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VOL. 91 


NO. 1 

(ISSN 0013-8797) 


of the 





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- 

ihonomus Germar (Coleoptera: Curculionidae) 88 

ELLSBURY, M. M. and G. T. BAKER — Biological and morphological aspects of the egg stage 

and oviposition of Languria mozardi (Coleoptera: Languriidae) 9 

sociated with a coleopterous leaf-bud gall on Xylopia aromatica (Annonaceae) 81 

HANSSON, C — New World species of Hokopelte and lonympha (Hymenoptera: Eulophidae), 

with descriptions of two new species 59 

HELLENTHAL, R. A. and R. D. VKICE-Geomydoecus (Mallophaga: Trichodectidae) from 

the Texas and desert pocket gophers (Rodentia: Geomyidae) 1 

HEYDON, S. L. — A review of the world species of Nologlyptus masi (Hymenoptera: Pterom- 

alidae) 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- 

tidae) 29 

M ILLER, S. E.andV.O. BECKER - North American moths described by L. A. G. Bosc D' Antic 

(Lepidoptera: Noctuidae, P^ralidae) 22 

PAOLETTI, M. G. and B. R. STINNER— Two new terrestrial Isopoda (Oniscidea) from coralline 

cays of Venezuela's Caribbean coast 71 

ROCK, E. A. and D. JACKSON — Naturally occurring host sites for xylophilic Cecidomyiidae 

(Diptera) 66 

STARR, C. K.— The ins and outs of a tropical social wasp nest 26 

(Continued on back cover) 




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Title of Publication: Proceedings of the Entomological Society of Washington. 

Frequency of Issue: Quarterly (January. April, July, October). 

Location ofOffice of Publication, Business Oflice of Publisher and Owner: The Entomological Society of Wash- 
ington, c/o Department of Entomology, Smithsonian Institution, lOth and Constitution NW. Wash- 
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91(1), 1989, pp. 1-8 




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 truncatiis Wemeck and G. quadridentatus Price anS Emerson 
are redescribed and illustrated. The new species G. neotruncatus is described, with the 
type host being Geomvs personatus streckeri Davis. / t n m -i 

Key Words: lice, Geomys arenarius, Geomys personatus \ 

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 texamis 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- 

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- 


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 (ieomydoecus lice are 
included in Hellenthal and Price (1980). 

Geomydoecus truncatus Werneck 

Figs. 1-6 

Geoiuydoecus truncatus Wenxcck, 1950; 13. 
Type host: Geomys personatus personatus 

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 (SM W) 0. 1 1 0-0. 1 25 (23: 0.119 
± 0.0054), scape distal width (SDW) 0. 1 1 0- 
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; 11, 12-16 (26 
14.0 ± 1.18); III, 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: 1 1.5 
± 1.30); IV, 1 1-15 (26: 13.2 ± 1.08); V, 8- 
12 (25: 10.1 ± 1.15); 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 ± 0.05 1 1 ). 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 ± 

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 
± 0.003 1 ) long, respectively, with submar- 
ginal seta positioned near inner marginal 
seta (Fig. 2). PW 0.340-0.385 (23: 0.356 ± 
0.0 1 1 9). Abdominal tergal setae: I, 2; II, 14- 
18 (23: 16.3 ± 1.05); III, 20-25 (23: 22.4 
± 1.38); IV, 23-30 (23: 26.2 ± 1.95); V, 
24-28 (23: 25.6 ± 1 .08); VI, 2 1-25 (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. 1 00. 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 ( 1 9: 
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- 
13 (23: 1 1.5 ± 1.04); IV, 12-17 (22: 14.4 
± 1.50); V, 10-14 (22: 12.0 ± 1.21); VI, 8- 
13 (22: 10.8 ± 1.60); VII, 6-10 (22: 7.7 ± 
1.16). Subgenital plate with 1 8-23 (23:21.2 
± 1 .53) setae, with distribution and lengths 
as in Fig. 1, with 1 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 it 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 is 


Figs. 1-6. Gcoinydoirus Iruncalus. 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 \ entral genitalia. 6, Male dorsal (left) — 
ventral (right) view. Measurements are in millimeters. 


easily distinguished from all other Gcomyd- 
oecus by its uniquely shaped parameral arch; 
no other described species of this genus has 
the distinctive mediopostcrior flattening. 
The female is not as readily diflbrenliated, 
but the combination of the genital sac struc- 
ture, dimensions, and chaetotaxy features 
should separate it. 

Wcrneck (1950) described G. truncatus 
from a series of six males taken o^ Geomys 
pcrsonatus from Padre Island, Texas. This 
locality would make the host G. p. pcrsona- 
tus. the only pocket gopher that Hall ( 1 98 1 ) 
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. iriin- 
catus may also occur there, but, conversely, 
we arc 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 Gconiys p. strcckeri 
Davis and the other from G. p. fallax Mer- 
riam. Price and Emerson (1971) had spec- 
imens only from G. p. strcckeri and named 
them Geomydoecus truncatus. Numerous 
subsequent collections from Gconiys p. fal- 
lax and the determination that these were 
different from the G. p. strcckeri 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 scries from 
Gcomys p. fallax. 

Gconiys p. fallax also has Geomydoecus 
texanus texanus occurring on it. Of the six 
gophers of this host taxon that yielded G. t. 
texanus. only one also had G. truncatus. This 
one gopher had 12 G. truncatus and only 
one specimen of G. t. 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 

Material examined. — 2 3, Paratypes of 

Geomydoecus truncatus. ex Gcomys pcrso- 
natus. Padre Island, Texas; 53 9, 57 <J, ex G. 
p. fallax, 9 gophers from 7 localities in San 
Patricio Co., Nueces Co., and Live Oak Co., 

Geomydoecus neotruncatus 
Hellenthal and Price, Nkvv Species 

Type host: Gcomys pcrsonatus strcckeri 

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 stemite II, 7-1 1 
(20: 9.1 + 1.02); VI, 8-12(19: 9.5 ± 0.90). 
Genitalia PAW 0.140-0.155 (20: 0.145 ± 

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, 8-1 1 (20: 9.7 ± 0.91); V, 
10-15(20: 13.0 ± 1.10); VI, 9-14 (20: 12.3 
± 1.22); VIL 8-1 1 (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 arc the temple width 0.435 (0.034), 
prothorax width 0.320 (0.085), and scape 
length 0.184 (0.109). For females, the best 


are temple width 0.466 (0.009), prothorax 
width 0.343 (0.081), and setae on stemite 
VII 8.69 (0.177). 

The males of both species key to G. trun- 
catiis in the first half of couplet 6 in Timm 
and Price ( 1 980), where G. neotmncatus 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. quadridentatiis 
Price and Emerson in couplet 9. Temple 
width under 0.466 and prothorax width un- 
der 0.343 will distinguish G. neotnincatiis 
from (/'. Iruncatus; both may be separated 
from G. quadridentatiis 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 9, ex Geo- 
niys personatus slreckcri. 14 mi (22.5 km) 
W Crystal City, Zavala Co., Texas, 
9.II.1953, KU-52238; in collection of the 
University of Kansas. Paratypes, ex G. p. 
streckeri: 9, 7 S, same as holotype; 22 9, 17 
S, same except KU-52239 or 10.11.1953, 
KU-52245, 52246; 13 9, 11 6, E Carrizo 
Springs, Dimmit Co., Texas, 4.1.1970, TT- 
9665, 9666; 6 9, 4 3, 13 mi (20.9 km) N or 
NE Carrizo Springs, Dimmit Co., Texas, 
17.1.1970, TT-10126, 10131; 1 9, I mi (1.6 
km) SW Carrizo Springs, Dimmit Co., Tex- 
as, 23. V. 1974, TAM-27613; 5 9, 4 <?, Car- 
rizo Springs, Dimmit Co., Texas, 
24.XI.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. 7-1 1 

Geomydoecus quadridentatus Price and 
Emerson, 1971: 240. Type host: Geoniys 
arenarius arenarius Merriam. 

Male.— Grossly as in Fig. 6. except an- 
tenna as in Fig. 10, and dorsal abdomen as 

in Fig. 1 1. 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); III, 14-23 (78: 18.7 ± 1.65) 
IV, 17-27 (78: 21.1 ± 2.03); V, 16-26 (78 

19.5 ± 1.90); VI, 1 1-19 (78: 15.0 ± 1.54) 
tergal and pleural setae on VII, 15-24 (80 
20.2 ± 1.69). Abdominal sternal setae: II, 
9-15 (79: 1 1.7 ± 1.49); III, 1 1-17 (79: 13.9 
± 1.39); IV, 11-19(80: 14.3 ± 1.62); V, 8- 
14 (80: 10.8 ± 1.42); VI, 6-1 1 (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 ± 

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 ± 0.005 1 ) and 0.040-0.050 (78: 0.045 
± 0.0036) long, respectively. PW 0.280- 
0.345 (80: 0.3 1 1 ± 0.0 1 20). Abdominal ter- 
gal setae: I, 2; II, 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, 1 6-26 (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, 0.085-0.1 15 (80: 0.102 ± 0.0069), 
with 0-6 (80: 0.9 ± 1.44) of these longer 


Figs. 7-1 1. Geomydoecus quadridenlatus. 7, Female dorsal abdomen. 8, Female genital sac. 9, Male ventral 
genitalia. 10, Male ventral antenna. 1 1. Male dorsal abdomen. Measurements arc in millimeters. 

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- 
16 (79: 11.9 ± 1.77); III, 11-17 (77: 14.3 
± 1.26); IV, 1 1-19 (79: 15.0 ± 1.75); V, 8- 
16 (78: 1 1.8 ± 1.51); VI, 7-12 (78: 9.6 ± 
1.02); VII, 6-1 1 (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 ± 0.01 15), with 0-4 (79: 2.1 ± 0.82) 
curved medioanterior loops, posteriormost 
loop, when present, situated 0.040-0.105 
(78: 0.07 1 ± 0.0 1 1 5) back from antenor sac 

Discussion. — Both sexes of (7. quadriden- 
latus are easily separated from G. truncatus 

and G. neotruncatus. Males of G. quadri- 
denlatus have conspicuously different gen- 
italia (Fig. 9 vs. Fig. 5), the antennal scape 
with a posterior process (Fig. 1 0), and dorsal 
abdominal chaetotaxy (Fig. 1 1) 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. 1 2). 

As originally described by Price and 
Emerson (1971), males of G. quadridentatus 
were said to have only four large genital sac 


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 6'. cjiiadndentatus 
studied. The presence of these smaller spines 
should not complicate proper identification, 
if other characters and host association are 

There is discussion among mam malogists 
as to whether Gcomys arcnaniis is a valid 
species apart from G. bursarius (Shaw). Also 
uncertain arc the relationships among up to 
five populations ofGeoniys 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. hrcvirosfris Hall; 4) a "river" popu- 
lation of gophers belonging to G. a. arc- 
narius; and 5) an "upland" population of 
G. a. arcnarius. 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 misidcntification, 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 Gcomys bursarius to support 
G. arcnarius as a separate taxon and suffi- 
ciently similar to each other to group all five 
gopher populations into G. arcnarius. 

Material examined. — 244 9, 267 S, ex 
Geomys arenarius arcnarius, 50 gophers 
from 1 7 localities in New Mexico, Texas, 
and Chihuahua; 67 2, 5 1 <5, ex G. a. hrcviros- 
tris. 1 5 gophers from 6 localities in New 



Fig. 12. Geographic distribution of OV'(W)n(/"f«(,v 
Iruihciliis (closed circles), (/. ncolnincaliis (open cir- 
cles), and G. quadndentalus (triangles). 

Mexico; 26 9, 22 3, ex Geomys, 7 gophers 
from 2 localities near Gran Quivera, New 
Mexico; 35 9, 28 3 ex Gcomys, 4 gophers 
near San Antonio, New Mexico. 


We wish to thank Sebastiao J. de Oli veira. 
Curator, Fundacjao Oswaldo Cruz, Rio de 
.laneiro, Brasil, for the loan of the Werneck 
type scries. Partial support for this study 
was supplied by a grant from the National 
Science Foundation to the University of 
Notre Dame (Grant No. BSR86- 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- 

Literature Cited 

Hall. t:. R. 14X1. The Mammals of North America. 
2nd Ed. Vol. \. John Wiley & Sons, New York, 
NY. XV + 600 + 90 pp. 

Hellenthal, R. A. and R. D. Price. 1980. A review of 
the (ic(ii»ydoccus suhcalifonucus Complex (Mal- 
lophaga: Trichodectidae) from rliDiuoniys pocket 
gophers (Rodcntia: Geomyidac). with a discussion 
of quantitative techniques and automated taxo- 


nomic procedures. Ann. Enlomol. Soc. Amcr. 73: 

. 1984. Disliihiilional associations among 

Geoinydoecus and I'hoinninydoccus lice (Mal- 
lophaga: Trichodcctidac) and pocket gopher hosts 
of the Thotnoiiiys bullae group (Kodentia: (ieo- 
niyidae). J. Med. Entomol. 21: 4.32^46. 

Price, R. D. 1975. The 6'«;/m'^«rM.v (Mallophaga: 
Trichodectidae) of the southeastern USA pocket 
gophers (Rodentia: Cieomyidae). Proc. I^ntomol. 
Soc. Wash. 77: 61-65. 

Price, K. 1). and K. C. Emerson. 1971. A revision of 
the genus (Icoinydoccus (Mallophaga: Trichodec- 
tidae) of the New World pocket gophers (Rodentia: 
Cieomyidae). .1. Med. Entomol. 8: 228-257. 

Price, R. D. and R. A. Hellenthal. 1975. A review of 
the Geoinydoecus Ic.xanu.s complex (Mallophaga: 
Irichodectidae) from (icimiys and I'appoiieninys 
(Rodentia: (ieomyidae)..l. Med. Entomol. 12:401- 

Timm, R. M. and R. D. Price. 1980. The ta.xonomy 
of Cieomydoecus (Mallophaga: Trichodectidae) 
from the Cieomys Iniisanus complex (Rodentia: 
Geomyidae). J. Med. Entomol. 17: 126-145. 

Werneck, F. L. 1950. Os Malofagos de Mamiferos. 
Parte II: Isehnocera (eonlinuavao de Trichodec- 
tidae) e Rhyncophthirina. Ed. do Inst. Osw. Cru/., 
Rio de .laneiro. 207 pp. 

91(1). 1989, pp. 9-14 




Michael M. Ellsbury and Gerald T. Baker 

(MME) USDA-ARS. Crop Science Research Laboratory, Forage Research LInit, 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 T. repens L., T. pratense L., T. hybridum L., T. resupinatian 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: Languria mozardi. sensilla, larva, mouth parts, antenna 

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 saliva 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, T. vesiculosum 

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 



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 T. repens 
L. (white clover), T. prateme (red clover), 
T. hybridum L. (alsike clover), T. resiipi- 
natitm L. (persian clover), T. striatum L. 
(pitts or striate clover), T. nigrescens Viv. 
(ball clover), 7". lappaceum L. (lappa clo- 
ver), and T. 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 intemodal location of ovi- 
position punctures was calculated for clo- 
vers in the host range study from integer 
values assigned to each intemode beginning 
with the lowermost intemode (#1) and in- 
creasing to the (last) intemode 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 in the same 
buffer and then post-fixed in 4% OsOj 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 ovemight. Six treated and 6 un- 
treated females were placed in petri dishes 
containing pieces of flowering clover stems 
with the second and third intemodes. The 
specimens were examined every 1 minutes 
for 1 h and the experiment was replicated 
3 times. 


Frequency distribution by intemode of 
oviposition sites on crimson clover con- 
taining 1 or 2 eggs is illustrated in Fig. 1. 
Oviposition punctures were most frequent 
(22/63 and 18/63, respectively) in the 2nd 
and 3rd intemodes of primary flowering 
stems. No eggs were deposited in lateral 
flowering stems. Of 78 stems examined, 1 1 
had 2 oviposition punctures and 4 1 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 (.V ± SE, N = 78). 

Oviposition by L. mozardi in 1 1 species 
of clovers during a no-choice test is sum- 
marized in Table 1. Lappa clover, T. lap- 
paceum. was the only species in which eggs 
were not deposited. The usual number of 
eggs per site is 1 or 2 for all clovers studied, 
although 3 eggs occurred in a single punc- 
ture on T. nigrescens and 4 eggs in a punc- 
ture on a stem of T. 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, T. pra- 

Behavior of the adult female during ovi- 



2 Eggs per Puncture 

5 10 15 20 -^ 


Fig. 1. Iniernodal frequency dislnbulion of Lan- 
guria mozardi egg punctures containing 1 or 2 eggs in 
stems of crimson clover. Tnfolium incarnaltim. 

position is similar on all Tnfolium 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 is 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 ^m (480- 

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. 20. 

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 ^m (69-74 nm) long and 20 
^m (18-21.5 ^lm) wide and lOsensilla trich- 
odea are on the apex (Fig. 3b, d). These 
sensilla vary in length from 23.5 ixxn (21- 
25 jxm) to 98 ixm (95-101 ^m) and they are 
either slightly curved or straight (Fig. 3d). 
Another group of 9 sensilla trichodea are 
situated on the eclal surface of each coxite 
near the base of the style (Fig. 3a, c). These 
trichoid sensilla range in length from 1 9 ;um 
(17.5-21 urn) to 71.5 Mm (69-74 Mm). 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 

Table 1. Frequency and internodal sites of oviposition by Languna mozardi in eleven Tnfolium species 
dunng a no-choice test. 

Ave Intemodal Location 

f'niolnirn Specii's' 

No Stems 

Eggs/Stem - 

Mean No. Nodes/Stem 

o1 tlviposuion Punctures 

* ■/■. subtcnancum 


.47 ± .16 

6.42 ± .55 

4.67 ± .69 

T. lappaceum 


9.14 ± .60 


*T. nigrescens^ 



7.00 ± .32 


*T. resupinatwn 


.23 ± .08 

5.65 ± .23 

4.17 ± .65 

*T. repens 


.38 ± .14 

7.96 ± 1.04 

3.40 ± 1.17 

*T. SI rial urn 


.44 ± .18 

6.25 ± .40 

3.00 ± .82 

T. prawnse 


.75 ± .33 

5.25 ± .37 

5.00 ± .50 

T. vesuulo.sum 


.69 ± .24 

10.50 ± .61 

8.27 ± .75 

*T. atexandrinum 


.23 ± .12 

12.85 ± .69 - 

7.67 + 1.45 

*T. hvhridiim 


.22 ± .18 

8.22 ± .78 ' 

4.00 ± 1.00 

*!'. mcarnalum 


.69 + 1.8 

6.50 ± .33 

4.22 ± 0.43 

' I'rifolntm species preceded by * arc previously unreported hosts. 

- .'Ml numbers are mean ± standard error. 

' Only one T. nigrescens plant used; 3 eggs laid in a single stem. 




250 H"" 


250 H*" 




450 H"" 


250 H"" 

15 r 




Fig. 2a-f. Egg of L. mozardi. a. External appearance oftheoviposition 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 tim (4-6 ixm) in 
length and are situated on a cuticular 
depression (Fig. 3e). There are pores (ca. 1 
Mm 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. 


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 intemodes. Sampling time 
for egg-infested crimson clover stems could 
be reduced by limiting samples to stem sec- 
tions including only the 2nd through 4th 

Previously reported hosts of L. mozardi 
in the genus Trifolium include only T. pra- 
teme (hrnxnev 1881, Folsom 1909) and T. 
vesiculosum (Knight et al. 1976). Since ovi- 
position by L. mozardi occurs in 7 addi- 
tional Trifolium species (Table I) these clo- 
vers also should be considered potential 
hosts for this insect pending confirmation 
of their suitability as hosts for larval de- 

The excavation for oviposition is similar 
to that described for L. mozardi by Girault 
( 1 907) but he does not mention the rim sur- 
rounding the opening internally. The egg 
shape of L. mozardi differs from Languria 



■«g K'^^i 

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. 

erythwcephalus Blatchley and Acropteroxys 
gracilis (Crotch) in that the end opposite the 
micropyie 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 
Thanasimiis diibius (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. 
o/7/('tV»5 (Szentesi 1976) and T. duhiiis (Bak- 
er and Nebeker 1986) but L. mozardi has 
no sensilla basiconica in this region. The 



number of scnsilla on the apex of the style 
of /.. mozanii (9-10) is similar to A. lythri 
(8-11) (Phillips 1978) but dilfers from A. 
obtectus (22-24) and T. dubius (23-25) 
(Szentesi 1976, Baker and Nebeker 1986). 
The short basiconic scnsilla 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 

The behavioral experiment indicates that 
the scnsilla on the ovipositor of L. mozardi 
are needed for the deposition of an egg in 
the plant substrate. Scnsilla on the ovipos- 
itor of other insects arc known to respond 
to chemicals such as salts, water, and amino 
acids (Rice 1976, 1977,Chadhaand Roome 
1980, Hood-Henderson 1982, Liscia et al. 
1982). The scnsilla 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. 

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. 64Ii of 
the Mississippi Agricultural and Forestry 
Experiment Station. Research contributes 
to the Southern Regional Project No. S-20 1 , 
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 duhiu.s 
(F.) (Coleoptera: Cleridae). Ann. Soc. Entomol. Fr. 
22; 49-52. 

Chadha. Ci. and R. Roomc. 1980. Oviposition be- 

havior and the sensilla of the ovipositor of Chilo 
patellus and SiXHloplcra lillorali.s (Lcpidoptcra). 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. 

111. Agric. Exp. Sta. Bull. No. 134. 197. 
Gcnung. W. G., R. Woodruff, and E. Grissell. 1980. 
Lanaiiria cryllmKcphalus: Host plants, immature 
stages, parasites, and habits (Coleoptera: Langu- 
riidae). Florida Entomol. 63: 206-210. 
Girault, A. A. 1907. Oviposition of LaM^wna 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 Mciasyrphus venabtesi and Eu- 
IK'odcs volucns (Diptera; Syrphidae). Can. J. Zool. 
60: 3187-3195. 
Iwala. R. and K. Nishimoto. 1982. Observations on 
the external morphology and the surface structure 
of l.yctus hmniwus (Stephens) (Coleoptera, Lyc- 
tidae) by scanning electron microscopy. Kontyu 
50: 10-22. 
Knight. W. E., O. W. Bamett. 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. Cmjar, A. Angioy, P. Pictra, and J. Stof- 
folano. 1982. 1 chemosensilla dell ovopositore 
in Tabanus nigmviltatus Macq.), Chrysops fuligi- 
nosus (Wied.) e Rhagolelis pomonella (Walsh.). 
Boll. Soc. Ital, Biol. Spmt. 58; 1325-1329. 
Phillips, W, M. 1978. Sensilla types from the ovi- 
positor of the tlea beetle Allica lyllin (Coleoptera; 
Chrysomelidae). Entomol. Exp. Appl. 24; 399- 
Piper, G. L. 1978. Life history of Acropuroxys gra- 
cilis (Coleoptera; Langunidae) on common rag- 
weed in northeastern Ohio. Ohio J. Sci. 78; 304- 
Rice, M. .1. 1976. Contact chemoreceptors on the 
ovipositor of Liicilia ciipnna (Wied.), the Austra- 
lian sheep blowlly. 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, Acaiuhoscclides obleclus Say (Coleoptera: 
Bruchidae). Symp. Biol. Hung. 16; 275-281. 
Wildcrnuith, V. L. and F. H. Gates. 1920. Clover 
stem-borer as an alfalfa pest. USDA Bull. No. 889. 

91(1), 1989, pp. 15-21 




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 (1 1 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: Languriidae, Languria mozardi, larva, mouth parts, antennae, sensilla 

The clover stem borer, Lani^unu mozardi 
Lalreille, is nearly ubiquitous in North 
America on a wide range of cultivated and 
wild host plants (Wildcrmuth and Gates 
192(J). Adults occur frequently in the annual 
clovers TrifoHum incarnatiim L. (crimson 
clover), T. vcsiculosuiu Savi (arrowleaf clo- 
ver), and 7'. alcxandnnuni 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 1 920). The im- 
pact of/,, 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- 

Wildermuth and CJates (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 (5th) instar larvae of 
/,. mozardi were reared on artificial diet 
(Rose and McCabe 1973). Use of the diet 
permitted verification of molting to ensure 
that 5lh instar larvae were examined. Ihe 
techniques for SEM and staining the spec- 
imens are given in Baker et al. (1986). All 
SEM observations are based on 8 last (5th) 
instar larvae. 



Figs. 1-5. Lahriini and mandibles of L. wor^viv/; I Distal row of sensilla chaetica on the labrum thai c.Mcnd 
to the mandibles. 2. Mandible, apical view. 3. Mandible, ectal surface. 4. Molar region. 5. Stria composed of 
several rows of stout, conical setae. C = campaniform sensillum; L = labrum; M = mandible; m = mola; P = 
prostheca; Sc = sensillum chaeticum; St = stria; 1,2,3 = mandibular denies. 


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. 

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 in 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). 



Figs. 6-12. Labial and maxillary regions of the mouth parts. 6. Labium, maxillae, and mandibles. 7. Labial 
palpus, apex. 8. Maxilla, mner surface. 9. Maxilla, outer surface. 10. Sensilla trichodea at base of maxillary 
dentes. 1 1. 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. 



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-segmcnted 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, 

The maxillary palpi are 3-segmented and 
the 3rd is the longest. There are 2 sensilla 
chaetica on each palpal segment and 1 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 is located on the inner sur- 
face of the last palpal segment (Fig. 1 1 ). 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 antcnnal 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. 1 3). The antennal sen- 

sory appendix is on the ventral side of seg- 
ment 2 near the intersegmental membrane 
between segments 2 and 3 (Fig. 1 5). 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. 


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 



Figs. 13-16. Larval antenna of L. mozardi. 13. Antenna, dorsal surface. 14. Antenna, ventral surface. 15. 
Intersegmental region between antennal segments 2 and 3. 1 6. Antenna, apex. Asa = antennal sensory appendix; 
C = campaniform scnsillum; Ps = placoid sensillum; Sb = sensillum basiconicum; Sir = sensillum tnchodeum. 

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 Acw- 
ptewxys gracilis (Newman) in the number 
and types of sensilla situated on the apex of 
the labial palpi. A. gracilis has 1 1-12 basi- 
conic sensilla on the apex of the palpus (Pip- 
er 1978), while L. mozardi has a total of 9- 
1 sensilla but there are 2 types of basiconic 

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 is found on Tribolium lar- 
vae (13 & 12) (Ryan and Behan 1973) and 



Hypera larvae (12 & 1 1) (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 1 966, 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 antenna) sensilla on 
L. mozardi (22) is similar to Tenehrio mol- 
itor L. (24 sensilla) (Pierantoni 1969) and 
Triboliuni 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 
antenna) 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. 


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. 6414 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 

Literature Cited 

Baker, G. T.. W. L. Parrott, and J. N. Jenkins. 1986. 
Sensory receptors on the larval maxillae and labia 
of Heliolliis zea and Heliothis virescens (Lepidop- 
tcra: Noctuidae). Int. J. Insect Morphol. Embryol. 
15: 227-232. 

Behan. M. and M, F. Ryan. 1978. Ultrastructure of 
anlennal sensory receptors of Tnboliuni larvae 
(Coleoptera: Tenebrionidae). Int. J. Insect Mor- 
phol. Embryol. 7: 221-236. 

Bland, R. G. 1983. Sensilla on the antennae, mouth 
parts, and body of the larva of the alfalfa weevil, 
Hypera postica (Gyllenhal) (Coleoptera: Curcu- 
lionidae). Int. J. Insect Morphol. Embryol. 12: 261- 

Bloom, J. W., R. Y. Zacharuk, and A. E. Holodniuk. 
1982a. Ultrastructure of the larval antenna of Te- 
nehrio 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 Tenehrio molitor L. (Coleoptera: Tenebrioni- 
dae): Structure of the blunt-tipped peg and palliate 
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- 
aca. NY, 701 p. 

Doane. J. F. and J. Klingler. 1 978. Location of CO,- 
receptive sensilla on larvae of the wireworms 
Agnotes Itncatus-obscurus and Limonius califor- 
iiictis. Ann. Entomol. Soc. Am. 71: 357-363. 

Guse, G. W. and K. Honomichl. 1980. Die digiti- 



formen Sensillen auf dem Maxillarpalpus von Co- 
leoptera II. Feinstrucktur bei Agabus hipuslulalus 
(L.) and Hydrobius fuscipes (L.). Protoplasma 103: 

Honomichl, K. 1980. Die digitiformcn Sensillen auf 
dem Maxillarpalpus von Coleoptera I. Verglei- 
chendlopographische Untcrsuchungdcs kuticular- 
cn Apparatcs. Zool. Anz. 204: 1-12. 

Klingler, J. 1966. Uber den Sitz der CO,-Rcceptoren 
bei der Larve von Otiorhynchus sukatus. 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: 1 185-1209. 

Mitchell, B. K.. 1978. Some aspects of gustation in 
the larval red turnip beetle, Enlomoscclis anien- 
cana. related to feedmg and host plant selection. 
Entomol. Exp. Appl. 24: .■(40-349. 

Peterson, A. 1951. Larvae of Insects, Part II. Edwards 
Bros. Inc., Ann Arbor, Ml, 416 p. 

Pierantoni, R. L. 1969. Mechano- and chemo-recep- 
tors in the antenna of Tenebno molilor in the larval 
stage. Proc. Elect. Micros. Soc. Am. 2: 246-247. 

Piper, G. L. 1978. VAk \i\sX0Ty oi Acropwroxys gra- 
cilis (Coleoptera: Langunidae) on common rag- 
weed in northeastern Ohio. Ohio J. Sci. 78: 304- 

Rose, R. I. and J. M. McCabe. 1973. Laboratory 
rearing techniques for the southern com root- 
worm. J. Econ. Entomol. 66: 398^00. 

Ryan. M. F. and M. Behan. 1973. The sensory re- 
ceptors of Tnbohum larvae. Physiol. Zool. 46: 238- 

Scott, D. A. and R. Y. Zacharuk. 1971. Fine structure 
of the antenna! sensory appendix in the larvae of 
Cleniccni dcslruclor (Brown) (Elatendae: 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- 
tors of larval Orthosoma bninneum (Forster) (Co- 
leoptera: Cerambycidae). J. Comp. Physiol. 88: 

Wildcrmuth, 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. 
Ill: 1-34. 

Zacharuk, R. Y., P. J. Albert, and F. W. Bellamy. 1977. 
LUtrastructure and function of digitiform sensilla 
on the labial palp of a larval elaterid (Coleoptera). 
Can. J. Zool. 55: 569-578. 

91(1), 1989. pp. 22-25 


Scon E. Miller and Vitor O. Becker 

(SEM) Bishop Museum, Box 19000-A, Honolulu. Hawaii 96817; (VOB) Centro de 
Pesquisa Agropecuaria dos Cerrados, Caixa postal 70-0023, 73300-Pianaltina, DF, Brasil. 

Abstract.— Three species of moths were described from "Caroline" by Bosc [1800]. 
Cnmihus ci(lspcr,Klllii.s i^osc was recognized by Franclcmont and Todd (1983) as Renia 
aclspcigillus (Nocluidae), but the other two names have remained unplaced. We recognize 
Ahicita cereella Bosc as conspecific with Nolo sorghiclla Riley, n. syn. (Noctuidae) and 
Pyralis sacculana Bosc as conspecific with Clydonoptewn tecomae Riley, n. syn. (Pyral- 

Key Words: nomenclature, Nearctic, moths, Noctuidae, Pyralidae 

The frenchman l.ouis Auguste (iuillaume 
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 ,1. C. Fabricius, went to the Mu- 
seum 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 centur>. because they were 
illustrated by C'oquebert (1801). However, 
no Lepidopteran types from the Bosc col- 
lection are now extant in the Paris museum 
(Zimsen 19(i4, P. Viette pers. comm.). 

Bosc [1800] described three species of 
North .Xinorican moths, but the paper was 
overlooked for over a century. The part of 
the journal in which Bosc's paper appears 
does not bear a date, but is headed "Prairial. 
an 8 de la Republique," meaning it was pub- 
lished in 1 800, the eighth year of the French 
Re\ olulion (an earlier pari bears the head- 
ing "Germinel. an 6 de la Republique (Mars 
1798)"). It appears that the first use of any 

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 [ 1 800], only 
one, ( 'rciDihus adspcrgillus. has been in- 
cluded in the North American literature 
(Franclcmont 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 

Two of the species, Crambus adspergillus 
and Pyralis sacriilana. were illustrated in 
color by Coquebert de Montbret ( 1 80 1 ), but 
no mention ofAliicita cereella was included. 


Renia adspergillus (Bosc) 

Figs. 1, 4 

Crambus adspergillus Bosc [1800]: 114, fig. 
2 [Type locality: United States, "Caro- 
line"']. -Coquebert 1801: 72, pi. XVII, fig. 








Figs. 1-3. Reproductions of illustrations from Bosc ([1800]: pi. VII), Fig. 1: Crambus adspergtUm. Fig. 2: 
Alucila ccrecila. Fig. 3; I'yralis sacculana. 

Figs. 4-6. Figures of specimens matching Hose's illustrations: Fig. 4: Renia adspergillus (Bosc); Fig. 5: Nola 
cercclla (Bosc); Fig. 6: Clydonoplcron 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 Franclcmont and Todd 

Renia rcstrictalis 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 asfJa- 
vipunctalis (Geyer) and larvalis (Grote). We 
follow Franclemont and Todd ( 1 983: 121), 

who know the group well and were the first 
revisers, in recognizing adspergillus as con- 
specific with larvalis. 

Nola cereella (Bosc), 
New Combination 

Figs. 2, 5 

Alucila cereella Bosc [ 1 800]: 1 1 5, fig. 4 [Type 

locality: United States, "Caroline"] 
Nola sorghiella Riley 1882: 187, pi. XI, fig. 

1 [Type locality: United States, Alabama 

and Florida], n. syn. 
Nola portoricensis Moeschler 1890: 118 

[Type locality: Puerto Rico], n. syn. 



Because ofthc small si/.c of the figure and 
the quality of printing of the plate, this was 
the most diiricult 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 ofthc head thai 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 nwliiu'lla group. The figure also resem- 
bles the pattern of some species of Elhniia 
Hucbner (Oecophoridae), especially /:. tri- 
furccllci (Chambers). 

We have two main reasons for consid- 
ering ccrcclla as conspecific with sori^hiella. 
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 tly (Maycliola dc- 
slnictor (Say), Diptera: CYx'idomyiidae), and 
Sito!r(\^a ccrcalclla (Olivier) (Lepidoptcra: 
Gelcchiidae), no other known grain pest re- 
sembles the figure ol' ccrcclla. Bosc's original 
description follows: 

"Elle est cendree. Les bords des ailes sont 
brans et paroissent renfles. On voit sur le 
disque dc chaquc ailc superieure deux points 
bruns. Les ailes posterieures sont d'un gris 

"Tette Alucitc est la teigne qui, sous Ic 
nom d' Hessian fiy, a fait, il y a environ douze 
ans, dc grandes ravages dans les bleds 
d'Amerique, et a menace de les etendre sur 
ceux d'Europe, ce qui a oblige le parlement 
d'Anglcterre 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 germc du grain, puis la farine. ne laissant 
que Tecorce. II n"y a qu'unc larve dans cha- 
que grain: elle est tres-fcconde ct les gen- 
erations se succcdenl rapidement. Au defaut 
de bled, elle attaque le mais, et se multiplie 
considerablement dans les greniers oil l"on 
conserve ce grain. 

"Le meilleur moyen pourdetruire la larva 
de cette .Alucite, c'est de faire passer les grains 
dans une etuve Ircs-chaude." 


Clydonopteron sacculana (Bosc), 

New Combination 

Figs. 3, 6 

Pyralis sacculana Bosc [1800]: 115, fig. 3A, 
3B [Type locality: United States, "Caro- 
line"]. -Coquebert 1 80 1 : 7 1 , pi. XVIL fig. 

Clydonopteron tccomae 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 
tcconiac. 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. Kayc and 
Lamont ( 1 927) 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 (Hampton 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 
is a label, added by Carl Heinrich. proposing 
this synonymy, based on Coquebert ( 1 80 1 ), 
but Heinrich never published his discovery. 


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- 
molog) Laboratory, U.S. Department of 



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 Venczolana. 
Bol. Entomol. Venez. 10: 1-336. pi. 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]. Descnption dc Irois 
espdcies de Lepidopteres do la Carolme. Bulletin 
des Sciences, par la Societe Philomathique, Pans 
[scries 1] 2(39): 1 14-115. pi. VII. [German trans- 
lation in Archiv fur zoologie und zootomie 3{ I ): 
188-190, 1802] 

Cashatt, E. D. 1969. Revision of the Chrysauginae 
of North America (Lepidoptera: Pyralidae) [ab- 
stract only]. Diss. Abstr. (B) 29: 4696. 

Coquebertde Montbert. A. J. 1801. Illustratioicono- 
graphica insectorum quae in Musaeis Parisinisob- 
servavit ct in lucem edidit Joh. Christ. Fabricius, 
praemissis ejusdem descriptionibus; accedunt 

species plurimac, vel minus aut nondum cognitae. 

Volume II, pp. [45]-90. pi. XI-XX. 
Franclemonl, 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. 1. 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. 
Kayc, W. J. and N. Lamont. 1927. A catalogue of 

the Trinidad Lepidoptera Heterocera (moths). 

Mem. Dept. Agr.. Trinidad and Tobago 3: viii -l- 

144 -I- 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. 
Moeschlcr. H. B. 1890. Die Lepidopteren-Fauna der 

Insel Portorico. Abh. Senkenb. Naturforsch. Ges. 

16: 77-360, I pi. 
Munroe, E. G. 1983. Pyralidae, pp. 67-85. //( 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. On a new pyralid infesting the seed 

pods of the trumpet vine. Clydonopteron lecomae, 

nov. gen., n. sp. Amer. Entomol. 3: 286-288. 
. 1882. Report of the entomologist, in Report 

of the Commissioner of Agriculture for the years 

1 88 1 and 1882. Washington, U.S. Government 

Pnnting Office. 
Zimsen, E. 1964. The type material of I. C. Fabricius. 

Copenhagen, Munksgaard. 656 pp. 

91(1), 1989, pp. 26-28 


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: nest, Synoeca, Vespidae, wasp 

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 1 925, Maid! 
1934, Richards and Richards 1951, R. S. 
Jacobson, pers. comm.; pers. obs.), and 
Reaumur (1722) intimated that it is usual 
for \ espula sp. (probably I '. i^crnianica 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 

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 

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- 
lobcd nest in Panama, and Buysson (1906) 
figured one from Mexico with six lobes, ap- 
parently the largest nest reported for the ge- 

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. 

Reaumur (1722) reported that Vespula 
workers consistently enter through one of 
the two nest-holes and leave through the 



Fig. 1 . Nine-lobed nest of Synoeca septenlrionalis 
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 arc 
about 20 mm long. 

Other. Although the quantitative study of 
behavior was nearly unknown in his time, 
we know that Reaumur made original ob- 
servations on ( 'espula 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 
tratfic. Indeed, if a colony has considerable 
traffic 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 1 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 7, 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. 

Lcll Hole Right Hole lolal 



1 14 






Table 2. Percentage breakdown of total traffic dur- 
ing the observation time. Based on Table 1 and the 
hole-choice results, as explained in text. 

Iillllclf Riphtllnk- 













idcnlical. Table 2 thus gives the percentage 
breakdowti of the Iralllc 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 ditVerence 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 llexibility in nest construction 
(Vecht 1967, Overal 1982), which may ac- 
count for this nest's two-holed condition. It 
makes little diUcrence here whether this 
originated and was maintained as an adap- 
tive response to extraordinarily large col- 
ony-si/e or through ordinar> 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 indi\ iduals have been summed to 
produce a meaningful phenomenon at the 
colony level. 


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 

Huysson, R. Du. 1906. Monographic des vcspides 
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, \'cspa cincia Fabr. .1. Bombay Nat. 
Hist. Soc. 30: 858-860. 

Jeanne, R. L. 1975. The adaptivencss of social wasp 
nest architecture. Ouart. Rev. Biol. 50: 267-287. 

Maidl. F. 1934. Die 1 ebcnsgewohnheitcn und In- 
stinktederstaatcnbildcnden Insektcn. Vienna: Fritz 
Wagner 823 pp. 

Overal. W.L. 1982. Acoustical behavior and variable 
nest architecture in Synoeca virginea (Hymenop- 
tera, Vespidac). J. Cicorgia 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 gucpes. 
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. \an der. 1967. Bouwproblemen van sociale 
wespen. Verb. K. Ned. Akad. Wetensch. (Afd. Na- 
tuurkunde) 76: 59-68. 

Wen/el, J. W. In press. Evolution of nest architecture 
in social vespids. In K. G. Ross, and R. W. Mat- 
thews, eds.. The Social Biology of Wasps. Ithaca. 
Cornell Univ. Press. 


91(1), 1989, pp. 29-34 


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. 

Ahstiacl.—lhe two North American species of the unusual genus Hyhnzon Fallen 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: taxonomy, Ichneumonoidea, ant-parasites 

The genus Hybrizon Fallen is one of the 
most peculiar and taxonomically confusing 
groups in the Ichneumonoidae. It contains 
seven species (five Palearclic, 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 
theBraconidae{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 Hyhnzon 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 ( 1 984) 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 

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 % of the specimens 
collected by Smith are n/nv (Ashmead); the 
other 'A are a distinct species which I thought 
was undescribed but now have identified as 
the previously unknown female of Jlavo- 
c//R7;«(Ashmead). I have provided descrip- 
tions and a key to separate the species be- 
low. Additional specimens were borrowed 



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 
Rijksmuseum van Naliiurlijke Histoire, 
Leiden, The Netherlands (C. van Achter- 

rhe 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- 
ihorpe 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 rilcyi and 
137:5 in jlavocinctus. Female biased sex ra- 
tios are predicted by the local mate com- 
petition (LMCl model of Hamilton (1967), 
". . . where females place otrspring in dis- 
crete patches of the resource (in this case, 
ant colonies), and those offspring mate ran- 
domly in their patch before female olfspring 
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 dispcrsers. Thus, random sweep- 
ing or llight intercept traps would produce 
mostly females, whereas collections made 
in ant colonies might yield a more balanced 
sex ratio. 

Family Paxylommatidae 

Pachylommaloidac Foerster, 1862: 247. 
Oldest family-group name (see Mason 
1 98 1 for discussion). 

Hybrizon Fallen 

Hybrizon Fallen, 1813, p. 19. No species. 
Type-species: Hybrizon latcbricola Necs, 
1834. Monotypic, hrst included species 
by Nees( 1834:28). 

Paxylomina de Brebisson. 1825: 23. Type- 

species: PaxyloDinia buccatiini de Brcb- 
bison. Monotypic. Synonymy by Wes- 
mael, 1835. 
Plancus Curtis, 1833: 188. Type-species: 
Plancus apicalis Curtis. Monotypic. Syn- 
onymy by Stephens, 1835. 
Eiirypterna Foerster, 1862: 247. Type- 
species: Paxylomma cremieri Romand. 
Monotypic. Synonymy by Marshall, 1891. 
L";Ypac7;i7t»/>;/?;a Ashmead, 1894: 58. Type- 
species: Wesmaelia rilcyi Ashmead. Orig- 
inal designation. Synonymy by Watana- 
be, 1935. 
Ogkosoma Haupt, 1913: 52. Type-species: 
Ogkosonia sclnvarzi Haupt. Monotypic. 
Synonymy by Strand, 1914. 
The names Paxyloma (Stephens 1835), 
/'avi7('/n/);r (Wesmael 1835), Paxyllonuna 
(C^urtis 1837), ra.vr//()//;a(Blanchard 1840), 
and Pachylomma (Ratzeburg 1848) are all 
to be considered emendations oi Paxyloiu- 
nia (see Shenefclt 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, Hy- 
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 1 
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 

Ocelli small, ocell-ocular distance at least equal to 
diameter of lateral ocellus, often greater (Fig. 9); 
first segment of radius in fore wmg shorter than 
first segment of discoideus and about '2 length of 
recurrent vein, branchial cell not as tall as disco- 
cubital cell (Fig. 1); head, thorax and abdomen 
usually entirely black rileyt (Ashmead) 

Ocelli larger, occll-ocular distance less than diam- 
eter of lateral ocellus, often less than half (Fig. 8); 


Figs. I, 2. Wings of //i/if/ro/i species. 1. // n/frMAshmcad). 2, 1/. Jlavoci ml us (Aiihmead) (scale = 0.5 mm). 

first segment of radius equal to or longer than dis- 
coideus and about -A 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 , . Ilavocmctus (Ashmead) 

Hybrizon rileyi (Ashmead) 

Figs. 1, 3, 4, 7, 9 

Wesmaelia rileyi Ashmead, 1 899: 64 1 . Ho- 
lotype female in U.S. National Museum, 
Washington, D.C. 

Female. Length of body, 2-3 mm. Color: 
head black, clypeus and mouthpains 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 1 1 flagello- 
meres. Thorax: pro and mesothorax smooth 
and shining; propodcum irregularly rugose, 
without any indication of median longitu- 
dinal carina. Abdomen: terga smooth and 
shining, terga I and 2 sometimes weakly 
striate at base (Fig. 4). Wings (Fig. 1 ): first 
segment of radius shorter than first segment 

of discoideus and about Vi 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 $S 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 

Eupachyloniniajlavocincta Ashmead, 1894: 
59. Holotype female in U.S. National 
Museum, Washington, D.C. 

Female. Length of body, 3.5^ mm. Col- 
or: head black, clypeus and mouth parts light 
yellow; antennal scape and pedicel yellow, 
fiagellum black; prothorax honey yellow; 



Figs. 3-9. Hybrizon species. 3, H. rileyi (Ashmead), habitus (scale = 2 mm). 4, H. nleyi. abdominal terga 
1-2 (scale = 500 ii). 5, H . flavocinctus (Ashmead). abdominal terga 1-2 (scale = 500 n). 6, H. flavocinctus. face 
(scale = 100 ^). 7. //. nUyi. face (scale = 200 n). 8, H. Jlavocinclus. vertex (scale = 200 m). 9, H. ritevi, vertex 
(scale = 200 jj). 

mesonotum dark brown or black, some- 
times with yellow longitudinal lines; scu- 
tcUum yellow with brown spot at base; 
mesopleuron var\ing from entirely brown 
to yellow; propodeum dark brown; tegula 
yellow; legs yellow, hind femur, tibia, and 
co.\a light brown; abdomen brown, terga 3 
and 4 yellow at base. Head: reticulate and 

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 1 1 flagello- 
meres. Thorax: pro- and mesothorax smooth 



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 -A 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 S6 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 ofjiavocinctus 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 1 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 ofjiavocinctiis. 

This species differs from rileyi in its larger 
size, larger ocelli, generally lighter body col- 
or, and wing venation. It is also very similar 
to the European huccatus (de Brebisson) 
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. 


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. 

Thanks also go to David Smith for oper- 
ating the Malaise traps that produced the 
specimens which motivated this study. 

Literature Cited 

van Achterberg, C. 1976. Hybrizonlinae or Hybri- 
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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 Brebisson, L. A. 1825. Pa.xylomme. Pa.xy/nnjDia. 
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Curtis, J. 1833. Charactersof some undescribed gen- 
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. 1837. A guide to an arrangement of Bntish 

insects. 2nd Edition. London, Weslley. vi. + 294 

Donisthorpe, H. St. J. K. and D. S. Wilkinson. 1930. 
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Fallen, C.F. 1813. Spec. Nov. Hym. Disp. Meth., p. 

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Gauld, I. D. 1984. An introduction to the Ichneu- 
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Hahday, A. H. 1840. Braconidae, pp. 61-65. //; 
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Hamilton, W. D. 1967. Extraordinary se.x ratios. Sci- 
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Haupt. H. 1913. Bcitragc zur Hymcnopteran-Fauna 
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Marsh, P. M. 1963. A key to the Nearctic subfamilies 
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91(1), 1989, pp. 35-50 


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: Diptera, Therevidae, Phycus. Ataenogera, New World, species descriptions, 

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- 
em 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- 

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, Schhngeria Ir\vin, and Par- 
apherocera Irwin on the other (Irwin and 

Lyneborg 1981), we elected to describe and 
diagnose their New World species here in a 
single publication. 


The morphological terminology used for 
the male terminalia was originally defined 
and described by Lyneborg ( 1 968) 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- 
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- 



ment system originally designed by Rauch 

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 

2) Boldface type: intermediate political 
unit (state or province outside the United 
States or county within the United States) 
and elevation expressed in 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, 

5) Numbers of specimens is followed by 
the sex designation M for male or F for 

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- 

Depositories. — Paratypes of Phycus 
fronimeri 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, Bemice 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, Maipii; FSCA, Florida State Collec- 
tion of Arthropods, Gainesville; IAS, Insti- 
tute of Agricultural Sciences, Tokyo; IE, In- 
stituto di Entomologia, Bologna, Italy; IML, 
Institute Miguel Lillo, Tucuman, Argen- 
tina; INHS, Illinois Natural History Survey, 
Champaign; INIA, Institute 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- 



bus: OSU, Oregon State University, Cor- 
vallis; PAS, Polish Academy of Sciences, 
Warsaw, Poland; RNHL, Rijicmuseum van 
Natuuriijke Historic, 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; LIM, 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 Phyciis 
frontalis n. sp. are deposited in the following 
museums; AMNH, American Museum of 
Natural History, New York; INHS. Illinois 
Natural History Survey. Champaign; 

USNM, LInited States National Museum, 
Washington, D.C. 

Key to New World Species of 
Ataenoger.4 and Phycus 

1. Maxillar> 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); stemites 5-8 in males, 
5-7 in females (Fig. 10) with numerous lan- 
ceolate setae ventrally. male genitalia with large. 

subtriangular hypandrium (Fig. 8) 

itacnogcra ahdommalis Krober 

- Maxillary palpus two-segmented (Fig. 18); eye 
margins divergent slightly from level of ocellar 
tubercle toward genae ( Fig. 1 7 ); discal cell trun- 
cate basally (Fig. 19); stemites 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 frominen n. sp. 

- Eyes separated 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 fronlalis n. sp. 

Superficially, the genus Ataenogera is 
morphologically similar to Phycus, the two 
genera are however quite distinct. The best 
distinguishing characteristics arc: (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 Ataenog- 
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 Ataenogera. 

Genus Ataenogera Krober 

Ataenogera Krober (1914: 31): Malloch 
( 1 932: 255). Type species: A. abdominalis 
Krober, by monotypy. 

