coleoptera: Chrysomelidae) from
Gen tralvAMericare ete scbececch:2hs.c, 5 s.cuesanebnipsenn e ratduclefelthate eiende ova ae Pear eerie Sees ae
STARR, C. K.—The ins and outs of a tropical social wasp nest ............. Est oke cg Bran
STEPHENS, G. T.—See KNISLEY, C. B.
STINNER, B. R.—See PAOLETTI, M. G.
STINNER, D. H.—See PURRINGTON, F. F.
STRICKMAN, D. and J. PRATT—Redescription of Cx. corniger Theobald and elevation of
Culex (Culex) lactator Dyar and Knab from synonymy based on specimens from Central
Americal (Dipteras: Cullicic ae) l Ryser: 3 sxe «aes ts ais aucsatte aveiehecsie ose « Weeecteta shee hated aes eee
TOGASHI, I.—Japanese sawflies of the family Blasticofomide (Hymenoptera: Symphyta) .
TOWNES, H. K.—North American species of S¢i/bops (Hymenoptera: Ichneumonidae)
TRIPLEHORN, C. A.—A new species of Neobaphion Blaisdell, from Idaho (Coleoptera: Te-
Nebrionidae)iiz.49- ese seh ee ee eis
VINSON, S. B.—See MAcKAY, W. P.
WARTHEN, J. D., Jr.—Neem (Azadirachta indica A. Juss): Organisms affected and reference
listrupoate: anit shh il ste ke A ee RE pec A on” ml Pre diesen AA Soe oN ook
WEATHERBY, J. C., J.C. MOSER, R. J. GAGNE, and H. N. WALLACE—Biology of a pine
needle sheath midge, Contarinia acuta Gagné (Diptera: Cecidomyiidae), on Loblolly pine .
71
421
203
66
206
441
367
346
VOLUME 91, NUMBER 4
WEBB, D. W. and M. E. IRWIN—Genera Ataenogera and Phycus in the New World (Diptera:
Hiberevad ae sieMy GLa) ecanase sea teyaieswaratenny ese beaj abet ati aynevetsas iets sverenetay peice erste ran screen ye caver te
WHARTON, R. A.—Final instar larva of the embolemid wasp Ampulicomorpha confusa
(Eliyvimenoptera)) cena wan oaeret ite sat neces 45 vata cave eit Otis mcrae era vente Zeta aan eee me
WHEELER, A. G., Jr.—Grypotes puncticollis (Homoptera: Cicadellidae), a Palearctic pine-
feeding leafhopper new to North America .... 0.0.0.0 0 200
WHEELER, A. G., Jr.—Megalonotus sabulicola (Heteroptera: Lygaeidae), an immigrant seed pred-
ator of Centaurea spp. (Asteraceae): Distribution and habits in eastern North America .......
WHEELER, T. A. and S. A. MARSHALL—<Apilotus martini, a new species of the Apilotus
beckeri group (Diptera: Sphaeroceridae) from caves in the Canary Islands ..............
WIRTH, W. W.—See LEE, K. M.
YOUNG, D. C., B. C. KONDRATIEFF, and R. F. KIRCHNER—Description of male Os-
trocerca Ricker (Plecoptera: Nemouridae) using the scanning electron microscope ....
ZAPPAROLI, M.—Notes on Pleurolithobius of Turkey (Chilopoda: Lithobiomorpha) ......
ZEIGLER, D. D.—Drumming behaviors of three Pennsylvania stonefly (Plecoptera) species . .
NOTES
APPEL, A. G.—Rapid and non-destructive gender determination of nymphal and adult Cryp-
tocercus punctulatus Scudder (Dictyoptera: Cryptocercidae) ............00..0.00000000..
BAKER, W. W.—See WILSON, N.
CARMEAN, D.—Sex and deposition of the holotype of Bareogonalos canadensis (Harrington)
(ivinenopteras dp onalyidae) Wem. yer cere ce rer etree tee retest cr ae trae cect nee
CURCIC, B. P. M. and R. N. DIMITRIJEVIC—Pedipalpal anomalies in Neobisium simoni
(L. Koch) and N. bernardi Vachon (Neobisiidae: Pseudoscorpiones: Arachnida) ..........
DIMITRIJEVIC, R. N.—See CURCIC, B. P. M.
