Attractant for a Beneficial Insect and Its

Parasitoids: Pheromone of the Predatory Spined Soldier Bug,

Podisus maculiventris (Hemiptera: Pentatomidae)1

J. R. ALDRICH, J. P. KOCHANSKY, ANDC. B. ABRAMS

Insect Physiology Laboratory, Agricultural Research Service,
U.S. Department of Agriculture, Bldg. 467, BARe-East, Beltsville, Maryland 20705

Environ.Entomol.13: 1031-1036 (1984)

ABSTRACT A mixture of (E)-2-hexenal, a-terpineol, linalool, terpinen-4-0I, and benzyl
alcohol attracts adults of the predaceous spined soldier bug, Podtsus maculiventris (Say).
The male-produced pheromone contains (+ loa-terpineol; however, (- )-a-terpineol is not
inhibitory to the bug, so artificial pheromone can be made with racemic a-terpineol. Four
parasitic species use the pheromone as a kairomone and two of these parasitoids are partially
inhibited by (- loa-terpineol. This is the first artificial long-range pheromone for a member
of the true bugs (Hemiptera: Heteroptera) and the first such pheromone for a nonsocial
predaceous insect.

SPINEDSOLDIERBUG ADULTSand immatures are
voracious predators of a wide variety of insects
throughout North America, preferring the eggs and
larvae of phytophagous species (McPherson 1982).
P. maculiventris has been repeatedly released in
Europe, but the species has apparently never be-
come established (Couturier 1938, Warren and
Wallis 1971). In North America, P. maculiventris
is the most widespread and probably the most ben-
eficial of the pentatomid predators (Asopinae), yet
little is known of how this predator behaves in
nature.
Descriptions by Dupuis (1952) of large dorsal
abdominal glands (DAG) opening underneath the
wings of male predaceous pentatomids prompted
one of us (J.R.A.) to examine the chemistry of these
glands. (E)-2-Hexenal, a-terpineol, and benzyl al-
cohol are major constituents in the male DAG se-
cretion of P. maculiventris, with lesser amounts of
Iinalool, terpinen-4-ol, and piperitol (Aldrich et al.
1978).
In order to clarify the biological role played by
the DAG secretion of male spined soldier bugs, we
have determined the optical rotation of the major
optically active component (a-terpineol), and have
field-tested artificial blends containing (+ )-a-ter-
pineol, (- loa-terpineol, and racemic a-terpineol.
Materials and Methods
A culture of P. maculiventris was maintained
in the laboratory on Tenebrio molitor L. larvae
and pupae (Rainbow Mealworms, Compton, Cal-
if.), as previously described (Aldrich et aI. 1978).
The culture was started with field-collected bugs
IThisarticlereportsthe resultsof researchonly.Mentionof a
proprietaryproductdoesnotconstitutean endorsementor a rec-
ommendationfor itsuseby USDA.
from Geneva, N.Y., and augmented with field-col-
lected bugs from Beltvsille, Md. The insectary was
kept at 28°C and 65% RH, on a photoperiod of
LD 16:8.
For determination of the optical rotation of
a-terpineol, a CHzCI2 extract of the DAG from 77
male bugs was prepared (Aldrich et al. 1978).
a- Terpineol was isolated, using a Varian 3700 gas
chromatograph, equipped with a thermal conduc-
tivity detector, from a 3% OV-275 Chromosorb
W-AW (100/120) column programmed from 60
to 160°C at 100C/min, using helium as the carrier
gas. a-Terpineol was trapped in a glass capillary
tube jacketed in dry ice as it eluted from the col-
umn. By repeated injections of the gland extract,
a total of 1.4 mg of the natural product was iso-
lated. The optical rotation was measured in an
ethanol solution with a 0.7-ml cell in a Rudolph
70 polarimeter (0. C. Rudolph and Sons Inc.,
Caldwell, N.J.).
The artificial blends were formulated to mimic
the composition of airborne extracts from live male
P. maculiventris. For airborne-trapping of the
pheromone, single adult males were confined in
an all-glass, closed system and air was drawn by
vacuum at 65 mil min over the insect and through
about 30 mg of an activated carbon filter (Amber-
sorb XE-348, Rohm and Haas, Philadelphia, Pa.).
The trapped volatiles were extracted from the fil-
ter with 200 j.L1 of CH2CI2 and analyzed using a
Packard 421 gas chromatograph, equipped with a
3% OV-275 Chromosorb W-AW (100/120) col-
umn programmed from 60 to 1200C at 10°C/min
using nitrogen as the carrier gas. The relative pro-
portions of the components were based on the per-
centage of areas of the gas chromatogram peaks.
The weighted average composition of several in-
dividual male determinations was 48.5% (E)-2-
hexenal, 46.1% a-terpineol, 4.5% benzyl alcohol,