F' --— Ss 

Figs. 1-15. Ataenogera abdominalis. — \. Antenna (5747). 1. Head of male, frontal view (5747). 3. Maxillary 

palpus, lateral view (5747). 4. Wing, dorsal view (5747). 5. Male tergite 8, epandnum, cerci, and ventral epandrial 

sclerite, dorsal view (5761). 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 (576/). 10. Female abdomen, termmal segments, lateral view with enlargement of ventral lanceolate 

setae (575^). 1 1. 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 (nghl) (5764). 15. Frontal plate 

(576-^). .Abbreviations: (AS) antennal sheath; (C) cercus; (DA) dorsal apodeme; (Dp) distiphallus; (DSp) dorsal 

spiracle; (EA) ejaculatory apodeme; (Epa) epandnum; (F) furca; (Gs) gonostylus; (Gx) gonocoxites; (Hpa) hy- 

pandnum; (Sj) sternite 8; (T,) tergite 8; (TJ tergite 9; (VES) ventral epandrial sclerite. Scale = 0.1 mm, unless 

otherwise indicated. 




Leptocera Krober ( 1 928: 1 1 7). Type species: 
L. gracilis Krober by monotypy. Preoc- 
cupied by Olivier (1813: 489). 

Zichcnia Krober ( 1 929: January: 434). New 
name for Leptocera Krober. 

Epileplocera Richards (1929: August: 171). 
Unjustified new name for Leptocera 

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 1.3-2.0 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; prostemum bare 
in and around central depression; pleural 
setae elongate, scattered on propleuron and 
dorsal fourth of anepistemum. 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 
Rj 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. 

Abdomen. — Slender, especially in male, 
longer than wide, laterally compressed; dor- 
sum convex, shining fuscous to black; setae 
sparse, short, stemites 5-8 in males, 5-7 in 
females (Fig. 10) with numerous lanceolate 
setae ventrally. Male terminalia (Figs. 5- 
9).— Tergite 8 (TJ wide, deeply constricted 
medially. Stemite 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. 1 1- 
12).— Tergite 8 (TJ 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. Stemite 
8 (Sk) 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 

Ataenogera abdominalis Krober 

Ataenogera abdominalis Krober ( 1 9 1 4; 3 1 ): 
Malloch(1932: 255). 

Leptocera gracilis Krober ( 1 928: 1 1 8). New- 

Hcnicomyia brevicornis Bromley (1934: 
361). New synonym. 

Derivation oimmt.— abdomen (Latin) = 
belly; alls (Latin) = pertaining to. 



Diagnosis.— Ataenogera abdoininaUs is 
the only recognized species in tliis 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, dcO, 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 1-^ with narrow, white posterior mar- 
gin; setae black, short, appressed on dark 
areas, white, moderately long on white pos- 
terior margin, stemites 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 1.8-2.2, 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. 


The frons shows variation similar to male. 
Length of antenna 1.3-1.5, 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, 0.7 1 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. 1 2, 0.08 mm. Fore and mid- 
dle femora show variation similar to males. 
Characteristics of terminalia (Figs. 11-12) 
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. 13-1 5). -Length 8.0 mm (N 
= 1), width 1.5 mm. Alar process not spi- 
nose. Labial sheath broad (Fig. 1 5), 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-VIL 

Type material.— A syntypic series of 3 males 
of Ataenogera abdominalis Krober is in the 
Museum fur 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 Heiucomyia hrevicornis 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- 

type, was collected at Parana, Brazil, in De- 

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, 16-XI-1966, L. 
A. Stange, IM. Mendoza: Potrerillos, 6-1- 
1927, F. and M. Edwards, IF. La Rioja: La 
Rioja, 1928, IF. Catamarca: 17 km NW 
Chumbicha, 1143 iti, 25-XII-1971, C. A. 
Pearson, IF. BOLIVIA: Beni: Rio Itenez 
opposite Costa Marques, Brazil, 1-3-IX- 
1964, J. K. Bouseman and J. Lussenkop, 
1 F; 4-6-IX- 1 964, 1 M; 30-3 1 -VIII- 1 964, 1 F; 
Rio Itenez, Pampa de Meio, 11-1 3-IX- 1 964, 
J. K. Bouseman and J. Lussenkop, IF. 
BRAZIL: Parana: 1912, IM IF; Para, Bak- 
er, IF; Santa Catarina: Nova Teutonia, 
27°8', 52°23', 16-XII-1947, F. Plaumann, 
IM; 1-1945, IF; 27°11', 52°23', 18-XII- 
1959, F. Plaumann, IF. BRITISH 
GUIANA: Kartabo, 20-V-1924, IF. COS- 
TA RICA: Guanacaste: La Pacifica, 4 km 
NW Cafias [Las Cafias], 1-9-IV- 1 974, P. A. 
Opler, IM 4F; 29-III-1974, IF. EL SAL- 
VADOR: Quezaltepeque, 2-XI-1977, MEI, 
IF. MEXICO: Morelos: 1 1.7 km S Yaute- 
pec, 17-VIII-1962, N. L. Marston, 2F; 
Puebla: 4.8 km NW Petlalcingo, 3-1 V- 1 962, 
F. D. Parker, IF. PANAMA: Canal Zone, 
Ancon, 4-V-1926, C. T. Greene, IF; 9-1 V- 
1926, IF. PARAGUAY: San Bernardino, 
IV, K. Fiebrig, IM; 5-1 V, IM; -/-/-, IM. 
TRINIDAD: St. Augustine, V-1959, F. D. 
Barrett, IF. URUGUAY: Minas: Arassu- 
ahy, X-1929, Thieman, IF. VENEZUELA: 
Barinitas: 10 km SE Barinitas, 1 -III- 1986, 
R. B. Miller, IM. 

Genus Phycus Walker 

Phvciis Walker (1850: 1): Irwin and Lyne- 
borg (1981: 260); Lyneborg (1978: 212). 
Type species: Xylophagus canescens 
Walker (1848: 129) by monotypy (= Xy- 
lophagus brunneus Wiedemann, 1824: 



Caenophaiws Loevv (1874: 415): Lyneborg 
(1978: 212). Type species: C. insignis 
Loew (1874: 415) by monotypy. Preoc- 
cupied by Caenophanes Foerster (1862: 

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: Phyciis nitidus 
Wulp ( 1 897: 1 37) by original designation. 

Caenophaniella Seguy ( 1 94 1 : 112): Lyne- 
borg (1978: 212). Type species: C. nigra 
Seguy ( 1 94 1 : 1 1 2) 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. 1 7, 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, dc O-I, sc 
1; vittae absent; mesonotal setae short, uni- 
form, erect; prostemum 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 Rj and R^ sub- 
equal; cell rj large, enclosed apex of wing. 

2.2-2.6 times as long as wide at apex; ceU 
m, usuaUy closed, petiolate; discal cell 
trunctae basally; m-cu/r-m subequal. Legs.— 
Fore coxae elongate, sparsely setose, with 3 
stifl" 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 (T^) comparative- 
ly wide and deeply to moderately constrict- 
ed medially. Stemite 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 (TJ 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. Stemite 8 (Ss) large, as long 
as wide, with short median incision on pos- 



Figs. 16-26. Phycus frommen. — \b. Antenna {5446). 17. Head of male, frontal view (5169). 18. Maxillary 
palpus, lateral view {5446). 19. Wing, dorsal view {5169). 20. Male lergite 8. epandnum. cerci, and ventral 
epandrial sclerile. dorsal view {5446). 21. Male ventral epandrial sclerite, ventral view {5446). 22. Male gono- 
coxiles and gonostylus. dorsal view {5446). 23. Male gonocoxites and gonostylus, ventral view {5446). 24. Male 
aedeagus. lateral view (5446). 25. Female terminalia. dorsal view (57^0). 26. Female termmalia, ventral view 
(5730). Abbreviations: (ApSg) apical segment; (BaSg) basal segment; (C) cercus; (DA) dorsal apodeme; (Dp) 
disliphallus; (EA) ejaculatory apodeme; (Epa) epandrium; (F) furca; (Gs) gonostylus; (Gx) gonocoxites; (SJ stemite 
8; (Stp) stipes; (Tg) tergite 8; (T,) tergite 9; (T,,,) tergite 10; (VA) ventral apodeme; (VES) ventral epandrial 
sclerite. Scale = 0. 1 mm, unless otherwise indicated. 

terior margin. Stemite 9 greatly modified, 
invaginated beneath stemite 8 to form in- 
ternal furca (F). 

Several Old World species ofPhycus have 
been observed walking along dead and fal- 
len tree trunks in riverine habitats. The New 
World species from westcm North America 
have been observed walking on rocks under 

Washingtonia palms or other vegetation in 
desert canyon bottoms (Irwin, personal ob- 

Phycus frommeri Webb and Irwin, 
New Species 

Derivation of name: this species is named 
in honor of Saul I. Frommer, University of 



California, Riverside, who collected most 
of the specimens. 

Diagnosis. — P. froinnieri can be distin- 
guished from P. frontalis by the following 
combination of characteristics: eyes (Fig. 1 7) 
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. 1 8) 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 , 
dc 0, sc 1. Postpronotal lobe concolorous 
with thorax; setae white, fine, scattered. 

Propleuron, anepistemum, katepistemum, 
meron dark brown, tomentum silver; ane- 
pimeron fuscous, glossy. Pleural setae white 
to pale yellow, abundant on propleuron, and 
scattered over anepistemum; 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 

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 
(Tk) 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 

Variation in males (N = 1 0). — 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. 1 2-0. 1 4, 0. 1 3 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. 1 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 



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 anepistemum, 
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, 1.0-1.3, 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 polifoliiim (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 

60 ■ 

SO ■ 


40 ■ 


30 ■ 







Fig. 27. Phycus Jronuiieri. — F\\g,hl penodicily 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 (5679) in loose soil under boul- 
ders, both in the state of Colima, Mexico. 
These were reared to adults in our labora- 

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 



Figs. 28-34. Phycus frommeri larva and pupa cxuvia (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 cxuvium, dorsal view 
(right), ventral view (left). 34. Frontal plate of pupal exuvium. Abbreviations: (ANT) antenna; (AS) antennal 
sheath; (DSp) dorsal spiracle; (M) mandible; (MP) maxillar> palpus; (MR) nietaccphalic rod. Scale = 0.1 mm, 
unless otherwise stated. 

ventral spines on segments II-Vll line, not 
fused basally. Abdominal spiracles on seg- 
ments I-VII moderately thick, tubular. 

Distribution. — The range of /Vnti/.s //•<>/;;- 
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 5169. 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: 
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, I M; 1 5-1 8-V- 1970, SIF, 23 M 5 F; 
1 8-23-V- 1 970, 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 
RM W, 2 M 2 F; 1 9-20-VI- 1 969, SIF RMW, 
1 M 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, I F;26-IV-3-V- 1970, SIF 
RMW, 1 M; 8-12-V-1970, SIF RMW, 7 M 

2 F; 1I-I3-V-1970, SIF RMW, 5 M; 13- 
15-V-1970, SIF RMW, 8 M 2 F; 23-25-V- 
1970. SIF RMW, 16M18F;25-26-V-1970, 
SIF RMW, 25 M 17 F; 1-4-VI-1970, SIF 
RMW, 8 M 4 F; 2 1-29-V- 1 973, A. B. Tabet, 
2 M; 5-1 3-VI- 1973, A. B. Tabet, 7 M 4 F; 
24-V- 1 969, MEI SIF, 44 M 1 2 F; 8-VI- 1 965, 
MEI, 1 M 1 F; 16-V-1969, MEI, 1 M 1 F; 
1 8-V- 1 969, MEI, 8 M 3 F; 2 1 -V- 1 969, MEI, 
1 1 M 2 F; 13-20-VI-1973, A. B. Tabet, 5 



M; 21 -VI- 1962, EIS, 1 F; 19-V-1964, MEI, 
1 M; 1 l-VI-1965, MEI, 1 F; 9-16-V-1973, 
A. B. Tabet, 1 M; 5 mi W Sage, 3-VII-1963, 
P. D. Hurd, 1 F; 2-VII- 1 963, E. I. Schlinger, 
1 F: Idyllwild, 27- VI- 1 956, M. S. Wasbauer, 
1 M; Carrizo Creek, 30-VI-1964, E. I. 
Schlinger, 1 F; Massacre Canyon, 300 yds 
up from Highway 79, 29-VII-1964, MEI, 1 
F. San Diego Co.: Culp Canyon, 1 2-VI- 1958, 
E. I. Schlinger, 12 F; Borego, l-V-1946, J. 
S. Perry, 1 M; 9.7 km E San Diego, 26-VI- 
1963, H. L. Griffin, 1 F, R. L. Langston, 1 
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-IV-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, 1 M; 
2.5 km E La Burrera, 549 m, E. M. and .1. 
L. Fisher, 1 M; 6.4 km NW Don Pancho, 
1 -VIII- 1964, MEI, 1 F; Las Barracas, ca. 30 
km E Santiago, 25-3 1 -III- 1 982, P. DeBach, 
1 M; 1-6-IV-1982, P. DeBach, 2 M IF; 13- 
18-IV-1982, P. DeBach, 1 F; 1 9-24-1 V- 
1982, P. DeBach. 1 M; 25-30-IV-1982, P. 
DeBach, 1 M; 7-1 2-V- 1982, P. DeBach, 4 
M 2 F. Sinaloa: 86.9 km S Culiacan, 164.6 
m, 23-IV-1969, MEI, 1 M; "Las Escondi- 
das," 106.2 km N Mazatlan. 137 m. 22-IV- 
1968, MEI. 16 M 25 F. Colima: Los Tern- 
panes, 12-III-1985, R. B. Miller and L. A. 
Stange, 1 F reared from larva; Rio Salado, 
7 km S Colima, 14-III-1985, 1 F reared 
from larva. 

Phycus frontalis Webb and Irwin, 
New Species 

Derivation of name: //o/?/ (Latin) = brow; 
alls (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 

level of antenna! bases; posterolateral ex- 
tensions of gonocoxites in ventral view nar- 
row, attenuate (Fig. 39). 

Description of male holotype (5179).— 
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 
vcntrally; pedicel, length 0. 1 2 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. 



Figs. 35-40. Phyctis fronlalis {5 J SO). 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. 3^. 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 anepistcmum, on ventral third of 
katepistemum, 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 



1.4 times head depth; scape, length 0.41- 
0.44, 0.43 mm, 2.6-2.8, 2.7 times width, 
2.4-3.0, 2.7 times length of pedicel; pedicel, 
length 0.14-0.18, 0.16 mm, 0.8-1.0, 0.9 
times width; flagcllum, 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 oi Phycm fron- 
talis extends from Yucatan, Mexico to Cos- 
ta Rica. 

Specimens examined (4). — Holotype: 
male, Irwin specimen number 5179 
(AMNH), Mexico, Yucatan, Chichen Itza, 
VI-29. Paratypes as follows: MEXICO- 
Yucatan: Chichen Itza, VI- 1929, 1 M 1 F. 
COSTA RICA- La Suiza, 1924, P. Schild, 
1 F. 


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- 

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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Becker, T. 1923. Wissenschaftliche Ergebnisse der 
mil UnterstiJtzung der Academic der in Wien aus 
der Erbschafl Treill von F. Werner unlemommcn 
zoologischen Expedition nach dem angloagyp- 
tischen Sudan (Kordofan) 1914. VL Diptera. 
Denkschr. Akad. Wiss.. Wien 98: 57-82. 

Bez/,1, M. 1902. Neue Namen tur cinigc Dipleren- 
Gattungen. Z. Syst. Hymenopt. Dipterol. 2: 190- 

Bromley. S. W. 1934. Therevidae, pp. 360-361. //i 
Curran, C. H., ed.. The Diptera of Kartabo, Bartica 
District, British Guiana, with dcscnptions from 
other British Guiana localities. Bull. Am. Mus. 
Nat. Hist. 66: 287-532. 

Foerster. A. 1862. Synopsis der Familien und Gat- 
tungen der Braconen. Verh. Naturh. Ver. Preuss. 
Rheinl. 19: 225-288. 

Irwin, M. E. 1972. Diagnoses and habitat preferences 
of the immature stages of three South African 
species of the \cstomyza-gTO\if> (Diptera: Ther- 
evidae). Ann. Natal Mus. 21: 377-389. 

. 1976. Morphology of the terminalia and 

known ovipositing behaviour of female Therevi- 
dae (Diptera: Asiloidea), with an account of cor- 
related adaptations and comments on phyloge- 
netic relationships. Ann. Natal Mus. 22: 9 1 3-935. 

. 1977a. Two new genera and four new species 

of the Pheroccra-group from western North Amer- 
ica, with observations on habitats and behavior 
(Diptera: Therevidae: Phycinae). Proc. Entomol. 
Soc. Wash. 79: 422-451. 

. 1977b. A new genus and species of stlletto- 

flies from southwestern North Amenca with close 
affinities to Chilean and Australian genera (Dip- 
tera: Therevidae: Therevinae). Pan-Pac. Entomol. 
58: 287-296. 

. 1983. The hoharii species group of the genus 

Pherocera (Diptera: Therevidae; Phycinae). Pan- 
Pac. Entomol. 59: 113-139. 



Irwin. ME. and L. Lyneborg. 1981(1980]. The gen- 
era of Nearctic Therevidae. Bull. 111. Nat. Hist. 

Surv. 32: 193-277. 
Krober, O. 1914. Beitrage zur KenntnisderTherevi- 
den und Omphraliden. Jahrb. Hamb. Wiss. Anst. 

31: 29-74. 
. 1928. Neue und wenig bekannte Dipteren 

aus den Familien Omphralidae, Conopidae und 

Therevidae. Konowia 7: 116-134. 
. 1929. Neue Beitrage /,ur Kenntnisder Therc- 

viden und Tabaniden (Dipt.). Dtsch. Entomol. 

Ztschr. 1928: 417-^34. 
Loew, H. 1874. Diptera nova a Hug. Theod. Chris- 

topho collecta. Z. Ges. Naturw. Berlin. N. F. 9: 

Lyneborg, L. 1968. A comparative description of the 

male terminalia in There\'a Latr., Diatmeura Rond., 

and Psiloccphala Zett. (Diptera, Therevidae). 

Entomol. Meddel. 36: 546-559. 
. 1972. A revision of the .\'<wo»n'ra-group of 

Therevidae (Diptera). Ann. Natal Mus. 21: 297- 

. 1976. A revision of the therevine stiletto- 

flies (Diptera: Therevidae) of the Ethiopian region. 

Bull. Br. Mus. Nat. Hist. (Entomol.) 33: 189-346. 
. 1978. The Afrotropical species of Phycus 

Walker (Diptera: Therevidae). Entomol. Scand. 9: 

. 1983. A review of the Palaearctic genera of 

Phycinae (Insecta, Diptera, Therevidae). Steen- 
strupia 9: 181-205. 
Malloch, J. R. 1932. Rhagionidae (Leptidae), Ther- 
evidae, Scenopinidae, Mydaidae, Asilidae, Lon- 
chopteridae, pp. 199-283. In British Museum 

(Natural History), Diptera of Patagonia and South 

Chile. Part V, Fascicle 3. 
Olivier, A. G. 1813. Premier memoire sur quelques 

insectesquiattaquent lescereales. Mem. Soc. Agric. 

Seine 16: 477-195. 
Rauch, P. A. 1970. Electronic data processing for 

entomological museums, an economical approach 

to an expensive problem. Ph.D. dissertation in 

entomology, Ltniversity of California, Riverside. 

78 pp. 
Richards, O. W. 1929. Systematic notes on the Bor- 

bondae ( Diptera), with descriptions of a new species 

of Leplocera (Limosina). Entomol. Month. Mag. 

65: 171-176. 
Seguy, E. 1 94 1 . Dipteres recueillis par M. L. Chopard 

d'Alger a la Cote dTvoire. Ann. Soc. Entomol. Fr. 

109(1940): 109-130. 
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for entomological labels. Bull. Entomol. Soc. Am. 

19: 164-168. 
Walker, F. 1848. List of the specimens of dipterous 

insects in the collection of the British Museum. 1: 

1-229. London. 
. 1850. Diptera, 1: 1-76. /« Saunders, W. W., 

ed., [1856] Insecta Saundersiana: Or characters of 

undescribed insects in the collection of William 

Wilson Saunders. London. 
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demia Christiana Albertina aditurus analecta en- 

tomologica ex Museo Regio Havmensi. 60 pp., 1 

pi. Kiliae [Kiel]. 
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Ceylon. Termeszetr. Fiizet. 20; 136-144. 

91(1), 1989. pp. 51-54 





Ecology Program, School of Life Sciences, University of Delaware, Newark, Delaware 

Abstract. — "Wt 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 T. angmtipennis 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 T. angitstipennis 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 Tenodera sinensis is more of a habitat generalist than 
the other two species. 

Key li'ords: Mantodea, Mantidae, Tenodera, Mantis, food limitation, sympatry, pred- 

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. 1 978, 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 (Ei- 
senberg et al. 1981, Matsura and Marooka 
1983. Hurd and Eisenberg 1984, Hurd and 
Rathet 1 986). 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 

field is qualitatively the same for each 

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 ( 1 983) 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). T. angustipennis (Saus- 
sure), Mantis religiosa (Linnaeus). These 
three morphologically similar species hatch 
at different times, differ in body size (T. 
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 



I I I I I I I I I 1 I I I 
50 50 

Fig. 1. Sizes of mantid nymphs in Iwo habitats on 
19 July 1986. Open bars represent frequency ot'nymphs 
of various sizes found at site AG; shaded bars are for 
site CHRY. Ts = Tenodcra sinensis. Ta = T. angiis- 
tipennis. Mr = Mantis religiosa. One-way ANOV.^: F 
= 24.945; df = 5, 72; P < 0.001. LSD comparisons 
indicate T. sinensis is significantly larger than the other 
two species; no differences between sites. 

site and interspecific difierences, 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 (Phleiim pratcnse) 
and Canada bluegrass (Poa comprcssa) with 
patches of goldenrod (Solidago spp.) and 
thistle (Cirsiuin 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- 

We censused mantids in both habitats on 

























>— 1 

















1 I 1 1 1 1 1 1 1 

1 1 1 1 1 1 1 1 1 1 

1 1 1 1 1 1 1 1 1 1 


50 50 


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. 
angustipcnnis. Mr = Mantis religiosa. One-way AN- 
OVA: F = 25.714; df = 5, 84; P < 0.001. LSD com- 
parisons indicate T. sinensis is significantly larger than 
the other two species; site differences indicated by as- 

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 T. an- 
gustipcnnis (Fig. 1). This interspecific dif- 
ference was expected from earlier results 



Table 1. Numbers of individuals. % adult and adult sex ratio for three mantid species in two habitats (AG 
and CHRY) in July and August of 1 986. Mr = Manlis rcligiosa. Ts = Tenodera sinensis. Ta = T. angustipcnnis. 


July A' 

August ;V 

M. religiosa 

T. angustipcnnis 

T. sinensis 





























(Rathet and Hurd 1983) and published size 
differences for adults (Gumey 1950, Heifer 
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 still 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. angiistipennis 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 rrtonth at site 
AG for these two species than at site CHRY. 
Some adult M. religiosa at CHRY were con- 
siderably larger (75-80 mm) than previ- 
ously published records for this species 
(about65mm)(Gurney 1950, Heifer 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 T. 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 T. 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- 
giistipennis could be better indicators of 
habitat quality in terms of resident arthro- 
pod prey availability than T. sinensis late 
in the season. A detailed examination of the 
diets of these species in the field is needed 
to test these ideas. 


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 
8506 181, and is contribution #111 from the 
Ecology Program, University of Delaware. 

Literature Cited 

Eisenberg, R. M., L. E. Hurd. and J. A. Hartley. 1981. 

Ecological consequences of food limitation for adult 

mantids {Tenodera andtjolia sinensis Saussure). 

Am. Midi. Nat. 106: 209-218. 
Gumey, A. B. 1950. Praying mantids of the United 

States, native and introduced. Smithsonian Inst. 

Annu. Rep. 1950: 339-362. 
Heifer. 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 



fiowers to the I'nncss ol an insect predator, Tcn- 
odera sinensis Saussure (Orthoplcra: Mantidac), 
in an old field community. Entomologist (U.K.). 

Hurd. L. E. and R. M. Eisenberg. 1984. Experimental 
density manipulations of the predator icnodcra 
sinensis (Orthoptera: Mantidae) in an old-field 
comnuinity. 1. Mortality, development and dis- 
persal ol'juvenile mantids. J. Anim. Ecol. 53: 269- 

Hurd, L. E., R. M, Eisenberg, and .1. O. Washburn. 
1978. Eflbctsofe-xperimentally manipulated den- 
sity on field populations of the Chinese mantis 
(Tenodcra andijotia sinensis Saussure). Am. Midi. 
Nat. 99: 58-64. 

Hurd, L. E. and I. H. Rathet. 1986. Functional re- 

sponse and success in juvenile mantids. Ecology 
67: 163-167. 

Matsura, T., T. Inoue, and V. Hosomi. 1975. Eco- 
logical studies of a mantid, I'aralenodera angus- 
lipennis 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 
angiislipennis (S.). Oecologia 56: 306-312. 

Rathet, I. H. and L. E. Hurd. 1983. Ecological rela- 
tionships among three co-occurring mantids, 
Tenitdera sinensis (Saussure), T. angiislipennis 
(Saussure), and Manlis religiosa (Linnaeus). Am. 
Midi. Nat. 110: 240-248. 


91(1), 1989, p. 54 

Ixodes downsi (Acari: Ixodidae) from Peru 

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. Anowa g. geofl'royi 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). /. downsi has 
not been recorded since the original descrip- 

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 
5. caripensis found on the tloor of a cave 
near Tingo Maria. Department Huanuco. 
Peru, on 1 1 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 

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 Ixodes 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. 


91(1), 1989, pp. 55-58 




Ecology Program, School of Life Sciences, University of Delaware, Newark, Delaware 

Abstracl. — Mantids are members of a guild of generalist predators in arthropod com- 
munities. Three species commonly coexist in old fields: Tenodera sinensis Saussure, T. 
angiislipennis 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 T. sinensis 
hatch first, and enter the second instar as T. 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: Mantodea, Mantidae, Tenodera, Mantis, bitrophic niche, predators, sym- 

Praying mantids (Mantodea: Mantidae) 
are members of a guild (sensii 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- 

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 (Gumey 1950); its duration 
of residence in Delaware is uncertain (Rath- 
et and Hurd 1983). All three species are 
sympatnc m Asia (Jing-jin et al. 1981). 

We have been studying various popula- 
tions of mantids, especially T. sinensis, in 



northern Delaware over the past decade (Ei- 
senberg and Hurd 1977, Hurd et al. 1978, 
Eisenberg et al. 1981, Rathet and Hurd 1 983, 
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 (Hartley 
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 T. 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 (Cirsiuni 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). 


Tenodera sinensis began hatching after 1 8 
days of incubation, and hatching continued 
for 1 1 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 ( 1 977). 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 < O.OI). 

Tenodera angustipennis did not start 
hatching until T. sinensis 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- 



ference in hatching pattern (Fig. 1). Hatch- 
ing time was intermediate, and overlapped 
with both Tcnodera species. Unlike Teno- 
dera spp., all but one ootheca produced 
nymphs on consecutive days, as many as 
five days in a row. As with T. sinensis. 
nymphs from site CHRY were larger (6.1 
mm, SD = 0.3 1 ) than nymphs from site AG 
(mean = 5.8 mm, SD = 0.20) (t = -3.78, 
df= 58, P < 0.001). 


We found that under identical laboratory 
conditions of a "common garden" experi- 
ment, eggs of T. 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 T. 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 T. angustipen- 
nis and M. religiosa begin to hatch, the ear- 
liest T. 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. 

50 n 


P 50- 


O 30 


O 10-1 


Mantis religiosa 


Tenodera angustipennis 

—1 — 1 — I — I — I — r 

Tenodera sinems 





T i — I 


25 30 


Fig. 1. Timing and patlemofegg 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 T. 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 T. 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 T. sinensis. The difference in nymph 



size may well be an indieation of diircrences 
in feeding eonditions 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 alfected. 
This would indicate that adults of these two 
species were more food limited at site AG 
than at site CHRY. We are currently testing 

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 /'. sinensis, at least, is tlexible in this 
regard (Hurd and Rathet 1986). We cannot 
as yet olfer a satisfactory explanation for the 
ditference in hatching patterns for M. reli- 
giosa (unless ecotypic); however, roughly the 
same pattern dilFerence 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. 


This work was supported by NSF Grant 
BSR 8506181. This is contribution #110 
from the Ecology Program, University of 

Literature Cited 

Bartlcy. J. A. \9^i. Prey selection and eaplure b\ the 
Chinese nianlid (Icnodcni sinensis Saussure). 
Ph.D. dissertation. Dniversity of Delaware. New- 
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Eisenberg. R. M. and L. E. Hurd. 1977. .An ecological 
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Eisenberg, R. M., L. E. Hurd. and J. A. Hartley. 1 98 1 . 
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mantids {Iciutdera arulifolui sinensis Saussure). 
Am. Midi. Nat. 106: 209-218. 

Gurney, A. B. 1950. Praying mantids of the United 
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Hurd, L. E. 1985. Ecological considerations of man- 
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. 1988. Consci|uences of divergent egg phe- 
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Hurd, L. E. and R. M. Eisenberg. 1984a. Experi- 
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Ienodera sinensis (Orthoptera: Mantidae) in an 
old-lield community. I. Mortality, development 
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53: 269-281. 

. 1984b. Experimental density manipulations 

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Hurd, L. E., R. M. Eisenberg, and J. O. Washburn. 
1978. Etl'ecls of experimentally manipulated den- 
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Hurd, L. E. and I. H. Rathet. 1986. Functional re- 
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Jing-jin, Y., X. Chong-hua. Y. De-fu, and L. Ying-mei. 
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.loncs. F. M. 1933. .Another oriental mantis well es- 
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Laurent. P. 1898. A species of Orthoptera. Entomol. 
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Rathet. I. H. and L. E. Hurd. 1983. Ecological rela- 
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Midi. Nat. 110: 240-248. 

Roeder, K. D. 1936. Raising the praying mantis for 
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91(1), 1989, pp. 59-65 




Christer Hansson 

Department of Systematic Zoology. Lund University, Helgonavagen 3, S-223 62 Lund, 

Abstract.— T\iQ genera Holcopeltc Forster and lonympha 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. lonympha is represented by two species in the Americas: /. came (Walker) 
from Canada, Ecuador and Peru, and /. ochus (Walker) from Canada. The hosts are 
unknown for all four species. 

Key IVords: Eulophidae, Holcopelte. lonympha. New World, taxonomy 

Forster (1856) described Holcopelte to in- 
clude Elachistus obscurus (Forster, 1841). 
Later, he described H. fulvipes ( 1 86 1 ), which 
was later synonym ized with H. obscura by 
Boucek & Askew (1968). Thomson (1878) 
included Holcopelte as a species-group in 
the genus Derostenus; he also described D. 
sulcisciita 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. Erdos (1958) described Horis- 
menits 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 stelteii 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 

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- 
cek & Askew 1968, Boucek 1969). 

lonympha was described by Graham 
(1959) to include Entedon came 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; 



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 FoTsier, 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 scutelium (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- 

The American species difter 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 

1 . Females 2 

- Males 3 

2. Antcnnal 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 

along hind part of scutelium); cubital hair-line 
offorcwingalmost straight (Fig. 1); propodeum 
sculptured and with 4 plicae (Fig. 1); petiole 

conical without a dorsal shield 

//. amcncana 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. II); petiole quadrangular (shape 
transverse to slightly elongate) with a dorsal 
shield that covers petiolar foramen (Figs. 10, 

11) //. 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); (lagellum infuscate, segments 

2.0-2.5 X as long as wide 

//. amencanci 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 X (apical segment 5.0 x) as long as 

wide and slightly narrowed medially 

H. huggerli 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 x 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. — f^/?;fl/e' Scape yellowish- 
white, more or less infuscate in apical part, 
remaining antenna brown. Face, frons and 
occiput golden-green, occasionally bluish. 
Thorax var\ing from blackish to brownish, 
usually also with a slight golden-green tinge 



Figs. 1-9. Ilnkopclw spp. 1,9//. anwricana new species, habitus. 2, Same, lateral view of head. 3. Same, 
frontal view of head. 4, 6 H amcncana. lateral view of head. 5, Same, frontal view of head. 6, 9 II. huggerli 
new species, lateral view of head. 7, Same, frontal view of head. 8, 6 II. huggerli, lateral view of head. 9, Same, 
frontal view of head. Scale representing 0.30 mm. 



Figs. 10-14. \0-\2, 9 Holcopellehuggerli new species. 10, Side view of petiole. 1 1, Dorsal view of propodcum 
and petiole. 1 2, Base of forewing. 1 3-14, 9 lonympha tarm' (Walker). 1 3, Frontal view of head. 14, Left mandible. 
Scale to the left representing 0.25 mm (Fig. 10), and 0.50 mm (Figs. 1 1-13); scale to the right 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. Gastcr 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 , II about 2 X , III 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 x 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 comers of petiolar fo- 
ramen, reaching halfway 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 



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: 1 mm. Scape apically expanded, 
flagellar segments I-III about 2 x , IV 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 9 labelled: 
"USA: West Virginia, Greenbrier Co., 10 
miles E. Richwood, Summit Lake, 
29.vii.1983, leg. L. Huggert," in LUZM. 
Paratypes: 2 9 with same label as holotype; 
1 9 "USA: West Virginia, Greenbrier Co., 
Richwood, Summit Lake, 27.vii.1983, leg. 
L. Huggert"; 2 9, 1 3 "USA: West Virginia, 
Pocahontas Co., Falls of Hills Creek, 
22.vii.1983, leg. L. Huggert"; 3 9 "USA: 
West Virginia, Pocahontas Co., Falls of Hills 
Creek. Monongahela Forest, 26.vii.1983, 
leg. L. Huggert"; 1 <5 "USA: West Virginia, 
Pocahontas Co., Dogway Road, Monon- 
gahela Forest, 26.vii.1983. leg. L. Huggert"; 
1 9 "Canada: Quebec, Bouchette, Lac Rod- 
dick, 12.ix.1982, leg. L. Huggert." These 
paratypes deposited: 3 9, 1 <5 in CH, 5 9, 
1 $ in LUZM, 1 9 in USNM; 1 9 
"N.S.C.B.H.N.P., Skyline Trail, 25.vii. 
1983," "Birch PG 635802, B.R.L Survey"; 
1 9 "Prince Edward L, Sand Banks Prov. 
Pks., 25.vii.1982, L. Masner"; 1 9 "St. Law- 
rence Is. Nat. Park, Ontario, Thwartway Is.," 
" 1 9.vii. 1 976, L. Masner. Code 42 1 2-7"; fi- 
nal three paratypes in CNC. 

Distribution. — Canada (Nova Scotia. 
Ontario, Prince Edward Island, Quebec) and 
the United States (West Virginia). 

Holcopelte huggert i. New Species 

Figs. 6-12 

Diagnosis. — Petiole quadrate (9), or 1.5- 
3.0 X as long as wide (S) with a thin dorsal 
shield that covers petiolar foramen; cubital 

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 x 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. 

DescnpXion. — 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, H, III and V about 
3.8 X, and IV about 3.4 x 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 Vi of meso- 
scutum to anterior 'A of scutellum, some- 
times absent on mesoscutum. Notaular 
depressions like anicncana. Midlobe of 
mesoscutum separated from scutellum by a 
narrow furrow. Dorsellum short, 5.9 x as 



wide as long, with surface convex. Shape of 
wings hke 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 
halfway 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 X 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 x as long as wide. Malar 
space 1.2 X 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 9 labelled: 
"US/^: West Virginia, Greenbrier Co., 10 
miles E. Richwood, Summit Lake, 29.vii. 
1983, leg. L. Huggert" in LUZM. Paratypes: 
2 2 with same label as holotype; 1 9 "USA: 
West Virginia, Pocahontas Co., Falls of Hills 
Creek, Monongahela Forest, 26.vii.1983, 
leg. L. Huggert"; 1 9 "Peru: Huanucu, To- 
cache, 2.ii. 1 984, leg. L. Huggert"; 1 9 "Peru: 
Cusco, Machu Pichu, 21.xii.l983, leg. L. 
Huggert"; 2 9 6 <? "Peru: Cusco, Aqua Cal- 
iente, 28.xii. 1983, leg. L. Huggert"; 1 9 "Ec- 
uador: Napo, Lumbaqui, 10-1 1 .iii. 1 983, leg. 
L. Huggert"; 1 9 "Ecuador: Pichin, Rio Pal- 
enque, 4.ii.l983, leg. L. Huggert"; 2 9 "Ec- 
uador: Pichin, Puerto Quito, 2.iii.l983, leg. 

L. Huggert"; 4 9 2 <5 in CH, 6 9 3 <5 in LUZM, 
1 9 1 <? in USNM. 

Distribution. — The United States (West 
Virginia), Ecuador and Peru. 

Genus lonympha Graham 

lonympha Graham, 1959: 199. Type- 
species: Entedon ochus Walker, 1 839: 2 1 , 
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 lonympha 
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 /. came with 
2-5 setae on propodeal callus, 7-9 setae in 

lonympha came (Walker) 

Figs. 13-14 

Entedon came V^a\keT, 1839: 123. 
lonvmpha came (Walker), Graham, 1959: 


Material. -CANADA: 1 9 Ontario, Ron- 
deau Prov. Park. 26. viii. 1 982. ECUADOR: 
1 9 Pichin, Tinalandia, 800 m, 7.ii.l983. 
PERU: 1 9 Cusco, Ollantaytambo, 19.xii. 
1983; 1 9Cusco, Machu Pichu, 21. xii. 1983. 
All leg. L. Huggert ( 1 9 in CH, 3 9 in LUZM). 
Type material of E. came in BMNH (not 

Remarks. — Most European specimens of 
this species have dark coxae and remaining 
parts of legs predominantly infuscate. Three 



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 /. came 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. lonympha carne was previously 
not recorded from the Americas. 

lonympha ochus (Walker) 

Entedon ochus Walker, 1839: 21. 
lonympha ochus (Walker). Graham, 1959: 

Material. -CANADA: 1 9 British Co- 
lumbia, N. Vancouver, 31.viii.l960, S. M. 
Clark (CNC). This specimen agrees well with 
European specimens. Type material of /. 
ochus in BMNH (not seen). 

Distribution. — West Palearctic (Boucek 
«fe Askew 1 968) and Canada (British Colum- 

bia), lonympha ochus was previously not 
recorded from the Nearctic Region. 


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. Pan II. J. Linn. Soc. (Zool.) 25: 108-188. 

. 1904. Classification of the chalcid llies or the 

superfamily Chalcidoidea, with descriptions of new 
species in the Carnegie Museum, collected in South 
America by Herbert H. Smith. Mem. Cameg. Mus. 
I(I-IX): 225-551. 

Boucek, 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. Telrastichinae). Index of Ento- 
mophagous Insects. Lc Francois, Pans. 254 pp. 

Erdos, J. 1958. Eulophidae in Hungana recenter de- 
tectae. Acta Zool. Hung. 3: 205-223. 

Forster, A. 1841. Beitrage zur Monographic der 
Pteromalinen Nees. Aachen. 46 pp. 

. 1856. Hymenoptcrologische Studien. II. 

Chalcidiae und Proctotrupii. Aachen. 152 pp. 

. 1861. EinTaginden 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. 1 987. 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. 

91(1), 1989, pp. 66-70 


Emily 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. 

^torac?.— 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 ^ 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 

Key Words: wood loving, tree damage, squirrel activity 

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 1 900, 
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. 


Storm and squirrel damage.— Two sites 
in northeast Ohio were monitored for two 

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 cecidomyiids. 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 



■*l — I 1 r~T 1 1 — I — I — I 1 i 1 1 1 — T — ^■^— 1-:3^ ^ 

u.o o.r> III i.r, 2.0 2.5 ;i.ii :i :> lo i.r. 5.(1 r..r. d.o 6.5 7.» 7.r. u.o ii.r. ')X) ■)..■; iii.ii 


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 

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 
n 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 alhidum (Nutt.) Nees) branches 
with diameters varying from 3 mm to 20 
mm were offered to a population of 300 
caged Xylodiplosis longistylus Gagne fe- 
males for one week. The branches were held 
at room temperature in plastic bags for 14 



0.r> ID 1.5 2.0 2.5 :i.() 3.5 4.0 4.5 5.0 5.5 0.0 6.5 7.0 7.5 8.0 8.5 9.0 


Fig. 2. Number and size of live branches broken from hosl trees by squirrel activity. 

days. Each was then soaked in a separate 
container of water, and the number of larvae 
emerging from both exposed ends was re- 

Period of suitabiHty 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 

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 

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 (Sciiinis nigra) are known to build 
nests in many of the hardwoods that are also 
hosts to xylophilic midges. Uhlig(1955), in 




o _ 



0.9 _ 
0.8 _ 
0.7 _ 
0.4 _ 
0.3 _ 





3. Relationship of number of larvae collected to branch diameter: Sassafras albiduin. 

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 

>75 M d 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 >15 n d/mm- 
in each end, and it yielded over 1 700 larvae. 

Preliminary field data indicate that actual 
utilization of vessels is much less than in 
laboratory samples. A 1 2 mm d ash 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- 



Table \. Relationship of host sample diameter to 
length of attractiveness. 

Hosl Plant 





Juglans nigra L. 



Fraxinus amehcana 



Carya ovala (Mill.) K. Koch 



Quercus alba L. 



Ulmus ainencana L. 



Populus detloides Bartr. 



v. amencana 



Sali.x babylonica Marsh 



Sassafras albidum (Nutt.) Nees 



F. amerkana 



F. amerkana 

2 to 4 


F. amerkana 

1 to 2 


F. amerkana 

0.6 to 1 


* 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, in- 
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. Monographic des Cecidomyides 
d'Europe et d'Algene. Ann. Soc. Entomol. Fr. 69: 
181^72 and pis. 15-^4. 

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: 

Shorten, M. 1954. Squirrels. Collins, London. 212 

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. 

91(1), 1989, pp. 71-80 


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. Biichnerillo litoralis Verb, 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 

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 
1936,1 940, 1 942. 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 

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 oi creepers (Schmalfuss 
1984, Paoletti 1987). 



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. 3 A, C). 





532 - 




0492 - 







y =13 8493'0142e x 
r'.O 2107 

0412 - 


p. 01942 

Length (mm ) 

Fig. I . Regression for male dimensions of Metas- 
tenoniscus neotropicalis n. sp. 

Length (mm) 

Fig. 2. Regression on female dimensions of Me- 
lastenoniscus neotropicalis n. sp. 

Head: with three posterior main tubercles 
and two median main tubercles (Figs. 3A, 
B, C); eye with 3-4 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. 43), 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- 

eonite from 1 to 4 has medial costa well 
developed (Taiti and Ferrara 1981 Figs. 
ICand 2C); 

5. telson shorter and distinctly three-lobed; 

6. uropod exopods shorter and with wider 

Habitat. Found under Coccolobis uvifera 
(L.) Jacq. (Polygonaceae) (uva de plaja) lit- 
ter, eating decayed litter tissue (Paoletti 1 987 
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. Stenonisciis 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 



Fig. 3. Metastenoniscus neolropicaUs n. sp. Female: A dorsal view. B head frontal view, C lateral view, 
pleon and telson, E and F second antennal flagellum article from dorsal and ventral views. 


Vandel 1962) in Florida Cays, Tavemier 
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), Tylos niveus Budde-Lund. 
and Annadilloniscus ellipticus (Harger, 



Armadilloniscus caraibicus n. sp. 

Type locality. Coralline cays (Cayos) of 
Parque Morrocoy, estado Falcon, Venezue- 
la: Cayo Sombrero, one gravid female 
31. XII. 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 



Fig. 4. Metastcnoniscus 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. 1 1 F, H) and 
digitiform trichomes. 



Fig. 5. Metastenoniscus neotropicalis n. sp. Male, A and B pleopod I endopods, C plcopods I exopod, D 
pleopods II endopod. Armadilloniscus caraibicus n. sp. Male, E pleopods I, F pleopods II, G VII male pereopods. 

Dimensions: length males: 1.92-2.46 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 

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 

1 so- 

y - 0.0437 

0,4449 X 


r'=0 37 

ns - 

p. 0,15 





094 - 





23 - 

y = 0302 •0 4546 X 




A y/^ 


A jr 


11 - 

j^ A 





204 219 2 34 249 1.1 19 2 7 

Length (mm) Length tmmi 

Fig. 6. Armadilloniscus caraibicu.'in.ip. Regression Fig. 7. Armadilloniscus caraibicus n.sp.Regxession 

of male dimensions. of non gravid female dimensions. 



y ,07228'02364 X 
r'. 205 
p^ 0397 

2 82 

3 24 

3 38 

296 3 10 

Length (mm) 

Fig. 8. Annadilloniscus caraibicus n. sp. Regression 
of gravid female dimensions. 


y = 21322-01178 x 
r'.0 564 
p = 00912 


58 6 7 76 

Pulli per lemale 

Fig. 9. Annadilloniscus caraibicus n. sp. Regression 
of gravid female length and pulli per female. 

Fig. 10. Armadillomscus caraibicus n. sp. Female: A and B pleon; C and D head and pereon. 



Fig. 1 1. Armadilloniscus caraibicus n. sp. Female: A-D body surface in difTerent perspectives; E head; F 
pereonite ornamentation; G and H pleon and peculiar tubercle ornamentation. 

anterior part (Figs. IOC, D, 1 1 A, C, D, E). 
Eye with 4-5 visible ommatidia; antenna, 
without enlargements, holds four flagellar 
articles, the second with three and the third 
with two aesthetascs on the inferior surface 

(Figs. 12C, D); antennule characteristically 
three-articulated and apically bifurcated; it 
bears aesthetascs (5 on specimens exam- 
ined) (Figs. 12A, B). 
Body ornamentation: pereon with four 



Fig. 1 2. Annadilloniscus caraibkus 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. 



main medial ridges and four shallow lateral 
ridges of tubercles (Figs. IOC, 1 lA-E);pleon 
supporting two main ridges of tubercles 
(Figs. lOA, B; IIB, D, G). 

Uropods: endopod somewhat longer than 
basis, holding a distal spike of 3-4 aesthe- 
tascs (Figs. lOB, IIG). Male pereopods I 
and VII slightly modified (Fig. 5G). Pleo- 
pods: male pleopods vary little from other 
members of the genus (Figs. 5E-F). 

Affinities. The development of tubercle 
ribs on the head, pereon and pleon are the 
features ofAnnadilloniscus caraibiciis 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. cownacapitalis 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 Armadillouiscus quadh- 
ro;7»s 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 Armadillouiscus. 

Distribution and Discussion. Anuadil- 
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 is 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. 


We are indebted to Carlos Bordon for field 
collection and discussion. A. L. Dreon, C. 
Furlan. U. Arezzini, C. Friso assisted with 

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 Slate M useum of Natural 
History. Arkiv for Zool. 12: 537-569. 

Arcangeli, A. 1957. 11 genere Annadilloniscus Ulj. e 
gli Scyphacidae. Atti Ace. Sc. Torino 91: 1-30. 

Brian. A. 1957. Descrizione di Neosanfilippia vene- 
zueiana n. gen., n. sp. di Isopodo terreslrc trog- 
lobio. Ann. Mus. Civ. St. Nat. Genova 69: 352- 

Ferrara, F. and S. Taiti. 1981. Isopodi terrestri delle 
isole Adamane. Boll. Mus. Civ. St. Nat.. Verona 
8: 459^92. 

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 culicular 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: 9^8. 

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. Contribucional conocimientode 
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 terrestnal 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: 477^88. 

. 1972. Ecology and systeniatics of terrestnal 

isopod crustaceans from Bermuda. Crustaceana. 
Supp. 3, pp. 79-99. 

. 1977. Terrestrial isopod crustaceans from St. 

Catherines Island. Georgia. Georgia J. Sci. 35: 151- 

. 1983. Disposition of three species of Onis- 

coidea from Western Atlantic seashores. Proc. Biol. 
Soc. Wash. 96; 440-45 1 . 



. 1984. Three new and five other species of 

Oniscoidea from BeUze, 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; 

Strinati, P. 1971. Recherches biospeleologiques en 

Amenque du Sud. Ann. de Speleologie 26: 439- 

Taiti, S. and F. Ferrara. 1981. Melastenomscus oscllai 

genere e nuova specie di isopodo terrestre dell'isola 

di Bali. Boll. Mus. Civ. St. Nat.. Verona 8: 443- 

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- 

gique beige 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, 

— . 1 972. Les Isopodes terrestres de la Colombie. 
Studies on the Neotropical Fauna 7: 147-172. 

— . 1973. Les Isopodes terrestres de la Melanesie. 

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. 
. 1 942. A second supplement to the American 

land and fresh-water isopod Crustacea. Bull. Amer. 

Mus. Nat. Hist. 80: 299-329. 

91(1), 1989, pp. 81-87 


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, 

Abstract.— We describe the ant fauna (1 1 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-occuiTcnce. 

Key Words: Annonaceae, ants, Brazil, coleopterous gall, community ecology, habitat 
selection, insect galls, Minas Gerais, resource partitioning, Xylopia aro- 

Plant galls represent an important re- galls. Mani ( 1 964) divides the locatari into 

source for many species other than the gall 33 categories according to their ecological 

formers. Owing to their localized concen- niches. The locatari often represent a vast 

tration of nutritive tissues and their marked fauna with the most numerous categories 

succulence, galls provide favourable breed- consisting of parasites and predators. For 

ing sites for a variety of species (Brandhorst example, Stegagno (1904) reported 177 

1962, Mani 1964, Shorthouse 1973, Yu- species associated with cynipid galls on 

kawa 1983). Galls protect their inhabitants Quercus in Italy; of these, 138 are parasites 

from inclement weather (Felt 1940, Uhler and predators. 

1951, Sandlant 1979; but see Baust et al. Species that inhabit galls after emergence 

1979) and natural enemies (Askew 1961, of gall formers and inquilines are the "suc- 

1980, but see Price et al. 1986, 1987). Use cessori" (Mani 1964, Yukawa 1983). Most 

of galls ranges from species that open galls of these are plant-nesting ants and myr- 

simply to prey upon gall formers and in- mecophilous insects, such as aphids and 

quilines to those that depend exclusively on coccids. Others include mites, spiders, 

gall tissues for food and shelter. thrips, collembola, bees, and wasps (re- 

Beauvisage (1883, cited in Mani 1964) viewed by Mani 1964). The most common 

applied the term "locatari" to species, other ant genera associated with old galls are: 

than the gall formers, associated with insect Camponotus, Cataulacus, Crematogaster. 



Fig. 1. Location of the Ecological Station of Parapitinga in Tres Marias, Minas Gerais, Brasil. 

Colobopsis, Lasius. Leptothorax, and Olop- 
5/5 (Walsh 1864, Patton 1879, Wheeler 1910, 
Ping 1920, Sturtevant 1925, Mam 1964, 
Torossian 1971a, b). Despite their impor- 
tance and dominance within the successoin, 
little is known about the ecologies of these 
ants other than compendiums ol 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 

Material and Methods 

One of us (G.W.F.) collected galls from 
the Ecological Station of Parapitinga, Tres 
Marias, Minas Gerais, Brazil during one 
sample period in mid-May, 1984. The sta- 
tion is a continental island located in a man- 

made lake (Tres 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- 



Fig. 2. Coleopterous leaf-bud galls found on Xylopia aromatica. Gall diameter ranged from 5.0 to 
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 

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 1 969, 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. 