HALSTEAD, J. A.—Hymenoptera associated with a California population of the Russian thistle
biological control agent, Coleophora klimeschiella Toll (Lepidoptera: Coleophoridae) ... .
HOFFMAN, K. M.—Taxonomic status of Mantispa sayi, Mantispa fuscicornis, and Mantispa
unleri (Neuroptera: Mantispidae)..2......-.s.6a0ee.6..0ee caves len sseeuee cosas
LASALLE, J. and J. A. MCMURTRY~—First record of Thripobius semiluteus (acienopie
Eulophidae) from the New World ...............0 000 cece cece eee eee ee
MacKAY, W. P. and S. B. VINSON—Rediscovery of the ant Gnamptogenys hanmaat ie
menoptera: Formicidae) in Eastern Texas .................... ey ore retens Sve ee
McMURTRY, J. A.—See LaSALLE, J.
ROBBINS, R. G.—Ticks of the subgenus /xodiopsis: First report of Ixodes woodi from man
and remarks on /xodes holdenriedi, a new junior synonym of Ixodes ochotonae (Acari:
TXOGIAAS) Sve eee ost ie nee ies HbA ae eee aarti ae
WILLIAMS, C. E.— Black walnuts Taelans nigra: A new host ccord for Colaspis fiosd (Co-
leopterasi@hrvsomelidae) Gere. series 2 ence euesele de <rcue stele ele ueteneterele cece eeu ier ;
WILSON, N. and W. W. BAKER— Teaties aay nsi (Acar edie) or Peru ...
BOOK REVIEWS
ADAMSKI, D.—The Guild Handbook of Scientific Illustration
BURGER, J. F.—Outlines of Entomology
DAVIS, D. R.—Lepidoptera Anatomy .
GAGNE, R. J.—Asa Fitch and the Emergence of . a ierean Eniomolony
GERBERG, E. J.—Silent Spring Revisited
McCAFFERTY, W. P.—The Mayflies of Florida, Revised! Edition”
NORDEN, B. B.— Bees of the New Genus Ctenoceratinia in Africa South of the Sahara ( cee
menoptera: Apoidea)
257
389
583
286
640
641
490
293
294
493
296
124
664 PROCEEDINGS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
PAPE, T.—Ecological Methods with Particular Reference to the Study of Insect Populations,
SecondlRevised VEGI omg di ayircsteretecctseue cic ays she ate ber eseve eres evar celsuetocn ere neen Meroe te aero 126
PICKENS, L. G.—Control of Arthropod Pests of Livestock: A Review of Technology ........ 125
ROBBINS R.G:— ihe: FleasiopithesPacificsNorthwest) ss. andes sac esseaen eee oe eee 486
STEYSKAL, G. C.—Catalog of the Heteroptera, or True Bugs, of Canada and the Continental
UNILTEAIS TALES a2 inp, SAR es, bein avec fe aad ee eee 649
STEYSKAUHGHEGOINOCiiuIdae waa oak era CE een Dee EERE? Peer 649
WHEELER; O8Di—Insect=FungusclnteractiOns cre se teieiele)sie~ -rretaie erate) oloketete) eereete rita 645
OBITUARIES
WALLENMAIER, T. E.—Frederick William Poos, Jr. 1891-1987 ...................0005- 298
WHITES RSE:—Donaldoyce Borror l907=19 88m e ete mcr oe ete rere seitoeieaittle emieeeeteree 304
MISCELLANEOUS
ENDANGERED! SREGIBSIN © iI GE Are cies ca. ceseetecepeneps cnet sieredce teens aecese omctctatsfetercyaynystseeeesaosters 496
INSTRUCTION TOYAUTHORS: . 72.25. .) : PO ICR Ae nee a RISO a 318
MEMBERSHIPAIS Taisen cso cecuoiertacen acter cre aac aore noe arcuate ice reais keine cena ie 311
NEW MEMBERS FOR: 1988 2... 05 h.0.0.00 Bee oer rae ONERS Ach to keek coe CR MORE ee TE 306
SOCIETY MEETINGS AND REPORTS OF OFFICERS FOR 1988 ...............2..08+ 307
SOCIETY@MEEMINGS & x, chierreeta ccs caer n tetera i onmen tiatanrenh nome eroremets Papen aeoitisc rs “n 651
TABLE OF CONTENTS, VOLUME 90 .. : A ROTC TOA SOOT OCG 128
TABLE OF CONTENTS, VOLUME 91 ees Leste ho ae ee ene O5 9.