Table 1. Ant species associalcd with coleopterous leaf-bud galls on Xylopia aromalka (Annonaceae). 

i.X) Individuals 

Gall Diameler 

Anl Species 

NumlKr of (\)lonics 



{.?) mm 

Azura huolor 





Azieca sp. 





Lcplolhorax wilda 





Leptolhorax sp. 





Pseudomyrmex lla\tdus 





Pscudomyrmex sp. A 





Pseudomyrmex sp. B 





'/.acryploccrus pallens 





Zaayptocenis pusdius 





Zacryptoccrus sp. \ 





Zacryploccnis sp. V. 





We compared the mean diameters of oc- 
cupied galls among the si.\ most common 
species of ants by one-way analysis of vari- 

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 X, the 
Poisson parameter, from the sample. We 
compared the observed distribution to the 
expected distribution with a Chi-square 
goodness-of-fu test. 


Wecollected 1 Mgallsof 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 1 7 galls, Leptolhorax wilda which 
occupied 1 3 galls, and Zacryploccnis 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 Zacryploccnis 
piisillus which contained 1 29 adults and lar- 

The internal structure of the galls was 
highly modified in some cases, unmodiiied 

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 Azicca sp. and Zacryploccnis 
pusillus exhibited extensive modifications 
when compared to unmodified galls. Galls 
inhabited by Pseudoiuyrmcx Jlavidus were 
unmodified. Modified galls typically con- 
tained larger colonies than did unmodified 

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, 1 3.6 mm 
(F, ,,, = 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'm = 72.1, P < 

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 (Fs^4 = 2.74, P < 0.05). Az- 
ieca sp. typically occupied the largest galls 
(.V = 16.4 mm), Zacryptocenis pallens the 
smallest (.v = 12.9 mm). 

We collected 65 colonies distributed 




r—" — I 


Fig. 3. Cross-sections of galls showing internal modifications by ants. Gall (A) contained Azteca sp., 
Lcplolhora.\ sp.. (C) Zacryplocems pallens. (D) Zacryptoccms pusillus. and (E) Pseudomynnex flavidus. 


among 1 14 galls. We estimated X, the Pois- 
son parameter, as 65/1 14 = 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- 

ference between the observed and expected 
distributions (x", = 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 

6 7 9 11 13 15 17 19 21 2J 


Fig. 4. Relative frequency histograms of gall di- 
ameter for Ihc entire set of galls (foreground) and for 
the subset of galls that were occupied by ants (back- 


p<0 001 













-•• . •• • 

160 180 

— I 

Fig. 5. Relationship between the logarithm of col- 
ony si/e (number of individuals) and gall diameter. 
The relationship is Y= -0.929 -i- 0.122X(F,„ = 20.1. 
P < 0.001; R- = 0.24). 



Table 2. Distribution of colonies among galls. Ex- 

Table 3. Patterns of co-occurrences among ant 

pected values are based on a Poisson distribution with species. 
A = 0.57. 















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- 
lens exhibited a slight tendency to share galls. 


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 

Joint Occurrences 


Azteca bicolor 
Azteca sp. 
Lcptothorax wilda 
Leptothorax sp. 
Fseudomyrmex fla vidus 
Fseudomynnex sp. A 
Pseudomyrmex sp. B 
Zacryptocerus pallens 
Zacryptocerus pusdius 
Zacryptocerus sp. A 
Zacryptocerus sp. B 





















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- 

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 pusiltus 
never shares a gall, although approximately 
four joint occurrences are predicted. Fowler 
et al. (1985) provide evidence that Z. pusit- 
lus 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 


We thank Helcio R. Pimenta and Ivo das 
Chagas for field assistance. We also thank 


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 Xylopia aro- 
niatlca, and W. E. Claric 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- 
Tres Marias, MG provided logistical sup- 
port. This work was supported by a spe- 
cialization fellowship from SEMA/CNPq (n° 
10.6193/84) to G. W. Femandes. 

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Wheeler, W. M. 1910. Ants. Columbia University 
Press, New York. 

Yukawa, J. 1983. Arthropod community centered 
upon the ncolitsea leaf gall midge, Pscucloasphon- 
dylia lU'olit.seae Yukawa (Diptera: Cecidomyiidae) 
and its host plant, NeolUsea .sc ntva (Blume) Koldz, 
(Lauraceae). Memoirs of the Faculty of Agricul- 
ture 19: 89-96. 

91(1), 1989, pp. 88-111 


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 Anthonoiniis oniatus species group are A. ornatus 
Blanchard, A. sii^nalipcnnis Blanchard, A. chilicola Clark, new species, A. kuscheli Clark, 
new species, A. araucanus Clark, new species, A. blanchard! Clark, new species, and A. 
berberidis Clark, new species. These arc hypothesized to constitute a monophylctic 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 .(. ornatus and .1. kuscheli are also described and illustrated. 

Key Words: Anthonomus, Anthonominae, PAUP, phylogeny, Berberis, Chile, Argentina 

The Anthonomus ornatus group includes 
.4. ornatus Blanchard, A. signatipcnnis Blan- 
chard, and live 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 .1. 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 .-1. 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 

completely known laxonomically. Mean- 
while, the descriptions, illustrations and keys 
presented in this paper should facilitate 
identification of the species in the .4. ornatus 

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, 



CWOB Collection of C. W. O'Brien, Tal- 
lahassee, Florida, USA; 

HAHC Collection of H. and A. Howden, 
Ottawa, Ontario, Canada; 

MCZC Museum of Comparative Zoology, 
Cambridge, Massachusetts, USA 
(S. R. Shaw); 

MNHN Museum National d'Histoire Nat- 
urelle, Paris, France (H. Perrin); 

MNNC Museo Nacional de Historia Nat- 
ural, Santiago, Chile (M. Elgueta 

MZSP Museu de Zoologia, Universidade 
de Sao Paulo, Sao Paulo, Brazil (U. 
R. Martins); 

NZAC Division of Scientific and Indus- 
trial Research, Auckland, New 
Zealand (G. Kuschel); 

TAMU Texas A&M University, College 
Station, Texas, USA (H. R. Burke); 

USNM National Museum of Natural His- 
tory, Washington, D.C., USA (D. 
R. Whitehead). 

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 ( 1 947) and Thomas (1957). Where 
differences in terminology exist between the 
two systems, the Thomas terms are placed 
m parentheses following those of Anderson. 

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. 1 9-30). 
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 vitla; pleuron with 
small anteromedian patch of pallid, broad 
scales that is feebly developed in small spec- 
imens. Elytra: 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. 42^5). The species in 
the.-l. ornatus group are known mainly from 



^"^^L a^% 

Figs. 1-6. Habitus of adults (lines represent 1 mm): 1) .-1. signalipennis. male, Quinta Pittet. Magallanes, 
Chile, lateral view; 2) the same, dorsal view; 3) A. ornalus, male, Rio El Ganso, Seno de Otway, Magallanes, 
Chile, lateral view; 4) the same, dorsal view; 5) A. kuscheli. female, Chepu, Chiloe, 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 
( 1 85 1 : 387) stated that both A. signalipennis 
and A. ornatus were found in "la provincia 
de Coquimbo." These records, along with 
records of .4. signalipennis 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 
















—'■ ■ A. 

Figs. 7-12. Habitus of adults (lines represent 1 mm): 7) A. araucanus. male, Laguna Malleco, Malleco, Chile, 
lateral view; 8) the same, dorsal view; 9) A. chilicola. male, 20 km. E Manzanar, Malleco, Chile, lateral view; 
10) the same, dorsal view; 1 1) .-1. herbertdis. male, Frutillar, Llanquihue, Chile, lateral view; 12) the same, dorsal 

temperate plant species (not including Ber- 
beris, the known hosts of the members of 
the .4. ornatus group), as occurring in these 
moist refugia. 

Plant associations. Six of the 7 species of 
the A. ornatus group are known to be as- 
sociated with one or more of the following 
Berberidaceae: Berberis buxifolia Lamarck; 

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 .1, sig- 
natipennis and A. ornatus have been col- 





Figs. 13-16. Pygidium, adult female, dorsal view (line represents 0.25 mm): 13) A. signatipennis. Quinta 
Pittet, Magallanes, Chile; 14) .-(. ornatus. Lago Frio, Aisen, 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, 
Aisen, Chile; 18) A. blanchanh. El Coigo, Curico, Chile. 

lected together, on the same date, at several 
locahties. One of these records indicates that 
specimens of the two species were taken on 
the same day on B. buxifolia. Larvae of one 
of the species, A. ornatus, are known to de- 

velop in flower buds, but the developmental 
site oi A. signatipennis is unknown. Adults 
of the two species have also been taken at 
the same locality on B. ilicifolia. but one 
day apart. Adult A. blanchardi have been 









Figs. 19-30. Aedeagus, adult male (line represents 0.5 mm): 19) A. signatipemus. Estancia Fenlon, Magal- 
lanes, Chile, dorsal view; 20) the same, lateral view; 21) .-1. ornaliis, Trapatrapa, Concepcion, dorsal view; 22) 
the same, lateral view; 23) A. ornatus. Lago Frio, Aisen, dorsal view; 24) the same, lateral view; 25) .-1. blanchardi. 
El Coigo, Cunco. Chile, dorsal view; 26) the same, lateral view; 27) .-1. kuscheli. Pichmahuel. Malleco, Chile, 
dorsal view; 28) .(. araucanus. 20 km. E Manzanar, Malleco. Chile, dorsal view; 29) A. chilicola. 20 km. E 
Man/.anar, Malleco, Chile, dorsal view; 30) A. hcrbcridis. Frutillar, Llanquihue, Chile, dorsal view. 




Figs. 31-37. Third inslar larva o( Anihonomus kuscheli. 31) lateral view; 32) head capsule, frontal view; 33) 
labium and right maxilla, ventral view; 34) epipharvnx; 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. 

Figs. 38^1. Pupa oi Anihonomus ornalus. 38) head, rostrum and ventral view of prothorax; 39) prothorax, 
dorsal view; 40) terminal abdominal segments, dorsal view; 41) 9lh abdominal segment; lateral view. Line 
accompanying Fig. 38 = 1 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, A. kuscheli 
and A. berberidis, have been collected on B. 
danviiii at the same time and locality. 

Discussion. No observed characters of 
adults of the members of the A. ornalus 
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. ornalus 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 .4. kuscheli are distinct in 
possessing three rather than four epipharyn- 

Fig. 42. Map of southern South America showing the 
Fig. 43. Map of southern South America showing the 
Fig. 44. Map of southern South America showing the 

Fig. 45. Map of southern South America showing the 

(triangles) and .(. berhendis (squares). 


distribution o( A. signalipennis. 

distribution of .-1. ornatus. 

distributions of.), kmchcli (circles) and .1. clulicola 

distributions oi A. hlanchardi (circles), A. araucanus 



geal sensory pores. Larvae of Coccotonis 
sciitellaris (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 A. ornatus group. 
The pupae of members of the A. 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. slupulosus Champion, A. ghseis- 
quamis Champion, A. unipttstulatus Cham- 
pion and A. niibiloidcs Fall (Burke 1968). 

The association of the species in the A. 
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 A. 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 Willmk 
(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 ( 1 934) 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) 


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 2 

2. Sutural elytral interstnae with acute, overlap- 
ping apical extensions; metatibia of male with 
ventral margin strongly concave in distal -A, 
with enlarged, excavated apical mucro; aedea- 
gus asymmetrical (Figs. 29, 30) 3 

2'. Sutural elytral interstnae 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) 4 

3. Sternum 5 of male with large posteromedian 
prominence (Fig. 11); pallid annuli around 
posterolateral elytral maculae weakly devel- 
oped (Fig. 12) herhendis 

y . Sternum 5 of male without 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 offemale 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 
offemale with shallow emarginations (Fig. 1 8) 
or deep incisions (Fig. 1 7) on each side of me- 



dian prominence; acdeagus with apicolateral 
prominences (Figs. 21, 23, 25); pygidiuni of 
female without apicomedian prominence (Fig. 
14) 6 

5'. Elytral pattern weakly developed, except for 
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. 1 7); acdeagus (Figs. 2 1 , 23) about 

as broad at broadly rounded apex as at base, 

with acutely pointed apicolateral prominences 


6'. Sternum 3 of female (Fig. 18) with small, shal- 
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 

Figs. 1, 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 
del. 19](MNHN). Blackwelder 1947: 839. 
Schenkling and Marshall 1934: 59. Wib- 
merand O'Brien 1986: 204. 

Recognition (Figs. 1, 2). Adults of .-I. 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. 1 9, 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 

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 A. 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. 
19, 21, 23). 

Adult male. Length: 3.28-3.64 mm (.y = 
3.40, n = 10). Width: 1.46-1.60 mm (.v = 
1.53, n= \0). Rostrum: \Qx\%\h 1.30-1.51 x 
(.V = 1.40, n = 10) pronotal length; distal 
portion finely punctate, length 18-30% (.v = 
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 ofstemum 5 slightly 
depressed behind transverse impression. 
Legs: protibia with slight ventral marginal 
prominence at proximal 'A; mctatibia nar- 
row, ventral margin slightly, broadly con- 
vex in middle -A; metatibial mucro short, 
straight, acute, not excavated. 

Adult female. Length: 3.40-3.72 mm (.v 
= 3.52, n = 10). Width: 1.56-1.66 mm (.v 
= 1.61, n = 10). Rostrum: length 1.25-1.47 
X (.Y = 1.36, n = 10) pronotal length; distal 
portion subcylindrical, shallowly punctate. 



glabrous, length 22-29% (x = 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- 

Distribution (Fig. 42). The 336 adult 
specimens of .-J. signatipennis Qxaraxn^d are 
from the following localities. ARGEN- 
TINA. Neuquen: San Martin de los Andes. 
Rio Negro: Norquinco. Santa Cruz: Caiia- 
don Leon; Lago Argentina. CHILE. Aisen: 
Chile Chico; Coyhaique; 10 km. N Puerto 
Ibafiez. Bio-Bio: Los Barros, Laguna Laja; 
Volcan Copahue. Caiitin: Cherquenco. 
Chiloe: Castro; Chepu; Dalcahue. Concep- 
cion: Salto de Laja. Llanquihiie: Lepihue, W 
of Puerto Montt; Puerto Varas; Los Muer- 
mos; MauUin; 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. Niible: Las Cabras, Cordillera 
Chilian; Shangrila, 75 km. E Chilian. Osor- 
no: Laguna la Copa, Parque Nacional de 
Puyehue. Santiago: Melocoton; 4 km. W 
Portillo; Quinta San Ramon. Taica: 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, Chiloe, Quinta Pittet and Puerto 
Williams, Magallanes; on B. chilensis ai Sal- 
to de Laja, Concepcion and Algarrobo, Val- 
paraiso; on B. darwinii at Laguna la Copa, 
Osomo, and on B. ilicifolia at Chile Chico, 
Aisen, Chile. 

Anthonomus ornatus Blanchard 

Figs. 3, 4, 14, 17, 21-24, 38-41, 43 

Anthonomus ornatus QXanchard 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 
1883] [anthonomus/ biplagiatus/ Fairm] 
(MNHN). Kuschel 1950: 17. Wibmer and 
O'Brien 1986: 204. 

Anthonomus redtenbacheri Blackwelder 
1947: 839. Replacement name iov A. bi- 
plagiatus Fairmaire (1884) not Redten- 
bacher (1867). 

Recognition (Figs. 3, 4). Adults of .4. 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. 2 1-24) symmetrical, with 
rounded apex and acute lateral promi- 
nences; pygidium of female (Fig. 14) with 
weakly developed subapical promi- 

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 .-I. 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 



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. 2 1 ), 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 -A in dorsal view (Fig. 
23) and the lateral plates are correspond- 
ingly distended dorsally in lateral view (Fig. 
24). This variation is 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 (.v = 
2.90, n = 10). Width: 1.16-1.60 mm (x = 
1.39, n= \0). Rostrum: \mgx\\ 1.40-1.70 x 
(.V = 1.60, n = 10) pronotal length; distal 
portion finely punctate, length 23-32% (.y = 
28. n = 10) of total rostral length. Prothorax: 
pronotum without distinct subapical fascia 
and dorsolateral vittae of pallid scales. Ely- 
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- 

ia narrow, straight, with slight ventral mar- 
ginal prominence; metatibial mucro large, 
curved, excavated. 

Adult female. Length: 2.48-3.32 mm (.v 
= 2.98, n = 10). Width: 1.12-1.58 mm (.v 
= 1.41, n = 10). Rostrum: length 1.59-1.76 
X (.Y = 1.68, n = 10) pronotal length; distal 
portion slender, shining, glabrous, length 29- 
37% (.Y = 35, n = 10) of total rostral length. 
Legs: metatibial mucro minute. 

Larva (2 specimens from Frutillar, Llan- 
quihue. Chile, collected November 2, 1 983, 
by G. Kuschel, ex flower buds Berberis hitxi- 
folia). The third instar larva oi A. ornatus 
resembles that of A. kuscheh 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. kuscheh. 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 ffower buds of Berberis 
buxifoha). 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 Vi 
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 



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. Mesorwtum: 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. Laterostemal and sub-laterostemal 
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- 

Distribution (Fig. 43). The 235 adult 
specimens of .•^. ornatus examined are from 
the following localities: ARGENTINA. 
Neuqiien: 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. Aisen: Chile Chico; Coy- 
haique; Lago Escondido; Lago Frio; 10 km. 
N Puerto Ibafiez; Quellon. Bio-Bio: Los 
Barros, Laguna Laja. Chiloe: Chepu. Con- 
cepcion: Trapatrapa. Linares: Parral. Llan- 

quihiie: Frutillar; Rio Peiion, 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. Niible: Cordillera 
Chilian, Las Cabras, Las Trancas, 70 km. E 
Chilian; 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- 

Plant associations. Specimens of A. or- 
natus were collected on Berberis biixifolia 
at Chepu, Chiloe, Frutillar, Llanquihue, and 
Quinta Pittet and Puerto Williams, Maga- 
llanes; on Berberis ilicifolia at Chile Chico, 
Aisen, 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]; I 
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: 



86007 Farellones/ P. Santiago/ XII-25- 
1968] [under dung L &/ C. W. O'Brien]; 1 
female [Chile Santiago/ Peiialolen/ 25-1- 
1975/ Coll: M. Beeche]; 1 female [Altos de 
Vilches/ Talca A280 mts./ 25-26-XI-1970/ 
J. Solvericens]. Total paratypes, 7 (CWOB, 

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 '/s, with rounded apex and 
short, acute apicolateral prominences; 
pygidium of female (cf Fig. 14) with 
weakly developed subapical promi- 

They are distinguished from adults of the 
closely allied A. ornatus by characters of the 
male genitalia and of sternum 5 of the fe- 
male. The aedeagus of .4. blanchardi (Figs. 
25, 26) is constricted and narrowed in the 
distal -h 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 (.v = 
2.58, n = 2). Width: 1.30-1.34 mm (x = 
1.32, n = 2). Rostrum: length 1.43-1.55 x 
(.V = 1.49, n = 2) pronotal length; distal 
portion finely punctate, length 32-35% (.v = 
34, n = 2) of total rostral length. Prothorax: 
pronotum without distinct subapical fascia 
and dorsolateral vittae of pallid scales. Ely- 

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 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 (.v 
= 2.81, n = 6). Width: 1.10-1.50 mm (.\- = 
1.34, n = 6). Rostrum: length 1.54-1.69 x 
(.V = 1.62, n = 6) pronotal length; distal 
portion slender, shining, glabrous, length 30- 
40% (.V = 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; Penalolen. Talca: Altos de Vilches. 

Plant associations. The paratype of A. 
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 Spkcies 

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, 1 
female [Argentina/ Rio Negro/ L. Nahuel- 
huapi/ Llao Llao] [24 Nov 1950/ W. Witt- 



mer] [Coll. Kuschel] [Entomology/ Divi- 
sion/ D.S.I.R./ New Zealand]; 1 male 
[Argentina/ R. Negro/ L. Nahuclhuapi] [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/ 22-1-1 971/ 
F. Silva G.] [Ex. Berberis/ buxifolia]; 1 male, 
2 females [CHILE: P./ Aisen, 7 km./ W. 
Coyhaique/ 1-21-1968] [at night L. &/ C. 
W. O'Brien]; 1 female [CHILE: P./ Aisen, 
1 5 km./ W. Coyhaique/ 1-21-1 968] [Collec- 
tors: L &/ C. W. O'Brien]; 1 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, 1 female (CHILE/ Chiloe/ Chepu/ 
22. 10.58] [s/ Berberis/ darwini] [Coll./ Kus- 
chel] [Entomology/ Division/ D.S.I.R./ New 
Zealand]; 1 male. 1 female [CHILE/ Chiloe/ 
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.I.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- 1 967] [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/ omatus/ 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]; 1 male, 1 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]; 1 male, 2 females [CHI- 
LI/ Panquipulli/ Valdivia/ 1 2 Jan 1 944] [G. 
Kuschel/ Collection] [Entomology/ Divi- 
sion / D.S.I.R./ New Zealand]. Total para- 
types, 95; (CACA, CWOB, MNNC, NZAC, 

Recognition (Figs. 5, 6). Adults ofA. kus- 
liu'li 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 A. 
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 



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- 
natiis 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 A. ornatus 
(Fig. 14). 

Adult male. Length: 2.72-3.00 mm (.v = 
2.86, n = 10). Width: 1.22-1.32 mm (.v = 
1.26, n= 10). /?05/n/m.- length 1.31-1.59 x 
(.V = 1.46, n = 10) pronotal length; length 
of distal portion 24-33% (.v = 28, n = 10) 
of total rostral length. Prothorax: pronotum 
without distinct subapical fascia and dor- 
solateral vittae of pallid scales. Elytra: 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 'A; metatibia narrow, 
straight, with slight ventral marginal prom- 
inence at proximal 'A; metatibial mucro 
large, curved, excavated. 

Adult female. Length: 2.60-3.12 mm (.v 
= 2.88. n = 10). Width: 1.12-1.40 mm {x 
= 1.26, n= \0). Rostrum: \englh 1.38-1.61 
X (.V = 1.49, n = 10) pronotal length; distal 
portion subcylindrical, shallowly punctate, 
glabrous, length 29-38% (.v = 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 Herberts 
durwini). Body (Fig. 3 1 ): curved; length 3. 1- 
5.4 mm (n = 8). Asperities rounded to sub- 

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 0.5 1-0.54 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 'A length of 
frons. Epicranial suture (coronal suture) less 
than 'A length of head capsule. Four pairs 
of frontal setae; setae 1 and 3 short, seta 4 
ca. 2 X 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 1 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 1 short, 
about 'A 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 1 slightly longer and some- 
times located closer to anterior margin of 
frons than 2. One pair of clypeal sensilla 
that arc located closer to seta 1 than to 2. 
Three pairs of labral setae; setae 1 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 



of Stout median cpiphanngcal spines pres- 
ent between labral rods (tormac). 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 slightly 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 1 slightly longer than other two; 
seta 2 much shorter than others. Mala (la- 
cinial lobe) bearing 1 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 emarginatc dorsally (Fig. 36). .Ab- 
dominal segments 1-7 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 1. Epipleurum (dorsopleurum) 
bearing one long and one short seta. One 
short pleural (\entropleural) seta and pedal 
setae ca. same length. Two minute pedal 
setae present. Anus subterniinal; surround- 
ed by four lobes, each bearing minute seta. 
Seven pairs of minute setae borne on seg- 

ment 9. Spiracles bicameral; air tubes with 
6 annuli. 

Pupa (5 specimens from Frutillar, Llan- 
quihue, collected October 31, 1983, by G. 
Kuschel, e.\ flower buds of Berheris dar- 
wiiii). The pupa of ^. kiiscfieli 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 A. orna- 

Distribution (Fig. 44). The type series of 
A. A:w.V(:7?c// consists of adult specimens from 
the following localities. .ARGENTINA. Rio 
Scgro: Llao Llao, Lago Nahuel Huapi. 
CHILE. Aisen: 1 km. W Coyhaique, Rio 
Correntoso. Bio-Bio: Pemehue. Caittin: 
Volcan Llaima. C7;/7o(;v Chepu. Llanquihue: 
Frutillar. Malleco: 19 km. E Manzanar, 
Cordillera Nahuelbuta. Pichinahuel, 20 and 
38 km. SE Victoria. I'aldivia: Panquipulli. 

Plant associations. Specimens of .'I. kiis- 
chiii were collected on Berheris darwiru at 
Chepu, Chiloe, and Frutillar. Llanquihue; 
and on B. biLxifoha at Rio Correntoso, Aisen, 
Chile. Larvae and pupae were taken from 
tlovvcr 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, 


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 



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/ 1 500 m] [ 1 6 Jan 1 944/ 
G. Kuschel] [Coll./ Kuschel] [Entomology/ 
Division/ D.S.I. R./ New Zealand]; 1 female 
[Bio-Bio/ La. Laja/ Los Barros/ 1 500 m] [ 1 3 
Jan 1945/ G. Kuschel] [Coll./ Kuschel] 
[Entomology/ Division/ D.S.I.R./ New 
Zealand]; 1 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/ 1 100 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.I.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; 

Recognition. Adults of .^. aiancanus 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. herberidis. 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; .-1. araucanus lacks the apical 
extensions of the sutural elytral interstriae 

and the large prominence on sternum 5 of 
the male. In addition, adults of.-I. araucanus 
have the aedeagus symmetrical rather than 
asymmetrical as in adult A. 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: \cr\gx\\ 1.33-1.64 x 
(.Y = 1.44, n = 10) pronotal length; length 
of distal portion 23-32% (.y = 27, n = 10) 
of total rostral length. Prothorax: pronotum 
without subapical fascia and dorsolateral 
vittae of pallid scales, middorsal vitta in- 
distinct. Elytra: 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. Abdomen: sternum 
5 with slight apicomedian prominence. Legs: 
protibia with slight ventral marginal prom- 
inence at proximal 'A; metatibia narrow, 
ventral margin slightly curved in distal -A, 
with slight prominence at proximal 'A; me- 
tatibial mucro large, curved, excavated. 

Adult female. Length: 2.00-2.44 mm (.y 
= 2.30, n = 10). Width: 1.02-1.22 mm (.y 
= 1.08, n= 10). /?o^/n//H.- length 1.00-1.52 
X (.Y = 1.37. n = 10) pronotal length; distal 
portion subcylindrical, shallowly punctate, 
glabrous, length 25-34% (x = 30, n = 10) 
total rostral length. Abdomen: 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- 

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. 



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.xii./ 1976 H. F. How- 
den] [beating] (HAHC). Paralypes: Chile, 3 
males [CHILE, Malleco Prov./ 20 km E 
Manzanar/ 1 100 m 19-21.xii./ 1976 H. F. 
Howden] [beatmg] (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 %, apical mucro 
enlarged, excavated; aedeagus (Fig. 29) 
asymmetrical, constricted medially, with 
slight subapicolateral prominences. 

Like adults of A. kiischeli. 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 A. 
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 A. 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). 

Adult male. Length: 3.40^.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% (.v = 25, n = 4) of 
total rostral length. Prothorax: pronotum 
without distinct subapical fascia and dor- 
solateral vittae of pallid scales. Elytra: 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 '/i; meta- 
tibia narrow, slightly curved, with slight 
ventral marginal prominence at proximal 
'A; metatibial mucro large, curved, exca- 

Adult female. LInknown. 

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. 11, 12, 30,45 

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). Paratvpes: ARGENTINA, 1 male, 
1 female [ARGENTINA/ El Bolson, Rio/ 



Negro/ IX-20-1963/ A. Kovacs]. Chile, 1 
male [Chili] [Deyr.]; 2 males, 4 females 
[CHILE: 19 km. S. of Ancud,/ Is. Chiloe/ 
Site C2. El. 120 m. 14-11-79/ Valdivian Rain 
Forest/ Ashworth, Hoganson, Gordon] [on 
Berberis darwinii] [Anthonomus/ omatus/ 
Blanchardj; 5 males, 6 females [CHILE/ 
Chiloe/ 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/ 11-5-1968] [Collectors: L &/ C. W. 
O'Brien]; 1 female [Dalcahue/ Chiloe, 
CHILE); 1 female [CHILE/ Dalcahue/ Chi- 
loe/ 10-20 Nov 1957] [G. Kuschel/ Collec- 
tion] [Entomology/ Division/ D.S.I.R./ New 
Zealand]; 1 male [Quell6n-Chi/ loe 1-1955/ 
J. S. Vargas]; 1 female [CHILE: Chiloe/ Dal- 
cahue/ 17-31.1.1962/ Luis Pefia] [H. & A. 
Howden/ Collection]; 1 male. 1 female [Pto 
Montt/ 15.1.71]; 1 female [Chile: 12 km. 
N./ Puerto Montt/ P. Llanquihue/ II-8- 
1 968] [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]; 1 male, 3 
females [CHILE, 2 km./ S. Frutillar/ P. 
Llanquihue/ 280' II-3-1968] [Collectors: L 
&/C. W.O'Bnen]; 1 male, 1 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.I.R./ New Zealand]; 1 male [CHILE/ 
Frutillar/ 28 Dec 1944/ G. Kuschel] [G. 
Kuschel/ Collection] [Entomology/ Divi- 
sion/ D.S.I.R./ New Zealand]; 1 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/ 1 7,-Dic. 1972/ Coll: L. E. 

Pena]; 1 male [CHILE/ Volcan Osomo/ 
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, Osomo 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, Osomo Pro./ Site 
17, El. 520 m. 21-XII-77/ Valdivian Rain 
Forest/ Ashworth, Hoganson, Mooers] [on 
Berberis darwinii] [Anthonomus/ omatus/ 
Blanchard]; 3 males, 8 females [CHILE: La- 
guna Espejo/ P. N. de Puyehue, Osomo Pro./ 
Site 1 7A. El. 520 m. 1 0-1-79/ Valdivian Rain 
Forest/ A. C. Ashworth, J. W. Hoganson] 
[on Berberis sp.j [Anthonomus/ omatus/ 
Blanchard]; Total paratypes, 87; (AMNH, 

Recognition (Figs. 11, 12). Adults of .-I. 
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 -A, 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 A. berberidis where the 



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 .1. chilicola and .-1. hcrhcridis. 
but is weakly developed in adult A. hcrhcr- 
idis. which have a prominent midbasal 
macula (cf Figs. 9, 12). The large apico- 
mcdian prominence on sternum 5 of the 
male is unique to adults of A. berberidis. 

Adult male. Length: 2.52-3.04 mm (.v = 
2.83, n = 10). Width: 1.12-1.44 mm (.v = 
1.28, n= \G). Rostrum: \er\g\\\ 1.02-1.48 x 
(.? = 1.32. n = 10) pronotal length; length 
of distal portion 22-31% (.v = 25, n = 10) 
of total rostral length. Prothorax: pronotum 
without subapical fascia and dorsolateral 
vittae of pallid scales; middorsal vitta in- 
distinct. Elytra: 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 'A; metatibia nar- 
row, ventral margin strongly curved in dis- 
tal -A, with distinct prominence at proximal 
'/t; metatibial mucro large, curved, exca- 

Adult female. Length: 2.16-3.16 mm (v 
= 2.87, n = 10). Width: 0.92-1.36 mm (.v 
= 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% (.v = 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 
.1. hcrhcridis consists of adult specimens 
from the following localities. ARGEN- 
TINA. Rio Negro: El Bolson. CHILE. Chi- 
loe: 19 km. S Ancud; 13 km. S Castro; Che- 
pu; Dalcahuc; Puerto Montt; 12 km. N 
Puerto Montt; Quellon. Llanquihue: Frutil- 
lar; 2 km. S Frutillar; Maullin. Osorno: La- 
guna la Copa and Laguna Espejo. Parque 
Nacional de Puvehue; Volcan Osorno. 

Plant associations. Specimens of .4. ber- 
beridis were collected on Berberis darwini 
19 km. S Ancud and at Chepu, Chiloe, 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- 

Etymology. The name of this species is 
derived from that of the plant genus with 
which it and several other A. ornatus group 
members are associated. 


The classification of the subfamily An- 
thonommae has been called "chaotic" 
(Burke 1 976). Some 500, mostly New World, 
named species are assigned to the subfam- 
ily, but recent rcvisionary 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 A. ornatus 
group, it was necessary to attempt to iden- 
tity characters unique to the group by ex- 
amining a broad, though incomplete, array 
of anthonomine taxa. Further studies may 
re\eal that some of these characters are more 
widely distributed. 

The most detailed comparisons included 
those of characters of the A. 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. 
gidaris. unipustulatus. and vcnustus groups, 
the species of the Anthononnis subgenera 
Anthotnorphus, Anthonomocyllus and An- 











bla nchard i 

k uscheli 

ara ucanu s 




Fig. 46. Phylogenetic tree depicting hypothesized relationships of the species of the A. ornatus group 

thonGworphiis, the genera Atractomcnis, 
Loncophorus. Omogoniis and Pseudantho- 
noiniis. 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 A. ornatus 
group, nor are there indications that any of 
them is the sister group of the A. ornatus 

As a result of the comparisons made, 13 
characters (listed below) were determined 
to be unique to adults of the A. ornatus 
group. The distribution of these characters 
among the seven species in the group is de- 
picted in Table 1. Presence is indicated by 
a score of "1," absence by a score of "0," 
"missing," because the female of A. chili- 

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. 
























































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. ornatm 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- 

2) Elytra with large, posterolateral mac- 
ulae (Figs. 3-12). These are weakly de- 
veloped in A. araitcaniis 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 A. 
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 oi A. ornalits and A. blan- 
chardi and in that oiA. chilicola and A. 

8) Parameres of tegmen of male genitalia 
close together or fused at base. 

9) Metatibia of male with ventral margin 
strongly concave in distal -A, 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 A. ornalus (Fig. 14) and 
A. blanchardi. 

11) Sternum 5 of female with apicome- 
dian prominence. This is a simple 
prominence in A. 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 A. ornatus (Fig. 17). 

12) Sternum 5 of female incised or emar- 
ginate(Figs. 17, 18). 

1 3) Pygidium of female with apicomedian 
prominence (Fig. 16). 


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. 1 7-88 1 496P and 
Texas Agricultural Experiment Station T.A. 

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checklist of the weevils (Curculionoidea senus lalo) 
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Indies (Coleoptera: Curculionidae). Mem. Amer. 
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Using Parsimony. LIsers Manual, Illinois Natural 
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the study of bark beetles (Coleoptera: Scolytidae). 
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checklist of the weevils (Curculionidae sensu lato) 
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91(1), 1989, pp. 112-123 


Steve L. Heydon 

Department of Entomology, NHB; Mail Stop 165, Smithsonian Institution, Washing- 
ton, D.C. 20560. 

Abstract. — Until the present review, Notoglyptus Masi was a monotypic genus containing 
the single species A', vircsccns 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 Notoglyptus from the New World are described: A', bidentatus, N. 
luteicrus, N. nesiotes and A', 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: Hymenoptera, Pteromalidae, Notoglyptus, Nearctic, Palearctic, Neotropics, 

Masi (1917) described the genus Noto- 
glyptus and included two species. A', vires- 
ccns 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 A', nigcr 
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- 
gasterinc Pteromalidae, 1 came across spec- 
imens belonging to four undescribed species 
of Notoglyptus from the New World— .V. /)/- 
detuatus n. sp. from South America, A', tzcl- 
talesn. sp. from Central America and north- 
ern South America, A', luteicrus n. sp. from 
the continental Nearctic region south to 
Venezuela, and A', nesiotes n. sp. from the 
Caribbean and coastal southeastern U.S. 

Graham ( 1 969) placed Notoglyptus in the 
Sphegigasterini (Miscogasterinae), probably 

because the elongate Tl is found in other 
genera he placed there such as Crypt o- 
pryiuna Forster and Novit-kyaiius Boucek. 
Genera with an elongate Tl occur sporad- 
ically throughout the Pteromalidae, and I 
feel its common occurrence in these three 
genera is due to convergence (Heydon 1 988). 
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 A', bidentatus 
and A', tzeltales. 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. 1 1). However, 
using this character state as direct evidence 
for a close relationship between Halticop- 



tcra and Notoglyptu.s is complicated because 
A', lutcicrus 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 Notoglyptiis. 

The characters defining Nologlyptus are 
reviewed in the Discussion section follow- 
ing the generic description below. Noto- 
glyptu.s may have evolved in South America 
since that is where the most primitive species 
(N. bidciUatiis and A', tzcltalcs) are found. 


This study is based on examination of 67 
specimens from the museums whose acro- 
nyms are given in 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 ( 1 969), 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 
sensillac, the lower ocular line is LOcL, the 
antennal funicular segments are Fl, F2, . . , 
F6, and the gastral terga are Tl, 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 
A', niger Masi (original designation). Ga- 
han and Fagan, 1923: 98. Peck, Boucek, 
and Hoffer, 1 964: 36 (key). Graham, 1 969: 
124 (key), p. 140. Boucek, 1976: 15. 
Dzhanokmcn, 1978: 77 (key), p. 80. Fa- 
rooqi and Subba Rao, 1985: 259, 31 OF, 
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 

A', hidcntatus (Fig. 1); gena! concavities well 
developed, extending '/2 to 'A malar length. 
Antenna inserted near or below LOcL; with 
2 anelli, 6 funicular segments, and 3-seg- 
mcnted 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), A', tzel- 
tales, and N. virescem but not present in N. 
lutcicrus (Fig. 10) and N. ncslolcs (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 A', ncsiolcs (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. tzcltalcs (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. 
hidcntatus; terminal two segments lamel- 
lately expanded in A', lutcicrus (Fig. 1 1). 

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; 



Figs. 1-7. 1-2, ^'olof;ly|>llls huicntalus n. sp., male. 1, Amcrior view of head. 2, Fore wing. 3, Notoglypliis 
hiU'icnis n. sp., female. 3. Antenna. 4-5, Notoglyptus nesiotes n. sp., female. 4, Fore wing. 5. Antenna. 6-7, 
Notoglypliis tzellales n. sp., female. 6, Fore wing. 7, Anterior view of head. 

Tl almost completely covering the entire 
length of gastcr, 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 is easily seen in com- 
parison with other closely related misco- 
gasterine genera such as Ualticoptera 
(compare Figs. 8, 10, 12 with 13). The 
unique fovea in the center of the scutellum 
so prominent in the type species is clearly 
of no more than specific value when looking 
at this genus on a worldwide basis. 

The species ol'Noioglypliis can be divided 
into two distinct species-groups. Notoglyp- 

liis hidenlaliis and N. izellales 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- 
toglypliis luleicnis. N. nesiotes, and N. vi- 
rescens form the second species-group. 
These species are relatively small, have the 



Figs. 8-13. S-9. Nolojflyplus hidenlalus n. sp., male. 8, Dorsolateral view of mesosoma. 9, Frenuni, dorscllum, 
and propodcum. 10-1 1, Notoglyplus luleicrus n. sp., male. 10, Dorsolateral view of mesosoma. 1 1, Maxillary 
palps. 12, ,\'i>ioglyplus iiesiows n. sp., female. 12, Dorsolateral view of mesosoma. 1 3, Ilalncopiera sp., female. 
1 3, Dorsolateral view of mesosoma. Scale bar = 0. 1 mm. 

body color vei7 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- 



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- 
iuni. 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 yv. bidentatus and N. 
tzeltales; yet, in the characters separating 
the species-groups, A^. nigrescens belongs 
with A', luteicrus and N. nesiotes. 

Key TO World Species of 

Notoglyptus Masi 

1. Pronotum lacking horizontal collar, sloping 
downward immediately from antenor margin 

of mesoscutum (Figs. 10, 12) 2. 

- Pronotum with short horizontal collar sepa- 
rated from dccli\itous neck by a weak to 
strongly developed transverse canna (Fig. 8) 3. 

2. Scutellum with a pair of diverging foveae at 
base (Fig. 12). Dorsellum canniform. Female 
antenna with funicular segments 5-6 quadrate 
to elongate (Fig. 5). (West Indies and coastal 
southeastern LInited States) . nesiotes Heydon 

- Scutellum with sculoscutellar 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 setatc (Fig. 2). Body length 2 mm or 
more. (South Amenca) Indenlalus 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 

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 2 x 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 (46:2 1 ); clypeus 
separated from face by deep furrowlike ca- 
rina, anterior margin produced and biden- 
tate; genal concavity shallow, extending '/a 
malar distance; frons regularly concave be- 
tween eyes; eye height l.4x length (16.5: 
12.0), 1.1 X malar length (16.5:15.0), eye 
length 1.5 X 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 x 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 2 x as large as first; club without 
micropilosity. Mesosoma (Fig. 8) length 
1.6 X width (59:36); collar with weak an- 
terior transverse carina; scutellum regularly 
rounded, scutoscutellar sulcus continuous 



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 x 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: 

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 

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 X 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 

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 A', izeltales 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 A^. hidentatiis 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 1 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 ■ 1 983. 1 <5, Panama (4200 
m, Quito-Baeza Road), 14-17 II 1982, 3 <5. 

Notoglyptus lutekrus. 
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 'i; pedicel brown: flagellum dark 
brown. Mandibles yellow, teeth reddish yel- 
low. Maxillary palps yellow. Legs yellow, 
pretarsi dark brown. Wing veins pale brown. 



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 x height 
(27:2 1 ), 2.2 x length (27: 1 2); 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 X length (13.0:5.5), 2.2 x malar 
length ( 1 3:6), 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 flagcllum 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 
1.4x width (28:20); collar undeveloped; 
scutellum regularly rounded, scutoscutellar 
sulcus continuous furrow; dorsellum length 
'/: frenal length; spiracles round, 1 x own 
diameter from anterior margin of propo- 
deum; nucha acarinate anteriorly. Fore wing 
length 3.1 X 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 1 seta on left 
wing; speculum open posteriorly; 1 row of 
distinct admarginal setae present. Petiole 
length 1.1 X width (4.5:4.0); median carina 
present. Gaster ovate, length 1.4 x width 
(29:21); height 0.86x 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, Fl-6, club as 14:3:1: 

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. 1 1 ). 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). Notoglyplus luteicrus has 
spots of metallic coloration located in sim- 
ilar places to those in A^. nesiotes but the 
boundaries of the spots on the head are dif- 
fuse in A^. luteicrus and sharp in TV. nesiotes. 

Etymology. The specific name comes from 
the Latin words luteus. 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- 



ed as follows (AMNH, CNC. INHS, SEC, 
USNM): Canada. ALBERTA: Lethbridge, 
5- VII- 1956, (swept from brome) 2 2; Lost 
River, Onefour ( 1 mi. WNW Wild Horse), 
3 VI • 1 956, 1 9. ONTARIO: 1 3 mi. N Belle- 
ville, 27-V- 1970, 1 9. United States. CAL- 
IFORNIA: Rancho Santa Fe, 14 I- 1959, 
(alfalfa field) 1 9. COLORADO: Fort Col- 
lins, V- 1894, (alfalfa) 1 <5. FLORIDA: Col- 
lier Seminole State Park (Collier Co.), 25- 
26V- 1978, 1 9; Fort Ogden, 8-IV- 1952, 1 
9. IDAHO: Hansen, 29- V- 1930, {Sisymbri- 
um altissimiim and Descuraima sophia) 1 
9. INDIANA: Hovey Lake (Posey Co.), 3 
VI • 1 98 1 , 1 3. ILLINOIS: Dixon Springs Ag- 
ricultural Research Station (Pope Co.), 29- 
31 VII 1980, 1 S. LOUISIANA: Cameron 
Parish, 4 VIII 1969, (soybeans) 1 9. 
MARYLAND: Patuxent Research Center 
(Pnnce Georges Co.), I VIII - 1 982, 1 9. NEW 
JERSEY: Ramsey, 3 1 VII 1 9 1 8, 1 9. NEW 
MEXICO: Elmendorf, 21 -VII- 1936, 1 9. 
UTAH: Richfield, 18-V 1954, I 9; Utah 
Lake (Utah Co.), 1 9. Mexico. VERACRUZ: 
Jalapa, III-IV 1965, 1 9. Venezuela. POR- 
TUGUESA: 1 km N Biscucuy, 9 - VI - 1 98 1 
(sweeping Baccharis), I S. Country? San Ra- 
fael Jicoltepec, 1 9, 3 3. 

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 

Sculpture: Body sculpture weak, face co- 
riaceous, mesoscutum (Fig. 12) weakly al- 

veolate mesally, petiole granulate, body 
nearly smooth otherwise. 

Structure: Body length 1.4 mm. Head 
ovate in anterior view, width 1.4 x 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 '/3 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 1 x own 
outside diameter above LOcL; combined 
length of flagellum and pedicel 1.0 x head 
width (23:22); relative lengths of scape, ped- 
icel, anelh, 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 x -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 x width (3.5: 
4.0); median carina present. Gaster ovate, 
length 1.4 X width (25:18); height l.Ox 
width (18:18); T2-7 protruding from be- 
neath Tl. 

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 



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. luteicrus. It also resembles N. 
hitcicrus 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 A^. lu- 
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 A', 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 IX 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. 

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 till 
pretarsus black. Wing veins reddish brown. 

Sculpture: Clypeus granulate; head, 
mesosoma, scutellum regularly alveolate; 
frenum, dorsellum, propodeum smooth; 
petiole finely strigulate dorsally; gaster 

Structure: Body length 1 .6 mm. Head (Fig. 
7) broadly ovate in anterior view, width 
1.3 X height (32:24), 2.1x length (32:15); 
clypeus separated from face by distinct sul- 
cus, anterior margin produced but truncate; 
antennal scrobes shallow; genal concavity 
extending '/: way to eye; eye height 1.3 x 
length (14:11). 1.8 x malar length (14:8), 
length 3.7 X 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 X 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 
1.4x width (34:24); collar developed, an- 
terior edge rounded; scutellum regularly 
convex, scutoscutellar sulcus continuous 
furrow: dorsellum length about Vi frenal 
length; spiracles ovate, < 1 x own diameter 
from anterior margin of propodeum; carina 
bordering nucha anteriorly. Fore wing (Fig. 
6) length 2.2 x 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; 



speculum open posteriorly; no distinct ad- 
marginal setae. Petiole length 1.6 x width 
(8:5); lacking median carina. Gaster ovate, 
length 1 .7 X width (30: 1 8); deep, height 1 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, Fl-6, club 
as 1 2:3: 1 :2:2:2:2:2:2:5; relative widths of Fl , 
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 

Variation. The color of jV. 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 A', 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. 
bidentatiis in having metallic legs and the 
anterior margin of the clypeus produced; 
although in A', tzeltales (Fig. 7), the clypeus 
does not have the median emargination 

present in N. bidentatiis (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- 

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 1 male para- 
types were collected as follows (CNC, 
USNM): Costa Rica. Cartago (1500 m), 
VIII- 1980, 1 9; Heredia, 10VIIM975, 2 
9. Mexico. CHIAPAS: San Cristobal de las 
Casas, 27IV1969, 2 9. 12VI1969, 1 9. 
MICHOACAN: Huetamo de Nufiez, 7 III 
1972, 1 9, 1 <?. QUERETARO: 10 mi. E San 
Juan Del Rio, 30 VII 1954, 2 9. Guate- 
mala. GUATEMALA: Guatemala City, IX- 
1959. 1 9. 

Notoglyptus virescens Masi 

Notoglyptiis 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, 9, 
BMNH Hym. 5.873 (examined). 

Notoglyptus niger Masi, 1917: 181. Erdos, 
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 



apically; remainder brown, pedicel pale 
ventraily. 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 

Structure: Body length 1.3 mm. Head 
ovate in anterior view, width 1.3 x height 
(23.5:18.0), 2.0x 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 '/z way to eye; eye height 

1 .4 X length (11:8), 2.8 x rnalar 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 1 x 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, club as 11.0:3.0:1.0:2.0:2.5:2.5:2.5:2.5: 
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 X width (28:19); horizontal collar de- 
veloped, anterior edge carinate; scutellum 
with distinct discal fovea, scutoscutellar sul- 
cus foveolate medially; dorsellum length Vi 
frenal length; propodeum with spiracles cir- 
cular, 1.5 X 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 1 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 X width (21:16); height 1.1 x 
width (17:16). 

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, Fl-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 X 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 is intermediate in morphology with- 
in the genus since it has the developed hor- 
izontal pronotal collar like A', 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. luteicnis and A^. nesiotes. 

Distribution. This species is found from 
southern Europe to India, Japan, and south- 
em Africa (Boucek 1976, 1977). There is a 
male in the Canadian National Collection 
from Mount Pakenham, Ontario. 

Material Exammed (BMNH, CNC). In- 
dia. Delhi, IMRI, 1 IV- 1977, (on grass) 1 
9. Zimbabwe. Salisbury, 1 S. Japan. KYO- 
TO: Kibune, 6- VIII 1980, 1 9. Canada. 
ONTARIO: Mount Pakenham, IIVI- 
1969, 1 6. 


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- 



tomological Collection (SEC), Lawrence, KS; 
and Dr. E. E. GrisscU, United States Na- 
tional Museum (USNM), Washington, D.C. 
1 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, 
.L Sherrod (INHS) for assistance with the 
illustrations, the staff at the Center of Elec- 
tron Microscopy at the University of Illinois 
at LJrbana-Champaign for use of the facil- 
ities, and Molly Scott (INHS) for preparing 
the plates. 

Literature Cited 

Boucck, Z. 1961. Beitriige zur Kcnnlnis der Ptero- 
maliden-fauna von Mittcleuropa, mit Beschrei- 
bungcn ncuer arten und Gattungcn (Hymenop- 
tera). Acta Entomol. Mus. Natn. Pragac 34: 55- 

. 1976. African Pteromalidac (Hymenoptera): 

New taxa, synonymies and combinations. J. Ento- 
mol. Soc. South. Afr. 39: 9-31. 

. 1977. A faunistic review of the Yugoslavian 

Chalcidoidca (Parasitic Hymenoptera). Acta 
Entomol. .lugosl. 13(Suppl.): 1-145. 

Boucck. Z., B. R. Subba Rao. and S. I. Farooqi. 1978. 
A preliminary review of Pteromalidac (Hymenop- 
tera) of India and adjacent countries. Orient. In- 
sects 12: 433^68. 

Dzhanokmen, K. A. 1978. [Identification of the in- 
sects of the European part of the USSR. Vol. 3. 
Hymenoptera. Second part. Pteromalidac.] Opre- 
dehteh Faunc USSR. No. 120: 57-228. 

Erdos, J. 1948. .'\dditamenta ad cognitionem faunae 
C'halcidoidarum (Hymenoptera) in alveo Carpa- 
Ihorum. IV. 11. Pteromalidac. Frag. Faun. Hung. 
II: 36-51. 

Farooqi. S. I. and M. G. R. Menon. 1973. New com- 
bination proposed for a pteromalid species O.vv- 
sychus sauteri (Masi) comb. n. (= Dmannus sau- 
teri Masi). Entomol. Newslet. 3: 55. 

Farooqi, S. I. and B. R. Subba Rao. 1985. Family 
Pteromalidac. pp. 254-263. In Subba Rao, B. R. 
and Hayat. M., eds.. The Chalcidoidca (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 Pteromalidac, pp. 279-306. /« 

Subba Rao, B. R. and Hayat. M., eds.. The Chal- 
cidoidca (Insecta: Hymenoptera) of India and the 
adjacent countries. Part II. A catalogue of Chal- 
cidoidca 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 Chalcidoidca or chalcid-flies. U.S. 
Natn. Mus. Bull. 124: 1-173. 

Graham. M. W. R. de V. 1969. The Pteromalidac of 
northwestern Europe (Hymenoptera. Chalcidoi- 
dca). Bull. Bnt. 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: Pteromalidac). 
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. Boucck, and A. Hoffer. 1964. Keys to 
the Chalcidoidca of Czechoslovakia. Mem. Ento- 
mol. Soc. Canada 34: 1-120. 

91(1), 1989, p. 124 

Book Review 

Bees of the New Genus Ctenoceratina in 
Africa South of the Sahara (Hymenoptcra: 
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 
1 3 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 pauiyi 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- 
abilitv 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- 

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- 
molog}'. Smithsonian Institution. National 
Museum of Natural History- 105. Wash- 
ington. D.C. 20560. 

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 1 5 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 2 1 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 

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 
(i.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 

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. Bcltsville. Maryland 

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 

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 
Httle 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 9-1 1 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 1 3 which deals with di- 
versity, species packing and habitat. The last 
chapter discusses productivity and energy 

'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- 

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 



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. 

whether as a research ecologist, a conser- 
vationist or an applied entomologist. 

Thomas Pape, Zoological Museum, 
Universitetsparken 15, DK-2100 Copenha- 
gen, Denmark. 


91(1). 1989, p. 127 


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 

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. 1 964. 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, Atta texana 
(Buckley), is 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 

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. 

William P. MacKay and S. Bradleigh 
Vinson, Department of Entomology, Texas 
A&M University, College Station, Texas 


91(1), 1989, pp. 128-132 


of the 



Volume 90 



President -Elect 

Recording Secretary 

Corresponding Secretary 




Program Chairman 

Afemhership Chairman 

Delegate to The IVashington Academy of Sciences 

F. Eugene Wood 

Christian Thompson 

Richard G. Robbins 

John M. Kingsolver 

Norman E. Woodley 

Hiram G. Larew 

Anne M. Wieber 

Warren E. Steiner 

Geoffrey B. White 

Manya B. Stoetzel 

Published by the Society 


Table of Contents, Volume 90 


BAKER, G. T.-See CHAN, W. P. 
B.ARNES. J. K. — Notes on the biology and immature stages of Poecilographa decora (Loew) 

(Diptera: Sciomyzidae) 474 



BOLDT, P. E., W. WOODS, andT. O. ROBBINS -Phytophagous insect fauna of Baccharis 

sarothroides Gray (Asteraceae) in .Arizona and New Mexico 207 

BURGER, J. F. — .A new genus and two new species of Pangoniini (Diptera; Tabanidae) of 

zoogeographic interest from Sabah, Malaysia 12 


CARPENTER, J. M. and M. C. D.AY — Nomenclatural notes on Polistinae (Hymenoptera: 

Vespidae) 323 

CARROLL, J, F.— Worker size and piracy in foraging ants 495 

VOLUME 9 1 , NUMBER 1 1 29 

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 (Coleoptera: Curculionidae) 269 

CHEMSAK, J. A. and C. FELLER — New species of Cerambycidae from Twin Cays, Belize 
(Coleoptera) 179 


CLEMENT, S. L. andJ. MIMMOCCHI — Occurrence of selected flower head insects oi Cen- 
laurea solstitialis in Italy and Greece 47 

CLEMENT, S. L., T. MIMMOCCHL R. SOBHIAN, and P. DUNN-Host specificity of adult 
Euslenopus hirtus (Walt!) (Coleoptera: Curculionidae), a potential biological control agent of 
yellow starthistle, Centaurea sohlilialis L. (Asteraceae, Cardueae) 501 

DAVIS, D. R. and E. G. MILSTREY — Description and biology of Acrolophus pholeter. (Lep- 
idoptera: Tineidae), a new moth commensal from gopher tortoise burrows in Florida 164 


DIATLOFF, G. and W. A. PALMER — The host specificity and biology of Ansloielia ivae 
Busch (Gelechiidae) and Loiita hacchanvora Pogue (Tortricidae), two microlepidoptera se- 
lected as biological control agents for Bacchans halimfolta (Asteraceae) in Australia 458 

DREA, J. J. and R. W. CARLSON-Establishmenl of Cybocephalus sp. (Coleoptera: Nitidu- 
lidae) from Korea on Unaspis euonymi (Homoptera: Diaspididae) in the eastern United 
States 307 

DROOZ, A. T. and H. H. NEUNZIG — Notes on the biology of two Phycitines (Lepidoptera: 
Pyralidae) associated with Toumcyella pint (Homoptera: Coccidae) on pine 44 



EMERSON, K. C. and R. D. PRICE — A new species of Haemalomyzus (Mallophaga: Hae- 
matomyzidae) off the bushpig, Polaniochoerus porcus, from Ethiopia, with comments on lice 
found on pigs 338 



GOEDEN. R. D. — Gall formation by the capitulum-infesting fruit fly, Tephnlis stigmalica 
(Diptera: Tephntidae) 37 

GOTT, K. M.-See NEAL. J. W., Jr, 


HALSTEAD, J. K. — Belaspidia longicauda. new species, the first Nearctic Belaspidia (Hy- 
menoptera: Chalcididae) 87 

HANNA, R. B.-See LANE. M. A. 

HANSON. P. E. and J. C. MILLER — Notes on the biology of Caenocephus aldrichi Bradley 
(Hymenoptera: Cephidae) 204 

HANSSON. C — A revision of the genus Meslocharis and a review of the genus Grahamia 
(Hymenoptera: Eulophidae) 28 

HARMON. J. D. and M. H. ROSS— Eflects of malathion and diazinon exposure on female 
German cockroaches (Dictyoptera: Blattellidae) and their oothecae 248 

HEYDON. S. L. — A review of the Nearctic species of Cryploprymna Forster. with the descrip- 
tion of a new genus. Polstonia (Hymenoptera: Pteromalidae) 1 

HEYDON. S. L. and\L. E. GRISSELL— A review of Nearctic A/erw/w!« Walker and Toxeuma 
Walker (Hymenoptera: Chalcidoidea: Pteromalidae) 310 


HUANG. Y. M.—Aedes (Slvgomyia) josiahae. a new species of the simpsoni subgroup (Diptera: 
Culicidae) 155 

JOHNSON. N. F. — Species of Australian Telenominae (Hymenoptera: Scelionidae) of A. P. 

Dodd and A. A. Girault 229 

JOHNSON. N. F. and F. fi\N —Telenomus species (Hymenoptera: Scelionidae) associated with 

the eggs of Zygaenidae (Lepidoptera) 244 





KURCZEWSKl, F. E., E. J. KURCZEWSKI, and M. G. SPOFFORD- Nesting behavior of 
Aportnellus wheeleri Bequaert and A. taeniolatus (Dalla Torre) (Hymenoptera: Pompilidae) 294 


LANE, M. A., F. E. KURCZEWSKI, and R. B. HANNA-Antennal sensilla and setae of 
Evagetes parvus (Hymenoptera: Pompilidae) 428 

MacDONALD, J. F. — New synonyms pertaming to Chelifera and generic key for North Amer- 
ican Hemerodromiinae (Diptera: Empididae) 98 


MARSHALL, S. A. andU. J. S. MONTAGNES-Ccrop/era longicauda. a second North Amer- 
ican species in the kleptoparasitic genus Ceroptera Macquart (Diptera: Sphaeroceridae) . . 189 

MATHIS, W. N. — First record of the genus Procanacc Hendel from North America, with the 
description of a new species (Diptera: Canacidae) 329 

MATH IS, W. N. — First record of the shore-fly genus Placopsidella Kertesz from North America 
(Diptera: Ephydridae) 334 

MATHIS, W. N. and T. ZATWARNICKI- Studies of the systematics of the shore-fly Inbe 
Dagini (Diptera: Ephydridae) 106 





NAKAHARA, S— Generic reassignments of North American species currently assigned to the 
genus Serkotlmps Haliday (Thysanoplera: Thripidae) 480 

NEAL, J. W., Jr. — Unusual oviposition behavior on evergreen azalea by the Andromeda lace 
bug Stephanitis lakeyai (Drake and Maa) (Heteroptera: Tingidae) 52 

NEAL, J. W., Jr. and K. M. GOTT— Evidence for multivoltinism in Prodiplosis platani Gagne 
(Diptera: Cecidomyiidae), a leaf curl midge of American sycamore 201 


NORTON, R. A.. W. C. WELBOURN, and R. D. CAVE- First records of Erythraeidae 
parasitic on onbatid mites (Acari, Prostigmata: Acari, Orbatida) 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 msecl fauna associated with 
Baccharis halimifolia L. in the eastern United States 216 


PETERSON. B. V., M. VARGAS, V., and J. RAMlREZ-PEREZ-.S;/)i»/n«» (Hcmicnetha) 
hieroglyphicum (Diptera: Simuliidae). a new black fly species from Costa Rica 76 


POGUE, M. G. — Revision of the genus Lorita Busck (Lepidoptera: Tortricidae: Cochylini), 

with a description of a new species 440 


QLUCKE, D. L. ]. — Digonogaslra: The correct name for Nearctic Iphiaulax of authors (Hy- 
menoptera: Braconidae) 196 


RAMIREZ-PEREZ. J.. B. V. PETERSON, and M. VARGAS. \ .- Mayacnephia salasi (Dip- 
tera: Simuliidae), a new black fly species from Costa Rica 66 


REEVES, R. M. — Distnbution and habitat comparisons for Carabodes collected from conifer 
branches with descriptions of brevis Banks and higginsi n. sp. (Acari: Orabatida: Cara- 
bodidae) 373 

ROBBINS. R. K. — Comparative morphology of the butterfly foreleg coxa and trochanter (Lep- 
idoptera) and Its systematics implications 133 


ROBBINS. R. K. — Male foretarsal variation in Lycaenidae and Riodinidae, and the systematic 

placement of 5n'-v infernalis (Lepidoptera) 356 


ROBINSON. W. H. and N. BAO-The pest status oi Periplaneta Juligmosa (Serville) (Dic- 

tyoptera: Blattidae) in China 40 1 

ROSS, M. H.-See HARMON, J. D. 

SANTIAGO-BLAY. J. A. andi. MALDONADO-CAPRILES-Observationson the true bugs 

Emcsa tenerrima. a possible spider mimic, and Ghilianella bonncana (Hemiptera: Redu- 

viidae: Emesinae) from Puerto Rico 369 

SITES, R. W.-See ZACK, R. S. 


SPECKER. D. R. and^. T. .lOHNSON-Biology and immature stages of the rhododendron 

gall midge. Clinodiplosis rhododcndn Felt (Diptera: Cecidomyiidae) 343 


STAINES. C. L.. Jr. — A review of the species ofAcritispa LJhmann (Coleoptera: Chrysomelidae: 

Hispinae) 193 

STARK. B. P.. S. W. SZCZYTKO. and B. C. KONDRATIEFF-The Culliis decisus complex 

of eastern North America (Plecoptera: Perlodidae) 91 

STRICKMAN, D. — Redescription of the holotype of Culex (Cule.x) pens Speiser and taxonomy 

ofCulex {Culex) sligmatosoma Dyar and Thriambus Dyar (Diptera: Culicidae) 484 


TRIPLETT. D. C. and A. R. GITTINS — Nestmg. matmg and foraging habits of Melissodes 

(Melissodes) tepida tepida Cresson m Idaho (Hymenoptera: Anthophondae) 462 

TURNER. W. J. — Lectotype designation for /r/);p;ic/»W;/»;efa Wheeler and Melander (Diptera: 

Empididac) 62 


variation and systemalics of four taxa of neotropical walking sticks (Phasmatodea: Phas- 

matidae) 422 





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-.Jp/ew/ia helix (Lepidoptera: Psychidae), a 

Palearctic bagworm moth in North America: New distnbution records, seasonal history, and 

host plants 20 

WHITFIELD. J. B — Taxonomic notes on Rhyssalini and Rhysipolini (Hymenoptera: Bracon- 

idae) with first Nearctic records of three genera 471 

WILKERSON. R. C — Notes and rcdescriptions of some Anopheles series Arribalzagia holo- 

types (Diptera: Culicidae) in the British Museum (Natural History) 411 



ZACK. R. S. and R. W. SITES — A new species of Donaceus Cresson (Diptera: Ephydridae) 

from Malaysia 101 




DAVIDSON. R. L. — Note on the habitat of Plerosiichus (Pseudomaseus) tenuis (Casey) (Co- 
leoptera: Carabidac) with six new slate records 260 


GAGNE, R. J. — A polynomial Riley name in Cecidomyiidae (Diptera) and implications of 

such names for Cynipidae (Hymenoptera) 394 

HALSTEAD, J. A.— A gynandromorph of Hockeria rubra (Ashmead) (Hymenoptera: Chalcid- 

idae) 258 


McCAFFERTY, W. P. — Neotype designation for Raploheptagenia cruenlata (Walsh) (Ephem- 

eroptera: Heptageniidae) 97 

NAKAHARA. S. — A new synonym and revised status in Apterothrips (Thysanoptera: Thrip- 

idae) 508 

PAVUK, D. M. and R. N. WILLIAMS — A sesiid host record for Pteroconnus chasmodops 

(Hymenoptera: Ichneumonidae) 105 

PLAKIDAS, J. D.— The newly discovered spring crown gall of Asphondylia rudbeckiaecon- 

spiciia (Diptera: Cecidomyiidae) on Rudbcckia lacimata (Asteraceae) in Pennsylvania . . 393 
RATHMAN, R. J„ J. F. BRUNNER, and S. J. HDLBERT-Feeding by Medelera species 

(Diptera: Dolichopodidae) on aphids and eriophyid mites on apple. Status domestica (Ro- 

saceae) 510 


VOEGTLIN, D. and M. B. STfOEYZEL-Hyadaphis tatancae (Homoptera: Aphididae): 10 

years after its mlroduction into North America 256 



AMRINE, J. W., ]T.-Bees and Their Keepers 262 

ARNAUD. P. H., h.-The Bombyliidae of Deep Canyon 124 

CARROLL, J. F.-The Behavioral Ecology of Ants 396 

FLINT, O. S., ]t. — Revision of the Caddisfly Genus Psilolreta (Trichoptera: Odontoceridae) . . 400 

FOOTE, B. A. — Manual ofNearctic Diptera, Volume 2 125 

HODGES, R. W.-Sphingtdae Mundi (Hawkmoths of the World) 127 

KIERANS. J. E.-7V;c Ixodid Ticks of Uganda 398 

KJNGSOLVER, J. M.-The Metallic Wood-boring Beetles of Canada and Alaska: Coleoptera: 

Bupreslidae 397 

NORTON, R. A.—Acarology. Mites and Human Welfare 524 

SMILEY, R. L.— Historical Perspective and Current World Status of the Tomato Russet Mite 

(Acari: Eriophyidac) 122 


KROMBEIN, K. V. and?. M. MARSH-Carl Frederick William Muesebeck. 1894-1987 513 







Miscellaneous Publications 

Cynipid Galls of the Eastern United States, by Lewis H. Weld $ 5.00 

Cynipid Galls of the Southwest, by Lewis H. Weld 3.00 

Both papers on cynipid galls 6.00 

Identification of Alaskan Black Fly Larvae, by Kathryn M. Sommerman _ 1.00 

Unusual Scalp Dermatitis in Humans Caused by the Mite Dermatophagoides, by Jay R. 

Traver _ _ 1.00 

A Short History of the Entomological Society of Washington, by Ashley B. Gumey 1.00 

Pictorial Key to Species of the Genus Anastrepha (Diptera: Tephritidae), by George C. 

Steyskal 1.50 

Taxonomic Studies on Fruit Flies of the Genus Urophora (Diptera: Tephritidae), by George C. 

Steyskal 2.00 

Memoirs of the Entomological Society of Washington 

No. 1. The North American Bees of the Genus Osmia, by Grace Sandhouse. 167 pp. 1939 $15.00 

No. 2. A Classification of Larvae and Adults of the Genus Phyllophaga, by Adam G. Boving. (om ^f 
95 pp. 1942 print) 

No. 3. The Nearctic Leafhoppers, a Generic Classification and Check List, by Paul Wilson Oman. 

253 pp. 1949 15.00 

No. 4. A Manual of the Chiggers, by G. W. Wharton and H. S. Fuller. 185 pp. 1952 15.00 

No. 5. A Classification of the Siphonaptera of South America, by Phyllis T. Johnson. 298 pp. 

1957 15.00 

No. 6. The Female Tabanidae of Japan, Korea and Manchuria, by Wallace P. Murdoch and Hirosi 

Takahasi. 230 pp. 1969 15.00 

No. 7. Ant Larvae: Review and Synthesis, by George C. Wheeler and Jeanette Wheeler. 108 pp. 

1976 11.00 

No. 8. The North American Predaceous Midges of the Genus Palpomyia Meigen (Diptera: Cera- 

topogonidae), by W. L. Grogan, Jr. and W. W. Wirth. 125 pp. 1979 12.00 

No. 9. The Flower Ries of the West Indies (Diptera: Syrphidae), by F. Christian Thompson. 200 

pp. 1981 10.00 

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 1 1.00 

No. 1 1. A Systematic Study of the Japanese Chloropidae (Diptera), by Kenkichi Kanmiya. 370 pp. 

1983 18.00 

No. 1 2. The Holarctic Genera of Mymaridae (Hymenoptera: Chalcidoidae), by Michael E. Schauff. 

67 pp. 1984 _ 5.00 

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 18.00 

Back issues of the Proceedings of the Entomological Society of Washington are available at $25.00 per volume 
to non-members and $13.00 per volume to members of the Society. 

Prices quoted are U.S. currency. Postage extra except on prepaid orders. Dealers are allowed a discount of 10 
per cent on all items, including annual subscriptions, that are paid in advance. All orders should be placed with 
the Custodian, Entomological Society of Washington, c/o Department of Entomology, NHB 168, Smithsonian 
Institution. Washington, D.C. 20560. 


(Continued from front cover) 

WEBB, D. W. and M. E. IRWIN-Genera Ataenogera and Phycus in the New World (Diptera: 

Thcrevidae: Phycinae) 35 


MacKAY. W. p. and S. B. VINSON — Rediscovery of the ant Gnamptogenys hartinani (Hy- 

mcnoptcra: Formicidae) in Eastern Texas 127 

WILSON, N. and W. W. BAKER- /.vtx^ra downsi (Acari: Ixodidae) from Peru 54 


NORDEN, B. ^. — Becs of the New Genus Ctenoceratinia in Africa South of the Sahara (Hy- 

menoptera: Apoidea) 1 24 

PAPE, T. — Ecological Methods with Particular Reference to the Study of Insect Populations. 

Second Revised Edition 1 26 

PICKENS, L. G. — Control of Arthropod Pests of Livestock: A Review of Technology 125 


'L. 91 

APRIL 1989 

NO. 2 
(ISSN 0013-8797) 


of the 




APPEL, A. G. and R. C. SPONSLER- Water and temperaturg%latioJ 

lophagous cockroach Cryptocercus punclulalus Scudder (DictyoptNafedj^fiCfi***^"^"- ■ ^■'^ 
ASQUITH, A. and D. ADAMSKI— Description of the predaceous larva of Pseudogaurax sig- 

nutus (Loew) (Diptera: Chloropidae) 185 

DIETRICH. C. H. — Surface sculpturing of the abdominal integument of Membracidae and other 

Auchenorrhyncha (Homoptera) 143 

GAGNE. R. J. and P. E. BOLDT— A new species of Neotasioptera (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 Chrysopcrta comanche (Banks), a green lacewing 

of the mfilabris species group (Neuroptera: Chrysopidae) 133 

KELLEY, R. W. — New species of micro-caddisflies (Trichoptera: Hydroptilidae) from New 

Caledonia, Vanuatu and Fiji 1 90 

KNISLEY, C. B., D. L. REEVES, and G. T. STEPHENS- Behavior and development of the 
wasp Picromhnis rufiventris hyalinaliis Krombein (Hymenoptera: Tiphiidae), a parasite of 
larval tiger beetles (Coleoptera: Cicindelidae) 179 

LaSALLE, J. — Notes on the genus Zagrammosoina (Hymenoptera: Eulophidae) with description 

of a new species 230 

MACKAY. W. p. and S. B. VINSON — Two new ants of the genus Solenopsis (Diplorhoplrum) 

from eastern Texas (Hymenoptera: Formicidae) 175 

MILLER, T. D. — First Ncarctic record of the genus Nordlandena (Hymenoptera: Eucoilidae), 

with descriptions of two new species 158 

NICKLE, D. A. and M. S. COLLINS— Key to the Kalotermitidae of eastern United States with 

a new Neotermes from Florida (Isoptcra) 269 

PURRINGTON, F. F. and D. H. STINNER-Wasp parasites of the burdock seed moth, Metz- 
ncria tappclla Zeller (Cjelechiidac): New host record for Hracon mellitor Say (Hymenoptera: 

Braconidae) 203 

{Continued on hack cover) 




Organized March 12, 1884 


F. Christian Thompson President 
Jeffrey R. Aldrich President-Elect 
Richard G. Robbins, Recording Secretary 
John M. Kjngsolver, Corresponding Secretary 
Norman E. Woodley, Treasurer 

Warren E. Steiner, Jr. Program Chairman 

Geoffrey B. White, Membership Chairman 

Anne M. Wieber, Custodian 

F. Eugene Wood, Past President 

Rebecca F. Surdick 

Hiram G. Larew, Editor 
Robert D. Gordon, Associate Editor 

Publications Committee 

T. J. Henry 

B. V. Peterson 

George C. Steyskal 

Louise M 

Honorary President 
Curtis W. Sabrosky 

Honorary Members 
Alan Stone 

Theodore L. Bissell 

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91(2), 1989, pp. 133-142 




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 Chiysoperla comanche. a common green lacewing 
from western North America, is described and compared to that of its apparent closest 
relative, C. ntfilabris. 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. 
nifilabris. reflect their close relationship and phylogenetic distinctness from the Chryso- 
peiiu canwa group. 

Key Words: Chrysopidae, Chrysoperla, courtship, acoustical, communication 

The green lacewing taxon Chrysoperla 
Steinmann has recently been accorded full 
generic status (Semeria 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 (Scmcria 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 et al. 1981, Adams pers. 
commun.). The better studied of these, the 

caniea group (Tauber and Tauber, 1987), 
includes the common North American 
species C plorabunda (Fitch), C. downesi 
(Banks), and C niohave (Banks). The other 
group, known as the riijilabris group, is 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 nifi- 
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 



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 Chrysopeiia may 
also use vibrational calls in this way. but 
only the common C. rufilabris has been ana- 
lyzed in any detail (Henry 1980a). Here, I 
describe the peculiar trcmulating 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. conianche is 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 coiuan- 
che at two sites in California, 220 km apart, 
during late September, 1987. Twelve were 
obtained in the earh 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) 

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 
1 40 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- 



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 DIE file conversion) to programs 
like Asyst Software Technologies' ASYS- 
TANT+'" for statistical analysis or Jandel's 
SIGMAPLOP- 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- 
mal 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- 
inanche, 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 7 1 male and 89 female vol- 
leys at 8 different temperatures. These fea- 
tures were specifically chosen to facilitate 
comparison with Chrysopcrla rufilabris (Ta- 
ble 3; Figs. 1, 2, 5; Henry 1980a). To com- 

pare the slopes of the temperature regres- 
sions of difllerent 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 diflfered 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. 


Mating in C. comanche. as in other lace- 
wings of the genus Clvysoperla, 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 of a 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 secondar>' 
bursts of vibration. The fundamental or pri- 




0.00 2.66 5.32 7.98 10.64 13.30 


Fig. I. Digitized oscillograph of the shortest re- 
pealed unit (SRU or volley) of the call of Chirsoperia 
comanclie. 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- 

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 

200 300 400 500 


Fig. 2. Digitized oscillographs of half-second frag- 
ments of songs produced at two different temperatures 
by a solo female Chrysoperia cniuaiichc. 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. 

30 n 

= -0.925« + 39 68 
! = 0.11. N = 150 

20 21 22 23 24 25 26 27 28 


Fig. 3. Effect of temperature on volley duration for 
all calls of all males and females of Chrysoperia cn- 
inanchc. 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- 
tion 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 2 1 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 

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. comanchc duets were not 
"polite," in the sense of nonoverlapping. 
In fact, the beats that were produced from 
the acoustical interaction between two su- 



Table 1 . Principal characteristics of the songs of individual males of Chrysopeiia 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 I. 



inal Vibration 1 

FrcgucncN (strokes/ 



Frequency (maxima/second) 



Inilial (.M 

Midi (B) 

Mid2 (C) 

End (D) 

Initial (A) 

Mid (CI 

End (D) 







12. X4 





± 1.35 

± 1.37 

± 2.20 


± 0.43 

± 1.47 

± 5.65 



















± 1.13 




± 0.45 

± 0.00 

± 4.19 


















± 4.16 

± 2.41 

± 3.00 

± 3.11 

± 1.64 

± 1.02 

± 1.44 

± 4.17 


















± 3.13 

± 2.33 

± 1.21 

± 2.69 


± 0.75 

± 1.77 

± 3.58 



















± 0.73 

± 0.47 



± 0.37 

± 1.61 

± 5.43 


















± 5.91 

± 2.19 

± 1.52 

± 5.91 


± 0.94 




















± 0.00 

± 3.40 

± 1.78 

± 0.76 

± 4.24 


± 3.02 

± 10.52 


















± 5.12 

± 1.08 

± 1.24 

± 2.01 

± 2.36 

± 0.68 

± 1.53 

± 3.29 


















± 2.45 

± 2.28 


± 3.20 



± 1.42 





































periniposed 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 1 5 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. rufilahris 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 



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. 


Abdominal Vibration F 

requency (strokes 



Frequency (inaxima/second) 



Initial (A) 

Midi (B) 

Mid: (C) 

End (D) 

Initial lA) 

Mid (C) 

End (D) 










± 1.96 

± 1.13 

± 1.40 


± 0.58 


± 6.15 


















± 2.80 

± 1.56 


± 3.37 

± 2.72 

± 0.33 


± 4.60 


















± 3.46 

± 1.60 

± 1.02 

± 2.96 

± 3.54 

± 1.34 

± 2.10 

± 2.67 


















± 2.94 

± 1.14 

± 0.89 

± 3.33 

± 2.87 


± 1.63 

± 3.85 


















± 4.31 

± 1.69 

± 1.14 

± 3.35 

± 1.28 

± 0.48 

± 1.45 

± 3.27 



















± 1.75 

± 1.60 

± 1.91 




± 6.05 


















± 4.10 

± 2.25 

± 1.33 

+ 2.51 


± 0.32 

± 1.11 

± 2.67 


















± 1.65 

± 2.66 

± 1.36 

± 1.99 

± 1.81 


± 1.16 





































beside each graph. Vollcy'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- 

biinda and Chrysoplella 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 



i 704 


2 21 - 


S 18 


V = 1,13« - 9.43. r2=0.85. N = tS7 

20 21 22 23 24 25 26 27 28 


Fig. 4. EfTecl 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 1 5 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 

0.00 1.00 2,00 3.00 4.00 5.00 6.00 


Fig. 5. Digitized oscillograph of the shortest re- 
peated unit (SRU or volley) of the male "long call"" of 
Chrysoperia 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. 

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). 


The song of Chrysoperia conianche 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 1 5 
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. plorahuucla. Its 
volleys, produced several per SRU, can be 
as long as eight seconds in individuals from 
the Kofa Mountains of Arizona (unpub- 

lished data). The only lacewing species that 
seems to exceed C. comanche in volley 
length is an undescribed relative of C cor- 
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 

A second remarkable feature of the C. 
conianche caW is the modulation of the pri- 
mary abdominal \ibration 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. conianche. For example, a sibling 
species of C downesi from the redwood for- 
ests of California (Founder's Grove) repeats 
its short volleys at nearly 1 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 1 983a, 



Table 3. Pnncipal 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 Mentonc, California (lower section). The corresponding features of the songs of 
Chrysoperla rn/ilcihns 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 

.Abdominal Vibration 

l-rcqiicncy (strokes 


Modulation 1 

Frequency (max 



Initial (A) 

Midi (B) 

Mid2 (C) 


Initial (A) 

Mid (C) 

End (D) 


All Males: 









(N = 9) 

± 5.50 

± 2.94 

± 2.35 

± 5.24 

± 3.23 

± 1.21 

± 2.56 

± 5.76 









All Females: 









(N = 8) 

± 4.57 

± 2.14 

± 1.67 

± 3.68 

± 2.39 

± 0.82 

± 1.76 

± 4.97 









All Isabella: 



86.42 *** 






(N= 13) 

± 4.38 

± 2.38 

± 2.04 

± 4.48 

± 2.50 

± 1.22 

± 2.37 

± 5.03 









All Mentone: 



84.71 *•* 






(N = 4) 

± 6.57 

± 2.78 

1 1.37 

± 3.87 

± 3.17 


± 2.17 

± 6.83 


















(N = 17) 

± 5.17 

± 2.53 

± 2.03 

± 4.49 

± 2.79 

± 1.01 

± 2.14 

± 5.53 










n, a 

n a 









+ 1.11 


± 1.23 

(N = 10, 28 C) 





1985a). In contrast, C. coinanclw's signals 
consist oflong. continuous volleys, with no 
pauses in primary abdoininai \ ibration 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. conianche's closest 
North American relative, C. riifilahris, 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. rufilahris (com- 
pare Figs. 1 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- 



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 Chrysoperia comanchc. Corresponding values for Chrysoperla plorahunda (Henry 1 983b) and for Chrysopwlla 
minora (Henry and Johnson in press) are also tabulated. N/a = not applicable. 


1 Vibration Frequency (slrokcs/sccond) 

Modulation Frequency (maxima/second) 



Initial (A) 

Midi (Bl 

Mid: (C) 


Initial (A) 

Mid (C) 

End (D) 


Chrysoperla comanche 









(N= 17) 

Chn!Soperla plorahunda 









(N = 10) 

Chrvsopiella minora 









(N= 15) 

proximately the same rate: 21 Hz in C. co- 
manche. and 17 Hz in C. rufilahris. 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. riifilabris. First, 
C. rufilahris has sexually dimorphic calls, 
which were not found in C. comanche. Male 
C. rufilahris 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. IF and G in 
Henry 1980a). Second, the song of C rufi- 
lahris is devoid of any primary frequency 
structure analogous to that of C. comanche: 
only lower frequency oscillations, corre- 
sponding to C. comanche'^ modulation fre- 
quency, are apparent (Fig. 5, inset). It is 
possible that C. rufilahris. unlike other lace- 
wings that tremulate, strikes the substrate 
with its abdomen, because considerable 500 
Hz noise is carried on the 1 7 Hz amplitude 
peaks (Fig. 5, inset). Whatever the cause, 
the songs of the two species are distinctive 
and "sound" difl'erent from one another 
through headphones, despite their several 
shared characteristics. 

Whether or not C. comanche and C. rufi- 
lahris can interbreed is unknown. The two 
species overlap geographically, thus provid- 
ing the potential for hybridization: C. rufi- 
lahris 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. comanchc 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 

The phylogenetic position of another 
common member of the rufilahris 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. rufilahris or C. 
comanche, and its song has not been found 
to exhibit any of the features shared by C. 
rufilahris and C. comanchc. 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 rufilahris-comanche lineage. 
The reproductive behavior of the fourth rel- 
ative of these species, C. externa, has not 
been investigated. 



My Studies of C. coinanchc and its rela- 
tives suggest that singing behavior has taken 
a different evolutionary direction in the ru- 
filahris group of Chrysoperla than it has in 
the carnca group. It also provides evidence 
for separating Chrysoperla into two genera, 
to avoid the confusion of species groups. In 
both groups, singing is 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 caniea group, while the three 
members of the rufilahris group have ac- 
quired long, overlapping calls that may in- 
volve drumming on the substrate rather than 


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 Chr>sopidae 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: 

Bram, R. A. and W. E. Bickley. 1963. The green 
lacewings of the genus Chrysopa in Maryland 

(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 hydrolysatcs of yeast and chemically 
defined diet upon the fecundity of Chrysopa car- 
nca Steph. Vest. csl. Spol. Zool. 30: 219-227. 

Henry. C.S. 1979. Acoustical communication during 
courtship and mating in the green lacewing Chry- 
sopa carnca (Neuroptera: Chrysopidae). Ann. 
Entomol. Soc. Am. 72: 68-79. 

. 1980a. Acoustical communication in Chry- 
sopa rufilahris (Neuroptera: Chrysopidae), a green 
lacewing with two distinct calls. Proc. Entomol. 
Soc. Wash. 82: 1-8. 

. 1980b. The courtship call of C/)n'5<)/)a (/oiv- 

nesi Banks (Neuroptera: Chrysopidae): Its evolu- 
tionary significance. Psyche 86: 291-297. 

. 1980c. The importance of low-frequency, 

substrate-bome 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 cornea 
(Neuroptera: Chrysopidae). Syst. Entomol. 8: 293- 

. 1983b. Temperature-induced changes in the 

calls of the green lacewing. Chrysoperla plorabun- 
da (Neuroptera: Chrysopidae). Psyche 90: 343- 

. 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: 


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. 

Semeria, Y. 1977. Discussion de la validite 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 C/(n'- 
sopa carnca (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. 

91(2), 1989, pp. 143-152 


Christopher H. Dietrich 

Department of Entomology, Box 76 1 3, North Carolina State University, Raleigh, North 
Carolina 27695-7613. 

^fo/raf/.— 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; 1) 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: Homoptera, Auchenorrhyncha, morphology, integument, surface sculptur- 
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 intcgumental 
sculpturing, proposing the use of relative 
rather than absolute measures of size of the 

individual elements of sculpturing (e.g. 
punctate vs. puctulate). He further proposed 
the prefixing of terms for sculpturing visible 
only at magnifications greater than lOOx 
with "micro-" (e.g. micropunctate). While 
such terminology is useful at the descriptive 
level, it 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 diflfer- 
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 



Fig. 1. Detail of abdominal integument of a cicadellid, Dracadacephala sp. showing microtrichia (m) and 
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 ( 1 960) and Giin- 
thart (1977) documented the presence of 
minute spherical excretory granules called 
brochosomes covering the bodies of some 
leafhoppers. Presently, such observations 

have not been applied to systematic studies 
and the integument of body parts other than 
those mentioned above has been virtually 

Materials and Methods 

Most of the materials examined were dried 
museum specimens (see Table 1 for a list 
of taxa examined). All are deposited in the 
North Carolina State University Insect Col- 
lection (Department of Entomology, North 
Carolina State LIniversity, 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 505T 
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 



Figs. 2-10. Abdominal scnsilla ofauchenorrhynchous Homoptera. 2-6, Sensilla coeloconica. 2, Enchenopa 
sp. 3, Cymbomorpha amazona. 4, Acidalis tartarea. 5, Oncometopia orbona. 6, Philaenus sp. 7-9, Sensilla 
papulosa. 7. Cymbomorpha amazona. 8. Aconophora cullellata Walker. 9, Hypsoprora coronata. 10, Sensillum 
placodeum, Cemrodontus 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, sec 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 hexanc. 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 aetaiionid, but not on any 
of the other specimens examined— see Ta- 
ble 1.) 

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 fuiiy- 
sclerotized adult, were embedded in Spurr 
medium (Hayat 1986), sectioned with an 
ultramicrotome, slide mounted, stained with 



Table 1. Summary of observations on the abdom- 
inal mtegument of selected auchenorr\nchous Ho- 
moptera. (Arrangement of higher taxa follows that of 
Evans (1977) except where otherwise noted.) 

Table 1. Continued. 

Scnsilla Features* 





Stictopclta sp. 

t, c 



Hyphinoc sp. 





Eurymela sp. 


m, b 

Proterpia sp. 

t, c 




Ledra sp. 




Evacanthus sp. 

t, c 

m, b 

Acutalis tartarea (Say) 

t. c 


Idiocerus sp. 

m, b 


Oncometopia sp. 

t, c 


Spissistdus festinus (Say) 

t, c 


Draeculacephala sp. 

t, c 

m, b 

Cyphonia sp. 

t, c 




Membracidae (sensu 

I 'anduzeea arquaia (Say) 

t. c 


Deitz 1975). 



Atymna querci (Fitch) 

I, c 



Archasia auriadata 

Tylocentrus sp. 




t, c 





Tragopa sp. 

t, c 


Goniolomus sp. 

t, c 




Entylia carinata 



t, c, pa 


Aconophora spp. 

t, c, pa 


Polyrhyssa sp. 

t, c, pa 




Talipes appenduidata 

t, c 



(da Fonseca) 

Microcentnis caryac 



t, c 


Alchisme sp. 

t, c, pa 



Hoplophonnn sp. 

t. c, pa 



Ochropepla sp. 

t, c. pa 


Hetcroiwtm sp. 

I, c 


Platycotis Yittata 

Biturritiidae (= Lampropteridae 


t, c, pa 


sensu Evans 1948) 

Potnia sp. 

t. c, pa 


Tropidaspis sp. 

t, c 


Stalotypa sp. 

t. c. pa 


Bdunitia sp. 

t. c 


Umbonia crassicnrnis 


(Amyot & Serville) 

t. c. pa 


Aetalion reticidatiim 



t. c 

d, b 

Campylenchm talipes 

Darthula hardwickn 


t, c 





Enchenopa sp. 

t, c 





Hypsoprora coronata 

Prosapia biancta (Say) 




t. pa 




Phdaenus sp. 

t, c 


Cenlrodonlus atlas Goding 

t. c, pi 






Tibicen sp. 

I, c, pi 


Cymbomorpha ama-ona 



t, c, pa 




Pintalia sp. 



Prncyrta sp. 

t. c 




Table 1. Continued. 



Libunuella oniala (Sl31) 

Alphma glatica (MetcalO 

Metcalfa pruinosa (Say) 

* KJiY: a = simple acanlhae; b = brochosomes; c = 
sensilla coeloconica; d = toothed acanthae; m = mi- 
crotrichia; pa = sensilla papillosa; pi = sensilla pla- 
codea; t = sensilla trichodea; — = absence of above 

methylene blue, and examined under a 
compound microscope. 


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 Cicadoidca; see 
Table 1). 

Richards and Richards (1979) classified 
the cuticular protuberances of insects into 
four major groups based on their cellular 
structure: 1) multicellular undifl^erentiated 
(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 mcmbracids. They are described 
as follows; A. Sensilla. Four kinds of sen- 
silla-like structures were found and arc 
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 

in pits approximately 1-20 ^^m 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 ^m in diameter (Fig. 10). They were 
found on only two of the specimens ex- 
amined (see Table 1) but eventually may 
be found to be common in some groups (e.g. 

Sensilla papillosa are previously unde- 
scribed structures, presumed to be sensilla, 
consisting of groups of papillae 2-5 jum 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. 1 1) 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 1 corresponds to an individual 
epidermal cell and is therefore derived from 
an acanthus (sensii Richards and Richards 

There is often a wide range of variation 
in the relative amount of development of 



Figs. 11-15. 11-12. Cellular structure of the cuticular protuberances of a membracid, Archasia auriculata. 
1 1, Distribution of microtnchia (m)and sensilla tnchodea (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 Enchcnopa sp. (Membracidae) illustrating variation in the integument 
sculpturing from anterior (left) to posterior (right). 14-15. Microtextures of membracid integument. 14, Micro- 
granulate, Acutalis tarlarca. 1 5, Microrugose, Atyinna guerci. 

acanthae and microtrichia on individual 
specimens. The genera! 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- 

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). 


The observations documented here dem- 
onstrate the need to study the fine-structure 



Figs. 16-21. Comparisons of foveae on the abdominal terga of three membracids (lateral views). 16-17. 
Gnniotoimis tncornigcr. 16. Terga III and IV. 17, Detail, tergum IV. 18-19, Talipes appcnciiculcita. 18, Terga 
Ill-V. 19, Detail, tergum IV. 20-21, Ilypsoprora coronata. 20. Terga III and IV. 21, Detail, tergum III. 

of integumental sculpturing if such features 
are to be used in syslematics. Simply char- 
acterizing integument features as textures 
ma\ be misleading with regards to homol- 

thc individual fine-structural elements. Fur- 
thermore, examination of the integument 
fine-structure yields a wealth of morpho- 
logical information that is potentially ap- 

ogies among such characters at the level of plicable to phylogenetic studies. These 



Figs. 21-21 . Variation of non-sensor> protuberances on abdominal lergum IV in two membracid tribes. 22- 
24, Hoplophorionini. 22. Ochropcpla sp. 23, Potnia sp. 24. Alchisme sp. 25-27, Aconophorini. 25, Aconophora 
cullellala Walker (t, sensillum trichodeum). 26, Aconophora grisescens. 27, Aconophora sp. 

points are best illustrated with examples 
from the membracid taxa surveyed. 

The species Gonioloinus tricorniger StSI, 
Talipes appendiciilata (da Fonseca), and 

Hypsoprom cownata (Fabricius) all have 
large pits on the abdominal terga bearing a 
superficial resemblance to one another (Figs. 
16, 18, and 20). However, the fine-structure 



of these pits differs greatly: G. triconiiger 
(Fig. 17) has simple inornate pits, T. ap- 
pendiculata (Fig. 1 9) has a single coeloconic 
sensillum in each pit, and H. coronata (Fig. 
2 1 ) 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 (sensii 
Deitz 1975) are compared. In the genus 
Ochwpepla (Fig. 22). for example, the pro- 
tuberances are maximally developed as 
small acanthae, each bearing one or two 
acute projections. In the genus P<Hnia (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 Alchlsmc (Fig. 24) are each ap- 
parently homologous to a single acanthus 
of Ochropepla. 

A similar trend occurs among the species 
of the genus Aconophora. The non-sensory 
protuberances of Aconophora cultcllata 
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. 


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, 

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. 
1 1529 of the Journal Series of the North 
Carolina Agricultural Research Service, Ra- 
leigh, NC 27695. 

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membracid pronotum (Homoptera). II. Histology. 
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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: 14.3-148. 

Zacharuk. R. Y. 1985. Antennae and scnsilla. pp. 1- 
69. In Kerkut. G. A. and L. I. Gilbert, cds., Com- 
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91(2), 1989, pp. 153-157 




A. G. Appel and R. C. Sponsler 

Department of Entomology and Alabama Agricultural Experiment Station, Auburn 
University, Alabama 36849-5413. 

Abstract . — VQvctni total body water content, cuticular permeability (CP), rate of water 
loss, and critical thermal maxima and minima were determined for the xylophagous 
cockroach Cryplocercus pimctulatus. There was no difference between female and male 
C. pimctulatus for any of the measured variables. Mean percent total body water was 75% 
and cuticular permeability was 44.1 jugcm - 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: Dictyoptera, Cryplocercus punctulatus, temperature sensitivity, water re- 
lations, desiccation 

The primitive xylophagous cockroach, 
Cryplocercus punclutatus 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 (Comwell 1 968). 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 Blalla and Periplanela. 

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 1 984). The logs containing colonies 

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, 



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 gravimctrically 
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 Mg of water lost 
per unit surface area (cm') per unit time (h) 
per unit saturation deficit (mm Hg) in a 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 = 1 2 • 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 respirator\' mass losses. Hourly 
mass loss and % TBW was used to calculate 
% TBW lost over time. 

Temperature relations experiments.— 
Critical thermal maxima (CTMax) and 

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 l°C/minand 100% 
RH. Adult female and male C. punctulatus 
were used for CTMax and CTMin experi- 
ments. To determine CTMax, a 1 30-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 1 5-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 
mctathoracic coxae. The jar-probe appa- 
ratus was placed on a hot plate that was 
adjusted to increase the temperature inside 
the apparatus at l°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 l°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 



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 

Statistical analysis.— A completely ran- 
domized design and a Student's /-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 /-tests (P < 0.05) were 
used to determine significance. Data are ex- 
pressed as means ± SE. 

Results and Discussion 

Masses of C. punctidaim 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 aduU females (829.76 ± 15.78 mg) and 
males (78 1 .83 ± 1 7.93 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 of a 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. punclulalus 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, Reticiditermes jlaxipes Rol- 
ler (both Rhinotermitidae), ranged from 62.3 

Fig. 1 . Cumulative % TBW lost by female (O) and 
male (•) Cryptocercus punctulams during desiccation 
at 30°C and 0-2% RH. 

to 75.9 (Sponsler and Appel unpublished 

Percent TBW loss of female (F = 32,502. 1 , 
P = 0.0001) and male (F = 2340.1, P = 
0.0001) C. puiiclulatiis 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. punctiilatus. 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, Macketal. 1988). 

Cuticular permeability determines the rate 
of water loss. There was no relationship be- 
tween the CP of female or male C. punc- 
tiilatus and initial mass (F = 0.12; df = 1; 
P = 0.74 and F = 0.00; df = \\ P = 0.99, 
respectively). Similarly, there was no rela- 
tionship between CP and % TBW for males 
(F = 1.92; df = \: P = 0.20). The CP of 
females, however, declined linearly (F = 
6.47; df = 1; Z' = 0.03) with % TBW (CP = 
-0.91 %TBW + 1 1 1.69; r' = 0.45). Appel 
et al. (1986) found that water loss of adult 
male Periplaneta fuliginosa (Serville) was 



Table 1 . Initial masses, percent total body water (% TBW), cuticula permeability, and mortality of female 
and male C. punclulatus [x ± SE; n = 20]. 


Initial Mass (mg) 

% TBW 

C iiticular Permeability* 

Mortality at 24 h 


596.69 ± 78.20 
476.11 ± 67.70 

74.87 ± 1.39 
76.01 ± 0.66 

43.52 ± 1.89 
44.73 ± 3.25 


' /jgcm - 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. punclulatus (Table 
1). The CP of C. punctulatus (44.1 ng 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 ^g cm - h ' mm Hg ', re- 
spectively (Appel el al. 1983). The CPs of 
worker C. fonnosanus and R. flavipes were 
also similar to C. punctulatus (37.5 and 27.8 
Mg cm"' 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 
Mg 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 

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 CTMax 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 often 
cockroach species, representing three fam- 
ilies, could be divided into four statistically 
distinct groups: <48.1°C, 48.7-49. 1°C, 
49.3^9.5°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 

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 1 3.2°C 
and there was no difference between stages 
or species. Although collected at the same 
time of year as these termites, the CTMin 

Table 2. Critical thermal temperatures (°C) of adult C. punclulatus [x ± SE (minimum-maximum); n = 10]. 





39.6 ± 0.28 (39.0-10.5) 
40.9 ± 0.21 (39.7^0.9) 

-8.5 ± 0.52 (-10,0—7.2) 
-7.1 ± 0.55 (-9.0—7,1) 



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- 
catia and B. ohentalis indicated that the 
CTMin of these cockroaches was much 
higher (ca. 1 0°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 > 1 7% 
moisture are at saturated humidity and do 
not contribute to significant body water loss. 
Since all of these groups inhabit moist areas, 
it is 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. 


We thank George W. Folkerts, Depart- 
ment of Zoology and Wildlife Science, Au- 

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. Rusl. 1983. 
Comparative water relations and temperature sen- 
sitivity of cockroaches. Comp. Biochem. Physiol. 
74A: 357-361. 

. 1986. Water relations of the smokybrown 

cockroach, Pcnplaneta 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: 

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: 

Comwell. 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 invcsligata and in Penplaneta 
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 LIniv. Agric. Exp. Sta. Mem. 389: 

Nalepa, C. A. 1984. Colony composition, protozoan 
transfer and some life history characteristics of the 
woodroach Cryptocercus punclulalus Scudder 
(Dictyoptera: Cryplocercidae). Behav. Ecol. So- 
ciobiol. 14: 273-279. 

Rust. M. K., D. A. Reierson. and R. H. Schcffrahn. 
1979, Comparative habits, host utilization and 
xeric adaptations of the southwest drywood ter- 
mites, Incisilermes frulicaviis Rust and Inctsi- 
lenncs /)»/io/(Hagen) (Isoplera: Kaloterniitidae). 
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SAS Institute. 1 982. 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 Cryplocercus punclulalus Scudder. Z. 
Tierpsychol. 61: 315-333. 

91(2), 1989, pp. 158-163 




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: Distribution, southern Africa, western North America, Cynipoidea, 


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- 
er} 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 1 mm total length) and the predom- 
inately reflective integument. Terms for sur- 
face sculpture follow Harris (1979) and 
morphological terms are after Richards 
(1977), Quinlan (1978) and Nordlander 

Type material will be deposited in the 
following institutions: British Museum 

(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 

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 



protruding, forming three spine-like keels 
(Figs. 1, 2, 4). Mandibles bidentate; inner 
tooth two-thirds length of outer tooth (Fig. 
1). Antennae 1 3-segmcnted, 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 1 (Fig. 2); lobe over 
propodeal spiracle wing-like, nearly 2 x 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 

hairs between the carinae, and densely pu- 
bescent laterally (Fig. 5). Legs normal and 
moderately pubescent. 

Gaster with segment 1 obscured by seg- 
ment 2 when viewed laterally; segment 1 
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- 

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, 1 female), 20-VI-83 (1 male), 15-VII- 
83 (1 male), 20-VII-83 (3 females), 23-VII- 
83 (1 male, 1 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. Nordtandcna nierickeli. I , Frontal view of female head (scale line 75 Aim). 2, Lateral view of female 
head and thorax (scale line 80 Mm). 3, Pronotal plate of male (scale line 30 ^lm). 4, Dorsal view of female facial 
projections (scale line 30 Mm). 5, Posterior view of female thorax (scale line 150 mhi). 6, Lateral view of female 
scutellum (scale line 35 Mm), (mb = median bridge; Ic = lateral carinae; mpl = metapleural lobe.) 

Figs. 7-12. Nordlandcria navajoae. 7, Frontal view of male head (scale line 75 Mm). 8, Anterior view of male 
head and thorax (scale line 100 Mm). 9, Dorsal view of female head (scale line 105 Mm). 10, Pronotal plate of 
male (scale line 87 Mm). 1 1. Lateral view of female thorax (scale line 135 Mm). 12, Posterior view of female 
thorax (scale line 150 Mm), (mb = median bridge; rl = metapleural ridge I; st = scutellar tubercle; mnp = 
metanotal plate.) 







and 2 female paratypes to be deposited in 
the California Academy of Sciences; 1 male 
and 1 female paratype to be deposited in 
the British Museum (Natural History); 1 
male and 1 female paratype gold coated on 
scanning electron microscope (S.E.M.) stubs 
and 2 uncoated males and 1 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 1 absent on the metapleuron, and 
the well developed lobe over the propodeal 

Nordlanderia navajoae 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. 1 1); 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- 

Gaster with segment I obscured by seg- 
ment 2 when viewed laterally; segment 1 
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 



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 scuteliar surface 
is less well defined. 