PUBLICATIONS FOR SALE BY THE
ENTOMOLOGICAL SOCIETY OF WASHINGTON
MISCELLANEOUS PUBLICATIONS
Gynipid| Galls of the Easter United ‘States; by Lewis Ho Weld ===" 2 as ee ee
(GynipidiGallssof the'Southwestiby, Mewis: Hy Weldt ae see ee ee eee
Botwupapers onicynipidvpalls 26 eee te ee ee Rien A oe a Se ee
Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman aoe eee
Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R.
Traver.
A Short History of the Entomological Society of Washington, by Ashley B. Gurney.
Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C.
TSS Kall es ee eer Or Pee ROR Bi. cee Mee eS a ae ON BE ae
Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C.
Steyskal
MEMOIRS OF THE ENTOMOLOGICAL SOCIETY OF WASHINGTON
No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939...
No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving.
SISSY 0) aa CY) Se a a a
No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman.
PAoyshiys) oe CY: poe NE eg et eee ont Se ae ES ah Ae eed ee ne ee
No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952
No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp.
NYS) pe eee eee Eee Re LT ee te te ee ee ee
No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi
diakahast-250);ppi 9 G9 te sme eee oo eee Fae, WE ee ee ee ee 6 hte
No.7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp.
NEFF G eo es sate te Seis hak eter ie OR re DO a ER, ee ee: Se AS on RE tl BE
No. 8. The North American Predaceous Midges of the Genus Pa/pomyia Meigen (Diptera: Cera-
topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 occ ccccccccsseeseeeneeeeeee a:
No. 9. The Flower Flies of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200
TD a SS Ne ee ee ge A ere ee ee Pe ee
No. 10. Recent Advances in Dipteran Systematics: Commemorative Volume in Honor of Curtis W.
Sabrosky. Edited by Wayne N. Mathis and F. Christian Thompson. 227 pp. 1982.00.00...
No. 11. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp.
No. 12. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff.
OTEK Wa eee Le ee ee pee Eee eal Ee A ee 8 co ee
No. 13. An Identification Manual for the North American Genera of the Family Braconidae (Hy-
menoptera), by Paul M. Marsh, Scott R. Shaw, and Robert A. Wharton. 98 pp. 1987
11.00
12.00
10.00
11.00
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Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume
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Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10
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the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian
Institution, Washington, D.C. 20560.
CONTENTS
(Continued from front cover)
TOWNES, H. K.—North American species of Sti/bops (Hymenoptera: Ichneumonidae)
WHARTON, R. A.—Final instar larva of the embolemid wasp Ampulicomorpha confusa (Hy-
menoptera)
WHEELER, A. G., Jr.—Megalonotus sabulicola (Heteroptera: Lygaeidae), an immigrant seed
predator of Centaurea spp. (Asteraceae): Distribution and habits in eastern North America
WHEELER, T. A. and S. A. MARSHALL—<Apilotus martini, a new species of the Apilotus
beckeri group (Diptera: Sphaeroceridae) from caves in the Canary Islands
ZEIGLER, D. D.—Drumming behaviors of three Pennsylvania stonefly (Plecoptera) species
NOTES
HALSTEAD, J. A.—Hymenoptera associated with a California population of the Russian thistle
biological control agent, Coleophora klimeschiella Toll (Lepidoptera: Coleophoridae) ....
HOFFMAN, K.M.—Taxonomic status of Mantispa sayi, Mantispa fuscicornis, and Mantispa
uhleri (Neuroptera: Mantispidae)
LASALLE, J. and J. A. MCMURTRY —First record of Thripobius semiluteus (Hymenoptera:
Eulophidae) from the New World
WILLIAMS, C. E.—Black walnut, Juglans nigra: A new host record for Colaspis favosa (Co-
leoptera: Chrysomelidae)
BOOK REVIEWS
ADAMSKI, D.—The Guild Handbook of Scientific Illustration
STEYSKAL, G. C.—Catalog of the Heteroptera, or True Bugs, of Canada and the Continental
United States
STEYSKAL, G. C.— Noctuidae
WHEELER, Q. D.—/Jnsect-Fungus Interactions
MISCELLANEOUS
SOCIETY MEETINGS
TABLE OF CONTENTS, VOLUME 91
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