Material examined. — U.S.A. Arizona. 
Apache Co., 1 mi. south of Ganado, T. D. 
Miller and F. W. Merickel collectors; 20- 
VI/ l-VII-85 (2 females. 1 male). 

Types. — Female holotype to be deposited 
in the California Academy of Sciences; 1 
male and 1 female paratype gold coated on 
S.E.M. stubs deposited in the University of 

Etymology. — This species was named for 
the Navajo Indians upon whose tribal lands 
this species was discovered. 

Remarks. — This species is distinguished 
from Dierickcli by the distinct striations on 
the cheeks and mesoplcuron, the setiferous 
tubercles on the scuteliar plate, metapleural 
ridge 1 present, the narrow median bridge 
of the pronotal plate, absence of lateral pro- 
podeal carinae, and the differences in an- 
tennal characters. 


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 Nordlandcria species ex- 
amined so far, including the undescribed 

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. 


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 is 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 

Hams, R. A. 1979. Aglossary of surface sculpturing. 
Occas. Pap. Ent., State Calif. Dep. Food Agnc. 
28. 31 pp. 

Nordlander. G. 1982. Syslematics and phylogeny of 
an interrelated group of genera within the family 
Eucoilidac (Insecta: Hymenoptera, Cynipoidea). 
Dissertation. Univ. of Stockholm. Dept. of Zo- 
ology. Stockholm. 32 pp. 

Qumlan, J. 1978. Hymenoptera: Cynipoidea; Eu- 
coilidae. Handbk. Ident. Br. Insects. Royal Ent. 
Soc. London. 8(lb): 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- 

91(2), 1989, pp. 164-168 


Richard D. Goeden 

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

Abstract.— Ne^^ host-plant records from California are reported for the tephritids, Nea- 
spilota achillcac Johnson, A', aenigiua Freidberg and Mathis, N. albiseta Freidberg and 
Mathis, N. appendiculata Freidberg and Mathis, A^. callistigma Freidberg and Mathis, N. 
stecki Freidberg and Mathis. and A', viridcscens Quisenberry. Neaspilota stecki is initially 
reported from California. The host plants of these tephritids are mainly in the tribe 
Astereae of Asteraceae, although A^. achilleae was reared for the first time from the western 
United States and from capitula ofStephanomeria virgata Bentham in the tribe Cichoreae. 
Besides new records from 1 3 species in six genera of Astereae for A', viridescens, another 
record for this tephritid from Lepidospartum squamatum (Gray) Gray is the first from 
the tribe Senecioneae for any Neaspilota. 

Key W 'ords: Diptera. Tephritidae, host plants, Neaspilota, subgenus Neorellia, Asteraceae 

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 

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 
histor>' 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 Stephanomena virgata 
Bentham collected in southern San Diego 
County as follows: 2 S and 2 9, SE of Barrett 
Junction, 8-X-1987; 7 3 and 6 9, Bratton 



Valley, 20-X-1987; 2 S, Deerhorn Valley, 
20-X-1987. This is a new host-plant genus 
for A', achilleae and the first rearing record 
for this fly from the western United States. 

Freidberg and Mathis (1986) list four 
species o{ Aster, three of Chi ysopsis, three 
of Ehgeron, four of Hieraceum. and one 
species each of Prenanthcs, Sencocarpus. 
and Trilisa as hosts of A', 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 A^. achil- 
leae from one sample of Stephanomeria 
cichoriacea Gray, three samples of Steph- 
anomeria e.xigua Nuttall, nor six samples 
of Stephanomeria paucijlora (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 A^. 
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 A^. 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 A', 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. 

Seaspilota aenigma Freidberg and Math- 
is.— Only five males have previously been 

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 (5 and 6 2, SE end of Kingston (Mountain) 
Range, NE San Bernardmo Co., 27-V- 1 982; 
Lessingia lemmonii Gray, 1 9, Lockwood 
Valley, Los Padres Nat. Forest, Ventura Co., 
at ca. 1 700 m elev., 3 1 -VIII- 1 986; 1 <?, Mo- 
jave River Forks, San Bernardino Nat. For- 
est, SW San Bernardino Co., 1 3-VIII- 1 987; 
Machaeranthera tortifolia (Gray) Cronquist 
and Keck, 2 <?, along Death Valley Road, 
Last Chance Mountain Range, at 1 1 30 m 
elev., Inyo Co., 9-VI-1987. 

Freidberg and Mathis (1986) reported its 
host plants as Haptopappus gooddingii (A. 
Nelson) Munz and Johnston, and possibly 
H. hartwegi (Gray) Blake. I have not yet 
sampled H. gooddingii or reared A', aenigma 
from 1 7 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 (Goedm 1987). Freid- 
berg and Mathis (1986) noted that the ma- 
jority of known hosts of Neaspilota spp. in 
the subgenus Neorellia. with which the pres- 
ent report is concerned, belong to the As- 

Neaspilota albisela 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 A^. 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 A', aenigma. Freid- 



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 a common mode of resource sharing 
in other genera of capitula-feeding Tephriti- 
dae, e.g. Tntpanea 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: Caret hrogyne 
filaginifolia (Hooker and Amott) Nuttall. 1 
9, N of Thomas Mountain, San Bernardino 
Nat. Forest, Riverside Co., 29-IX-1982. 3 
6 and 5 9, Big Sandy Bluff, Sierra Nat. For- 
est, Fresno Co., 235000 23-VIII-1988; Ma- 
caeranthera canescens (Pursh) Gray. 2 S and 
10 9, Onyx Peak, San Gorgonio Mountains, 
San Bernardino Nat. Forest, SE San Ber- 
nardino Co., 22-V- 1 987. Both hosts are As- 

Neaspilota brunneostigma Doane.— 
Neaspilota Ivunneostignia, as revised b\' 
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 Haplopappiis reneliis (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 <? 
and 1 9, Coyote Wells. E of Ocotillo. 
100-m elev., SW Imperial Co., 6-XI-1986; 

3 $. Sentenac Canyon, San Diego Co., 
7-X-1987; 10 <3 and 5 9, Mountain Springs 
Pass, SE San Diego Co., 7-X-1987. I have 
reared other tephritid genera and species, 
but not A^. callistigma from one to five sam- 
ples each of 1 5 other species of Haplopap- 
piis; therefore, this tephritid may be nearly 

Neaspilota stecki Freidberg and Math- 
is.— Heretofore known only from 6 males 
swept in New Mexico (Freidberg and Math- 
is 1 986). One male and 2 females were reared 
by me from capitula of Acamptopappus 
sphaerocephaliis (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 A', 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: A. eatonii (Gray) Howell, 6 S and 
13 9, Perazo Meadow. NW of Truckee, Ta- 
hoe Nat. Forest, 2200-m elev., Plumas Nat. 
Forest, Plumas Co., lO-IX-1986; A. scro- 
piiloriini Gray, 46 $ and 42 9, Cowhom Val- 
ley, Inyo Nat. Forest, NE Inyo Co., 30-VI- 
1982; Chrysothamnus teretifolius Durand 
and Hilgard, 53 S and 53 9, Mountain 
Springs, SW Imperial Co., 6-XI- 1 986; Erig- 
eron argentatus Gray, 3 5 and 1 9, White 
Mountain, 1630-m elev.. Inyo Nat. Forest, 
Inyo Co., 18-VI-1986; E. divergens Torrey 
and Gray, 7 S and 1 1 9, Seven Oaks, along 
Santa Ana River, San Bernardino Nat. For- 
est, SW San Bernardino Co., 5-VII-1983; 
Gutierrezia sarothrae (Pursh) Britton and 
Rusby, I <5, Mountain Springs, SW Imperial 
Co., 6-XI- 1 986: Haplopappiis cooperi (Gray) 
Hall, 284 $ and 280 9, Walker Well, Walker 
Pass, Kern Co., 21-V-1986; H. ericoides 
(Lessing) Hooker and Amott subsp. hlakei 
C. B. Wolf, 1 <? and 3 9, Orcutt, Santa Bar- 
bara Co.. 12-XI-1980; H. laricifolius Gray, 
1 3 (5 and 1 2 9, NW of Kessler Peak at S end 



of Ivanpah Mountains, NE San Bernardino 
Co., 21-X-1982; //. liiiearifolliis de- 
CandoUe, 33 S and 20 9, N Meadow Creek, 
Sequoia Nat. Forest, Tulare Co., 21 -V- 1986; 
H. palmcri Gray, 3 S and 4 9, N of Barrett 
Junction, San Diego Co., 20-X-1987; H. 
propinquus Blake. 27 S and 35 9, along 
Kitchen Creek, Cleveland Nat. Forest, San 
Diego Co., 14-X-1981; Lepidospartmu 
sqiiamatuiu (Gray) Gray, 1 1 S and 10 9, NW 
of Stepladder Mountains, SE San Bernar- 
dino Co., 22-X-1982: Machaeranthera ca- 
nescens (Pursh) Gray, 1 S and 4 9, Keystone 
Canyon, Inyo Nat. Forest, Inyo Co., 1-IX- 
1982; M. gracilis (Nuttall) Gray, 2 <5 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 (5 and 13 9, Obsidian Butte, Imperial 
Co., 7-XII-I983; 4 <3, Calipatria, Imperial 
Co., 28-XI-1984; 17 <5 and 28 9, NW of 
Blythe, Riverside Co., 2-XII-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 {Frcidberg 
and Mathis 1986). I have confirmed Lepi- 
dospartiini squainatum as a host as follows: 
7 S and 7 9, 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 A'. iv>- 
idescens with oligophagous and monopha- 
gous congeners has been shown for several 
generalist Triipanea species, including 
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 

Neaspilota wilsoni Blanc and Foote. — In 
staling that host plants of ,V. wilsoni are un- 

known, Freidberg and Mathis ( 1 986) missed 
my rearing record for this tephritid from 
Haplopappus squarrosus Hooker and Ar- 
nott subsp. griiideloides (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 1 6 other 
species of Haplopappus. many of which 
yielded other Tephritidae. 


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 A^. wilsoni; 
nearly monophagous N. callistignia: oligo- 
phagous N. aenigma and N. appendiculata 
as well as general feeders, A^. 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 A^. aenigma. N. albiseta. 
N. appendiculata, N. callistigma. N. stecki. 
N. viridescens. and A', 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). 




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- 

Literature Cited 

Bush.G. L. 1974. Mechanisms of sympatric host race 
formation of the true fruit flies (Tephritidac). 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 
Tephritidac of California. Bull. Calif Insect Surv. 
7. 115 pp. 

Freidberg, A. and W. N. Mathis. 1986. Studies of 
Terelliinae (Diptera: Tephritidac): A revision of 
the genus Xeaspilola 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: Tephritidac). Proc. Entomol. Soc. Wash. 
85: 399^00. 

. 1985. Host-plant relations of TrMpawea spp. 

(Diptera: Tephritidac) in southern California. Proc. 
Entomol. Soc. Wash. 87: 564-571. 

. 1986. New records of Tephritidac (Diptera) 

from Santa Cruz Island, California. Pan-Pac. 
Entomol. 62: 326-328. 

. 1987. Host-plant relations of native Uro- 

phora spp. (Diptera: Tephritidac) 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: Tephritidac). Calif Dep. Agric. Bur. Ento- 
mol. Occas. Pap. 19. 172 pp. 

91(2), 1989, pp. 169-174 




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; (PER) Grassland, Soil and 
Water Research Laboratory, Agricultural Research Service, USDA, 808 East Blackland 
Road, Temple, Texas 76502. 

Abstract.—^ new species, Neolasioptera rostmta Gagne (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: gall midges, Baccharis. saltwillow 

A new species of gall midge, Neolasiop- 
tera rostrata, is 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 A"'. 
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 
clal. 1986). 

Materials and Methods 

Anatomical terminology in the species 
description follows the Manual of Nearctic 
Diptera (McAlpinc et al. 1981). Some spec- 
imens were permanently mounted in Can- 

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. .W 
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 



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 lOOx 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 5. hal- 
iniifolia 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., 

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 R5 and costa; anterior 
half of tergites 1-7 brown, posterior half 
silvery-white. Antenna with 1 1 flagello- 
meres in <J (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; anepistemum with scales 
on dorsal half to two-thirds; katepistemum 
bare; anepimeron with 9-12 setae and 0-2 
scales. Wing length: (J, 1.2-1.6 mm (n= 10); 

9, 1.4-1.6 mm (n = 10). Length R5 to re- 
mainder of wing: 3, 0.55-0.60; 5. 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. 
Stemites 2-6 with setae inside periphery, 
scales elsewhere; stemites 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 <?, vestiture as in <5; 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. 
Stemites 2-7 similar to 1-6 of 3, stemite 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 = 

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 

Specimens examined. — Holotype: $, 
FLORIDA, Dodge I., Miami, emerged XI- 
1970 from seedheads oi Baccharis glome- 
rulijlora. collected XI- 1970, C. E. Steg- 
maier, Jr. Paratypes: FLORIDA: 1 <?, same 
data as holotype. MARYL.AND: 6 S, 6 9 

Figs. 1-10. Neolasioptera roslrala (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. 







\ -^ 


Fig. 1 1. Bacchans halimifolia. Left, flowering branchlet and leaf, 1 x . Center, flower heads, one in sagittal 
section to show receptacle swelling, 2 x . Right, flower m sagittal section to show swollen receptacle and curled 
larva, 6 x . 

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 <5, 3 9, Ft. Leaton 
St. Pk., Presidio Co., reared from flower re- 
ceptacle galls on Baccharis neglecta. col- 
lected VII- 10- 1986, T. O. Robbins; 10 last- 
instar larvae, Indianola, Calhoun Co., from 
flower receptacle galls on Baccharis hal- 
imifolia. XI-10-1987, P. E. Boldt; 1 5, Lib- 
erty, Liberty Co., Hwy U.S. 90, from flower 
receptacle galls on Baccharis halimifolia. 
X-10-1986, P. E. Boldt; 1 9, Port Lavaca, 
Calhoun Co., from flower receptacle galls 
on Baccharis halimifolia, X-28-1987, P. E. 

Boldt; 1 <?, 1 9, Waller, Waller Co., reared 
X-1 1-1986 from flower receptacle galls on 
Baccharis halimifolia, collected XI- 1 987, P. 
4 9. 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 TV. rostrata are sim- 
ilar to those of Neolasioplera lathami Gagne, 
which causes stem galls on Baccharis spp. 
from New York to Texas (Gagne 1971). The 

Table 1. Mean percentage (±SD) of galls of Neolasiopiera rostrata in male and female flowers oi Baccharis 
halimifolia at three locations in Texas, 1987. 


Pon La 










September 24 
October 14 
October 28 
November 10' 

1.0 ± 2.0 
35.0 ± 11.4 

30.0 ± 8.3 
11.0 ± 6.0 

8.0 ± 3.2 

5.0 ± 3.6 
20.0 ± 7.3 

16.0 ± 8.6 

6.0 ± 7.6 

10.0 ± 9.4 

8.0 ± 11.2 

5.0 ± 5.2 
51.0 ± 8.4 

2.0 ± 2.4 

12.0 ± 8.9 

32.0 ± 10.8 

' No male flowers remaining on plant. 



principal difference between adults of these 
species is the length of the mouthparts. The 
labrum and labella of A', wstrala are very 
long, reaching almost to the distal end of 
the outstretched third palpal segment (Figs. 
4, 5). The labrum and labella of A', 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 A^. rostrata and A', 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 A', rostrata (Fig. 2). 
Further, TV. rostrata has four terminal pa- 
pillae (Fig. 1) instead of the six found in N. 

Biological Notes 

Neolasioptera rostrata produces an en- 
largement of the receptacle of either male 
or female flowers of fl. halimifolia (Fig. 1 1). 
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 gal! 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 (.y ± 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 5. 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- 

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 A', 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 1 5 1 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 of a 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 ( 1 7.7% 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- 


We thank the following individuals for 
their help with this publication: Deborah 
Leather Roney drew Fig. 1 1 and inked Figs. 



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. rangclands; Proposals for biological control 
and ecological studies, pp. 49-68. In 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. 

Gagne, R.J. 1971. Two new species of North Amer- 

ican Neolaswptera from Bacchans (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 
Stales 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. 
11: 20. 

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. 


91(2), 1989. pp. 175-178 


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 Diplorhoptrum as it is presently defined. These 
two species have potential importance as natural enemies of founding queens of the 
imported fire ant, Solenopsis invicta Buren. 

Key Words: Formicidae, red imported fire ant, Solenopsis invicta predation, biocontrol, 

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 

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 

Solenopsis (Diplorhoptrum) subterranea 

MacKay and Vinson, 

New Species 

Figs. 1-4 

Description (worker). — Head length 
(HL— anterior median border of clypeus to 

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 
metastemal 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 = H W x 1 00/ 
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 



Figs. 1^ Holotype of S. suhlerranea: 1 . Full face view showing representative section of punctures; 2. Lateral 
view of mesosomai 3. Lateral view of petiole and postpetiole; 4. Dorsal view of petiole and postpetiole. 

Figs. 5-8. Holotype of 5. puncliceps: 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- 
ameter than the hairs which arise from them 
(Fig. 2); petiole and postpetiole rounded (Fig. 
3), width of postpetiole greater than that of 

petiole (Fig. 4); gaster smooth and shining 
with scattered punctures not much greater 
in diameter than hairs which arise from 



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 (Mexico), the Field Museum 
of Natural History, the Florida State Col- 
lection of Arthropods, Universidade de Sao 
Paulo, Museo Argentino de Ciencias 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- 
em Texas, on N side of Ferrill Creek Rd., 
3.94 K E of tumoff 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 1 7 Aug. 1 987, 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- 

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. 1 6-0. 1 8 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 

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 

Distribution. — Known only from the type 
locality, 10 K N Kurten, Brazos Co. TX 
(same type locality as S. subterranea). 

Biology.— Captured in a subterranean trap 



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 sublerranea is ap- 
parently most closely related to, and could 
be confused with 5. tcunesseensis 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. suhterranea. uneven in length 
with a range of 0.04-0.08 mm in S. ten- 
nesseensis). Solenopsis tennesseensis is 
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. suhterranea. This new 
species differs from S. krockowi Wheeler as 
it is much smaller (S. krockowi: 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 5. krockowi are uneven 
in length and range from 0.03-0.13 mm. It 
differs from 5'. 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 5. 
salina (0.04-0.08 mm) and are greatly un- 
equal in length. 

Solenopsis piincticeps 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. pcrgandei 
Forel which has similar punctures, but the 
postpetiole from above is not round in shape, 
as it is in 5. pcrgandei. It can be distin- 
guished from 5. krockowi by the coarser 

punctures on the dorsum of the head, and 
the diameter of the propodeal spiracle which 
is about twice that of the spiracle of 5. 
krockowi. It is easily separated from S. suh- 
terranea by the hairs on the propodeum 
which are much longer and unequal in 

We suspect these species are important 
natural enemies of the founding queens of 
the imported fire ant, as are others in the 
subgenus Diplorhoptruni (Lammers, 1987). 
The fire ant density at the type locality of 
the two new species is 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 


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 invicla 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. 

91(2), 1989, pp. 179-184 





C. Barry Knisley, Darren L. Reeves, and Gregory T. Stephens 

Department of Biology, Randolph-Macon College, Ashland, Virginia 23005. 

Abstract. — PtcroDibrus rufiventris hyalinatus was found attacking third instar larvae 
of three Cicindcla species in Arizona. Its parasitic behavior is generally similar to that 
described for tropical species of Pterombrus. 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: Cicindela, larvae, parasite, parasitic behavior, Pterombrus, tiger beetle, 
Tiphiidae, 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 Parish 1973). 
Pterombrus is primarily tropical in distri- 
bution and the limited knowledge about it 
comes from studies of P. cicindclicidus and 
P. iheringi in Brazil (Williams 1928) and P. 
piceus in Costa Rica (Palmer 1976). These 
studies included field observations and notes 
of parasitic behavior and development 

Pterombrus rufiventris is the onI> known 

United States species of the genus. Krom- 
bein ( 1 949) 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 



species (Knisley 1987, Knisley and Juliano, 
1988) and factors that influence rates of 
parasitism (Knisley, in prep.). 


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 Pteroin- 
brus were counted to determine relative 
abundance and seasonal activity. Wasp be- 
havior was observed and recorded. 

Laboratory studies.— The behavior of 
Pterombms was studied under laboratory 
conditions of summer photoperiod (13L: 
1 ID) 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 comer. An acrylic tube (2.5 x 
18 cm) filled with soil and containing an 
active third instar of C. pukiira (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 oi Pterombms 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 

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. manuka larvae and 
at a nearby (400 m) saltbush flat where there 
were several hundred C. pulchra. Pterom- 
bms 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 m 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- 
riitha. but in the laboratory some second 
instars of these species were attacked. Palm- 
er (1976) found that P. piceiis parasitizes 
primarily second instars of Pseudoxychila 
tarsaHs Bates. 

Searching behavior. — We observed the 
searching behavior of 1 5 wasps in the field. 
Female wasps walked rapidly over the 


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. Ptewmbrus 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. ihehngi (Wil- 
liams 1 928) 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, Pte- 
wmbrus initiated attacks on larvae in 4 to 
38 minutes (.v = 16.2). In the field, wasps 
were observed to search for 1 5-30 minutes 
without finding a larva. 

Ptcroinbms 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 

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 
1 983). 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 ihenngi 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 



that are necessary for successful oviposi- 

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 Pterombnis and Methocha is 
variable. Any movement bN 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 Pterombnis. 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 2-4 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 

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 1 0-20 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 



Table 1 . Duration and size (length in mm) of developmental stages of P. r. hyalmalus reared m the laboratory 
at 24-28°C. 


Egg length (mm) 

Duration egg stage (days) 

Larval length (mm) on day after hatch: 







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. i: 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 

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 1 3. 5 mm and those collected 
at the sand ridge sites with the smaller C. 
luarutha 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 1 3.2 days (SD 
= 1.6, range 10.2-1 7.3) (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 



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 


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 (DEB81 19256 and BSR8415949). 

Literature Cited 

Adlerz, G. 1906. Melhoca ichneiimonides Lair., dess 
lefnadsaUoch utvecklingsstadier. Arkiv for Zoo- 
logie, Bd. 3, No. 4, p. 1-48, 1 pi. Uppsala and 

Bouwman, B. E. 1909. Uber die Lebenweise von 
Melhoca ichneumonides Latr. Tijdsehr. voor Ent. 
52: 284-294. 

Burdick, D. J. and M. S. Wasbauer. 1959. Biology 
of Methocha caUfornica Westwood (Hymenop- 
tera: Tiphiidac). Wasmann .lour. Biol. 17: 75-88. 

Champion. H. G. and R. J. Champion. 1914. Ob- 
servations on the life-history of Melhoca ichneii- 
monidcsLalT. Ent. Monthly Mag., Ser. 2, 25: 266- 

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 of a new species (Hy- 
menoptera, Thynnidae). Kontyu 10: 67-89. 

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: I 191-2000. 

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 Cicindeta (Coleoptera: 
Cicindelidae). Trans. Amer. Entomol. Soc. 110: 

Krombein, K. v. 1949. A new subspecies of ftfrow- 
hrus rtifiwmns (Cresson). Pan-Pacific Entomol. 25: 

Mury Meyer. E. J. 1983. An analysis of survivorship 
and foraging methods in larvae of three sympatnc 
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 Melhoca ichneumonides Latr. 
Trans. Entomol. Soc. London Ser. 4, 26: 591-597. 

Palmer, M. K. 1976. Notes on the biology of P/erom- 
bnis I'iceus Krombein (Hymenoptera: Tiphiidae). 
Proc. Entomol. Soc. Wash. 78: 369-375. 

Williams, F. X. 1916. Notes on the life-history of 
Melhoca slygia 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. Pleromhrus. a wasp-enemy of the larva 

of tiger beetles. In 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. Fansh. 1973. Predatory be- 
haviour in the ant-like wasp Methocha slygia (Say) 
(Hymenoptera: Tiphiidae). Anim. Behav. 21: 292- 

91(2), 1989, pp. 185-189 


Adam Asquith and David 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: larval morphology, ultrastructure, spider-parasite 

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 maclans (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 Stelcocercllus laliscta (Lamb) (Kirk- 
Spriggs 1986) and Pseudogaurax anchora 
(Loew) (Howard 1916); in the latter how- 
ever, only the anterior spiracles and pupar- 

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 pcn- 
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 bufler. After dehydration in eth- 
anol, specimens were critical point dried. 




Figs. l-3a. Pseudogaurax signalus. larva. 1. Lateral view. Line scale = 0.25 mm. 2, Ventral view. Lme scale 
= 0.25 mm. 3. Spinules on anteroventral portion of Irst abdominal segment (enlargement of area indicated by 
arrow in Fig. 1), Line scale = 10 ^m. 3a, Spinule pattern on 4th abdommal segment (ventral view). Line scale 
= 0.05 mm. 

mounted on stubs with silver paint and 
coated with gold-palladium in a Polaron 
eSlOO sputter coater. The ultrastructure of 
the larva was studied with an Hitachi HH- 

S-2R scanning electron microscope at an 
accelerating voltage of 20 kV. 

For study of the cephalopharyngeal skel- 
eton, larvae were cleared in 20% hot potas- 



Fig. 4. Pseudogaurax signalus. lar\a, cephalopharyngeal skeleton (lateral view). D Com = Dorsal Comu, 
Hyphar = Hypopharyngeal Sclerite, Md = Mandible, Pastm B = Parastomal Bar, V Com = Ventral Comu. 
Line scale = 0.12 mm. 



Figs. 5-1 1. Pseudogaurax signatus. larva. 5, Thorax, and Irsl 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 ^m. Line scale for 
Figs. 8-10 = 10 Mm. A = Antenna, ASp = Anterior Spiracle, FR = Frontal Rami, MxP = Maxillary Palpus, 
MxR = Maxillary Ring, PSp = Posterior Spiracle. 1 1 . Posterior spiracle (posterior view). Line scale = 0.05 mm. 




slum hydroxide, mounted on microscope 
slides in glycerine, and examined with an 
Olympus compound microscope. 


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; maxillar\ 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 

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 
comu of tentoropharyngeal sclerite broader 
and longer than dorsal comu, 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 l);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. 


The larva of Pseudogaurax signatus is 
comparable to that of F. 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 
off. 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 1981a), 



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, Yamadaetal. 1981). 

The spinules found on the body of P. sig- 
iialus 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. 
signalus are similar to those of the necroph- 
agous chloropid Conioscinella hinkleyi 
(Malloch) (Norrbom 1983), and a preda- 
ceous species Steleocercllits 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. 


We thank Curt Sabrosky, Systematic 
Entomology Laboratory, USDA, LI.S. Na- 
tional Museum, for providing us with larvae 
of P. anchom 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. 

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91(2), 1989, pp. 190-202 




Robert W. Kelley 

Biologist, Enwright Laboratories. 25 Woods Lake Rd., Greenville, South Carolina 29607. 

Abstracl.—New species of micro-caddisfly genera Oxyethira Eaton, Paroxyethira Mose- 
ly, Acriloptila Wells and Ca/cdonolrichia 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: caddisfly, new species. New Caledonia, Hydroptilidae 

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: Hydropti/a Dalman, Oxyethira 
Eaton, Orthotrichia Eaton. Orphninotrichia 
Mosely, Maydenoptila Neboiss, Xuthotri- 
chia Mosely, Hellyethira Neboiss, Auslra- 
trichia Wells, and Tricholeiochiton (KJoet 
& 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 Oxyethira, two new species 
of Caledonotrichia and six new species of 
Acritoptila. This brings to 1 7 the number of 
species, distributed among four genera, of 
micro-caddisflies known from the island. 

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 in- 
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 V\ ells, 1982 

Type species: Acritoptila glohosa 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 



Figs. 1-8. Male terminalia ofAcnioplila Wells, lateral and dorsal views. A. ainphapsis: 1, lateral. A. chtasma: 
2, lateral; 3, tcrgum X (dorsal). ,1- cniiihr. 4, lateral. A. disjuncla: 5, lateral; 6, tergum X (dorsal). .1- glo.ssocercus: 
7, lateral. A. planichela: 8, lateral, ap, apico-mesal process; bp, bilobed process; la, inferior appendages; sg, 
subgenital processes; tX, tergum X; vp, ventrolateral process. 

new species share with the Austrahan 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 Spec ies 
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 scleroti/cd 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. 



1 5 disjuncta 


1 9 planichela 

Figs. 9-20. Male terminalia of Acriloplila Wells, aedeagus and ventral views. A. amphapsis: 9, ventral; 10, 
aedcagus. .1. cliiasma: 1 1. ventral: 12. aedeagus. A. crinila: 13, ventral; 14, aedeagus. .-1. disjuncta: 15, ventral 
with inset of caudal view of inferior appendages; 16, aedeagus. A. glossocercus: 17, ventral; 18, aedeagus. .1. 
plaiuchcia: 1 9. \ entral; 20. aedeagus. ap. apico-mesal process; bp, bilobed process; ej. ejaculatory duct; ia, inferior 
appendages; lAe, 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. 

Paraty pes.— 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 

Figs. 2, 3, 11, 12 
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. 

Female. — Unknown. 

Etymology. — Named for the membra- 
nous mesal area dividing tergum X into right 
and left halves. 



Holotype. — Male. New Caledonia: 
mountain stream up Boulari River, light 
trap, 3 Nov. 1958, C. R. Joyce. 

Paratypes. — Same as holotype, 26 S. Pla- 
teau du Dogny, 20 Nov. 1958, 1 <5. 

Diagnosis. — This species is most closely 
related to the New Caledonian A. 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. 

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 
oiA. 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. 1 3), the setose bilobed 
process (Fig. 13) and the non-dilated cau- 
dolateral processes of tergum X (Fig. 4). 

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 latcrad 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 is 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 6. 
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 



sclcroti/.cd and bilobcd; one lateral seta; bi- 
lobed process willi short lobes. Aedeagus: 
single apical sclerotizcd 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. 

Paratypcs. — None. 

Diagnosis. — A sister species of . I. criiiita. 
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 Spkciks 
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 tlared with 
ventral flat, heavily scleroti/ed, claw-shaped 
processes. Inferior appendages: fused into 
broad plate with pair of ventral protuber- 
ances and single spine at each lateral mar- 
gin. Scleroti/ed subgenital structure venlrad 
of aedeagus, with narrow scleroti/ed band 
connecting it to tergum .\. Aedeagus: sim- 
ple, elongate, lacking titillator (Fig. 20). 

Female. — Unknown. 

Etymology. — Named for the Hat claw of 
tergum X. 

Holotype. — Male. New Caledonia: 
mountain stream up Boulari Ri\er, 3 Nov. 
1958, C. R.Joyce. 

Paratypcs. — Same data as holotype, 2 6. 

Diagnosis. — This is the most divergent of 
the New Caledonian species of AcritopriUi. 
The subgenital structures are difficult to 
homologi/e 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 

other New Caledonian Acritoptila are un- 

Genus Caledonotrkhia Sykora, 1967 

Type species: Calcihvioinchia lilicsi Svkora, 

As noted by Wells (1983), males oiCale- 
(lonotrichia closely resemble those of .\fay- 
clcnoptila Neboiss. Indeed, if only genitalic 
characters are analyzed, the two genera could 
be synonymous. .-Xdult males of Calcdono- 
incliia 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- 

Caledonotrichia charadra, 

Nkw Spkciks 

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: bilobcd 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. — Unknow n. 

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. 

Paratypcs.- Same data as holotype, 31 $. 

Diagnosis.— This species is closely relat- 
ed to C. minor Sykora. particularly in the 
shape of the inferior appendages and ae- 



Figs. 21-32. Male terminalia of Caledonnlrichia Sykora and Paroxyethira Mosely. C. charadra: 21, lateral; 
22, ventral; 23, aedeagus. C. extcusa: 24, lateral; 25. ventral; 26, aedeagus. P. diimagnes: 27, lateral; 28, ventral; 
29, aedeagus. P niiinspina: 30, lateral; 31, ventral; 32, aedeagus. ap, apio-mesal process; as, aedeagal sheath; ej, 
ejaculalory duct; ia, inferior appendages; iae, inferior appendage extensor muscle; lal, infenor appendage lower 
lobe; iau. inferior appendage upper lobe; ms, membranous sac; sg, subgenital process; ti, tilillator; vl, ventral 

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 vcntrally directed tooth and 

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, 1 $. 

Diagnosis. — C cxlcnsa is distinctive, with 
the lobes of the inferior appendages longer 
than wide. It is likclv a sister to the distinct 



group composed oflhc other species in the 


Genus Oxyethira Katon, 1873 

Subgenus 7>/(7;<\i,7(7;c (Neboiss, 1977) 

Tvpe species: Oxvethira albiceps (Mac- 
Lachlan, 1862). 

Two species of Oxyethira from New Cal- 
edonia are placed in subgenus Trichoi^lem' 
Neboiss along wiili ilic New Zealand (). al- 
biceps and the Australian brevis Wells, co- 
liimba (Neboiss). micnica Wells, irlaimii- 
lala Wells and rctnuta 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; aedcagus with re- 
curved sub-distal spinous process and 
subgenital processes widely separated and 
partly fused with each pleuron of segment 
IX. Males of <9. calcdonicnsis n. sp. are sim- 
ilar to O. brevis in ha\ ing a short titillator. 
but seem to be more primitive than all Aus- 
tralian and New Zealand species in the re- 
tention of a scleroti/ed 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 Daclylotri- 
cliia Kelley but is probably not homologous. 

The t\pe species of I'richoi^lene was in- 
correctly identified as O. cohiniba (Neboiss) 
in Kelley (1984). It should be O. albiceps 
(MacLachlan). A redescription of the sub- 
genus was gi\en in Kellev (1984). 

Oxyethira caledoniensis. 

New Spkcies 

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 scleroti/alion. Subgenital process- 

es; widely separated, narrow, connected by 
thin mesal scleroti/ed strip; dorsolaterally 
fused to pleuron IX; bilobed processes short. 
Acdeagus; 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, 


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; acutch pointed, 
connected mcsally by curved sclerotized 
band and connected dorsolaterally to pleu- 
ron IX; bilobed processes short. Acdeagus; 
titillator lacking; scleroti/ed, recurved pro- 
cesses at apex. 

Female. — Unknown. 

Ftymology. — Named for its island habi- 

Holotype. — Male. New Caledonia; 
mountain stream up Boulari Ri\er. light 
trap, 3 Nov. 1958, C. R. Joyce. 

Paratypes. — Same data as holotype, I 5. 

Genus Oxyethira F.aton, 1873 
Subgenus Pacificotrichia, n. subgen. 

Seven new species oiOxyethira 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 DaDipfitrichia Mose- 
ly. Similarities between the males include 
the shallowly excised venter VIII (Fig. 62), 
the deeply excised dorsum VIII (Fig. 37), 
fused Rj and Rj forewing veins, configura- 



Figs. 33-41. Male terminalia of Oxyelhira Eaton, lateral views. 33, O. caledoniensis. 34, O. insularts. 35, O. 
dorsennus. 36, O. indorscnnus. 37, O inclasmu. 38, O. oropedion. 39, O. scutica. 40, O. efalensis. 41, O. fijiensis. 
ap, apico-mesal process; bp, bilobed process; la, inferior appendages; sg, subgenital processes; si, 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, arc like 
the genitalia of the minima group of sub- 
genus Dampfitrichia. the only other known 
Australasian group ofOxycthira 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,, 
inclorsennus 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 mcsodistal 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 arc distinguished by 
venter IX which extends into segment VII 
(Fig. 40), subgenital processes which are not 
connected antcromesally (Fig. 63) and an 
aedeagus with a tapered dorsal sclerotized 



42 43 44 45 46 47 48 49 



Figs. 42-55. .\cdeagi and female terminalia of Oxyelhira Eaton. Figs. 42^9. aedeagi. 42. O. caledoniensis. 
43. O. insularis. 44. O. dorsennus. 45. O. iiwlasma. 46. O. oropedion. 47, O. scutica. 48, O etatensis. 49, O 
fijiensis. Figs. 50-53, female terminalia. O. oropedion: 50, lateral; 51. ventral. O. sculica: 52. lateral; 53. ventral. 
O. efaensis: 54. lateral; 55. ventral, ap. apodcme IX; ej, ejaculatorv- duct; hi. horizontal lamella; sp. spcrmathecal 
process; ss, spermathecal sclerite; ti. titillator; tX, tergum X; vl, ventral lobe. 

process and a membranous \ cntra! tube (Fig. 
49). Females bear short apodemes and cerci 
(Fig. 54). 

Subgenus Pacificotrichia, n. subjjen. 
Type species: Oxyethira oropedion, n. sp. 

Spur formula: 0-2-4 or 0-3-4. 

Forewings: Rj and Rs fused. 

Male. — Length 2.1-2.4 mm. Antennae 
24-30 segmented; sparse placoid sensilla. 
Segment VIII: v enter shallow K 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 distalh (Fig. 63): bilobed process short 
(Fig. 62). Aedeagus: titillator absent (Fig. 

Female.— There is too much variability 
among females to describe subgeneric char- 

Range. — New Caledonia; Vanuatu; Fiji. 






Figs. 56-63. Male terminalia of O.xyelluia Ealon, ventral views. 56, O. caledoniensis. 57, O. insulans. 58, 
O Jorscnniis. 59, O melasma. 60, O oropedion. 61, O. scutica. 62, O. efatensis. 63, O. fijiensis. bp, bilobed 
process; ia, inferior appendages; mr, mesal ridge; sg, subgenital processes; si, seial 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- 

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. 

Paraty pes. — Same data as holotype, 1 6. 
New Caledonia, Plum. 20-60 m, 23-25 Mar. 
1958, 1 S. 

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. 



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. 

Paraty pes. — Same data as holotype, 5 S. 

Diagnosis. — This species differs from O. 
(lorsciinus. n. sp. only in the shape of dor- 
sum VIII (Figs. 35, 36) and the spur for- 

Oxyethira melasma. 

New Spkcies 
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: sctal lobes 
lacking. Subgenital processes: with ventral 
black spot at fused apex: elongate bilobed 
process with mesaiiy 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. 

Paraty pes. — Same data as holotype, 1 i. 
New Caledonia, Honailu Ri\ cr headwaters, 
26 Oct. 1958, 1 S. 

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: 
wideh separated; setal lobes forming setose 

membranous bridge between inferior ap- 
pendages. Aedeagus: without apical pro- 

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, I S. 

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 apodcmes 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 $. 
Same locality as holotype, 17 Nov. 1958, 

Diagnosis. — Males of this species may be 
recognized by the minute inferior append- 



ages and triangular (ventral view) subgenital 
processes, in addition to the shape of the 
aedeagal process. Oxycthira sciUica is the 
only species of subgenus Pacificotrichia with 
paired black teeth along the posterior fused 
margin of the subgenital processes. This 
character is common among species of other 
neotropical subgenera of Oxyethira. 

efatensis Group 

Oxyethira efatensis, 
Nkw 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, sclcrotized, attenuate band 
subtended by membranous tube (Fig. 48). 

Female. — Length 2.4 mm. Antennae 21 
segmented. Segment VIII: tergum short with 
anterior margin heavily sclcrotized. Seg- 
ment IX: lacking. Spermathecal process 
teardrop-shaped; horizontal lamella dis- 
tinctly sclcrotized. 

Etymology. — Named for locality of col- 

Holotypc. — Male. Vanuatu: Efate (NW). 
Maat, Ambryn Village, 3M., 18 Aug. 1957, 
light trap, J. L. Cressitt. 

Paraty pes. — Same data as holotypc, 4 S. 
Same locality as holotypc, 19 Aug. 1957, 
5 3. 

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- 

ment LX: 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 
sclcrotized blade-like process and one 
membranous process enclosing ejaculatory 
duct (Fig. 49). 

Female. — Unknown. 

Etymology. — Named for collection local- 
ity of holotypc. 

Holotypc.- 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, 1 924 

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 oi 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 Spec ies 
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: 



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 S. 
Same locality as holotype, 17 Nov. 1958, 
165 <5. 

Diagnosis. — Paroxyclhii a 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 S. 

Same locality as holotype, 17 Nov. 1958, 

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. 


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 

Literature Cited 

Eaton, A. E. 1873. On the Hydroptilidae, a family of 
Trichoptera. Trans. Entomol. Soc. London 1873: 

Kelley, R.W. 1984. Phylogeny. morphology and clas- 
sification of the micro-caddisfly genus Oxyethira 
Eaton (Tnchoptera: 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 Tnchoptera). Trans. R. Soc. N.Z. 55: 670-673. 

Neboiss, A. 1977. .\ 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. 
lilies in New Guinea and New Caledonia. Pac. 
Insects 9: 585-595. 

Wells. A. 1982. Tncholeiochilon Kloet & Hincks and 
new genera in the Australian Hydroptilidae (Tn- 
choptera). Aust. J. Zool. 30: 25i-270. 

1983. New species in the Australian Hydrop- 
tilidae (Trichoptera). with observations on rela- 
tionships and distnbutions. Aust. J. Zool. 3 1 : 629- 

91(2), 1989, pp. 203-205 





Foster Forbes Purrington and Deborah H. Stinner 

Department of Entomology, The Ohio State University, The Ohio Agricuhural Research 
and Development Center, Wooster, Ohio 44691. 

Abstract. — Bracon mellitor parasitizes the burdock seed moth, Metzneria lappella. This 
is a new host record for this wasp, best known as a parasite of the boll weevil, Anthonomus 
grandis. Agathis ntalvacearuDi. 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 Hyssopiis thymus, the paramount Nearctic wasp parasite of the European 
pine shoot moth, Rhyacionia huoliana. 

Kev Words: Braconidae, burdock, distribution, Gelechiidae 

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 univoltinc 
moth is found wherever common burdock, 
A. minus (Hill) Bemhardi, 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 (]uha\a 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 (12-11-86), 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- 

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 et al. 1 969, Cross and Chesnut 1971, 
Krombein et al. 1979, Cuda and Burke 
1 983). It is an important parasite of the boll 
weevil, .Anthonomus grandis Boheman (Co- 
leoptera: Curculionidae) (Adams et al. 1 969, 
Sturm and Sterling 1986, Pcncoe 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 .\f. lappella. In the Pale- 
arctic, -). malvacearum is also found on the 
moths, Af. carlinella Stainton (Gelechiidae), 
Evetria resinella L. (Gelechiidae), and Co- 
leophora otitae Zeller (Coleophoridae) 
(Shcncfelt 1970). North American distri- 
bution records are Quebec, New England, 



Long Island, and west to North Dakota (Ju- 
hala 1967, Krombein et al. 1 979). We found 
A. mahaccantm on M. lappella in burdock 
collected at Portland, OR (Reed College, 25- 
XII-85, col. C. B. Purrington), Bellfountain, 
OR (10-1-86), 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 Fylcs ( 1 899) 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. mahaceanim. 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 Hyssopiis ihymus Girault 
(Hymenoptera: Eulophidae), a multivol- 
tine, gregarious, larval ectoparasite from M. 
lappella in burrs obtained at Mattapoisett, 
MA (12-11-86), Wooster, OH (lO-IV-86), 
and Kent, WA (2-1-86). These new collec- 
tion records establish a coast-to-coast North 
American distribution for the M. lappella- 
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 (Schiffermuller) (Tortrici- 


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, M. 

lappella and A. mahacearum, as well as 
eclectic elements from native fauna. 


We thank P. M. Marsh (USDA-Beltsville) 
for identifying .-I. mahacearum 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 110-88. 

Literature Cited 

Adams, C. H., W. H. Cross, and H. C. Mitchell. 1969. 
Biology of Bmcon metlitor. 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, Anihonomiis grandis: 
1. An annotated list. Ann. Entomol. Soc. Am. 64: 
Cuda, J. P. and H. R. Burke. 1983. Trwhobans brid- 
nelli. a new host for Bracon nwllitoi: Southwest. 
Entomol. 8: 65-66. 
Fyles.T. W. 1899. Noteson 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) Bemh. and .-1. lappa L. Can. J. Plant Sci. 
60: 621-634. 
Gunther. E. 1973. Ethnobotany of western Washing- 
ton. Univ. Washington Press. Seattle. 71 pp. 
.luhala. C. 1967. Notes on parasitic Hymenoptera 
associated with a gelechiid moth, Metzneria lap- 
pella. in the common burdock, and description ol 
a new species of.igalhis (Braconidae). Ann. Ento- 
mol. Soc. Am. 60: 95-97. 
Krombein. K. V.. P. D. Hurd. .Ir.. 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 



parasites in easlern North Dakota. Ann. Entomol. 
Soc. Am. 63; 942-945. 

. 1979. Biology of the hyperparasitic wasp 

Pcnlampus 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. 

Shenefclt, R. D. 1970. Braconidae 3: Agathidinae. 

pp. 342-343. In Ferriere, C. and J. van der Vecht. 

eds., Hymenopterorum Catalogus, W. Junk, Dor- 
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. 

91(2), 1989, pp. 206-229 






Louis M. Roth 

Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts 
02138. (Correspondence: Box 540, Sherborn. Massachusetts 01770.) 

y4fo/racY. — Diagnostic characters for the cockroach genus Margattea Shelford are given. 
Theganosilpha Kumar and Princis is sy nonymized with Margattea. Margattea paracey- 
lanica, n. sp. from the Krakatau Islands is described. The following Margattea species are 
redcscribed: ceyluiuca (Saussure), luitihata (Shelford) (previously considered a synonym 
of ceylanica). perspicillaris (Karny), anceps (Krauss), contingens (Walker) [= huineralis 
(Walker)], and longcalata (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 Balta Tepper: aiirea Hanitsch, setifera (Hanitsch), 
parviila (Bolivar), cra.s.sivenosa (Bolivar), and longicereata (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: Cockroaches, Blattellidae. taxonomy, Krakatau Islands 

Princis (1969: 862, 1971: 1 162) listed 44 
species of .Margattea Shelford. It is 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 re^■ision 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 

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 dilhcult 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 Balta Tepper: 
.Margattea aiirea Hanitsch (HDEO), .\/. se- 
tifera Hanitsch (NRSS), M. parvula Bolivar 
(BMNH), M. crassivenosa Bolivar (BMNH), 
and M. longicereata 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 



cockroaches were taken and I will report on 
these elsewhere. Only one species of Mar- 
gallea 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. ccylanica 
(Saussure), M. contingens (Walker) [= M. 
humeralis (Walker)]. M. aiiccps (Krauss), 
and A/, nimhata (Shelford). Kuc/unga He- 
bard has been considered a junior synonym 
of Margatlca. and I have concluded that 
Theganosilpha Kumar and Princis also is a 
junior synonym and therefore will rede- 
scribe their type species [Kuchinga 
longcalata (Brunner) and Theganosilpha 
perspicillans (Karny)]. 

The following museums and their cura- 
tors or collection managers loaned me spec- 

(ANIC)— Australian National Insect 
Collection, CSIRO, Canberra, 
A.C.T., Australia; Dr. D.C.F. 
(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)— Bemice 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 Naturellc, 

Geneva, Switzerland; Dr. 

Bemd Hauser. 
(NRSS)— Naturhistoriska Riksmuseet, 

Stockholm, Sweden; Per Inge 

(RNHL)— Rijksmuseum van Natuurlijke 

Historic, 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: Blalta ceylanica Saussure, 
Rehn 1931: 302; Bey-Bienko 1938: 121; 
1950: 145; Princis 1969: 862. 

Kuchinga Hebard 1 929:39, 4 1 . Type species, 
by selection: Phyllodroniia longcalata 
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: Theganoptery.x 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). Tcgmina 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 branches in reduced wings) 
and no incomplete branches, apical triangle 
small (Figs. 6, 13). Abdominal tcrga unspe- 



cializcd, or w itli a group of setae near llie 
posteromedial margin of eightli torgum 
(Figs. 24. 28); this speciali/alion may be 
hidden under the se\enth tergum. Antcro- 
ventral margin of front femur Type B; or B, 
(with 1 to 4 pro.\imal spines) (Fig. 34) (both 
types may occur in tiie same species, dif- 
fering between the sexes), rareh T>pe C,; 
all 4 pro.ximal tarsomcres with puh illi, tar- 
sal claws symmetrical with inner margins 
generally denticulate (Figs. 7. 20. 32: the 
serrations often subobsoictc 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 s>mmetrical 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 phallomcrc 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 ']'lici;aii(Ksilphii from Ihcgcin- 
oplery.x were hind wings reduced in both 
sexes, eighth tergum of male weakly spe- 
cialized, antero\entral margin of front fe- 
mur Type B, (some large proximal spines 
followed b\ a row of piliform spinules and 
terminating in 3 distal spuies), inner mar- 
gins of tarsal claws finely denticulate. Asa- 
hina(1979: 11 9) accepted rhci^unosllplnias 
a valid genus and correctly pointed out that 
it was closer to Margcittea than to Thegau- 

Rehn (1931) slated that the front femur 
of M. ceylanica was Type B. and that the 
tarsal claws were equal with internal mar- 
gins unspeciali/ed (i.e. not denticulate). Bey- 
Bienko (1938: 121) believed that the tarsal 
claws of U. ceylanica and \f. longcalaui are 
equal and unspccializcd, but later (Bey- 

Bienko 1958: 587) correctly stated that the 
former has barely visible notched tarsal 
claws. The claws of .\/. longcalata also have 
subobsoictc serrations. .Asahina ( 1 979: 11 9) 
pointed out that since some species of Mar- 
gattca have toothed tarsal claws, this char- 
acter cannot be used to distinguish it from 
Thcganosilpha. 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 A/, ceylan- 
ica and .)/. longealata arc unspccializcd. The 
male of Theganosilpha perspicillaris has a 
setal tuft on the eighth tergum. There are 
man\ examples ofgenera in w hich the males 
of a genus ha\e or lack abdominal tergal 
glands (used in courtship behavior), and in 
those species that ha\ e them the specializa- 
tions may be found on one or more seg- 
ments. For example, species ofBlalte/la may 
have glands on T7 only, or on T7 and T8, 
and the morphology of the glands may differ 
considerabh between ditlcrent species (Roth 
1985a). In species of Parcoblatta Hebard, 
the males have glands on Tl only (6 spp.), 
Tl and T2 (4 spp.), or they are absent (2 
spp.) (Hebard 1917). 

.Asahina (1979: 1 19) separated Thegano- 
silpha from Margattea as follows: In The- 
ganosilpha the front femur is T\pe B,, the 
tegmina are narrowed apicalh, somewhat 
nail shaped, rather coriaceous, and the hind 
wings are rudimentai"\ but sometimes one 
half the length of the tegmina. In Margattea 
the front femur is Type B., the tegmina are 
not narrowed apicalh 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 w ith 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 



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 T. per- 
spicillaris (fig. 25 in Asahina 1 979) 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 1 943: 
39). Bey-Bienko (1938: 121) claimed that 
Margatlea 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 difl"erence 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 
Thcgannsilpha. and I am synonymizing the 
latter genus. The diagnostic characters for 
the genus Margattina Bey-Bienko are so 
similar to those oi 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 Margaitea 

The genitalia of male cockroaches are 
strongly asymmetrical and one of the more 
conspicuous structures is a hooklikc phal- 
lomcre (Fig. 29) although in some species 

(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- 
dophyllodromiinae in Vickerx 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 Phyllndroniica (Loholainpra) nion- 
tana Chopard and several other species of 
the "/>70«/fl/7a -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. monlana 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 niinbata 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 sufiicicnt to use the 
position of the genital hook in placing blat- 
tcllids 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 arc 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. 



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 lougealata (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 
Rchn and Lophoblatta arid Albuquerque 
from South America (see figs. 1-6 in 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 in 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 calcium ox- 
alate crystals but the amount of this com- 
pound varies from dense to sparse to absent 
in species of Blattclla. 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- 

ella Roth (Roth 1984) have an ootheca that 
is typical of species of Blattclla. but it is 
retracted and carried internally until the eggs 
hatch (Roth 1 982). The egg cases of Stayella 
lack calcium oxalate as do species of Bla- 
beridae, all of which incubate their eggs in- 

The oothecae ot M. longealata 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 M. 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 longealata because the male 
and female genitalia of these 2 taxa are so 
much alike (Figs. 5 1 , 57-59, 61,63, 68-70). 
The male genitalia (particularly the left and 
right phallomeres) of M. longealata 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 Kiichinga 
(type species, longealata) would be resur- 
rected. For the present I am placing them 
in a species group of Margattea. 

Species Groups of M.argattea 

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- 



Perspicillaris species group. — Eighth ab- 
dominal tergum specialized (Figs. 9, 24. 28. 
43). Intcrstyiar 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. uimbata. 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- 

Key to Some Males of 
Indo-Pacirc Margattea* 

1. Eighth abdominal segment without a tergal 
gland 1 

- Eighth abdominal segment with a tergal gland 

2. Subgenital plate with lateral comers produced 
giving the appearance of accessory stylelike 
structures (Figs. 51. 55. 56. 69) y 

- Lateral comers of subgenital plate rounded, not 
stylelike (Fig. 1) ceylanica 

3. Head brown with a yellowish orange band be- 
tween antennal sockets (Fig. 65). Pronolal disk 
with a broad dark brown macula (Fig. 64) 

* Includes only males discussed in this paper. I have 
seen specimens only ofnimlmia and paraceylanica from 
the Krakataus. The earlier records of ceylanica. con- 
(mfion. and anccps from these islands arc question- 

- 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) 


4. Intcrstyiar region a subrectangular plate with 
short robust spines on posterior margin (Fig. 
29). Front femur Type B, (Fig. 34). Wings 
shorter than the tegmina, sometimes vestigial 

- Intcrstyiar region not as above. Front femur 
Type B,. Tegmina and wings fully and equally 
developed 5 

5. Tegmina with a dark brown macula on basal 
region (Fig. 41), sometimes extending the full 
length of the wing cover anceps 

- Tegmina without dark markings 6 

6. Apex of median genital phallomere terminat- 
ing in a pair of heavy spinelike structures (Figs. 

18. 23, 26) mmbata 

- Apex of median genital phallomere slender, 
acute (Fig. 10) paraceylanica 

Descriptions of Species of 

Margattea ceylanica (Saussure) 

Figs. 1-7 

Blatta ceylanica Saussure 1868: 355 (9); 
1869: 247; Walker 1871: 20. 

Allacta ceylanica (Saussure): Kirby 1904: 

Phyllodroinia ceylanica (Saussure): Shel- 
ford 1908b: 12; Dammerman 1948: 483. 

Margattea ceylanica (Saussure): Shelford, 
1911: 155: Rehn 1931: 302, lig. 1; Han- 
itsch 1933a: 232; 1933b: 310; 1934: 118; 
1936: 392; Bruijning 1947: 221; 1948:62; 
Bey-Bienko 1957: 899; Princis 1969: 862 
(references to generic combinations and 
records; incorrectly included M. niinbata 
as a synonym); Asahina 1979: figs. 69- 

Material examined. — Sri Lanka. 
(MNHG): holotype female of Blatta ceylan- 
ica. Feradenia, "Ceylan." (ANSP): The fol- 
lowing specimens were determined by He- 
bard as M. ceylanica (Sauss.) and many were 
labelled topotypes: Belihuloya, 1 3 (termi- 
nalia slide 398), l.ix.l928; Bibile Estate, 1 



Figs. 1-7. Margattea cevlanica (Saussurc) trom Sn Lanka. 1, .^-6. males from L'dahamulla. and 7. from 
Battaramulla, 2. female from Labugama: 1 , suhgemtal plate and genitalia (dorsal); 2. supraanal plate and genitalia 
(ventral); .V distal ends of median, aecessory median, and hooklike right genital phallomeres; 4. pronotum; 5, 
supraanal plate and paraprocts; 6. hind wing; 7, tarsal cla« s. ,-\bbre\ lations: a. left phallomere; b. median 
phallomere; c, accessory median phallomere; d. hooklike right phallomere; e, style; f, supraanal plate; g, paraproct; 
h. intercalary sclerite; i. first valvifer. Scales (mm): 1, 2, 0.5; 3, 0.25; 4, l.O; 5, 0.5; 6, 2.0; 7, 0.15. 

S. 26.vii.1929; Wellaway. 1 .^. 28.iv. 1931; 
Rakwana, 1 5, 3.V.1929; Horawupotana, 1 
S. 1 2, 13.x. 1924: Labugama. 1 <5. 2 9(1 with 
genitalia slide 404). 15-18. viii.1932; 
Woodside, Urugalla. 2 6, 22.iv.1924; Bat- 
taramulla, W.P.. 1 s. 20.V.1931; IMaha- 
mulla. W.P.. 1 3 (terminalia slide 399), 

Male. — Intcrocular space decidedly less 
than distance between anlennal sockets. 

Tegmina and \\ings fully de\eloped; hind 
wing with costal veins clubbed, cubitus vein 
\\ ith 2 complete and incomplete branches, 
apical triangle small (Fig. 6). Pronotum sub- 
oval (Fig. 4). Abdominal terga unspecial- 
i/ed. Antero\ entral margin of front femur 
Type B, (with 3 or 4 large spines on prox- 
imal halt); tarsal claws weakly serrated (Fig. 
7; serrations sometimes difficult to see un- 
der binocular microscope). Supraanal plate 



transverse, hind margin trigonal, apex 
sometimes weakly indented: paraprocts dis- 
similar (Fig. 5). Subgcnital plate almost 
symmetrical, styles similar, cylindrical, in- 
tcrstylar margin weakly convex (Fig. 1). 
Genitalia as in Figs. 1 and 3; hooklike right 
phallomere with a small apical spine; apex 
ofmedian phallomere enlarged, the swollen 
portion terminating in a transparent fila- 
ment; apex of accessory median sclerite nar- 
row and slightly curved; left phallomere with 
spinclikc 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 sclcrites greatly reduced. 

Measurements (mm) (9 in 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 

referred to M. nimbata (see remarks under 
that species), or M. 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 
Nkw Spkcies 

Figs. 8-14 

Holotype. — Male, Rakata, Krakatau Is- 
lands, 200 m, s. face, 24.viii.1985. Zool. 
Exp. Krakataus. 

Paratypes. — Krakatau Islands; Rakata, 
Zwarte Hoek, 850 ft., ridge, water trap. 1 S 
(terminalia slide no. 3), 1984; 6.09S 
105. 25E. under rocks. 1 (abdomen missing), 
1 9 (genitalia slide no. 4), 6.ix. 1 984; Rakata. 
s. face, 200 m, I 9, 400 m, 1 nymph, 
24.viii.1985; Panjang, 6.05S 105. 28E, ex 
litter, 1 9, 14.ix.l984, 1 9, beating, 
transition zone, sweep, 1 <?. 18.viii.l985. [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 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 



Figs. 8-14. Margatlea paraccylanica Rolh, paralypes from Rakata. 8-13, male: 8, supraanal plate and 
paraprocts (ventral); 9. abdominal terga 8 and 9; 10, subgenilal plate and genitalia (dorsal); 1 1, hooklike right 
phallomerc; 12, pronotum; l.V hind wing; 14. female, supraanal plate and genitalia (ventral). Abbreviations; a, 
supraanal plate; b, accessorv' median phallomere; c. median phallomere; d, right phallomere; e, left phallomere; 
f subgenital plate; g, style; h, paraproct; i, intercalary sclerite; j, 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. 1 1); 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).— 



Light brown. Head with dark band on ver- 
tex extending as faint stripes on occiput. 
Pronotal disic with light symmetrica! 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 

Measurements (mm) (9 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 ccylanica. 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 

Phvllodroniia ninihala Shelford 1907b: 31 
(<J 5); 1908b: 13; Hanitsch 1915: 57; 
1923b: 410. 

Margattea nimbata (Shelford): Hanitsch 
1928: 23; 1929a: 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- 

Kuchinga nimbata (Shelford): Hebard 1929: 

Material examined. — Lectotype (here 
designated). Male (terminalia slide 200). 
Kuching, N.W. Borneo [Sarawak], pres. 
1905 by Sarawak Museum, 15.xii.l898. 
Type Orth. 89 1/4 in (HDEO). Paralecto- 
types. Sarawak. (HDEO): 2 9 (genitalia slides 
201 and 203), same locality as lectotype, 
Dyak coll. pres. 1900 by R. Shelford, 
25.1.1900 and 27.vii.1900, Type Orth. 
89 3/4 and 89 4/4. 

Additional material. — Krakatau. (RNHL): 
1 S (terminalia slide 139) 1 9 (with rotated 
ootheca, keel to right. Fig. 25) (det. as Mar- 
gattea ceylanica by Hanitsch); 1 9, v. 1908, 
E. Jacobson (labelled M. ceylanica). 

Australia (Northern Territory). (BPBM): 
Holmes Jungle, Palm Cr.. 1 5 km NE of Dar- 
win, 5 m, 2 (? (1 with terminalia slide 468), 
1 (abdomen missing), light trap, 1 1 .iii. 1 96 1 , 
1 3. 14.iii.l961. .1. L. and M. Gressitt. 

Kei Island. (RNHL): 3 9, 1922, H. C. Sie- 
bers (labelled Phyllodromia ninibata Shelf, 
by Hanitsch). 

Christmas Island (Indian Ocean). (ANIC): 
National Park, 1 ,3 (terminalia slide 210), 2 
9, 3 nymphs, x.1983, L. Hill [ANPWS]. 

Thailand. (ZILS): 30 km south of Pak 
Thong Chai, 2 3, 1 9 (reared from 9 taken 
in a dry stream bed) (det. as M. ceylanica 
by Princis), 13.iv.l967, 1 3, 14.iv.l967, 2 9 
(on shrubbery at night), 26.iv.1967 (det. as 
M. ceylanica by Princis), 1 3 (reared from 9 
taken on shrubbery at night) [det. as Mar- 
gattea piinctulata (Brunner) by Princis], 
26. iv. 1967, L. M. Roth. 

Sarawak. (ANSP): Kapit. 1 9 (terminalia 
slide 405), 9.vii. 1910, Wm. Beebe [reported 
as Kuchinga nimbata (Shelford), by Hebard, 
1929:42]. (NRSS): Medan, 5 3. 1 (abdomen 
missing), Mjoeberg (labelled Plniloclromia 
or Margattea nimbata Shelf, by Hanitsch); 
Tjinta Radja. 1 9, Mjoeberg (labelled Phyl- 
lodromia ninibata Shelf, by Hanitsch). 

Borneo (Kalimantan). (NRSS): Tand- 
jong, Redeb, O. Borneo, 3 3 (1 with ter- 
minalia slide 67) (labelled Margattea cey- 
lanica Sauss., by Hanitsch), 2 9 [1 with 
rotated ootheca and labelled Margattea cev- 




. \f\ 

:\ >M 


Figs. 15-22. Margattea nimbata (Shelford). 15. 16, 18. 20. 21. male lectotype. 17, 19. female paralectotype, 
22. female from Kapit. Sarawak: 1 5. pronotum; 1 6. supraanal plate and paraprocls (ventral); 1 7, pronotum; 1 8, 
subgenital plate and genitalia (dorsal); 19. supraanal plate and genitalia (ventral); 20, tarsal claws and arolium; 
2 1 . right phallomere; 22. hmd wmg. Abbreviations: a. supraanal plate; b. paraproct; c. left phallomere; d. median 
phallomere; e. accessory median phallomere; f right phallomere; g, style; h, intercalary sclente; i, 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. mnibala Shelf.), b> 
Hanitsch]. (ZILS): Pelawan besar, 1 9 (del. 
as M. cevlanica by Princis). vi.l937, Mrs. 
M. E. Walsh. 

Java. (RNHL); Ardja Sari, Prcanger, 1 9 
(det. as .U. nimhaia 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 incom- 
plete branches, apical triangle small (Fig. 
22). Front femur Type B, (with 3-5 large 
proximal spines); puh illi 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 posteromcdially 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- 
\exly rounded, apex of hind margin weakly 



Figs. 23-26. Margallfd lumbala (Shelford). 23-25, from Krakatau: 23, male subgenital plate and genitalia 
(dorsal); 24, male, sctal modification on eighth abdommal tergum; 25, female with rotated oothcca attached to 
termmal segments (ventral); 26. male from Northern Temtory 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 intcrocellar band. Pronotal disk yel- 
lowish brown with symmetrica] 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- 



ing 3 terminal segments) pale (cereal color 
varies and the dark areas may be practically 

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) (5 in parentheses). 
Length, 7.2-9.0 (8.0-9.6); pronotum length 
X width, 2.0-2.5 X 2.5-3.3(2.2-2.5 x 3.0- 
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- 
tusifwns Walker, but Hanitsch (1931b: 392) 
was correct in stating that these 2 species 
are distinct. The female genitalia of nimbata 
(Fig. 19) and obtitsifwns (fig. 14L in Roth 
1985b) differ. I have reexamined the type 
female of Blalia ohtusifrons which has been 
placed in IKiichinga (Hebard 1929), Sym- 
/?/ocf (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 arc 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 1 985b)]. The tarsal claws 
of the male are minutely but distinctly 
toothed so I am placing obtusifrons in Mar- 

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: 3 10) 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. 1 

with 18, 23, 26, and Fig. 2 with 19). Mar- 
gattea nimbata is 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 

Theganoptervx perspicillaris Karny 1915: 
103 (6 9); Hanitsch 1927: 33; Shira'ki 1931: 
209, Princis; 1969: 1012 (sp. incertae se- 

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 5(1 with terminalia slide 
6), 1 9. 

Male. — Pronotum subparabolic(Figs. 33, 
35). Tegmina reaching slightly beyond end 
of abdomen. Hind wings vestigial, narrow, 
lateral, reaching between Tl 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; 



Figs. 27-35. Margaltea perspiciUaris (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. pronolum; 34, front femur (anterolateral); 35. 
lectotype. pronotum. Abbreviations: a. paraproct; b. median phallomere; c. accessory median phallomere; d. 
hooklike nght 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 rcflexed dorsad; styles 

tween them a rectangular plate, comers 
rounded, with short robust spines on its hind 
margin (Fig. 29). Genitalia as in Fig. 29; 
hooklikc right phallomere relatively small. 

similar, cylindrical, widely separated, be- without a preapical spine, apex acute, curved 



(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 of an interocular band. Pro- 
notal disk with distinct (Fig. 33) or faint 
(Fig. 34) markings, remaining portion hya- 

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) (9 in parentheses). 
Length, 9.3-9.5 (1 1.5); pronotum length x 
width, 2.7-3.1 x 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 (Phvllodynmia) aiiccps 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: 3 10; 1933a: 232; Dam- 
merman 1948: 483, 555; Bruijning 1948: 
63;Princis 1969: 864. 

Kiichinga anceps (Krauss): Hebard, 1929: 
42 (S). 

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 

worked at that time and that the specimen 
probably no longer exists.) 

Material examined. — Neotype (here des- 
ignated). Male, Tjibodas. Java, 1500 m, 
viii.1921, Karny: in (RNHL). 

Additional material. Java. (RNHL): same 
data as neotype, 2 S (one with terminalia 
slide 141), 1400 m, 1 6 (terminalia slide 
143), 3 9, viii.1921, 2 9, vi.l929, Kamy; 
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 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 



Figs. 36^1. Margaltea 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 antenna! 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) (9 in parentheses). 
Length, 8.5-10.0 (9.0-10.1); pronotum 
length X width, 2.2 x 3.0-3.1 (2.2-2.4 x 
3.1-3.2); tegmen length. 11.0-12.0 (I FO- 

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 



Figs. 42-46. Margaltea anceps (Krauss), male from Tjibodas, Java: 42. supraanal plate and paraprocts 
(ventral); 43, abdominal lergum 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 (9). 
Phvllodromia 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. 
Kiichinga contingens (Walker): Hebard 

Margattea contingens (Walker): Hanitsch 

1929a: 13; 1936: 392; Brmjning. 1948: 


Blatta humeralis Walker 1 869: 1 40 {$); Han- 
itsch 1915: 45. 
Phvllodromia humeralis (Walker): Kirby 

i 904: 9 1 ; Shelford ( 1 906) 1 907a: 490, pi. 

30, fig. 4 (synonymized with contingens); 

1908b: 13;'Hanitsch 1923a: 198. 
Kiichinga humeralis (Walker): Hebard 1 929; 

Margattea humeralis (Walker): Hanitsch 

1928: 21 (contingens and humeralis are 

distinct taxa); Bruijning 1948: 64; Princis 

1950: 174. 
Phvllodromia abrupta Hanitsch 1923b: 399, 

figs. 2, 3 (5). 
Margattea abrupta (Hanitsch) 1928: 21 

(synonymized with humeralis); Bruijning 

1948: 64. 

Material examined. — Holotype. Female 
(genitalia slide 204) of Blatta contingens. 



Figs. 47-52. MargaUea contingens (Walker), male from Long Petak, Borneo: 47, head; 48, supraanal plate 
and paraprocts (ventral); 49, pronotum; 50, hind wmg; 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 Ininieralis, 
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, 1 
(? (terminalia slide 145), 2 2(1 with genitalia 
slide 146), ix.l925, H. C. Siebers, M. O. 
Borneo Exp. 

Male. — Interocular space about the same 
as distance between the antennal sockets 
(Fig. 47). Maxillan. 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 incomplete branches, apical triangle 
small (Fig. 50). Front femur Type B, (with 
5 proximal spines). Eighth abdominal ter- 
gum unspcciali/ed. 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), 



Figs. 53-59. Margallea conlingcns (Walker), male holotype of Blalta humcralis Walker: 53, pronotum; 54, 
supraanal plate and paraprocts (ventral); 55, distal region of subgenital plate showing hind margin and styles 
(dorsal); 56, right posterior comer 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 asymmettical. 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 antenna! 
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 \ein (this mark may be reduced and 
only indicated at the base of the vein; Fig. 
52). Hind wing infuscated (Fig. 50). Ab- 
dominal tcrga light brown, edges pale. Ab- 
dominal sterna lighter, edges darker. Cerci 
pale dorsally and ventrally. 

Female. — Pronotum as in Fig. 60. Su- 



Figs. 60-63. Margallea spp. 60, 61, M. contingens (Walker), female holotype: 60, pronotum; 61, supraanal 
plate and genitalia (ventral): 62. 63, M longcalata (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. 6 1 ; 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) (9 in parentheses). 
Length, 7.4-9.6 (7.7-9.5); pronotum length 
X width, 2.2-2.4 x 3.1-3.2(2.3-2,7 x 3.0- 
3.5); tegmen length, 10.4-10.5 (1 1.0-12.0). 

Comments. -Shelford (1907a: 490) syn- 
onymizcd humcralis (r?) with contingens (9). 
Hanilsch initially agreed with him but later 
(Hanitsch 1 928: 27) considered both species 
distinct, basing his conclusion on the pres- 
ence of a brown streak along the radial vein 
of the tegmen o\' Iniincralis. 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 humeralis that I am cer- 
tain they are the same taxon. Also the fe- 
males from Long Petak are very similar to 
the 9 holotype of contingens from Singa- 

Although M. longcalata 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 





® „ 

Figs. 64-72. Margauea longealala (Brunner). 64, 65, female from 10 miles soulh of Kuching, Sarawak, 
pronotum and head. 66-72. male from Sandakan. Sabali: 66. hind wing; 67. siipraanal pUuc and paraprocts; 
68. lefl gcniial phallonicre; 69, subgenilal plalc and genitalia (dorsal); 70, hooklike right phallomere; 71. tarsal 
elaws and arolium; 72, right eorner of hind margin of subgcnital plate (dorsal). Seales (mm): 64. 65. 1.0; 66, 
2.0: 67. 0.5; 68. 0.15; 69. 0.5; 70-72. 0.15. 

are vcr\ simihif to tliat oi loniiCLilala (Fig. Phyllodroniia loiiiicalala Brunner: Kirby 


Margattea longealata (Brunner) 

Figs. 62-72 

Phyllodionua longe-alata Brunner 1898: 
205, pi. 16, fig. 9 (2). 

1904: 92; Sheltbrd 1908b: 13; Hanitsch 
1915:42; 1923b: 463: 1925: 83 (<?). 

Kuchinga longealata (Brunner): Hebard 

Margatlca longealata (Brunner): Hanitsch 
1933a: 232; Prineis 1969: 865 (the fol- 
lowing references erroneously listed by 



Princis under Margaltea longealata ac- 
tually refer to Balta longealata (Hanitsch) 
Hanitsch, 1930: 254; 1931a: 43; Bruij- 
ning 1947: 214— Princis correctly listed 
these references under Balta 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 9, 25.V.1900, 
Dyak coll., pres. 1900 by R. Shelford (det. 
as longealata by Shelford). (ANSP): same 
data as above, 1 6, 1 S (with ootheca), 
27.iii.1900, 1 <?, 30.iii.l900; 10 miles south 
of Kuching, I 9 (with ootheca), 27. vi. 1910, 

Sabah. (ANSP): Sandakan, Borneo, 1 <5, 
1 3(terminalia slide 401), 1 9 (genitalia slide 
403), 1 9 (carrying ootheca, wall of the 
ootheca on slide 400), Baker. [All (ANSP) 
specimens reported as Kiichinga longealata 
by Hebard(I929: 46).] 

Borneo. (RNHL): Long Petak, M.O., 450 
m. 6 <J, 10 9, ix.l925 or ix.-x.l925, H. C. 
Siebers, Borneo Exp. (NRSS): Pajau River, 
O. Borneo, 2 3, 2 9, 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 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 unspecializcd. Supraanal 
plate transverse, narrow, subrectangular, not 

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 ofcontlngens (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 contingcns (cp. Fig. 61). 
Ootheca as in Fig. 62; length, 5.5 mm; width, 
2.5; height, 1.5. 

Measurements (mm) (9 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, 7.9-1 1.9 (1 1.0- 

Comments. — The color markings of 
Margattea longealata are strikingly differ- 
ent from those of M. contingens. However, 
male structures (subgenital plate, styles, 
genitalia) and female genitalia are so similar 
in both taxa that one might be tempted to 
consider longealata a color morph, variant, 
or subspecies of contingent. Additional 
specimens from other localities should be 
studied to see how variable the color mark- 
mgs are. The unusual ootheca of longealata 
is discussed in the section on subfamily 



placement. It is likely that contingens has a 
similar ootheca. 


I thank the curators and collection man- 
agers listed earlier who sent me specimens. 
Dr. Syoziro Asahina for specimens of The- 
ganosilpha ogatai and T. satsumana. and 
Dr. I. W. B. Thornton for the Maigattca 
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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91(2), 1989, pp. 230-236 




John LaSalle 

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

Abstract.— Jhe genus Zai^rcininicisonui Ashmead is characterized. The genus Mirza- 
gra))iniosonia Girault is synonymized with Zagmmmosoina, and the type species, and 
only included species, M. liiwaticeps Girault, is transferred to Zagrammosoma. New 
Nearctic distributional records are given for Z. lineaticeps from Texas and California, and 
Z. niiriini from Washington. New host records are given for Z. lineaticeps from Kiejeria 
lycopersicella (Gclcchiidae) and Z. itiinini from Kiejeria lycopersicella and Phthorimaea 
operculella (Gelechiidae) and Phyllonorycter eltnaella (Gracillaridae). A new species of 
Zagraiuniosonia. Z. hobbesi LaSalle, is described from California. A key is given to 
separate the three species of Zagraniniosonia which have an entirely black metasoma: Z. 
niiruni Girault, lineaticeps (Girault), and hobbesi LaSalle, n. sp. 

Key Words: Hymenoptera, Euiophidae, MIrzagrammosoma, Zagrammosoma 

During a study involving MIrzagram- 
mosoma lineaticeps Girault, the type species 
and only included species in the genus Mlr- 
zagrammosoma Girault, I realized that this 
genus was a synonym of Zagrammosoma 
Ashmead. At the same lime, I came across 
specimens of a new species of Zagrammo- 
soma from Southern California. The pur- 
pose of this paper is to formally synonymi/c 
M irzagrammosoma with Zagrammosoma. 
characterize Zagrammosoma. describe this 
new species, and provide new host and dis- 
tributional records for Z. lineaticeps 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: AEL American Entomological Insti- 
tute, Gainesville, Florida; BMNH, British 

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- 
ton, D.C. 

Genus Zagrammosoma Ashmead 

llippocephalus Ashmead, 1888: viii. Type 
species HIppocephaliis multilineatus Ash- 
mead (monotypy). Preoccupied by HIp- 
pocephaliis Swainson 1839, in fishes. 

Zagrammosoma Ashmead 1904: 354, 393. 
Replacement name for llippocephalus 

Zagrammatosoma Schulz 1906: 142. Un- 
justified emendation. 

Atoposoma Masi 1907: 276. Type species 
Atoposoma varlegatum (monotypy). 

Mirzagrammosoma Girault 1915: 279. 
Type species MIrzagrammosoma linea- 
ticeps Girault (monotypy). N. syn. 



Figs. 1-6. 1 . Zagrammosoma miruin. 9. head. 2. Cirrospilusjlavoviridis, 9. head. 3. Z. mirum. 9, mesoscutum. 
4. C. flavoviridis. ?. mesosculum. 5. Z. hohbesi. 9, mesoscutum. 6. Z. hobbesi, 9, dorsum of mesosoma. Scale 
bars = 0. 1 mm. 

Diagnosis.— Sculellum with 4 setae (2 
pairs); submarginal vein with at least 3 dor- 
sal setae; funicle 2-scgmented; 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. 

Discussion. — Members of the genus Za- 
graniniosoina 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 



vein, and having 3 or more setae on its dor- 
sal surface; postmarginai vein present and 
well developed. Elachertini have complete 
notauli, Eulophini have the notauli incom- 
plete or absent. Zagrainmosonia is very 
close to Ciirospihis, as both have a 2-seg- 
mented funicle, and the postmarginai vein 
shorter than or equal in length to the stigmal 
vein. ZagraniDiosonia 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 Zagraniniosoina as a 
distinct genus, and provided a key to Nearc- 
tic species. He gave two morphological 
characters to support ZagraniDwsoma as 
distinct from Cirrospilus: Zagrammosonia 
species have an elongate head which is 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- 
podcum, while in Zagramniosoma 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 Zagrammosonia 
(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. lineaticcps Girault displays 
characters consistent with the definition of 
Zagrammosoma: the elongate head with a 

vaulted vertex; and the notaulus curving to 
meet the axilla. 

There are presently eight species of Za- 
grammosoma known in North America 
(Gordh 1 978); the inclusion of Z. lineaticeps 
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 . , 
Z. lineaticeps (Girault) 

I '. Forewing (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- 
ings 2 

2. Mesosoma dorsally with broad, median yel- 
low stripe (Fig. 7); venter of mesosoma yellow. 
Funicular segments both the same color, dark 
dorsally, brown to yellow ventrally 

Z. minim Girault 

2'. Mesosoma entirely black, or with only slight 
yellow markings, not with markings as above. 
First funicular segment dark, second funicular 
segment yellow (Fig. II) 

Z. hobbesi LaSalle, n. sp. 

Zagrammosoma lineaticeps (Girault), 

New Combination 

Fig. 8 

Mirzagrammosoma lineaticeps Girault 

1915: 279. Holotype 9, MEXICO, San 

Rafael, Jicoltepec (USNM) [Examined]. 

Diagnosis. — Z. lineaticeps can be distin- 
guished from the other species of Zagram- 



Dwsoma 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. lincaticeps 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) 

CALIFORNIA: San Diego Co., Batequi- 
tas Lagoon, 1 mi. E. Leucadia, 8.viii.l979, 
C. W. Melton (8 9, UCR). 

TEXAS: Cameron Co., Brownsville,, E. R. Oatman, ex. Kieferia ly- 
copersicella (6 9. UCR). 

Zagrammosoma mirum Girault 

Figs. 1, 3, 7, 9 

Zagrammosoma /?7/ra Girault 1916: 119. 
Holotype 9, USA, California, mountains 
near Claremont (LISNM) [Examined]. 

Diagnosis. — Z. mirum can be distin- 
guished from the other species of Zagram- 

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 in 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 ( 1 978), the species of 
Z. flavolineatiim Crawford (1913) is very 
similar to Z. mirum, differing only in co\- 
oraiion; flavolineatum is slightly lighter col- 
ored than mirum. In Jiavolineatum the 
metasoma is uniformly dark brown, but 
there are small but distinct yellow spots lat- 
erally on the metasomal tcrgites; the pro- 
podeum has the callus yellow, as opposed 
to the propodeum completely black or dark 
in Diirum; 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. flavolinealum 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 
Jiavolineatum would have precedence over 
mirum. Using the kev in this paper, speci- 
mens of jiavolineatum 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). 






Figs. 7-n. 7. Zagraminosoma nuruni. 9, body (from Gordh, 1978; length 2 mm). 8. Z. Imealiceps. 5, 
(brewing. 9. Z. minim, S, forewing. 10. Z. hobhesi. 9, forewing. 1 1. Z. hohbesi, 9, antenna. Scale bars = 0.2 mm. 

Dr. Michael Scliauff has kindly informed 
me that there are specimens of Z. minim in 
the USNM collection from Wenatchee, 
Washington, reared from Phyllonoryctcr cl- 

maella (Gracillaridae). Additional new host 
records for this species are: 

CALIFORNIA: Riverside Co., Corona, 
31.x. 1931, A. J. Baringer, ex. Kicferia ly- 



copersicalla (Gelechiidae) (1 9, UCR); Los 
Angeles Co.. Pasadena, viii.1915, J. E. Gray, 
ex, Phthorimaea operculella (Gelechiidae) 
(1 9, UCR). 

Zagrammosoma hobbesi LaSalle, 
Nkw Si'kcies 

Figs, 5, 6, 10, 11 

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. 1 1 ) 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. 

Head 1.0-1.2 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. 1 1 ) 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 scutcllum distinctly retic- 
ulate, mctanotum and propodeum lightly 
sculptured to smooth. Propodeum with me- 
dian carina; propodeal callus with 4-7 setae. 

Forewing(¥\%. 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 Ll-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^.0 times longer than wide, 
pointed apically. 

Male.- Length 1.1-1.5 mm. Differs from 
female only in genitalia. 

Distribution. — Known only from South- 
ern California. 

Material examined.- Holotype 9. CAL- 
IFORNIA, San Bernardino Co., Summit 
Valley, 14.V.1985, G. Gordh, on Eriodic- 
tyon (mounted on point, LISNM). 

18 9, 10 3 Paratypes. CALIFORNIA: as 
holotype (7 9, 2 $. UCR; 3 2, USNM); as 
holotype but on Haplopappus (2 9, 1 S, 
UCR); San Bernardino Co., Mojave River 



Forks, ~9 mi. S. Hesperia, 2.V.1985, J. D. 
Pinto (3 9, 4 6, LAS; 1 9, 1 <5: BMNH, CNC, 

Etymology.— The coloration of the face, 
yellow with numerous black stripes, is rem- 
iniscent of the face of a tiger. This species 
is named for Calvin's tiger friend, Hobbes. 


I thank G. Gordh and J. D. Pinto for 
supplying material of Z. hobbesi, M. E. 
Schaufffor loans of material from the USNM 
and for host information on USNM speci- 
mens, and G. Gordh for supplying the il- 
lustration of Z. minim. 

Literature Cited 

Ashmead, W. H. 1888. Descriptions of some un- 
known parasitic Hymenoptera in tlie collection of 
the Kansas State Agncultural College, received 
from Prof E. A. Popenoe. Bull. Kansas St. Agric. 
Coll. 3: I-VII (Appcndi.x). 

. 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. pis. 31-39. 

Boucek. Z. and R. R. Askew. 1968. Palcarctic Eu- 
lophidae (excl. Tetrastichinae). (Hym. Chalcidoi- 
dea.) Index of Enlomophagous Insects. Lc Fran- 
cois, Paris. 260 pp. 

Burks. B. D. 1979. Family Eulophidae, pp. 967-1022. 

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. 1 198 pp. 

Crawford, J. C. 1913. Descriptions of new Hyme- 
noptera, No. 6. Proc. U.S. Natl. Mus. 45: 241- 

De Santis, L. 1979. Catalogo de los himenopteros 
calcidoideos de america 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: 1 19-122. 

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- 

Graham, M. W. R. de V. 1969. The Pteromalidae of 
Northwestern Europe (Hymenoptera: Chalcidoi- 
dea). Bull. Bril. Mus. (Nat. Hist.), Entomol. Supp. 
16. 908 pp. 

Masi, L. 1907. Contribuzioni all conoscen/a dei Cal- 
cididi italiani. Boll. Lab. Zool. gen. agr. Portici I: 

Peck, O. 1963. A Catalogue of the Nearctic Chalci- 
doidea (Insecta: Hymenoptera). Can. Entomol., 
Suppl. 30: 1-1092. 

Peck, O.. Z. Boucek, and A. Hofler. 1964. Keys to 
the Chalcidoidea of Czechoslovakia (Insecta: Hy- 
menoptera). Mem. Entomol. Soc. Canada 34: 1- 

Schulz.W. A. 1906. Strandgut.Spoliahymenop. Pad- 
erborn: 77-269. 

91(2), 1989, pp. 237-247 


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: C. dentata, C. prolixa, taxonomy, neotropics 

The genus Coenochroa Ragonot, 1 887 was 
previously known from three species, all 
North American. C. hipunctella inhabits the 
Atlantic and Gulf coastal plains, while ////- 
bclla and californiella are mainly western, 
the former extending eastward in 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. Heppncr, ed., in 

Although a review of the genus would be 
premature, 1 have included a key to the five 

known species. It is useful to know that: a) 
only illibella has a white costal band, b) all 
but illibella 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 ( 1 968, 
1984) for illustrations and other informa- 
tion pertaining to North American species 
of Coenochroa. 

Key to Species of Coenochro.4 

Male genitalia with valve rounded, unmodi- 
fied, not dentate; forewing spot present or ab- 
sent {illibella): costal band present {illibella) or 
absent; Irons with central beak minute to 
prominent; North and Central America , . 2 
Male genitalia with valve dentate; (brewing with 
dark spot at lower outer angle of cell; costal 
band absent; frons wiih central beak minute 
(e.g. Fig. 17) or absent (Fig. 20). varying in- 

traspecifically; Brazil 4 

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) 



Figs. 1^. Adull moths and head profiles. 1-2, Coenochroa denlala. male paratype (LJSNM slide no. 57864). 
3-4, Coenochroa piolixa. male parat\pe (USNM slide no. 57867). Scale bar = 2 mm (Figs. 1, 3) 1 mm (Figs 
2, 4). 

Forewing with dark spot at lower outer angle 
of cell; costal band absent; frons with the cen- 
tral beak small to absent, not extending beyond 

rim 3 

Central beak of frons extending to rim; aede- 
agus with vesica bearing numerous small cor- 
nuti and single larger one; .Atlantic and Gulf 

coastal plains 

bipunctella (Barnes and McDuniiough) 

Central beak of frons minute, not reaching nm; 
aedeagus with a smgle long slender cornutus; 
western North America through Central Amer- 
ica californiella Ragonot 

Forewing light yellowish brown, darker ante- 
rior to cell; costa of valve with serrate flange 
(Fig. 24), vinculum with saccus very short (Fig. 

23); ductus bursae straight (Fig. 38) 

dentata Shaffer 

- Forewing pale yellow, not darker anterior to 
cell; costa of valve with single large tooth, sac- 
cus very long (Fig. 32); ductus bursae with loop 
(Fig. 41) pmlixci Shaffer 


Macerated and cleared genitalia were 
stained with 0.3% aqueous solution of mer- 
curochrome, mounted in Euparal, and pho- 
tographed with a Nikon AFM camera on a 
Nikon S-Kc II microscope set for Koehler 
illumination. All scanning electron micro- 
scope work was done with a Hitachi S-530 
SEM at 5 Kv. One head capsule for each of 
the two species was macerated in hot 10% 



Figs. 5-10. Scanning electron micrographs of denuded head capsules in frontal (top row), an terodorsal (middle 
row), and lateral (bottom row) views, all to same scale. 5-7. Coenochroa dciuala. paratypc (USNM slide no. 
57865). 8-10. Coenochroa prolixa. hololypc (J. Shatter slide no. 2239). Scale bar = 0.5 mm. 

KOH. cleaned, denuded of scales, mounted 
from 95% ethanol using silver paint, air 
dried, and sputter coated with gold. The 
dentata valve (Figs. 30-31) was similarly 
treated. All original photographs were taken 
on Kodak Technical Pan Film 2415. 

Coenochroa dentata Shaffer. 

New Spec IF.S 

Figs. 1, 2, 5-7, 11-13, 17-19. 

23-31, 38-40 

Diagnosis, — E.xternally ver>' similar to 

prolixa, but with forewing ground light yel- 



Figs. 1 1-16. Denuded head capsules showing ventral aspect (top row), mouthparts (middle row; g = galea, 
mp = maxillary palpus, pf = pilifer), and ventral view of frons protuberance. 1 1-13, Coenochroa dentata. 
paratype (USNM slide no. 57865). 14-16, Coenochroa prolixa. holotype. Scale bar = 250 ixm (Figs. II, 14), 50 
^m (Figs. 12, 15), 100 Mm (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 '4, 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- 


Figs. 17-22. Enlargments of frons protuberances in frontal (top row), anterodorsal (middle row), and lateral 
(bottom row) views. 17-19. Cocnochroa deniata. paratype. 20-22, Coenochroa prolixa, holotype. Scale bar = 
100 /urn (Figs. 17. 20). 200 ^m (Figs. 18, 19. 21, 22). 

ciput, patagium, and tcgiila light brown. 

Forevving radius about 8-9 mm: venation 
Nariable; R, usually free, sometimes stalked 
with Ri + s; R: always stalked with R^ + s, the 
common stalk variable in length; R, . 4 

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. 





-- J 









Figs. 23-31. Coenochroa dentata, male genitalia. 23-36, hololype (J. Shaffer slide no. 2254). 27. paratype 
(USNM slide no. 57865). 28-3 1 , 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 ^m (Fig. 30), 
25 Mm (Fig. 31). 



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 ape.x. 
M;+3 fused, long stalked with Cu,, from 
lower outer angle of ceil. 

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 acdeagus, 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 y4 as long as left; vesica unarmed. 

Female genitalia (Figs. 38-40) with ovi- 
positor triangular, about -A 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 lobesjoined by mem- 
brane roughened with numerous minute, 
closely-set cusps; this membrane extending 
anterior to ostium and posteriorly to ovi- 

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.— (?, labelled; "Rio Brilhante M 
Grosso, Brasil 22. I. 1971 Becker leg."; '"6 
genitalia on slide 2254 J. C. Shaffer"; 
"Holotype Cocnochroa dentata Shaffer" 

Paratypes. — 3 3, same locality as holo- 
type, dates; 23-1-1971 (USNM slide 57865), 
[USNM]; 25-I-I971 (USNM slide 57864). 
[USNM]; 25-1- 1971, Becker col. no. 1 3800, 
undissected [NMRJ]. 2 9, same locality as 
holotype, dates: 22-1-1971 (J. Shaffer slide 
2251), [NMRJ]; 23-1-1971 (USNM slide 
57866), [USNM]. All labelled; "Paratype 
Cocnochroa dcntala Shaffer." Specimen de- 
position given in brackets. 

Distribution. — Known only from the type 

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, 


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- 





Figs. 32-37. Coenochwa proli.xa. 32-35. 37. Holotype (J. Shaffer slide no. 2239). 36, Paratype (USNM slide 
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. 4 1 ). The single large 
costa tooth, elongate saccus, and ductus bur- 
sa loop are each unique to this species of 

Description. — Forewing radius about 8- 
9 mm; venation similar to C. dentata. 
Ground pale yellow throughout, set with 

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. 



Figs. 38^3. Female genitalia. 38^0, Coenochroa denlata. 41-43, Coenochroa prolixa. paratype (USNM 
slide no. 57868). 38. paratype. dorsal view (USNM slide no. 57866). 39. paratype. lateral view (J. ShafTer 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 ^m (Fig. 40), 10 yum (Fig. 42). 25 ^m 
(Fig. 43). 



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 -A; straight single row 
of about 8-10 setae extending from base of 
projection toward base of valve, row par- 
allel to and about 'A 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 '/j 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 dcnlaki, 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.— <?, labelled; "Rio Brilhante, 
Mato Grosso, Brasil 23-27. X. 1970 V. O. 
Becker col."; "<? genitalia on slide 2239 J. C. 
Shaffer"; "Holotype Coenochroa prolixa 
Shaffer"; [NMRJ]. 

Paratypes.— 3, same locality as holotype, 
date; 25-1-197(1), (USNM slide 57867), 
[USNM]. 6 9, same locality as holotype, 
dates: 25-X-1970 (USNM slide 57863), 
[USNM]; 27-X- 1 970, Becker col. no. 1 3304 
(J. Shaffer slide 2256) [NMRJ]; 21-1-1971 
(USNM slide 57868), [USNM]; 23-1-1971 
(USNM slide 57869), [USNM]; 25-1-1971, 
Becker col. no. 1 3925, (J. Shaffer slide 224 1 ), 
[NMRJ]; 25-1-1971 (J. Shaffer slide 2253), 
[NMRJ]. All labelled; "Paratype Coeno- 

chroa prolixa Shaffer." Specimen deposi- 
tion given in brackets. 

Distribution.— Known only from the type 

Etymology.— The specific epithet is an 
adjective derived from the Latin prolixa 
(stretched out) in reference to the elongated 

Apomorphies. — In dcntata 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- 

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 illibclla, and 
obvious differences in the frons structure are 
seen in the scanning electron micrographs 
of the two specimens illustrated herein (Figs. 
1 7-22). 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 


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 2 247 

The holotypc and selected paratypes are Literature Cited 

deposited in the National Museum, Rio de ^^^^^^^^ ^^^^ B ^^ ^j,^,^^,,^, ^^ Neotropical Lepi- 

Janeiro, Brazil [NMRJ], The remaining dopicra. Pan 2. E. j. Bnll. Leiden. In prep. 

paratypes are in the National Museum of Shaffer, J.C. 1968. A Revision of the Peoriinac and 

National History, Smithsonian Institution, Anerastiinae (Auctorum) of America North of 

Washington. D.C.. USA [USNM], The Mexico (Lepidoptera: Pyralidae). U.S. Natl. Mus. 

scanning electron microscope used in this !."" ,2°; Neotropical Pvrahd moths transferred 

study was supported in part by NSF Grant from Anerastiinae (Auctorum) to Phycitmac. Proc. 

No. BSR-85 1 11 48. Entomol. Soc. Wash. 86(2): 383-395. 

91(2), 1989, pp. 248-256 





Jay C. Shaffer and Eugene Munroe 

(JCS) Department of Biology, Cicorge Mason University, Fairfax. Virginia 22030; (EM) 
Granite Hill Farm, R.R. #2, Dunrobin, Ontario KOA ITO, Canada 

.'Ifovrar/. — Lectotypes are designated for three Zeller species: Notairha qiiatcrruilis. N. 
icincratalis, and A', nnisccrdalis. These and the holotypc oi N. cassusalis Walker, the type 
oi Notarcha, are redescribed and the wings, head profiles, and female genitalia illustrated. 

Key Words: Nutairhu iiiuilcnialis. .V. teineratalis, N. cassusalis, N. muscerdalis, lecto- 
types, African Pyraustinae 

In his revision of the Pyralidae Hampson 
(1898: 728) synonymi/ed seven names un- 
der Lygropia qiialernalis (Zeller). In re- 
searching our paper (in prep.) on the Cram- 
bidae of Aldabra Atoll, we found it necessaiy 
to reexamine this synonymy to determine 
the identity of an Aldabran species erro- 
neously identified in the literature as qua- 
tcnialls. On examining type specimens we 
disco\ered that most, perhaps all. of these 
seven names represent distinct species. The 
.•\ldabra species matches none of them and 
will be described as new . 

The purpose of this paper primarily is to 
designate lectotypcs for quaternalis and two 
related African species to provide stability 
for the names, and secondarily to redescribe 
and illustrate the three lectotypes and the 
holotypc of an additional species to separate 
previously confused forms and to facilitate 

The African species that Hampson syn- 
onymi/ed under qualcnialis are tcnwralahs 
Zeller, and cassusalis Walker. All three 
species are properh referred to Xotarcha 
Meyrick, 1884, for which cassusalis is the 

type species. In this paper we include the 
related A', nuisccntalis. not part of Hamp- 
son's synonymy, but of which we studied 
ZcUer's type. Noiarcha is a large genus with 
many undescribed species and deserving of 
extensive study. The scope of this paper is 
limited to delineating described African 

Type material referred to herein is in the 
collections of the British Museum (Natural 
History) [BMNH], and the Naturhistoriska 
riksmuseet, Stockholm [NHRM]. 

Key ro Included Species of Noiarcha 

1. Forcvving iinilormly yellow, with single dark 
diseal spot (Fig. 4) nuisccrcJalis 

- Forewing yellow with transverse lines or dif- 
fuse bands of darker yellow; with dark diseal 
spot and y 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) quawnialis 

- Labial palpus without dark markings 3 

.V Foreuing with anterior half of transverse pos- 
terior band m I'orni of diagonal line (Fig. 3) . 

- Forewing with broad diffuse bands, no lines 
(Fig. 2) cassusalis 



Notarcha quaternalis (Zeller) 

Figs. 1, 5, 9, 13-16 

Botys qtialcnialis Zeller, 1852, pp. 44-45. 

Diagnosis. — Among the described Afri- 
can species of Ihe Notarcha qitalcnialis 
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 is 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; forctarsus yellow, black 
markings on basal half and on distal third. 
[Meso- and metathoracic legs lost on type.] 

Forewing(Fig. 1) radius 1 1 mm; with four 
subcqual 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 forcwing. 

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 Mj and on 1st 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- 



Figs. 1-4. Wing patterns. I, .youircha quatcrnalis. lectotype; 2. N. cassusalis. holotype; 3, A'. temeraiaUs. 
lectot>pe; 4. ,V. musardalis. lectotype. Scale bar = 2 mm. 

Figs. 5-8. Head profiles of above specimens. 5, A', qualernulis; 6, A', cassusalis; 7, A", lemeratalis: 8, N. 
muscerdalis^ Scale bar = 1 mm (Figs. 5-8). 

formly finely scobinate, each scobination set 
in center of iiregular plate, many plates hex- 
agonal or nearly so; posterior part of bursa 
with irregular sclerite, spinose on both sides. 

its posterior end folded into a short knob- 
shaped pouch with spines facing outward, 
these spines numerous, slender, sharp 
pointed; signum absent. Ductus seminalis 



Figs. 9-12. Types with labels. 9, Notarcha quatcnndis. Icctotypc (1.6x); 10, N. cassusahs. holotype. insert 
shows reverse side of "Pt Natal" label (1.4 x); 11, A'. iemcratahs,\cclo\yps(\.(i^)\ \2, N. luiiscerdalis, leclolype 

from membranous posterior part of corpus 

Type locality. — Natal, South Africa. 

Lectotype, hereby designated, labelled: 
"Caffraria."; "197"; "Riksmuseum Stock- 
holm"; "9 genitalia on slide 1 870 J. C, Shaf- 
fer"; "Botvs quaternalis Lectotype by .1. 
Shaffer & E. Munroe, 1989" [NHRM]. 

Notarcha cassusalis (W alker) 

Figs. 2, 6, 10, 17-21 

Zehronia cassusalis Walker. 1859. p. 477. 

Diagnosis.— Among the described Afri- 
can species of the Notarcha quaternalis 
complex this species is externally similar to 
quaternalis. but lacks dark markings on the 
labial palpus, and has broader more diffuse 
transverse bands on the wings. The spinosc 

triangular plate at the entrance to the corpus 
bursae and the internally spinose ductus 
seminalis are each characteristic of this 

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 quaternalis. 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; tcgula 
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 



Figs. 13-21. Female genitalia. XTi, Notarcha quatenmlis. leclolype(20x); 14, ovipositor, enlarged (45 x); 15- 
16. armature at entrance to corpus bursae, two focal levels (50 x); 17, N. cassusalis, corpus bursae surface, 
enlarged to show surface texture (220 » ); 18, female genitalia (28 x ); 19, armature at entrance to corpus bursae 
( 1 1 " ); 20. rotated specimen showing origin of ductus seminalis (28 x ); 21, ductus semmalis. enlarged to show 
internal spines (220 x). 

yellow, brown spot at apex of first segment, 
smaller spot at apex of second segment, third 
segment yellow, fourth segment brown, fifth 
segment yellow, Midfemur rather uniformly 
light yellow, small dark brown spot at apex; 

midlibia nearly uniformly light yellow, 
lacking dark markings: midtarsus light 
brownish yellow. Metathoracic leg color- 
ation similar to that of mesothoracic leg. 
Forewing (Fig. 2) radius about 9 mm. 



Figs. 22-27. Female genitalia. 22, Nolarcha lenwratalis, lectotype (28 x ); 23. corpus bursae surface, enlarged 
to show surface texture (200 >). 24, Nolarcha luiisccrcia/is. lectotype (23>); 25, signum, enlarged (55 x); 26. 
signum and adjacent bursa surface, enlarged (220 x); 27, rotated specimen showing origin of ductus seminalis 

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 



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. 1 9); 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; "Hoiotype"; "Type"; 
"Pt Natal [& on reverse side] 57 3"; "Ze- 
bronia Cassusalis"; "9 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 oi' 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- 

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- 

scribed for A^. quaternalis. Eye diameter 0.6 
mm, black. Ocellus as described for A^. qua- 
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, difl^use bands, and 
broader diff'use 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 1 st 
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- 



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- 

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) is 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 Wahlbcrg's first journey (1838-1845) 
there as to southern Africa, only later (1853) 

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"; "9 genitalia on slide 1 87 1 J. C. Shaf- 
fer": "'Botys teincratalis Lectotype by J. 
Shaffer & E. Munroe, 1989" [NHRM].' 

Notarcha muscerdalis (Zeller) 
New Combination 

Figs. 4. 8, 12, 24-27 

Botys muscerdalis Zeller, 1852, pp. 43-44. 

Diagnosis.— This species is 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 A', 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 

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 



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 temcratalis above 
applies here also.) 

Lectotype, hereby designated, labelled: 
"Type"; "Botys muscerdalis Z. Caffr. 43. 
Led 85. Caffraria"; "Zell. Coll. 1884"; "9 
Pyralidae Brit. Mus. Slide No. 1 806 1 "; "5o- 
tvs muscerdalis Lectotype by J. Shaffer & E. 
Munroe, 1989" [BMNH]. 


We thank Michael ShalTer 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. Pan 1. Proc. Zool. Soc. Lond. 1898: 590- 
761, figs. 1-87, pi. 49, 50. 

Horn, Walther and Use Kahle. 1936. Uber entomo- 
logische Sammlungen. Ent. Beih. Berl.-Dahlcm 3: 
161-296, pi. 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: 

91(2). 1989. pp. 257-268 




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. 

.4^5/rar/.— Scanning electron micrographs of the male terminalia are given for all six 
species of the stonefly genus Ostroccrca Ricker: O. albidipennls (Walker), O. complexa 
(Claassen), O. diinicki (Frison), O. foersten (Ricker), O. pwlongata (Claassen), and O. 
iruncata (Claassen). Structures of the male genitalia previously not illustrated are de- 
scribed. Distribution records for each species are also reported. 

Key Words: Stonefly, Plecoptera, Nemouridae, Ostrocerca, scanning electron micro- 

Scanning electron microscopy (SEM) has 
been used recently to elucidate fine struc- 
tures of the male genitalia of stoneflics 
(Kondratieff and 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 (Kondratieff and Kirch- 

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 30 1 Freeze-Frac- 
ture Unit at 10 '' mv. Specimens were then 
mounted and sputter coated with gold in a 
Hummer V Sputter Coater. Specimens were 



Figs. 1^. Oslrocerca albidipennis. I , Epiproct, dorsal view, 312^.2, Hypoprocl. apex, 845 x . 3, Terminalia, 
ventral view, 120x. 4, Terminalia, ventral view, 241 x. Terms: hp, hypoproct; vs, ventral sclerite; la, lateral 
arms; il, inner lobe; c. cerci; v, vesicle. 

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. Flmt, Jr., Smithson- 

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 



of Comparative Zoology, Harvard Univer- 
sity (MCZ); and J. R. Voshell. Jr., Virginia 

Morphological terms of the following de- 
scriptions follow Baumann (1975). Bau- 
mann (1975) provides excellent illustra- 
tions of the male terminalia of O. tnmcata, 
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 
scleritc; la. lateral arms: ol, outer lobe; ml, 
middle lobe; il, inner lobe; c, cerci; and v, 

Ostrocerca albidipennis (Walker) 

Neinoiira alhidipoinis 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, lilies, 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 arc hooked medially and beak- 
like apically (Figs. 3 and 4). The figures pre- 
sented by Claassen (1923) as TV. 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 (A'. 
serrata). Bar Harbor, Maine, 8 VI 1921, C. 
W. Johnson (Cornell #1 199). Paratype: same 
data as holotype, 1 M (Cornell #1 199). 

Other specimens: CANADA: Ontario, 
Algonquin Park, Costello Lake, 26 V 1938, 
W. M. Sprules, 1 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, 1 M. 

1 F (CNC). Quebec, Harrington Lake, Gat- 
ineau Park, 27 V 1954, H. J. Huckel, I 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, 1 
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, 1 8 V 1939, W. B. Nutting, 
Warren, 21 VI 1941, T. H. Frison & H. H. 
Ross, 1 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. 

I M, 2 F(CSU). VERMONT: Mt. Hay Stack, 
2400', 28 V 1938, W. B. Nutting. 1 M 
(USNM). VIRGINIA: Giles Co., Mountain 
Lake, 2 VI 1 962, J. R. Vockeroth, 2 F(CNC); 
Grayson Co., springs into Lewis Fork, Co. 
Rt. 603, 4 mi. W of Troutdale, 9-30 VI 
1981, 16M, 5F, 3N, RFK&BCK(RFK), 

II 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, 1 M (RFK). 

Ostrocerca complexa (Claassen) 

NemouracomplexaClaassen, 1 937:43. Type 
locality: Artists Brook, Essex County, New 

Ostrocerca complexa, lilies, 1966: 217. 

The elaborate epiproct (Figs. 5, 6) of this 
species has never been completely illustrat- 



Figs. 5-8. Osirocerca coinplexa. 5, Terminalia, dorsolateral view, 194 » . 6, Epiproct. dorsal view, 212 x . 7, 
Basal cushion of the dorsal sclerite, 965 x . 8, Terminaha, ventral view, 1 63 x . Terms: hp, hypoproct; vs, ventral 
sclerite; il, inner lobe; c, cerci; v. vesicle. 

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 

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). 



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, 1 1 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, I M (USNM). VIRGINIA: 
Nelson Co., tributary to South Fork Tye 
River, Montebello Fish Culture Station, jet. 
Rt. 690 & Rt. 56, 3 V 1980, BCK, 1 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, Monongahcla National Forest, 16 V 
1981, RFK, 14 M, 6 F(RFK), same locality 
15 V 1986. RFK & BCK, 2 M, 1 F (BCK); 
Tea Creek, Tea Creek Campground, Mo- 
nongahcla National Forest, 21 V 1978, RFK 
& R. M. Meyer, 1 M (RFK). 

Ostrocerca dimicki (Prison) 

Nemoura dimicki Frison, 1936: 256. Type 

locality: Corvallis, Oregon. 
Ostrocerca dimicki, lilies, 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. 1 2). 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 II 1935, R. W. Prentiss (INHS). 
Paratypes: Cor\allis. drainage stream. 16 III 
1934,' E. E. Ball, 5 M (USNM); same data, 
2 F (INHS); Corvallis. Brooklane Creek. 28 
II 1935, R. W. Prentiss. 1 M (USNM). 

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 


Ostrocerca foersteri (Ricker) 

Nemoura foersteri Ricker, 1943: 70. Type 
locality: Reservoir Creek, Cultus Lake, 
British Columbia. 

Ostrocerca foersteri, lilies, 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, 1 1 F(CNC). CALIFORNIA: 
Humboldt Co., Willow Cr., Hwy 299, below 
Berry summit, 24 IV 1 987, 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 1 964. 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 

Ostrocerca prolongata, lilies, 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 arc long and apically 
hooked (Figs. 1 9-2 1 ). The cerci are long and 




Figs. 9-12. Oslrocerca dimicki. 9. Terminalia, dorsal view. 163x. 10. Epiproct. dorsal view, 462 x. 11, 
Ventral sclcrite, lateral view. 356 x. 12. Terminalia. ventral view. 120x. Terms: hp. hypoproct: vs. ventral 
sclerite; c. cerci; \ , vesicle. 

The allotype female is badly damaged and subgenital plate (as illustrated), and vena- 

thc terminalia are missing. As Hitchcock tion is \ery similar to the sympatric Soye- 

(1974) points out, the female illustrated by dina washingloni (Claassen). Females found 

Claassen (1923) may be misidentified. The in collections with male O. prolongala ex- 



Figs. 13-14. Ostrocerca foersleri. 13, Terminalia, dorsal view, 163> 
Terms: vs, ventral sclerite; il, inner lobe; c, cerci. 

14, Terminalia, dorsal view, 462 x. 

aniined during this study were similar to 
females of O. coiuplexa and O. albiclipcnius. 
And, because Ostrocerca prolongata was 
usually collected with the other two species, 
made positive association impossible. This 
species appears to be rare throughout its 

Material examined. — Holotype M: Bret- 
ton Woods, New Hampshire. C. W. John- 
son, 26 VI 1913 (MCZ). Paratype: Orono, 
Maine, 1 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 Tyc River, Rt., 687, 3 V 1980, BCK, 
2 M (VPI), WEST VIRGINIA: Tucker Co., 

Red Creek at Lancville, 26. 27 May 1973, 
O, S, Flint, 2 M (USNM). 

Ostrocerca truncata (Claassen) 

Nenioura truncata Claassen, 1923: 290. 

Type locality, Walden, Massachusetts. 
Ostrocerca truncata, lilies, 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 

The female has been illustrated by Bau- 
mann (1975), Hitchcock (1974) and Ricker 



Figs. 15-18. Osiukcixu proU)ngala. 15, Tcriiiinalia, dorsal view, IVOx. 16, Process of ventral sclerite, 655 x. 
17, Terminalia, lateral view. 156x. 18, Ventral sclerite, bifurcate process, 356 x. Terms: ds, dorsal sclerite; vs, 
ventral sclerite; c, cerci. 

(1965). This species was once considered 
the rarest of the Oslwccrca. But. we found 
it to be common and widespread. The Al- 
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 

Material examined. — ALABAMA: 
DeKalb Co.. spring seep. DeSoto State Park, 
River Trail, cabins 8 & 9, 17 V 1988, RFK 
con Falls, Naugatuck St. Forest. 1 7 VI 1 96 1 . 
S. W. Hitchcock, 1 M (USNM): Portland, 
20 VI 1 963, S. W. Hitchcock. 1 M (USNM): 

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, 10 IV 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, 
Whately, Whately Glen, 18 V 1938, J. F. 
Hanson, 1 M (USNM); OHIO: Athens Co., 
Canaan Township. 25 IV 1941, J. Walker, 
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, 



Figs. 19-21. Ostrocerca prohmgata. 19, Terminalia, lateral view, 137 x . 20, Terminalia, ventral view, 106 x . 
2 1 , 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 
1987. RFK, BCK, 6 F (BCK); Tazwell Co.. 
Station Springs Cr., Burkes Garden 3840', 
off Rt. 666, 20 V 1982, RFK & BCK, 48 

M, 14 F (RFK); Washington Co., spring to 
Brumley Cr., Clinch Mtn. at Low Gap, 
3600', Rt. 690. 23 V 1981, RFK & BCK, 
15 M, 8 F (RFK); Wythe Co., Stony Fk. of 



Figs. 22-25. Oslrocerca tnincala. 22. Terminalia, dorsal view. 178x. 23, Epiproct, dorsal view, 263 x. 24, 
Terminalia, ventral view, 120'<. 25, Terminalia, ventral view, 356 x. Terms: hp. hypoproct; ds, dorsal sclerite; 
vs, ventral sclerite; il, inner lobe; ol, outer lobe; c, cerci; v, vesicle. 



Reed Cr., Dark Horse Hollow Rcc. Area. 
Jefferson Nat. For., 29 IV 1978, RFK, 1 M 
(RFK). WEST VIRGINIA: Cabell Co.. 
Lusher Hollow of Mud River, near Bar- 
boursvillc, 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, 1 5 IV 1 973, RFK, 1 F (RFK): 
Pendleton Co., Spruce Knob, 4 mi. below 
summit, 16 V 1963, W D. Field & O. S. 
Flint. 1 M, 1 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 
LaneviUe, 26, 27 VI 1973, O. S. Flint, 1 M, 

3 F (USNM). 


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 (9. coniplcxa easily distinguishes 
this species; the serrate apex of the hypo- 
proct is only found in O. alhidipcnnis: and 
the ventral sclerite terminating in an elon- 
gate process is characteristic of O. pralon- 
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. 


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- 

ment of Agriculture, Agriculture Research 
Service, Orlando, Florida for his sugges- 
tions and help. Dr. Richard W. Baumann, 
Brigham Young LIniversity 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 slonclly fam- 
ily Nemouridac (Plccoplcra): A study of the world 
fauna at the generic level. Smithsonian Conlnb. 
Zool. 211. 74 pp. 

Claassen, P. W. 1923. New species of North Amer- 
ican Plecoptcra. Can. Entomol. 55: 257-263, 281- 

. 1937. New species of stoneflies (Plecoptcra). 

J. Kansas Entomol. Soc. 10: 42-51. 

Prison, T. H. 1936. Some new species of stoneflies 
from Oregon (Plecoptcra). Ann. Entomol. Soc. 
Amer. 29: 256-265. 

Hitchcock, S. W. 1974. Guide to the insects of Con- 
necticut: Part VII. The Plecoptcra or stoneflies of 
Connecticut. Bull. Stale Gcol. Nat. Hist. Surv. 
Conn. Bull. 107. 262 pp. 

lilies. .1. 1966. Katalog der Rezenten Plecoptcra. Das 
Ticrreich. 82. Walter dc Gruyter and Co.. Berlin. 
632 pp. 

Kondratieff, B. C. and R. F. Kirchner. 1984. Anew 
species of Nemouridac (Plecoptcra) from the Great 
Dismal Swamp, Virginia, USA. Proc. Entomol. 
Soc. Wash. 86: 578-581. 

. 1987. Additions, taxonomic corrections, and 

fauna! aflinilies of the stoneflies (Plecoptcra) of 
Virginia. USA. Proc. Entomol. Soc. Wash. 89: 24- 

Needham, J. G. and P. W. Claassen. 1925. A mono- 
graph of the Plecoptcra or stoneflies of .America 
north of Mexico. Thomas Say Found. Entomol. 
Soc. .-^mer. 2. 397 pp. 

Nelson, C. R. and R. W. Baumann. 1987. Scanning 
electron microscopy for the study of the winter 
stonefly genus Capma (Plecoplera: Capniidae). 
Proc. Entomol. Soc. Wash. 89: 51-56. 

Ricker, W. E. 1943. Stoneflies of southwestern British 
Columbia. Indiana L'niv. Publ.. Sci. Ser. 12. 145 

. 1952. Systematic studies in Plecoptcra. In- 
diana Univ. Publ.. Sci. Ser. 18. 200 pp. 

. 1965. New records and descriptions of Pie- 


coptera (Class: Insccta). J. Fish. Res. Bd. Can. 22: stoneflies (Plecoptera) of West Virginia. Enlomol. 

475-501. 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 

91(2), 1989, pp. 269-285 


David A. Nicicle and Margaret S. Collins 

(DAN) Systematic Entomology Laboratory, BBH, 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. 

Abstract. — \ new species of drywood termite of the genus Neotennes 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, Ncotcnues jouteli (Banks), on the basis of karyotypic and isozymatic 

Key Words: Neotennes, Kalotermitidae, termites 

Termites resembling Neolernws joulcli 
(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 Thome 1989, in press). When 
chromosomal, isozymatic, or behavioral 
studies are used in termite investigations, 
more species may be discovered in North 

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 leni^lli. 
in lateral view, the distance from the left 
ventral mandibular condyle to the occiput; 

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; eye 
diameter, the width of the eye of the imago 
measured from its anterior to posterior 
margin; postociilar 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; length and wi(/th of 
pronotum. in dorsal view, the medial length 
and greatest width of the pronotum; length 
and width of wing, the total length of the 
forewing, including the scale, and the great- 
est width of the forewing; total length, in 



lateral view, the length of the body from the 
tip of the face to the apex of the abdomen; 
k'lii^lli 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 kalotcrmitids 
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 
W/D, 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; Wiiii; L IT, the ra- 
tio of the wing length to wing width; Wing 
L/U'ingscale L. and Post nient urn 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 is 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], 


Neotermes luykxi Nickie and Collins 
Nkw Spkcies 

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. 2! August 1986. (Coll. P. 
Luykx). [NMNH]. 

Paratype ^el•ies. — 26 roldiers, 15 ima- 
goes, 20 nymphs. Florida: Broward Co., 
Hollywood, PL 439, (P. Luykx), 6 soldiers, 
5 imagoes, 4 nymphs [NMNH]; Same lo- 

Tablc 1. A list of chromosome numbers (2n) for 
Florida species of Kalotemiitidae (Luykx and Syren 
1979, Luykx pers. comm). 

Spi-L ics 

Chromosome Number 

Calcaritermes nearclicus 


Cryplokrincs hrcvis 

37 (3), 36 (9) 

Cryptolermcs cavifrons 


Incisitermes milleri 


Inasitermes sch warzi 


Incisitermes snydcn 


Kalolennes approximaliis 


Neolermes castaneus 


Neotermes luykxi 

45 (3), 44 (S) 

Neotermes joiileli 


caiity, PL 558 [morphotype sample], VIII- 
21-1986, 7 soldiers, 10 imagoes, 3 nymphs 
[NMNH. AMNH]; Florida: Broward Co., 
Dania, PL 389, Vl-20-1984, (P. Luykx), 3 
soldiers, 3 nymphs [NMNH]; PL 389C, 
same locality as PL 389, VI 1984, 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, Xll-6-1930, (Coll. E. M. 
Miller), 1 soldier, 1 nymph [NMNH]. 

Short-Headed soldier. — //('at/.- 1.21 x 
longer than wide, 1 .46 x longer than deep 
(Figs. IE, 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 1 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 13 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 



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Table 4. TERMINTE LONG-HEADED SOLDIERS: means (.v) and ranges of measurements (in mm) of 
morphological features of Neotennes and Incisttermes species from Florida. Numbers in parentheses after value 
mdicate size of sample. NA = not available for measurement. 







# antcnnal articles 








Head length 






2.41 (2) 


2.00 (2) 

Head width 








Head depth 









Mandible length 








Postmentum length 





2.34 (4) 




Postmentum width 



0.41 (10) 

0.48 (6) 





Width pronotum 









Length pronotum 




1.21 (6) 





Total length 









cluster situated a distance of less than di- 
ameter of cluster from antennal fossa. Man- 
dibles -A as long as head capsule; in lateral 
view, curving upward anteriorly; mandible 
dentition and shape of postmentum as in 
Fig. IE. 

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 x 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. — //cflf/.' 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 x 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 Neoternies. 
with M nearly as sclerotized as Rs, running 
closer to Rs than to Cu. Length of forewing 
9.9-10.8 mm (.v= 10.3 mm), 3.77 x longer 
than its greatest width (Fig. 3B). 




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Table 6. Ratios of morphological features of long-headed soldier termites from Florida. Numbers in paren- 
theses after value mdicatc size of sample. NA = not available for measurement. 







Head L/W 









Head L/D 



1.91 (10) 




2.29 (8) 


Head W/D 








Head L/Mandible L 







2.08 (8) 


Pronotum W/L 





2.00 (4) 




Postment. L/min. W 



6.41 (10) 


7.09 (4) 

6.00 (2) 

8.04 (8) 


Color: Head, pronotum, legs, and tergites 
of abdomen uniformly yellowish; clypeus 
whitish-yellow; labrum same color as head 
capsule. Stemites lighter yellow than ter- 

( 'analion: Twenty-two soldier paratypes 
are short-headed forms and four are long- 
headed forms (Figs. IB, 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, postmenlum 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.— So/^/f/v 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 oi' Incisitcrnics. 
Of the Florida species of Neotermes. luykxi 
and joutcli are more closely related and arc 
easily distinguished from castancus (Bur- 
meister 1839), a species with only one form 

of soldier. Neotermes castaneus is 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 A^. 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. luykxi is 1 .2 1 ; 
for long-headed soldiers, the ratio is 1.42 
for N. jouteli and 1.57 for A^. luykxi. The 
L/W ratio for N. castaneus is 1.32. The long- 
headed soldier of A. luykxi usually has both 
a narrower head and a longer, narrower 
postmentum than N. jouteli. Although A'. 
luykxi is generally smaller than A. 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[S] 44[2] for A', luykxi, 56 [both 
sexes] for N. jouteli) (Table 1). The mor- 
phometric difference that seems to be most 
useful in differentiating these two species is 
the ratio of the postmentum length to its 
minimum width. For short-headed soldiers 
this ratio is 4.41 for A', luykxi and 4.98 for 
N. jouteli. For long-headed soldiers it is 7.09 



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. 





Incisiicrnws Kahtermes 

milleri approximalus 

Head L/W 


1.21 (13) 

1.21 (17) 





Head L/D 







2.11 (4) 



Head W/D 








Head L/Mandible L 









Pronotum W/L 









Postmcntum L/niin. W 




4.41 (17) 






for .V. Iiiykxi. 4.62 for N. joutcli. and 6.41 
for A', castanctis. Xeptcrnics Itiykxi and .\'. 
joutcli also differ in isozyme composition 
(Luykx et a!., in prep.). 

Imago: Assignment of the imago of A'. 
liiykxi to Neoti'inu's 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 of all morphometric fea- 
tures listed in Tables 2 and 5, none of which 
by itself is useful in identifying an individ- 
ual specimen. XcoUtdics luyk.xi 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 A^ 
joutcli and A', luyk.xi have onl\ scattered 
short setae (Fig. 6A-I). Wing length, wing 
scale length, and head width are generally 

largest for A', castaneus and smallest for A'. 
luyLxi (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) o{N. jou- 
teli was 56 [both sexes] and only 45 [S] and 
44 [9] for A", luyk.xi. Luykx also found in- 
terspecific differences in a series of isozymes 
among five selected kalotermitids (includ- 
ing A', joutcli and ;V. luyk.xi). 

It has become mcreasingly 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, Gryllotalpa 
scptcnidecimchromosoniica Ortiz (1958), 
and more recently with a complex of other 
sibling species of Gryllotalpa: quindecim, 
sedecim, octodccim. and \7^//;// (Baccetti and 

Fig. 1 . Termite soldiers, head and pronotum, dorsal view. .Also figured beside each soldier head: postmenlum, 
ventral view. .■\. Ncolcrmcs joulcli. long-head (LH); B, ;V liiykxi. LH; C. .V. caslancus: D, S. jouleli. short-head 
(SH); E, N. luykxi, SH; F, Kalolermcs appro.ximatus; G, Calcanicrmcs iwarctwus; H, Incisitermes snyden; I, /. 
schwarzi, LH; J, /. schwarzi. SH; K, I. milleri. LH; L, /. milleri. SH. 



/. schwarzl LH 

/. schwarzl SH 

mlllerl SH 




/. snyderi 

C. nearcticus 

Fig. 2. Termite soldiers, head profile. A, .Xeoleniws castciiieus: B, N. joutcli. long-head (LH); C, N. jouleli. 
short-head (SH); D, N. luykxi. LH; E, N. luykxi, SH; F, Incisitennes schwarzi. LH; G, /. schwarzi. SH; H, 
Kalotermes approximatus; I, /. snyderi; J, /. millcii, LH; K, /. milleri, SH; L, Calcaritermes nearcticus. 

Capra 1978). These species, othei^vise mor- 
phologically very similar to Gryllolalpa 
gryllotalpa (L.), have chromosome comple- 
ments of 17, 15, 16, 18, and 14, respec- 
tively, compared with 12 for (7. 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 Incisitennes milleri (Emerson). /. 
snyderi (Light), /. schwarzi (Banks), and 
Neotermes jouteli, N. Inyk.xi 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. Grasse 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 5th 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 luyk.xi). 
All of these species are known to occur in 

Kalotermes appro.ximatus is 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 




N. iouteli 

W. luykxl 

C. cavifrons 

Fig. 3. Wings of alates of kalotermitids. A, Neotermes joulelt B, A', hiykxi; C, N. caslaneus: D, Incisilermes 
schwarzi; E, /. snyden; F, Crypioiermcs cavifrons; G, Calcarilermes nearclicus. (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 liicisiteiDies. 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 liiyk.xi is presently known only from 
Monroe. Dade, ard Broward Counties. 
Florida. Neotermes species tend to be more 
moisture-dependent than those of Incisi- 
termes (Collins 1 969). Neotermes jouteli may 
enter man-made structures, infesting wet 
wood or wood that becomes wet from time 
to time. Neotermes castaneus also has been 



Fig. 4. Scanning electron micrographs of morphological features of kalotermitid termites. A, Calcanlennes 
nearcliais. apex of left foretibia, lateral aspect. B, W-oicrmcs castaneus soldier, left mandible, dorsal aspect. C, 
D. Scoiermes castaneus. imago mandibles: C. left. D, right. E, F. Crvptotermes 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). 
Incisitennes 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 
{Taxodiuin spp.) in the United States, caus- 
ing extensive damage to untreated wood and 



Fig. 5. Scanning electron micrographs of morphological features of kalotermitid termites. A, C, E. Crypto- 
tennes brevis. soldier: A, dorsal, C, lateral, E, frontal aspect. B, D, P. Cryplotcrmes cavifrons. soldier: B, dorsal, 
D, lateral, F, frontal aspect. 

fence posts (Miller 1949). Incisitennes 
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 /. snydcri. Incis- 

ilcrnu's nilllcri is known only from the for- 
ested areas of the Florida Keys and .lamaica 
and is found in the sound wood of dead 

The remaining drywood termites in this 
region all have soldiers with phragmotic 
(stopper-like) heads. Calcarilcnncs has nu- 






i '^ L^jriitfl^B 











merous species in the Neotropics but only 
a single species, ncarcticus (Banks 1918), in 
the United States. Calcaritcnncs ncarctlciis 
is found in natural habitats in northern 
Florida but has not yet been recorded from 
man-made structures. In contrast, termites 
of the genus Cryptotermes are major pests. 
Cryptotermes cavifrons has been found in 
natural habitats throughout peninsular 
Florida, islands of the Caribbean and Cen- 
tral America (Araujo 1977), but Ciypto- 
IcTDies brevis, the more important pest 
species of the two, is known only from the 
dry wood of man-made structures in the 
United States and elsewhere in the world. 
It is a common household pest from North 
Carolina through Florida to Texas and is 
frequently found in furniture, picture frames, 
and other wooden items that have been 
transported to northern states from areas of 
infestation in the South and other warm re- 
gions of the world. 

The following identification key resolves 
some of the difficulties in determining ter- 
mites from the eastern United States. 

Key to the Ten Species of 

Kalotermitidae Occurring in 

Eastern United States 


1. Anterior edge of second marginal tooth of left 
mandible elongated, longer than straight line 
transect of posterior edge of first marginal tooth 
(Fig. 4E) 2 

- .Anterior edge of second marginal tooth of left 
mandible not elongated, subequal in length to 
straight line transect of posterior edge of first 
marginal tooth (Fig. 4C) 6 

2. Media (M) of forevving running midway be- 
tween radial sector (Rs) and cubitus (Cu) and 
extending to apex of wing (Fig. .^D, E) [Iiicis- 
ilcniics] 3 

- Media (M) of forewing curving anteriorly and 
joining radial sector at about two-thirds length 

of wing from suture (Fig. 3F) [Ciyploiennes] 5 

3. Body length 7-8 mm; body dark brown .... 
Incisilermes milleri 

- Body length greater than 10 mm, but usually 
not exceeding 16 mm; body yellow to yellowish 
brown 4 

4. Body length 15-16 mm; yellow-brown; wing 
length (including wing scale) greater than 10 
mm (Fig. 3D); head shape as in Fig. 6M-0 

Incisitennes schwarzi 

- Body length 10-12 mm; yellowish; wing length 
less than 10 mm (Fig. 3E); head shape as in 
Fig. 6J-L lucisitcniu's snyderi 

5. Body length 10-12 mm; head width behind 
eyes greater than 1.0 mm; antenna usually with 
16-18 articles Crypiolermcs brcvis 

- Body length less than 10 mm; head width less 
than 1.0 mm; antenna usually with fewer than 

16 articles Ciyptolernies cavifrons 

6. M vein of forewing unsclerotized, located mid- 
way between Rs and Cu; compound eye less 
than 0.35 mm in diameter; ocellus less than 
0.12 mm in diameter; head length/width ratio 
greater than 1 .0; body sooty black; wings short- 
er than body Kalolcrmes approximatus 

- M vein of forewing nearly as scleroti/ed as Rs, 
located closer to Rs than to Cu; without other 
above combination of characters 7 

7. Rs and M of forewing close together, without 
crossveins; Cu equidistant between M and anal 
margin (Fig. 3G) Calcaritermcs nearclicus 

- Rs and M of forewing wider apart, with several 
crossveins extending from Rs to costal border; 
Cu closer to M than to anal margin (Fig. 3C) 
[Neotcrmcs] 8 

8. Body length 15-16 mm; wing length greater 
than 12 mm; wing dark brown; forewing with 
numerous crossveins between Rs and M (Fig. 
3C); body darkly pigmented; head and prono- 
tum usually with both long and short setae (Fig. 
6A-C) Neotermes caslaneus 

- Body length less than 1 5 mm; wing length 9.0- 
1 1.5 mm; wing clear or only slightly pigment- 
ed; forewing with a few crossveins between Rs 
and M (Fig. 3A. B); body golden or yellow; 
head and pronotum with short setae only (Fig. 
6C^I) 9 

9. Mean total body length 8. 1 mm (range 7.8-9.0 

Fig. 6. Scanning electron micrographs of morphological features of kalotermitid termites. A, B, C, Ncotenncs 
castaneus. dorsal, dorso-lateral. lateral, respectnely. D. E. F, Neolcrmes joulcli. G, H, I. Neoternws luykxi. J, 
K. L. Incisilermes snvderi. M. N. O. Incisitcrnies schwarzi. 



mm); mean head length L44 mm, mean head 

width 1.66; eye diameter 0.4-0.5 mm 

Neotermes joiileli 

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 luyk.xi 


1. Head phragmotic, with front of face rising 
sharply at an angle of 50-90'" to longiludmal 
a.\is; mandibles small relative to size of head 

capsule (Figs. 2L, 5A-F) 2 

Head not phragmotic, with front of face slop- 
ing gradually at an angle less than 45° to lon- 
gitudinal axis; mandibles well developed . 4 

2. One anterior tibial spur on foretibia enlarged, 

well developed (Fig. 4A) 

Caicanlernies nearcticus 

- Tibial spurs on foretibia subequal in length 

[( ryinolcnnes] 3 

3. Dorsum of head capsule weakh concave; sur- 
face of forehead smooth or finely crenulaled 

(Fig. 5B, D, F) Cryptotennes cavifrons 

Dorsum of head capsule distinctly concave; 
surface of forehead deeply convoluted (Fig. 
5.A, C, E) Cryplolenncs hrevis 

4. Anterior margin of pronotum neither deeply 
concave nor incised 5 

- .Anterior margin of pronotum deeply concave 

or incised (Fig. IH-L) [Inci^itenncs] 10 

5. Third antennal article only slightly longer than 
fourth; hindfemur moderately innatcd. with 
L/W ratio greater than 3.0; head length greater 
than 3.4 mm; head depth greater than 1.7mm 
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 6 

6. Eyes unpigmented Kalotermes approximatus 

- Eyes pigmented, purplish, gray, or sooty black 

7. Postmentum length greater than 2.00 mm . 8 

- Postmentum length less than 2.00 mm . . 9 

8. Mean ratio of length of postmentum to its 
minimum width 7.09; head width 1 .9-2. 1mm; 
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] 

9. Head length ca. 2.3 mm (range: 2.0-2.9 mm); 

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: 1.8-2.1 
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] 

10. 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 11 

1 1. Head length greater than 3.0 mm; postmen- 
tum length greater than 2.4 mm 

Incisitermes schwarzi [long-headed] 

Head length less than 2.6 mm; postmentum 
length less than 1.9 mm 12 

12. Head length greater than 2.3 mm, and head 

width less than 1 .6 mm 

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 

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. IJ); 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. IH); antenna usually with fewer than 16 
articles Incisitermes snyderi [short-headed] 


We wish to thanlc 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. BBIL USDA, 7c National Mu- 
seum of Natural History, Smithsonian In- 
stitution. Washington. DC. Finally, we are 
indebted to Dr. Peter Luykx. University of 
Miami. Coral Gables, PL, for the specimens 
upon which this paper is based and for shar- 
ing information regarding the biology of the 
drywood termites of Florida. 



Literature Cited 

Araujo, R. 1970. Termites of the neotropical region. 

Chapter 12, pp. 527-571. In 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. Lc Specie Italiane del gcnere 

Gryllolalpa 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). Ll.S. Natl. 

Mus. Bull. No. 108: 1-128. 
Burmeister. H. C. C. 1839. Handbuch dcr Entomo- 

logie, 11(1): 757-1050. Berlin. 
Collins. M. S. 1969. Water relations in termites. 

Chapter 14, pp. 433-458. //; Krishna, K. and 

Weesner, F. M., eds.. Biology of Termites, vol. 1. 

Academic Press, New York and London. 
Grant, H. J., Jr. 1 965. A measuring device for use m 

insect systematics. Entomol. News 76: 249-251. 
Grasse, P.-P. and C. Noirot. 1958. La societe de Ca/- 

oleniies JIavicollis de la fondation au premier es- 
saimage. Comp. Rev. 246: 1789-1795. 

Haverty, M. I. and B. L. Thome. 1989. (In press.) 
Agonistic behaviour correlated with hydrocarbon 
phenotypes in dampwood termites, Zootennopsis 
(Isoptera: Termopsidae). Animal Behaviour. 

Luykx, P., D. A. Nickle, and B. I. Crother. (In prep.) 
A morphological, electrophorelic, and karyotypic 
assessment of the phylogeny of some lower ter- 
mites (Isoptera: Kalotcrmitidae): Lack of congru- 
ence among different data sets. 

Luykx, P. and R. M. Syren. 1979. The cytogenetics 
of Iiicisitcrmes 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 (iryllotalpa grylliHalpa 
(L.). Publ. Inst. Biol. Apl. Barcelona 27: 181-194. 

Snyder, T.E. 1933. CV;/a(/7?('/7);« in the LInited 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 Linited 
States, A Handbook. The National Pest Control 
Association. Elizabeth, NJ. 70 pp. 


91(2), 1989, pp. 286-287 


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 ce;ci. 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. 1 A). 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. IB). It is 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) Ciyinocercuspuiiclulafiis. Airow indicates the 
apicolateral emargination. 



subgcnital plate had shriveled, making the 
apical area character ambiguous. With this 
exception, the presence (female) or absence 
(male) of an apicolatcral emargination of 
the subgenital plate is a diagnostic character 
for gender determination in C. punctiilatiis. 
I thank W. E. Clark for help with mor- 
phological terminology and G. L. Miller for 

the illustration. This is Alabama Agricul- 
tural Experiment Station Journal Series No. 

Arthur G. Appel, Department of Ento- 
mology and Alabama Agricultural Experi- 
ment Station, Auburn University, Alabama 

91(2), 1989, pp. 287-288 

Sex and Deposition of the Holotype of 

Bareogonalos canadensis (Harrington) 

(Hymenoptera: Trigonalyidae) 

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 1 897 by Taylor ( 1 898. 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 ( 1 896) described the holotype 
as a male. Taylor ( 1 898) 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. //; Wytsman, Genera 
Insectorum 61: 24 pp.), has been perpetu- 
ated by others, including Carlson (1979. 

1 197-1 198. In Krombein et al. Catalog of 
Hymenoptera in America north of Mexico. 
Vol. I. Smithsonian: Wash., D.C. 1 198 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 
1 897) on which only females were collected. 
Schulz [1907b, (1906). fieri. Entomol. Zeit. 
5 1 : 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 S [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 9 
[Harrington's handwriting]/Zool. Mus., 
Berlin, [printed label]. 

Two autotypes at the Canadian National 
Collection (CNC, Ottawa), with locality and 



Table 1. Specimens of BarcDifonalos canadensis 
collected from Gabriola Island, B.C., Canada, in Oc- 
tober, 1897. 

D.iu- NuintxT and Sex 


2 1 Oct 9 females 

22 Oct 3 females 

3 males 

23 Oct 2 females 

24 Oct 4 females 

25 Oct 2 females 

1 male 

26 Oct 2 females 

27 Oct 1 female 

3 at Canadian National Col- 
lection (CNC) (1 with yel- 
low paralype label) 

1 H. Townes, Gainesville, 

2 at CNC 

I 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 

2 at CNC 

H. Townes, Gainesville, 

I at CNC 

I at Washington State Uni- 

CNC (With red holotype 

' This information from Taylor ( 1 898) 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 ofTaylor 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 Universitact, Berlin) for loans 
oi 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, Depaiinient of Ento- 
i}wlogv, Oregon State University. Corvallis, 
Oregon 97331. 

91(2), 1989, pp. 289-290 


Pedipalpal Anomalies in Neobisium simoni 

(L. Koch) and A^. bernardi Vachon 
(Neobisiidae: Pseudoscorpiones: Arachnida) 

Pseudoscorpions have been found with 
segmental anomalies involving the sclerites 
(tergites and sternites) (Curcic & 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 cancwides (Linnaeus) with the 
right foreleg fused basally with the pedipalp. 
In addition, Chamberlin ( 1 949. 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 A', carpaliciini 
Beier (Curcic. 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 et) 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 Curcic (1980). The aim of this 
note is 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 

In a collection of pseudoscorpions made 
by one of us (RND) at Passarole, near Mou- 
lis (Ariege), France, during July 1987, one 
female of Neobisium simoni (L. Koch) and 
one tritonymph of A', 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 is 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 ihe pcdipalps, aberrant 
female, (2) Right chela of the peciipalps, normal female. 



Figs. 3^. Neobisium hcrnardi Vachon. Scale line 
= 0.5 mm. (3) Right chela of the pedipalps, aberrant 
tritonymph, (4) Right chela of the pedipalps. normal 

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. 

(Figs. 3, 4). The movable finger of the right 
pedipalpal chela is reduced and attains two- 

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 
TV. hcrnardi 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 
(A^. carpatictim, N. simoni, N. hcrnardi). 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. Curcic and R. N. Dimitrijevic, 
Institute ofZoolog\\ Faculty of Science, 16. 
Student ski Trg, YV- 11000 Beograd. Yu- 

91(2), 1989, pp. 291-292 


Ticks of the Subgenus Ixodiopsis: First Report of 

Ixodes woodi from Man and Remarks on 

Ixodes holdenhedi. a New Junior Synonym of 

Ixodes ochotonae (Acari: Ixodidae) 

Recently, this laboratory received a par- 
tially engorged nymphal Ixodes tick that had 
been removed from the neck of a child (age 
and sex not stated) on 5 June 1 967 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 Ixodes 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 Ixodes 
woodi Bishopp, a member of the morpho- 
logically close-knit subgenus Ixodiopsis 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 /. woodi. 

In North America, the subgenus Ixodiop- 
sis is often referred to as the "Ixodes an- 
gustus group," after Ixodes angustus Neu- 
mann, a widespread parasite of cricetid 
rodents. Other Nearctic members of the /. 
angustus group include (besides /. woodi) 
Ixodes eastoiii Keirans and Clifford, to date 
known only from southwestern South Da- 
kota and the border area of northeastern 
Wyoming, where it parasitizes cricetine and 
microtinc rodents; Ixodes ochotonae Greg- 
son, a parasite of ochotonid lagomorphs and, 
to a lesser extent, cricetmes 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 Ixodiopsis is 
Ixodes holdenriedi Cooley, which is known 
only from the holotype and paratype, both 
females taken on the pocket gopher Tho- 
nioniys 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, /. ochotonae. 

Though Spencer (1963, Proc. Entomol. 
Soc. Brit. Columbia 60: 40) published a rec- 
ord of/, 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 /. 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 /. 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: 1 partially engorged 
9 from the scalp of a boy. Sandy River. 
Multnomah Co., Oregon. 7 August 1934 
(RML 1 1 8942); 2 partially engorged 92 from 



the arm of a 14-year-old girl. Forest Grove, 
Washington Co., Oregon, 8 August 1932 
(RML 118959); and 1 partially engorged 9 
"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, 1 1 July 1938 
(RML 118964). There are also 4 unpub- 
lished records in the National Tick Collec- 
tion itself 1 engorged 9 from a child. Kirk- 
land, King Co., Washington, August 1947 
(RML 24023); 1 engorged 9 from a young 
boy, Washington [State], 20 July 1956 (RML 
33925); 1 partially engorged 2 from the head 
of a young girl, Seattle, King Co., Washing- 
ton, ca. 2 August 1977 (attachment site be- 
came inflamed on 12 August) (RML 
105346); and 1 engorged 9 from an other- 
wise undefined human host, Juneau. Great- 
er Juneau Borough, Alaska, 1 August 1953 
(RML 118623). 

The nymph of /. woodi reported herein 
(RML 1 1 8594) may be described as follows 

(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 comua api- 
ces 0.297, width at level of comua 0.261; 
palpi 0.245 long, 0.09 1 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. Robbms, Department of 
Health and Human Services, Public Health 
Service, National Institutes of Health, Na- 
tional Institute of Allergy' and Infectious Dis- 
eases, % Department of Entomology, Mu- 
seum Support Center, Smithsonian 
Institution, Washington. DC. 20560. 

91(2). 1989, pp. 293-294 

Book Review 

Lepidoptera Anatomy. By John L. Eaton. 
Wiley-Interscicnce (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 a representative example. At a 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 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- 



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. Al- 
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 

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. Smithsornan Institution, Washington, 
DC. 20560. 

91(2), 1989, pp. 294-295 

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 in- 
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- 
hshments, 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- 

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. 



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 
a little help from well-placed friends, he won 
in 1854 the post of State Agricultural So- 
ciety Entomologist. Fitch continued in that 
capacity for 1 9 years, during which he wrote 
1 4 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 

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 

The book has a large page (10^4" by S'/:"), 
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, 1 5 orders, and 1 07 
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. Gagne, Systematic Ento- 
mology Laboratory. Agricultural Research 
Service, % USNM NHB 168, Washington. 
D.C. 20560. 

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 
+ 4 1 5 pp. ISBN 0-8 1 30-0845-X. $35.00/ 
hard, from 15 NW St., Gainesville, FL 

The original edition of The Mayflies of 
Florida by Lewis Berner was published in 
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- 

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 7 1 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 



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. 

W. P. McCafferty, Department of Ento- 
mology. Purdue University, West Lafayette, 
IN 47907. 

91(2). 1989. pp. 298-303 


Frederick William Poos, Jr. 

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 1 9 1 7 and from 1 9 1 7 to 1 922 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 com. 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 



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 com 
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- 
\olving 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- 

When, in 1 94 1 , the property on which the 
experimental farm was located was needed 

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. 



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 60S. 
Federal Building. Hvattsville. Maryland 

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-histor\ studies of 

three jointworm parasites. Jour. Econ. Ent. 21: 
405^26, Dec. 1921. 

and . A lamp for la,\onomic work 

in entomology. Jour. Econ. Ent. 14: 504-506, 1 
fig., Dec. 1921. 

and . Five new species belongmg 

to genus Hannolila Motschulsky (Isosoma Walk- 
er etAuct.). Ka. Univ. Science Bull. 14: 349-359, 
pis, 36-37, Oct. 1922. 

and . The wheat strawworm and 

its control. U.S. Dept. Agr. Farmers' Bull. 1323: 
l-IO, May 1923. 

Poos. F. W. A new pest of spinach in Virginia. 
Va. Truck Expt. Sta. Bull. 56: 491-497, July 1, 

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 com borer 
(Pyrausta niibila/is Hubn.) and two closely relat- 
ed species in northern Ohio. Ohio Jour. Science 
27: 47-94. March 1927. 
. An insect borer damaging roses. South- 




em 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. 
Oct. 1, 1927. 

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. 







■ 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. 1 1, 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): 1 45-1 50, Nov. 

. Leafhopper injury to legumes. Jour. Econ. 

Ent. 22: 146-153, Feb. 1929. 

21. . Leafhopper injury to legumes. (Author's 

Abstract). Jour. Wash. Acad, of Sciences 20(6): 
116-117, March 19, 1930. 

22. . A new method of distributing Empoasca 

fabae (Harris). Jour. Econ. Ent. 23(4): 770, Aug. 


23. and F. F. Smith. A comparison of ovi- 

position and nymphal development of Empoasca 
fc/'ac (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 
of some leafhoppers of the genus Empoasca. Jour. 
Agr. Res. 43: 267-285, Aug. I, 1931. 

25. Poos, F. W. and C. M. Haenseler. Injury to va- 
rieties of eggplant by the potato leafhopper, Em- 
poasca (abac (Harris). Jour. Econ. Ent. 24: 890- 
892. Aug. 1931. 

26. Poos, F, W. Biology of the potato leafhopper, 
Empoasca fabae (Harris) and some closely re- 
lated species oi Empoasca. Jour. Econ. Ent. 25(3): 
639-646, June 1932. 

27. and R. B. Deenier. Is the absorption of 

copper by certain crop plants influenced by cli- 
matic, soil, or other factors? Jour. Econ. Ent. 26: 
648. June 1933. 

28. Poos, F. W. Leafhoppers of the genus Empoasca 
(Homoptera. Cicadellidae) in Virginia. Proc. Va. 
Acad. Sci. 1932-1933: 36-37. 

29. . Four new species of Empoasca (Homop- 
tera, Cicadellidae). Proc. Ent. Soc. Wash. 35(8): 
174-179, 1933. 

30. and N. H. Wheeler. On the hereditary 

ability of certain insects to transmit diseases and 
to cause diseaselike injunes to plants. Jour. Econ. 
Ent. 27(1): 58-69, 1934. 

31. and H. L. Westover. "Alfalfa Yellows." 

Science 79(2049): 319, April 6, 1934. 

32. Poos, F. W. A stand used in photographing ob- 
jects from above. U.S.D..A.. Bur. Ent. FT- 16. May 


33. . A leafhopper injurious to peanuts. Pea- 
nut Journal and Nut World 13(8): 9, 13, June 7, 



34. Elliott. Charlotte and F. W. Poos. Overwintering 
ofAplanobacter stewarti. Phytopathology 25(1): 
32. January 1935. 

35. Poos, F. W. and Charlotte Elliott. Bacterial wilt 
of corn and its insect vectors. (.Abs.). Phytopa- 
thology 25(1): 32. January 1935. 

36. Poos, F. W. Under heading of notes and exhi- 
bition of specimens: Plants of Einpoasca fahac 
(Harris) and their probable significance. (Au- 
thor's abstract). Proc. Ent. Soc. Wash. 37(8): 170. 

37. . New host plants of the potato leafhopper, 

Empoasca fahae (Harris) and their probable sig- 
nificance. Jour. Econ. Ent. 28(6): 1072-1073. 

38. , 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. & Pit. Quar. ET-72. 3 pp.. 
Feb. 1936. 

39. and H. W. Johnson. Injury to alfalfa and 

red clover by the potato leafhopper. Jour. Econ. 
Ent. 29(2): 325-331. illus.. 1936. 

40. and Charlotte Elliott. Certain insect vec- 
tors of Aplanobaclor stewarl. Jour. Econ. Ent. 
29(2): 325-331. 1936. 

41. Hurd-Karrer. A. M. and F. W. Poos. Toxicity of 
selenium-containing plants to aphids. Science 
84(2176): 252, 1936. 

42. 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. 

43. 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. 

44. Poos, F. W. Insects in relation to diseases of ce- 
real and forage crops. Jour. Econ. Ent. 31(1): 24- 
38. 1938. 

45. 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. 

46. Poos. F. W. and J. W. Scrivener. A convenient 
cage for determining field populations of the po- 
tato leaftiopper. U.S.D.A. Bur. Ent. & PI. Quar- 
antine ET- 121. June 1938. 

47. Poos, F. W. Control potato leafhoppers on pea- 
nuts. Southern Planter 100(6): 15, June 1939. 

48. . Control potato leafhoppers on peanuts. 

Extension Entomologists 3(2): 4-5. June 1939. 

49. . Host plants harboring Aplanobaclor 

slewarli without showing external symptoms af- 
ter inoculation by Chaetocncma pulicana. Jour. 
Econ. Ent. 32(6): 881-882. 1939. 

50. Elliott. Charlotte and F. W. Poos. Seasonal de- 

velopment, insect vectors, and host range of bac- 
terial wilt of corn. Jour. Agr. Res. 60(10): 645- 
686, 1940. 

5 1 . Poos, F. W. The locust leaf miner. Chalcpus dor- 
salis Thumb, as a pest of soybean. Jour. Econ. 
Ent. 33(5): 727-728. 1941. 

52. 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. 

53. Poos. F. W. On the Causes of Peanut "Pouts." 
Jour. Econ. Ent. 34(5): 727-728, 1941. 

54. . Peanut "pouts" caused by thrips. The 

Southern Planter 102(12): 12, illus., 1941. 

55. . The potato leafhopper. A pest of alfalfa 

in the eastern states. U.S.D.A. Leaflet No. 229: 
1-8. 1942, revised 1952. 

56. . Control of hay insects in the eastern 

United States. Mimeographed by Bureau of 
Entomology & Plant Quarantine. U.S.D.A., 
March 1943. 

57. 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. 

58. Poos, F. W. and L. A. Hetrick. Tctralopha scor- 
Icalis (Led. ) A new insect pest of Icspede/a. Jour. 
Econ. Ent. 38(3): 312-315, 1945. 

59. Poos. F. W. DDT to control com flea beetle on 
sweet corn and potato leafhopper on alfalfa and 
peanuts. Jour. Econ. Ent. 38(2): 197-199. April 

60. . The control of tobacco thrips on seedling 

peanuts. Jour. Econ. Ent. 38(4): 446^48, Aug. 

61. 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 

62. Grayson, J. W. and F. W. Poos. Southern corn 
rootworm as a pest of peanuts. Jour. Econ. Ent. 
40(2): 251-256, illus.. 1947. 

63. 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. 

64. 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, 

65. Poos, F. W. and E. T. Batten. Control of peanut 
pests. Southern Planter 109(6): 16, 1948. 

66. 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. & PI. Quar. 
E-751m, 8 pp., July 1948. 

67. Poos. F. W. and Nancy H. Wheeler. Some ad- 
ditional host plants of three species of leafhoppers 



of the genus Empoasca. Proc. Enl. Soc. Wash. 
51(1); 35-38, 1949. 

68. 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. 

69. Poos, F. W. Insects injurious to alfalfa. Report 
of the 1 1 th Alfalfa Improvement Conference: 75, 

70. 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. & PI. Quar. E-793, 12 pp., January 

71. 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 

72. Ely, R. E., L. A. Moore, R. H. 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. 

73. Moore, L. A., R. H. Carter, and F. W. Poos. 
Insecticide studies with dairy cattle. Milk and 
Food Technol. Jour. 12(2): 103-104, March-Apnl 

74. Poos, F. W., T. N. Dobbins, E. T. Batten, and 

G. M. Boush. Tests with benzene hexachloride 
for the control of insects attacking peanuts. 1 946- 
1949. U.S.D.A. Bur. Ent. & PI. Quar. E-820, 16 
pp.. May 1951. 

75. Poos, F. W. Control of the garden webworms in 
alfalfa. U.S.D.A. Leaflet 304, July 1951. 

76. 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 com rootworm on peanuts. U.S.D.A. 
Bur, Ent. & PI. 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. 

79. 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. 

80. Poos, F. W. Relative importance of various in- 
sects harmful to alfalfa in Eastern Canada and 
the United Stales. Dept. of the 13th .Mfalfa Im- 
provement Conference, 1952: 34-36. 



8 1 . . Get that grasshopper! The Windrow 2(4): 

4-5, September 1952. 

82. Poos, F. W. and T. L. Bissell. The alfalfa weevil 
in Maryland. Jour. Econ. Ent. 46(1): 178, May 

83. 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. 

84. Poos, F. W. and W. L. Howe. Control of the 
southern corn rootworm on peanuts. LISDA Bur. 
Ent. and PI. Quar. EC-23, 7 pp., April 1953. 

85. 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. 

86. Poos, F. W. The meadow spitllebug— How to 
control it. U.S.D.A. Leaflet 341, July 1953. 

87. 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. 

88. Poos, F. W. Insects as related to seed production. 
South. Pasture and Forage Crop Impr. Conf Rpt. 
10: 36-38, 1953. 

89. 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. 

90. 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. 

91. 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. 

92. Beattie, J. H., F. W. Poos, and B. B. Higgins. 
Growing peanuts. LISDA Farmers" Bull. 2063, 
54 p.. May 1954. 

93. 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. 

94. 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. 

95. Poos, F. W., J. L. Allsion, and K. W. Kreitlow. 
The clover root borer (Hrlastirnis ohsciinis) as a 
vector of southern and northern anlhracnoses of 



red clover. Plant Dis. Rptr. 39: 183, Feb. 15, 

96. Yeomans, A. H. and F. W. Poos. Spray distri- 
bution with boom sprayers. U.S.D.A., ARS-33- 
8, 7 p.. June 1955. 

97. Poos, F. W. Studies of certain species of Chae- 
locncma. J. Econ. Ent. 48: 555-563, Oct. 1955. 

98. Poos, F. W. Some forage insect problems in the 
western United States, .loint Comt. Grassland 
Farming Proc. 1955: 41^3, 

99. Poos, F. W. Citation of merit. Bull. Ent. Soc, 
Amer. 9(4): 272, Dec. 1963. 

100. App, B, A,, J. McGuire, and F. W. Poos. Francis 
Marion Wadley 1 892-1 969. Proc. Em. Soc. Wash. 
72(2): 270-271, June 1970. 

101. Poos, F. W. Walter Harrison Larrimer, 1889- 
1970. J. Econ. Ent. 64(1): 345, Feb. 15. 1971. 

102. Vance, A. M., A. S. Hoyt, and F. W. Poos. Clyde 
Monroe Packard, 1 889- 1 97 1 . J. Econ. Ent. 65(5): 
1531, Oct. 16, 1972. 

91(2), 1989, pp. 304-306 


Donald Joyce Borror 

With extreme sadness I learned of the 
death of Dr. Donald i. 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 Shcp- 
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 1 930 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 
genera! 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 (195 5). Assistant Managing 
Editor of the Entomological Society of 
.America Annals ( 1 943^4), 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 



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 im- 
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- 
ciass 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- 

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. 

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'/: years that it 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 
15,1 988, the insect guide has sold 4 1 2,790 
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- 



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. Horror'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. 

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. 
Serv., USD A, % 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 1 988 

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 
Teny 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: 


Submitted by Geoffrey B. White, Member- 
ship Chairman, 944th Regular Meeting, 1 
December 1988. 

91(2), 1989, pp. 307-310 

Society Meetings 

941st Regular Meeting-May 5, 1988 

The 941st 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 Ihe 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 virginclla 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, ail subsequent specimens have been 
taken on Sulawesi by Project Wallace staff. 
Though a member of the clothes moth fam- 
ily Tineidae, Ischnuridea 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 affiliation, 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 is 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- 

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 



summer recess. In happy anticipation of this 
repast, the meeting was adjourned at 9:10 

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 

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 

Reed Biosystematics LInit, Smithsonian In- 

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 sensii "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. Pales 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. Pales 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 
1 9 September, Pales was at Plum Point, Cal- 
vert County, Maryland, where he collected 
a single Danaus ple.xippus (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 

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- 



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 

Richard G. Robbins, Recording Secre- 
tary, Geoffrey B. White. Membership Chair- 

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- 

ner, Jr.; Membership Chairman, Geoffrey 
B. White; Custodian, Anne M. Wicber. 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 

Gaye Williams exhibited elegant copper 
wire sculptures of a mosquito and a scor- 
pion, both crafted by the Kentucky artist 
Ken Cams. 

R. G. Robbins projected a series of color 
slides showing males and females of 10 Af- 
rican tick species in the genus A)nhlyomn]a 
that are known to be capable of transmitting 
heartwater (Coiffl'/va ruminantiuni), 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 in this pantropical 
family of some 1 200 species. Limacodid lar- 
vae are ontogenetically remarkable: emerg- 
ing from scale-like eggs, they arc polypha- 
gous but prefer late season, tannin-enriched 



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- 

Visitors were introduced and the meeting 
was adjourned at 9:20 p.m. Refreshments 

Richard G. Robbins, Recording Secretary 

Reports of Officers 

Treasurer's Report 

Assets: November 1. 1987 
Total Receipts for 1988 
Total Disbursements for 1988 
Assets: October 31, 1988 




















Net Changes in Funds 
Norman E. Woodley, Treasurer 

% 7J4132 



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. KJngsolver, Corresponding Sec- 

Editor'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- 

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 

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- 

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 

Bcchtel, 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 



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 
Brushwcin, J. R. 1987 Florida 
Bucno-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 
Coumeya, P. 1986 Texas 
Courtney, G. W. 1985 CANADA 
Covell. C. v.. Jr. 1986 Kentucky 
Craig, G. B. 1954 Indiana 
Cross, H. F. 1954 Georgia 

Curcic, B. P. M. 1982 YUGOSLAVIA 
Currado, 1. 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- 
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 
Dnimmond. 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 
Eisenbcrg, 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 
Fisk. F. W. 1968 (1988) Florida 
Flint, O. S.. Jr. 1961 District of Columbia 
Fluno, J. A. 1957 Florida 
Foote, B. A. 1958 Ohio 



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 Horida 
Gerberg, E. J. 1953 Maryland 
Ghorpadc, 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- 
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. 1 983 UNITED KINGDOM 

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 

Hcraty, J. M. 1986 Texas 

Hespenheide. H. A.. Ill 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 

Kasler, C. H. 1979 Michigan 

Keirans, J. E. 1984 District of Columbia 

Kelley, R. W. 1982 South Carolina 

Kennedy, J. H. 1977 Texas 

Kcthley, J. B. 1874 Illinois 

Kim, K. C. 1983 Pennsylvania 

Kingsolver, J. M. 1963 Maryland 

Kirchncr, R. F. 1981 West Virginia 

Kitayama, C. 1974 California 



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- 
Kurczewski, F. E. 1970 New York 
Lago, P. K. 1984 Mississippi 
Lamb, N. .1. 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 
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 Man land 
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 
Afason, 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 

McMurtry, J. A. 1986 California 
McPherson, J. E. 1985 IlHnois 
Mead, F. W. 1976 Florida 
Mendez, E. 1976 Florida 
Menke, A. S. 1969 District of Columbia 
Messersmith, D. H. 1965 Maryland 
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 
Miller, W. E. 1983 Minnesota 
Mitchell, R. T. 1949 (1978) Maryland 
Mitter, C. 1987 Maryland 
Mockford, E. L. 1955 Illinois 
Molineaux, M. J. 1986 Maryland 
Moore, T. E. 1950 Michigan 
Moron, M. A. 1985 MEXICO 
Morse, J. C. 1976 South Carolina 
Moser, J. C. 1973 Louisiana 
Moulton, S. R., II 1988 Maryland 
Muegge, M. A. 1988 Tennessee 
Munroe. E. G. 1986 CANADA 
Miinson, S. C. 1938 (1976) Maryland 
Murdoch, W. P. 1966 Pennsylvania 
Nakahara. S. 1968 Mar\'land 
Neal, J. W., Jr. 1982* Maryland 
Neff, S. E. 1969 Pennsylvania 
Nelson, C. H. 1969 Tennessee 
Nelson. G. H. 1949 California 
Nelson, R. H. 1933 Pennsylvania 
Neunzig, H. H. 1956 North Carolina 
Newhouse, P. 1986 Maryland 
Newkirk, R. A. 1968 Maryland 
Ng, Y.-S. 1986 Virginia 



Nichols. S. W. 1985 Iowa 

Nielsen, G. R. 1983 Vermont 

Norrbom, A. L. 1983 District of Columbia 

Norton, R. A. 1978 New York 

Nuhn, T. P. 1981 Virginia 

Nutting, W. H. 1973 California 

Oatman, E. R. 1980 California 

O'Brien, M. F. 1979 New York 

Oman, P. W. 1930 Oregon 

Orth. R. E. 1981 California 

Osbrmk, W. L. A. 1984 Florida 

Oswald, J. D. 1987 New York 

Pagac, B. P., Jr. 1984 Maryland 

Paggi. A. C. 1987 ARGENTINA 

Pakaluk, J. 1981 Kansas 

Palacios- Vargas. J. G. 1980 MEXICO 

Palmer, W. A. 1985 Te.xas 

Parker, C. R. 1977 Tennessee 

Parker, H. L. before 1918 (1962) FRANCE 

Parrish. D. W. 1963 (1987) Maryland 

Parsons, M. 1963 Ohio 

Pavuk, D. M. 1987 Ohio 

Pena-Guzman, L. E. 1980 CHILE 

Penny, N. D. 1982 California 

Petersen. J. L. 1981 Florida 

Peterson, R. V. 1952 District of Columbia 

Peyton, E. L. 1968 District of Columbia 

Phillips. W. G. 1955 Maryland 

Pinto. J. D. 1982 California 

Plakidas, J. D. 1986 Maryland 

Piatt. A. P. 1985 Maryland 

Pogue, M. G. 1980 District of Columbia 

Polhemus, J. T. 1964 Colorado 

Porter. C. H. 1984 Georgia 

Pratt. G. K. 1974 California 

Pratt, H. D. 1943 Georgia 

Price. P. W. 1986 Arizona 

Price. R. D. 1963 Minnesota 

Pulawski, W. J. 1975 California 

Purrington. F. F. 1987 Ohio 

QuickC. D. L. J. 1987 ENGLAND 

Rack. G. 1975 WEST GERMANY 

Rainwater. C. F. 1954 (1975) Maryland 

Rainwater. H. I. 1964 (1983) Maryland 

Ramos, J. A. 1947 PUERTO RICO 

Ramsay, M. J. 1968 Maryland 

Rathman, R. J. 1988 Arizona 

Raupp, M. J. 1982 Maryland 

Rawlins. J. E. 1974 Pennsylvania 
Reed, W. D. 1931 District of Columbia 
Reichart. C. V. 1946 Rhode Island 
Richardson. H. H. 1939 (1976) New Jersey 
Riddick, E. W. 1988 District of Columbia 
Riegel, G. T. 1952 Illinois 
Riley. D. R. 1984 Texas 
Robbins. R.G. 1979* Maryland 
Robbins, R. K. 1986 District of Columbia 
Robinson. H. 1963 District of Columbia 
Robinson, W. H. 1975 Virginia 
Rock, E. 1984 Ohio 
Rogers. T. E. 1988 Louisiana 
Rolston, L. H. 1973 Louisiana 
Root, R. B. 1984 New York 
Rosenheim. J. A. 1987 Hawaii 
Rosenthal. S. S. 1988 Montana 
Ross. E. E. 1983 California 
Ross, M. H. 1981 Virginia 
Roth. L. M. 1944 Massachusetts 
Rozen. J. G.. Jr. 1956 New York 
Ruiter. D. 1976 Colorado 
Rush, P. A. 1984 Minnesota 
RUSSELL, L. M. 1930 (1988) Maryland 
Rust. M. K. 1984 California 
SABROSKY, C. W. 1946 (1988) New Jer- 
sey (Honorary President) 
Sakimura, K. 1982 Hawaii 
Sands, D. P. A. 1984 AUSTRALIA 
Santana, F. J. 1966 Florida 
Saugstad. E. S. 1979 Maryland 
Scarbrough. A. G. 1971 Maryland 
Schaeffer. C. W. 1985 Connecticut 
Schaffner. J. C. 1984 Texas 
Scharf W. C. 1981 Michigan 
Schauff, M. E. 1980 District of Columbia 
Schmidt. C. H. 1969 North Dakota 
Schulte. J. 1987 Michigan 
Schwan, T. G. 1980 Montana 
Schwartz, M. D. 1985 New York 
Scudder, G. G. E. 1984 CANADA 
Sedlacek, J. D. 1988 Kentucky 
Sedman, Y. S. 1951 Illinois 
Selander. R. B. 1954 Illinois 
Shaffer, J. C. 1974 Virginia 
Shands, W. A. 1940 South Carolina 
ShefTcr, B. J. 1986 Alabama 
Shepard. H. H. 1927 Virginia 



Shewell, G. E. 1949 CANADA 
Shinohara, A. 1981 JAPAN 
Shockley. C. W. 1955 California 
Sholes, O. D. V. 1979 Massachusetts 
Shorthouse, J.D. 1986 CANADA 
Slater, J. A. 1949 Connecticut 
Sleeper, E. L. 1976 California 
Sloan, M. J. 1983 District of Columbia 
Smiley, R. L. 1964 Maryland 
Smith, C. F. 1967 (1986) North Carolina 
Smith, D. R. 1965* District of Columbia 
Solis, M. A. 1985 District of Columbia 
Sollers-Riedel. H. 1 938* District of Colum- 
Spangler, P. J. 1958* District of Columbia 
Spence. J. R. 1986 CANADA 
Spilman. R. E. W. 1950 (1977) Maryland 
Spilman, T. J. 1950 District of Columbia 
Spinelli, G. R. 1983 ARGENTINA 
Spofford, M. G. 1986 New York 
Staines, C. L. 1975 Maryland 
Starr, C. K. 1987 District of Columbia 
Stasny, T. A. H. 1986 West Virginia 
Steck, G. J. 1988 Mainland 
Stcffan, W. A. 1970 Idaho 
Stegmaier, C. E., Jr. 1965 Florida 
Steiner, W. E., Jr. 1979 Maryland 
Steinly, B. A. 1983 Illinois 
Stewart, R. D. 1985 Maryland 
Steyskal, G. C. 1947 District of Columbia 
Stoetzel, M. B. 1971 Maryland 
STONE, A. 1931* (1988) Maryland 
Stonedahl. G. M. 1982 New York 
Stribling, J. B. 1983 Virginia 
Strickman. D. 1987 District of Columbia 
Surdick, R. F. 1979 Virginia 
Sutherland, C. M. 1974 New Mexico 
Sutherland, D. W. S. 1973* Maryland 
Taft, S. J. 1979 Wisconsin 
Tennessen, K. J. 1982 Alabama 
Thomas, D. B., Jr. 1983 Texas 
Thompson. F. C. 1968* District of Colum- 
Thompson, J. V. 1953 (1985) New Jersey 
Tibbetts, T. 1955 Utah 
Togashi, K. 1983 JAPAN 
Townes,, G. F. 1956 South Carolina 
Townes, H. K. 1941 Florida 

Traub, R. 1947 Maryland 
Triplehorn, B. W. 1985 Virginia 
Triplehorn, C. A. 1972 Ohio 
Trumble, J. T. 1979 California 
Turner, C. E. 1985 California 
Turner, W. J. 1982 Washington 
Tyson, W. H. 1970 California 
Ulrich, H. 1978* WEST GERMANY 
Utmar, J. A. 1974* Maryland 
Valley, K. 1976 Pennsylvania 
Vasquez, A. 1957 Virginia 
Venables, B. A. B. 1983 Maryland 
Villegas, B. 1977 California 
Voegtlin, D. 1981 Illinois 
Vogt, G. B. 1947 Mississippi 
Wagner. M. R. 1986 Arizona 
Wahl. D. B. 1987 Florida 
Waldbauer, G. P. 1983 Illinois 
Wallenmaier, T. E. 1979 Maryland 
Waller, D. A. 1984 Virginia 
Wallis, R. C. 1948 Connecticut 
Walton, M. 1937 Maryland 
Waltz, R. D. 1988 Indiana 
Ward, R. A. 1975 District of Columbia 
Webb, D. W. 1981 IlHnois 
Webb, R. E. 1967 Maryland 
Ucbcr, N. A. 1941 (1981) Florida 
Weems, H. V., Jr. 1953 Florida 
Weinstein, P. J. 1986 Maryland 
Wendleton, D. S. 1965 Pennsylvania 
Wenzel, R. L. 1984 Illinois 
Werner, F. 1948 Arizona 
Wharton. B. 1981 Texas 
Wheeler, A. G.. Jr. 1974 Pennsylvania 
Wheeler, G. C. 1949 Florida 
Whitcomb. R. F. 1966 Maryland 
White. G. B. 1977 Maryland 
White. R. E. 1966 District of Columbia 
White, T. R. 1979 Georgia 
Whitehead. D. R. 1974 District of Colum- 
Whitsel, R. H. 1967 California 
Wiegmann. B. M. 1988 Maryland 
Wilkerson. R. C. 1988 Maryland 
Williams, G. L. 1984 Maryland 
Williams, H. B. 1977 District of Columbia 
Williams, M. L. 1971 Alabama 
Wills, W. 1977 California 



Wilson, N. 1957 Iowa 

Wirth, W. W. 1945 Florida 

Wojtowicz. J. A. 1981 Tennessee 

Woke, P. A. 1936 (1976) Maryland 

Wood, D. M. 1987 CANADA 

Wood, F. E. 1968 Maryland 

Wood, T. K. 1974 Delaware 

Woodley, N. E. 1984* District of Columbia 

Wooley, J. B. 1986 Texas 

Yonke, T. R. 1971 Missouri 

Young, A. M. 1983 Wisconsin 

Young. D. A. 1950 (1983) North Carolina 
Young, D. K. 1981 Wisconsin 
Zack, R. 1983 Washington 
Zeigler, D. D. 1986 Texas 
Zenner-Polania, I. 1977 COLOMBIA 
Zimmerman, E. C. 1965 AUSTRALIA 
Zolnerowich, G. 1987 Texas 
Zuccaro, A. E., .Ir. 1986 Mississippi 
Zungoli, P. A. 1978 South Carolina 

91(2), 1989, pp. 318-320 

Instruction to Authors for 
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Typing the Manuscript 

Type manuscripts on bond paper with 
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Submittal of the Manuscript 

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Names of Organisms 

The first mention of a plant or animal 
should include the full scientific name with 
the author of a zoological name unabbrevi- 
ated. Use only common names approved in 
Common Names of Insects and Related Or- 
ganisms (1983 Revision) published by the 
Entomological Society of America. Do not 
abbreviate a generic name when it begins a 
sentence: e.g. use ""Xus alhus is . . ." rather 
than "X. albus is . . ." 


When citations are made in the text, a 
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Keep the number of tables to a minimum. 
Prepare them as a line drawing or type them 
on a separate page with the caption at the 
top and footnotes below. 



Return Proof Co: 
John R. Doe 
315 State St. 
Merlden, CT 06&20 
(203) 555-1212 

A new species of Xus (Order: Family) 
injurious to hollies. Ilex spp. (Aquifollai 

John B. Doe and John Smith 

(JRD) Resident Biologist. 315 State St., Merlden, Connei 
(JS) Entomologist, City Parks, Hartford, Connecticut 065iO. 

Xus albus . a new species of . . Is described. Illustrated, 

and compared with ... 

t^ey Words ■ Distribution, ornamental shrub, damage, leaf rolle 

Figure Legends 
Figs. 1-A. Xus albus - 1, Habitus. 2, Male genitalia (lateral vleu). 
3. Larva. ^ , Pupa. 

Fig. 5. Damage to holly leaves. 

Literature Cited 
Doe. 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 

(Ilex spp.)- Proc. Entomol. Soc. Va. 38: 54-68. 
(The above citations are fictitious.) 

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[Continued fram 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: 
Blaltaria: Blattellidae) 206 

SHAFFER, J. C— Two new phycitine moths of the genus Coenochroa (Lepidoptera: Pyrahdae) 

from Brazil 237 

SHAFFER. J. C. andE. MUNROE— Type material of four African species of A^ora/r/ia Meyrick, 
with designations of lectotypes and changes in synonymy (Lepidoptera: Crambidae: Pyr- 
austinae) 248 

YOUNG, D. C, B. C. KONDRATIEFF, and R. F. KJRCHNER- Description of male Ostro- 

cerca Ricker (Plecoptera: Nemouridae) using the scanning electron microscope 257 


APPEL, A. G. — Rapid and non-destructive gender determination of nymphal and adult Cryp- 

locercus puncluhtus 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. hernardi Vachon (Neobisiidae: Pseudoscorpiones: Arachnida) 289 

ROBBINS. R. G.— Ticks of the subgenus Ixodiopsis: First report of Ixodes woodi from man and 

remarks on Ixodes holdennedi. a new junior synonym of Ixodes ochotonae (Acari: Ixodidae) 29 1 


DAVIS, D. K.- Lepidoptera Anatomy 293 

GAGNE, R. ]. — Asa Fitch and the Emergence of American Entomology 294 

McCAFFERTY, W. P. -The Mayflies of Florida. Revised Edition 296 


WALLENMAIER, T. E.-Frederick William Poos, Jr. 1891-1987 298 

WHITE, R. E.-Donald Joyce Borror 1907-1988 304 





VOL. 91 

JULY 1989 

NO. 3 

(ISSN 0013-8797) 


of the 





FORATTINI, O. P. andM. A. M. SALLUM — Redescription oi Culex (Melanoconion) delpontei 

Duret, 1968 and Cx. (Mel.) pereyrai Duret, 1967, from southern Brazil 473 

H USBAND, R. W. — Two new species of Coccipolipus (Acari: Podapolipidae) parasites of Chilo- 
conis spp. (Coccinellidae) from Vera Cruz and Morelos, Mexico and Florida and Wisconsin, 

U.S.A 429 

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- 
chidae) 468 

LEE, K. M., W. W. WIRTH, and K. L. CHAN— A new species of Dasyhelea midge reared from 

drains in Singapore (Diptera: Ceratopogonidae) 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 Aleurolulus (Homoptera: Aleyrodidae) 436 

NEUNZIG, H. H. — New records of Phycitinae from Bolivia including a new species of Peadus 

(Lepidoptera; Pyralidae) 355 

NEUNZIG, H. H. and N. A. LEIDY— A new species of Dioryctria (Lepidoptera: Pyralidae: 

Phycitinae) from the southeastern United States 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 

of ovoviviparity in Blattaria (Dictyoptera) 44 1 

SCHWARTZ, M. D.—Polymends castilleja. a new mirine plant bug from California and Oregon, 

with remarks on generic characters of the genus Polymeries Hahn (Heteroptera: Miridae) ... 46 1 

SHAFFER. J. C — Review of Goya Ragonot and description of a new species, G. simulala from 

Parana, Brazil (Lepidoptera: Pyralidae: Peoriinae) 398 

SHAFFER, J. C. and E. MUNROE — Type material of two African species of Herpelogramma 

and one of Pleurnpiya (Lepidoptera: Crambidae: Pyraustinae) 414 

SPANGLER, P. J.— A new species of Neotropical water bug, Paravelia biae, from Brazil (Het- 
eroptera: Veliidae) 360 

(Continued on back cover) 




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91(3). 1989, pp. 321-324 




H. H. Neunzig and Nancy Antoine Leidy 

Department of Entomology, North Carolina State University, Raleigh, North Carolina 

Abstract.— Dioryctria taedivorella. a new species of coneworm, is described, illustrated 
and compared with Dioryctria nierkeli Mutuura and Munroe. 

Key Words: Dioryctria. coneworm, distribution, loblolly pine 

Hedlin et al. ( 1 98 1 ), in their investigation 
of cone and seed insects of North America, 
included nine species of Dioryctria Zeiler 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 nierkeli. Having compared the type of 
D. nierkeli. on loan from the Canadian Na- 
tional Collection, with North Carolina spec- 
imens and other Dioryctria that are sup- 
posedly nierkeli. we concluded: ( 1 ) the name 
inerkeli should be restricted to certain pop- 
ulations of Dioryctria mainly feeding as lar- 
vae in slash pine (Finns elliotii Engelmann) 

in northern Florida, southern Georgia and 
southern Mississippi, and (2) the similar, 
more northern and more widespread species, 
associated with loblolly pine, is new. 

Dioryctria taedivorella, Neunzig and 

Leidy, New Species 

Figs. 1-6 

Diagnosis. —D/onr?/76( taedivorella is 
most similar to Dioryctria nierkeli. D. tae- 
divorella. however, is a darker species with 
the antemedial and postmedial lines less 
distinct. These lines are chiefly white in D. 
nierkeli and mostly gray in D. taedivorella. 
Also, the dorsum of the thorax of D. tae- 
divorella is mostly brown to reddish brown, 
whereas the dorsum of the thorax ofD. nier- 
keli is mostly pale gray with few or no brown 
or reddish brown scales. 

Description. — //faf/.' 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 



Figs. 1-6. Dioryclria laedivorella. 3.5x I. 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. Forcning: 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- 
tipped 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) essentiallv like those of Z). mer- 

Type material. — Holotype, <?, USA, N. 
CAROLINA, Lenoir Co., Kinston, US 70 
Byp & Neuse R., 1 -IX- 1983. N. A. Leidy, 
Finns taeda cone, emerg. 13-IX-1983, gen- 
italia slide 895 HHN, in USNM. Paratypes 
25 3, 51 9. USA, N. CAROLINA, Lenoir 
Co., Kinston. US 70 Byp & Neuse R., 28- 
VII- 1 983, 1 4-VIII- 1 983. N. A. Leidy. Finns 
taeda cone, emerg. 8-IX- 1 983. 1 7-IX- 1 983. 
6-X-1983. 12-X-1983. genitalia slides 872, 
883. 893. 903 HHN (1 3, 3 2); USA. N. 
CAROLINA. Robeson Co.. Lumberton. NC 
2 II 4 mi W of 1-95, J. B. Lattay Forest Tree 
Nur.. 27-VII-1983, N. A. Leidy, host; Finns 
taeda cone, emerg. 5-IX-1983, lO-IX-1983, 



V 1 1 1 /;v 

X-l 983. genitalia slides 885, 887. 891 HHN 
(3 3); USA. N. CAROLINA. Robeson Co.. 
nr Lumberton, J. B. Lattay For. Nur., NC 
211 4 mi W of 1-95, 22-VII-1985. N. A. 
Leidy & M. May nor. in Pinus tacda cone, 
emerg. 2-IX-1985 to 1 -X-l 985 (7 3, 8 5); 

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 9); 
USA, N. CAROLINA, Wayne Co.. Golds- 
boro, Claridge State For., SR 1326 1.5 mi 



N of US 70, 15-VII-1985. N. A. Leidy, in 
Piniis taeda cone, emerg. 13-IX-1985 (1 $); 
USA, N. CAROLINA, Onslow Co., nr 
Richlands, NC 24 at SR 1230, 16-VII-1985, 
N. A. Leidy, in Pimis taeda cone, emerg. 
21-IX-1985 (1 d)\ USA, N. CAROLINA. 
Onslow Co., nr Maysville, Hofmann For- 
est, 25-VI-1985, NA Leidy & DJ Lodge, in 
Finns taeda cone, emerg. 1 1 -IX- 1985 (1 S)\ 
USA, S. CAROLINA, Berkeley Co., Fran- 
cis Marion Seed Orchard, col. 23-27-VI- 
1983, 2nd year cones Pinm taeda (1 <J, 2 9); 
USA, GEORGIA, Putnam Co., col. 9-VII- 
1985, emerg. 16-IX-1985, Pimis taeda (1 
2); USA, ALABAMA, Greene Co., Wey- 
erhaeuser Co., col. 1985, Pinus taeda (2 3, 
10 9); USA, ALABAMA, Greene Co., Wey- 
erhaeuser Co., col. 15-IX-86, emerg. 26-IX- 
86, 3-X-86, Pinus taeda (9 9); USA, ALA- 
BAMA, Greene Co., Weyerhaeuser Co., col. 
15-IX-86, Pinus taeda (8 <S, 14 9); USA, 
MISSISSIPPI, Perry Co., Erambert Seed 
Orchard, col. 16-20-VI-1986, 2nd yr. cone 
Pinus taeda (1 9). Paratypes deposited in 

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. zinmierniani. 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 


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. 1 1919 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. 198 L 
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 Hiraiilicos, Mexico. 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- 

Neunzig, H. H., E. D. Cashalt. 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. 

91(3), 1989, pp. 325-341 




Nancy Antoine Leidy and H. H. Neunzig 

Department of Entomology, North Carolina State University, Raleigh, North Carolina 

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. ahietivorella (Grote), D. disclusa Heinrich, 
D. pygmaeella Ragonot, D. clarioralis (Walker), D. amatella (Hulst), and D. taedivorella 
Neunzig and Leidy. 

Key Words: Pyralidae, Phycitinae, Dioryctria, immatures, coneworm, taxonomy 

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- 

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 1 969a, 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 ( 1 97 l),Coulsonetal.'( 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 lar\ae of D. reniciilcllnides Mu- 
tuura and Munroe (as D. rcniculella Grote), 

and briefly diflerentiated it from D. ahieti- 
vorella (Grote) (as D. abietella Denis and 
SchiflTermuUer). 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. ahietivorella (as D. abie- 
tella). Neunzig et al. (1964), gave detailed 
descriptions of six species occurring in the 
southeastern United States: D. amatella 
(Hulst), D. eheli Mutuura and Munroe (as 
D. abietella), D. clarioralis (Walker), D. dis- 
clusa. D. taedivorella Neunzig and Leidy (as 
D. ziinmermani (Grote). and D. pygmaeella 
Ragonot. Schaber (1981) published on the 
larvae of D. taedae Schaber and Wood, 
however, larvae of D. taedivorella and D. 
yalesi 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. eheli. The early-instar lar- 
vae of D. reiiiculelloides were examined by 
Spies and Dimond using SEM (1985). 



Fig. 1. Part of mesothorax including D and SD 
pinacula of D. ilisdiisa. dorsolateral aspect. (ISOx). 
TP, tonofibnllar> platelets; A, area shown enlarged in 
Figs. 22-27. 

In this paper we update the desciiptions 
of the last-instar larvae of si.\ species by 
utilizing the conventional light microscope 
and the scanning electron microscope to 
reexamine the known diagnostic characters 
and to discover additional species differ- 

Materials and meXhods.—Dioryctria lar- 
vae were collected in North Carolina from 
PIuks 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) (Eraser'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 

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 i" 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- 
dri*-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. I). The terminology used for the 
mouthparts follows Crumb (1929) (Eigs. 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 (Eig. 2). The margins of 
these sclerites are not clearly defined in elec- 
tron micrographs but are generally visible 
under the light microscope and consistently 



Fig. 2. Epipharynx of ZJ. amatclla ( 1 80 x ); ESc, epi- 
pharyngeal scleriles; ESh. epipharyngeal shield. 

bare of spines, except occasionally on the 
mesal margin. 

Common names of most species follow 
the Entomological Society of America 
( 1 982). The name fir coneworm for D. abie- 
tivorclla is from Hedlin et al. (1981). Hedlin 
et al. ( 1 98 1 ) 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. elliotii. 
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 

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- 

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 SDl pinaculum form- 
ing ring, dark brown with large pale center. 



Eighth abdominal SDl pinaculum forming 
ring, medium brown with small pale center. 
Setae brown basally, pale distally. Thoracic 
legs light brown laterally, yellowish white 
mcsally. 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: SDl on me- 
sothorax 1.35 mm to 1.73 mm long. Integ- 
ument of mesothorax areolate-rugose ven- 
irade of D1&2 (Fig. 16); aerolate anterior 
to D1&2 (Figs. 16, 22). Abdomen: Seta D2 
of abdommal segment 2 approximately 'A 
10 '/2 dorsoventral diameter of segment. SDl 
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.lO: 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 
Eraser's fir. leaving a mass of frass, webbing, 
and needle fragments at the feeding site 
(Leidy and Neunzig 1986). 

Dioryctria disclusa Heinrich 
(webbing coneworm) 
Figs. 5, 11, 17, 23,29 

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 bufll"; pinacula of meso- 
thorax and metathorax pale brown to buff; 
pinacula of abdomen bufll', 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 SDl pinaculum forming ring, light 
brown with large pale center. Eighth ab- 
dominal SDl 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 



Figs. 4-9. Dinryclria spp. Epipharynx: 4. D. ahielivnrella (150x); 5. D. disclusa (150x); 6. D. pygmaeella 
(150x); 7. D. clanorahs (lOOx); 8. D. amaiella (lOOx); 9. D. laediwrella (lOOx). 



Figs. 10-15. Dioryctna spp. Hypophar> n.\. dorsal aspect: 1 0. D. ahtetivorella (200 x ); 1 1 . D. disclusa (200 x ); 
12. D. pygmaeella (200 x); 13. D. danoralis (150x); 14. £). amatella (150x); 15. D. laedivorella (150x). 



Figs. 16-21. Dioryctna spp. Part of mesothora.\ including D and SD pinacula. dorsolateral aspect: 16. D. 
ahietn-orella; 17. D. disclusa; 18. D. pygmacella; 19. D. clanorahs; 20. D. amaiella; 21. Z?. taedtvorella. (lOOx) 




Figs. 22-27. Dioryctria spp. Integument of mesothorax anterior to D1&2, dorsal aspect (see Fig.l for exact 
location): 22. D. abietivorella: 23. D. disclusa; 24. D. pygmaeella; 25, D. clanoralis: 26. D. amatella; 27. D. 
taedixorella. (1500x) 



Figs. 28-33. Dioryctna spp. Perianal region, eaudal aspect. 28. D. ahiclivorclla: 29. D. disciusa: 30. D. 
pygmueclla: 31. £). danoralis; 32. D. amau-llu; 33. D. laedivorella. (75 x) 



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. 1 1) 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: SDl 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 'A dor- 
soventral diameter of segment. SDl 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 Finns taeda. 

Comments. — The length of seta SDl of 
the mesothorax and abdomen is at consid- 
erable variance from that reported by Neun- 
zig et al. ( 1 964) and figured by Farrier and 
Tauber (1953). It is consistent, however, in 
larvae from several dates and localities col- 

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, 18, 24, 30 

Diorydria 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 SDl pinaculum forming ring, dark 
brown with large pale center. Eighth ab- 
dominal SDl pinaculum forming ring, dark 
brown with large pale center. Setae brown 
basally, pale distally. Thoracic legs dark 



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 premaxillulai^ 
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; mcntum brown, darker an- 
teriomesally; arms of mentum brown. Spin- 
neret slender, approximately 6 x as long 
as median breadth. Thorax: SDl on me- 
sothorax 1.25 mm to 1.60 mm long. Integ- 
ument of mesothorax areolate-rugose ven- 
trad of D1&2 (Fig. 18); tuberculate- 
squamiform anterior to D1&2 (Figs. 18, 24). 
Abdomen: Seta D2 of abdominal segment 2 
approximately 'A the dorsoventral diameter 
of the segment. SDl 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.lO: 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, 17-VIII-1978, L. Grimes; Bladen 
Co., Singletary Lake State Park (collected 
by permit) 23-VII-i985, N. A. Leidy. All 
larvae collected from Taxndiuin disticlniin. 

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- 

all length of 11.1 mm to 1 1.8 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, 13, 19, 25, 31 

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 loss 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 SDl 
pinaculum forming incomplete ring, pale 
yellow (brown) with large pale center. Eighth 
abdominal SDl pinaculum forming ring, 



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: SDl 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 D1&2 
(Figs. 19, 25). Abdomen: Seta D2 of abdom- 
inal segment 2 approximately '/y dorsoven- 
tral diameter of segment. SDl 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- 
5 1 . Perianal region (Fig. 3 1 ) with coarse pale 
(medium brown) spines. 

Material examined.— Two larvae. USA 
N. Carolma: Wake Co., 27-VI-1986, N. A. 
Leidy; Wayne Co., Goldsboro, 14-1-1987, 
J. A. Smith. Larvae collected from Pinus 

Comments.— Only two larvae of Z). clar- 
ioralis were obtained. One larva was reared 
by placing a gravid light-trapped female in 
a sleeve cage over a branch off. 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- 

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. 

Dioryctha 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 



brown. Anal shield pale to medium yellow- 
ish brown. Mesothoracic SDl pinaculum 
forming ring, dark brown with minute pale 
center. Eighth abdominal SDl 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. Hypopharyn.x (Fig. 14) with pre- 
maxilluiary 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 5 x as long as me- 
dian breadth. Thorax: SDl 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. SDl of segment 8, 0.77 
mm to 0.97 mm long. Crochets of abdom- 
inal prolcgs 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. 

Material examined.— Sixteen larvae. USA 
N. Carolina: Richmond Co., Rockingham, 
4-VIII-1985, 6-VI1I-1986, N. A. Leidy; 
Moore Co.. Southern Pines, 4-VIII-I985, 
14-VIII-1983, N. A. Leidy; Onslow Co., 
Richlands, 16-VII-1985, N. A. Leidy. Lar- 
\ae collected primarily from Pinus palus- 
iris. a few collected from P. taeda. 

Comments. — Although the perianal 
spines of D. aniatella (Fig. 32) are in size 
actually very similar to those of D. taedi- 
xorclla (Fig. 33), they appear more slender 
when viewed with the light microscope Jdc- 

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 A taeda were not observed to leave 
masses of pitch at the entr> hole. 

Dioryctria laedivorella Neunzig and Leidy 

(loblolly pine coneworm) 

Figs. 9. 15, 21, 27, 33 

Dioryctria laedivorella 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 margmally. Anal shield yellowish brown 



to reddish brown mesally, reddish brown to 
dark brown laterally. Mesothoracic SDl 
pinaculum forming ring dark brown with 
minute, pale center. Eighth abdominal SDl 
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 x as long 
as median breadth. Thorax: SDl on me- 
sothorax 0.62 mm to 0.87 mm long. Integ- 
ument of mesothorax tuberculate-rugose 
ventrad of D 1 &2 (Fig. 2 1 ); spinose anterior 
to D1&2 (Figs. 21, 27). Abdomen: Seta D2 
of abdominal segment 2 approximately 'i 
dorsoventral diameter of segment. SDl of 
segment 8, 0.70 mm to 0.82 mm long. Cro- 
chets of abdominal prolegs mostly biordi- 
nal, arranged in a 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-VI1-1985. N. A. Leidy; 
Granville Co., Lewis, 18-VII-1985. N. A. 
Leidy; Onslow Co., Richlands, 1 6-VII- 1985, 
N. A. Leidy; Robeson Co., Lumberton, 27- 
VII- 1983, N. A. Leidy; Lenoir Co., Kinston. 
14-VII-1985, N. A. Leidy; Robeson Co., 
Lumberton, 22-VII- 1985, Leidy & Maynor; 
Robeson Co., Lumberton, 9-VII- 1 986, Lei- 

dy & Hardin. Larvae collected from Pinus 

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. disci iisa (Fig. 5) were easily observed 
using SEM, but difficult to see with the light 
microscope, because they lack strong pig- 
mentation. Neunzig et al. ( 1 964), 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. ( 1 964) as "lacking distinct spines"; how- 
ever, SEM showed considerable spinulation 
(Fig. 30). Additionally, Neunzig et al. ( 1 964) 
separated D. amatella (Fig. 32) and D. tae- 
divorella (Fig. 33) (as zinimcrmani) 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. 



8), however, with only a few spines in the 
general area, there is no real indication of a 

The tonofibrillary platelets of the meso- 
thorax also appeared quite dissimilar de- 
pending on the method of observation. 
Those of D. laedivorclla (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- 
ciusa (Fig. 1 7) 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. tacdiwreUa 
(Fig. 27) is spinose, whereas it ranges from 
areolate in D. ahietivorclla (Fig. 22) and D. 
clarioralis (Fig. 25) to squamiform in D. 
discliisa (Fig. 23) and tuberculate-squami- 
form in D. pygmaeclla (Fig. 24). 

Mutuuraand Munroe(1972, 1974) placed 
the adults of Dioiyctria 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. ahietivorella\ au- 
ranticella group: D. disciiisa; baumhoferi 
group: D. pygmaeella; and zimmermani 
group: D. amatella. D. clarioralis and D. 
laedivorella 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 SDl 
pinaculum on the eighth abdominal seg- 

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. ahietivorclla (Fig. 22) and 
D. disclusa (Fig. 23). In addition, the pig- 
mentation of the Dl 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. 
pyginaeella has been placed in the baum- 
hoferi group. Mutuura and Munroe (1972) 
noted, however, that their placement of D. 
pygfiiaeela was tentative because of differ- 
ences in the male genitalia. Examination of 
the larval characters also casts doubt on the 
placement of D. pyginaeella 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- 
inaeella 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- 
inaeella. Although larvae of D. caesirufella 
were not available for this study, the dis- 
similarity of the larvae of D. pyginaeella 
and D. clarioralis suggest that D. pygmaecl- 
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 
North American Species of Dioryctria 

1. Tonofibrillary platelets of abdomen pale, dif- 
fering in texture from, or only slightly darker 
than, integument; pinaculum SDl of 8th ab- 
dominal segment forming ring with pale cen- 
ter; integument not spinose anterior to DI&2 

of mesothorax (Figs. 16-19, 22-25) 2 
- Tonofibrillary platelets of abdomen black, 
shiny; pinaculum SDl 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 

2. Pmacula Dl & D2 of abdomen brown, at least 
at base of setae; seta D2 of abdominal seg- 
ments long, 'A to Vi dorsoventral diameter of 
segment 3 



- Pinacula Dl & D2 of abdomen usually pale, 
appearing lighter than surrounding integu- 
ment; seta D2 of abdominal segments short, '/, 

to 'A dorsoventral diameter of segment 4 

3. Tonofibnilary platelets yellowish white, shiny, 
distinct from integument; pinaculum SDl of 
8th abdominal segment with small pale center; 
labrum with distal margin distinctly indented 
(Fig. 4); overall length approximately 20 mm 

- Tonofibrillary platelets indistinct; pinaculum 
SDl of 8lh abdominal segment with large pale 
center; labrum with distal margin weakly in- 
dented (Fig. 6); overall length approximately 

1 3 mm pygmaeella 

4. Pinaculum SDl of 8th abdominal segment with 
small pale center; pinaculum SDl of meso- 
thorax forming complete ring; seta D2 of ab- 
dominal segments approximately 'A dorsoven- 
tral diameter of segment; lingua and gorge of 
mcntum bare (Fig. 11) disclusa 

- Pinaculum SDl of 8th abdominal segment with 
large pale center; pmaculum SDl of meso- 
Ihorax forming incomplete ring; seta D2 of ab- 
dommal segments approximately '/, dorsoven- 
tral diameter of segment; lingua and gorge of 
mentum spinose (Fig. 13) clanoralis 

5. Penanal 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) tacdimrella 


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., 

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. 1 1471 
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. 

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91(3), 1989, pp. 342-345 


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: Chrysomelidae, Hispinae, beetle, Uroplatini 

The Neotropical Hispinae arc 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 (19 11). 
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- 

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. Eh tral 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 

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. Sciitellum: quadrate; micro- 
punctate. Elytron: 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- 


Fig. 1 . Elytra of Fossispa lutena. 



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 gsnus.—Fossispa lii- 
tena. new species. 

E\ymo\ogy. — Fossispa, fossa = ditch or 
trench plus ispa, Latin. The gender is neuter. 

Comparative no\Q%. — 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: 




vertex of 

not sulcate 







segments 1- 

thick, 1 not 

6 thin, 7-8 




than 7 


highly arched; 

not arched; no 








longer than 

about as long 


as wide 


three entire 

three entire 


plus one 

costa 2 does 


not unite 


with 1 & 3 

costa 2 
unites with 
1 &3 

Key to the Genera of Uroplatini 
WITH Clavate Antennae 

Antenna 7-segmented; last four segments 
combined not longer than the three preceding 


Antenna 7- or 8-segmented; last four seg- 
ments combined longer than the three pre- 
ceding combined 

2. Elytral costae irregular, tuberculate 

Physocuryna Guerin 

- Elytral costae regular Corynispa Uhmann 

3. First 5 antennal segments similar; segment 6 
widened; club as long as 4 precedmg segments 
combined; antenna 7-segmented Bruchia Weise 

- Antenna not as above 4 

4. Elytral costae irregular, tuberculate 5 

- Elytral costae regular 6 

5. Antennal segments 7 and 8 almost fused; seg- 
ments 1 to 6 thick icntispa Uhmann 

- Antennal segments 7 and 8 distinct 
Oclotonia Chevrolat (m part) 

6. Third tarsal segment not obviously bilobed 
Slenopodius Horn 

- Third tarsal segment obviously bilobed 7 

7. Each elytron on apical third with 8 rows 

of punctures 8 

- Each elytron on apical third with 10 rows of 
punctures 9 

8. Antennal segment 7 as wide as 8 

Ocloloina Chevrolat (in part) 

- Antennal segment 7 narrower than 8 
Parvispa LIhmann 

9. Base ofeach elytron with 10 rows of punctures 
Brachycoryna Guerin 

- Base ofeach elytron with 8 rows of punctures 

10. Expanded puncture rows on apical half of ely- 
tra not divided by a costa; vertex of head not 

sulcate Hcpialomispa 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; V!II expanded, hir- 
sute, rounded at apex; I-III glabrous; IV- 
VI with some setae; mouthparts ventrally 
directed. Pwnotinn: wider than long; com- 
pletely margined at sides; lateral margins 
parallel at apical -A, 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 






100 95 




75 70 











r \ ^ 





"--— J 
















X / 



2 » 





















'^ K X 




""""■"^ ^ 







r^ ^ 








\ Or 








• s 10 



Fig. 2. Distribution of Fossispa lutena. 

1.3 mm (avg. 1.1) (n = 10); length 0.6 to 
0.8 (avg. 0.7). ScuteUum: brown; quadrate; 
longer than wide; micropunctate. Elytron: 
(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, 'A length of IV, pad of setae 
beneath; IV long, widening at apex; claws 
divergent. Venter: black, prostemum punc- 
tate in middle, rugose at sides, truncate at 
apex; mesosternum smooth, shining in mid- 
dle, rugose at sides; several punctures around 
coxae; metastemum smooth, shining in 
middle, rugose at sides; abdominal sterna 
smooth in middle, rugose at sides, all with 
scattered setae. Total length: 2.6 to 3.4 mm 
(avg. 3.0). 

Type dsiia.-Holotype: MEXICO: Vera 



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. Cvmbopogon spp./ 64- 
65. JAMAICA: Portland Par., Millbank, 10 
August 1987, M. A. Ivie coir.; deposited in 
the M. A. Ivie collection; St. Ann Parish, 
Ocho Rios, 19 October 1985, J. A. Suey 
coir.; 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, lO/X/84, Seguva, Gillett, and 
Miranda/ on \tai oi Sida sp., coll. 78/ LPL 
1012; Bara Nieja, 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, lO/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). 

Etymology.— the specific ephithet, hitena 
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 Cvmbopogon spp. 

Immature stages unknown. 


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. 

Literature Cited 

Baly, J. S. 1885. Biologica Centrali-Americana. In- 
secla. Coleoptera. Vol. VI. Part 2. Phytophaga 
Hispidae. 1-124. 

Weise, J. 1911. Coleoptera: Phytophaga, family 
Chrysomelidae, subfamily Hispinae. In P. Wyts- 
man Genera Inseclonim fac. 1 25: 1-1 23. Brussels. 


91(3), 1989, pp. 346-349 




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, Pineviile, 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, Pineviile, Lou- 
isiana 71360; (RJG) Systematic Entomology Laboratory, PSL 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 Gagne 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 1 1 and September 17, 1984. 

Key Words: Contarinia sp., pine needle sheath midge, pine seed orchards, loblolly pine 

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 
Gagne, is found on slash pine, P. elliottii 
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 is un- 

Contarinia acuta was found in 1971 to 
cause needle droop and defoliation on lob- 
lolly pine at the Erambert Seed Orchard in 

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. 
1 976). Similar damage was reported in 1 975 
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. 



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. 

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 1 1.36 1 
plastic wash tubs and 0.24 1 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- 
lected on October 19, indicating that the 
late August generation was the overwinter- 
ing generation (Table 1). 

Population sampling conducted in 1984 



Table 1 . Comparison of mean lar\ al densities re- 
corded on the last three sampling dates at the U.S. 
Forest Service Stuart Seed Orchard, Pollock, LA (1983). 

Mean Number of Lar\ae per Fasciele 


.Aug. 2.1 

.Aug, 31 








— 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 1 8. 
The second, third, and fourth adult flights 
occurred between June 4 and June 1 8. 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 

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 
I si Instar 2nd Inslar 3rd Inslar 











































(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 ( 1 984), 
respectively (Table 3). A fourth generation 
prepupal peak was barely detectable due to 
a rapid collapse in the population. 



Table 3. Mean number of prepupae captured on I 
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 Number 
of Prepupae 
per 1 Sq Fl 



Number of 


per : Sq. Fl. 

May 29 









June 4 

















































July 2 



■ 4 




















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 Pycmotes eniargina- 
tiis 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 


The authors thank personnel at the U.S. 
Forest Service Stuart Seed Orchard for pro- 

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 ofPymotes (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. Contannia haeri (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. 

Gagne, R. J. and G. M. Beavers. 1984. Contannia 
spp. (Diptera: Cecidomyiidae) from shoots of slash 
pine [Piniis clliotti Engelm.) with the description 
ofa new species injurious to needles. Fla. Entomol. 
67(2): 221-228. 

Overgaard. N. A., H. N. Wallace. C. Stem, and G. D. 
Hertel. 1976. Needle midge (Diptera: Cecido- 
myiidae) damage to loblolly pines in the Erambert 
federal seed orchard, Mississippi. LISDA-FS. 
Southeastern Area, S&PF, HDM, Rep. No. 76-2- 
13. 11 pp. 

Skuhravy, V. 1973. "Needle blight" and "needle 
droop" on Piinis sdvestris L. in Europe and P. 
resinosa Ait. in North America (Diptera: Ceci- 
domyiidae). Z. Angew. Entomol. 72 (1972/73): 

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. 2 1 (3): 97- 

91(3), 1989, pp. 350-354 




A. G. Wheeler, Jr. 

Bureau of Plant Industry, Pennsylvania Department of Agriculture, Harrisburg, Penn- 
sylvania 17110. 

Abstract. — Gry pot es puncticolUs (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, Pimis sylveslris 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. puncticolUs 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. puncticolUs: Em- 
poasca perlonga Davidson & De Long and Gyponana geminata (Osbom). 

Key Words: Auchenorrhyncha, immigrant insect, Pinus sylvestris, Pinus mugo 

Grypotes puncticolUs (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 1 985 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. puncticolUs has been listed as a pest 
of P. brutia and P. nigra (Ossiannilsson 
1983, Lodos and Kalkandelen 1 985 and ref- 
erences therein). Metcalf s (1967) world cat- 
alogue may be consulted for references to 

additional European literature containing 
ecological notes. 

Herein, G. puncticolUs 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. puncticolUs 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- 



Fig. 1 . Known North American distribution of the Palearctic leafhopper Grypoles puncticoUis. Black dots 
indicate established populations; counties in which limited sui-veys for G. puncticoUis on Scotch pine were 
negative are represented by open circles (negative sites in Ohio are not shown). 

Ion (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 
Beime's (1956) leafhoppers of Canada and 
Alaska. With access to European literature 
and specimens, E. R. Hoebeke determined 
the species as the Palearctic G. piincllcollis. 
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. niiigo Turra. For all other collections list- 
ed below, P. sylvestris was the host; all col- 
lections were made by the author in 1988 
except for the 1 3 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, I Aug.; Mercyhurst College, 
Erie, 21 July, 1 Aug., 13 Sept.; North East, 
1 .'\ug. 



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 in 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, 
Pimis 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 

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. Hamilto