1031
1032 ENVIRONMENTAL
Table 1. P. maculiventri. adults and tachinid Oy parasiloids caughl on or near slicky traps bailed
terpineol-blend, (- )-a-terpineol-blend, or laboratory reared bugs
P. macu/lven tris
Experiment Lure"
Male Female
(+ )-a- Terpineol- 147a 164a
blend
Male 30b 25b
Female 0 0
Control 0 0
II ( + )-a- Terpineol- 227a 281a
blend
(- )-a- Terpineol- 35b 31b
blend
a Numbers in a column for each experiment are significantly different if followed by different letters (P < 0.01; X2 test).
b Numbers of males and females are significantly different if followed by an asterisk (P < 0.01; X2 test).
0.4% piperitol, 0.3% linalool, and 0.2% terpinen-
4-01. Because initial attempts to synthesize the iso-
mers of piperitol were unsuccessful, this compo-
nent was not included in the artificial blends tested.
A formulation of 354 JlI of (E)-2-hexenal, 216 JlI
of a-terpineol, 19.7 Jll of benzyl alcohol, 2.4 JlI of
linalool, and 2.0 JlI of terpinen-4-01 closely ap-
proximates the proportions of these components in
the natural airborne secretion and these volumes
were convenient for preparing the mixture.
Four mixtures differing only in the a-terpineol
content were prepared for field-testing. (+ )-a- Ter-
pineol (1.6% antipode contamination) and (- )-a-
terpineol (1.9% antipode contamination) were syn-
thesized from (+ loa-pinene (Aldrich Chemicals,
Milwaukee, Wise.) and (- loa-pinene (Tridom
Chemicals, Hauppauge, N.Y.) (Colonge and Cra-
balona 1959). One blend was prepared with (+)-
a-terpineol and another was prepared with (- )-a-
terpineol. Two mixtures were prepared, using ra-
cemic a-terpineol (Hercules Inc., Wilmington,
Del.); one contained 216 Jll of (±)-a-terpineol (1
vol) and the second contained 432 JlI of (± )-a-
terpineol (2 vol). The remaining components (Al-
drich Chemicals) were mixed in the same propor-
tions listed above for all the artificial blends. Ra-
cemic linalool and (+ )-terpinen-4-o1 were used
because these were the only commercially avail-
able standards of these compounds.
Blends made with either (+ loa-terpineol or (- )-
a-terpineol were field-tested in sticky traps from
30 April through 19 October 1982, in and around
deciduous woods at the Agric. Res. Cent.-East,
Beltsville, Md. One experiment (Experiment I), us-
ing sticky wing traps (Zoecon Corp., Palo Alto,
Calif.) compared the attractiveness of traps baited
with (+ loa-terpineol-blend, to traps baited with a
live male or female bug, and to unbaited control
traps. Traps baited with live insects contained a
laboratory-reared bug confined inside the trap in
a screen cage provided with water and T. molitor
pupae for food. A second experiment (Experiment
II) compared the attractiveness of (+ )-a-terpineol-
ENTOMOLOGY Vol. 13, no. 4
wilh (+ )-a-
Tachinid parasitoidsh
E. flava H. aurata
Male Female Male Female
863a 907a 22a· 235a·
15b· 104b· 5b· 26b·
0 0 0 0
0 0 0 0
1.231a 1,147a 14a· 298a·
53b· 112b· 5b· 207a·
blend with (- )-a-terpineol-blend. using sheet metal
plates (13 by 20 cm) coated with Tack Trap (An-
imal Repellents Inc., Griffin, Ga.). Five traps were
run for each treatment in the two experiments.
Ten microliters of the neat blend was applied dai-
ly to a rubber septum (5 by 9 mm) (Thomas Sci-
entific, Philadelphia, Pa.) in the middle of attrac-
tant-baited traps. Traps were hung on tree branches
about 2 m from the ground and at least 100 m
apart, alternating treatments. From April to Oc-
tober 1983, seven sheet metal sticky traps were
hung, rebaited, and monitored as in 1982. Spined
soldier bugs that did not land on the sticky portion
of a trap were reluctant to walk onto the sticky
material or, if entangled, sometimes crawled off
the trap. Therefore bugs within 1 m of a trap were
collected and counted as trapped bugs. The para-
sitic flies were highly susceptible to capture in sticky
traps so only those parasitoids actually caught in
traps were counted.
Artificial blends were made with racemic a-ter-
pineol and (+ loa-terpineol were field-tested in
1983, using slow-release formulations and a new
type of trap that captured the bugs and parasitoids
alive ..The new traps were made from transparent
cylindrical containers (20.2 by 19.7 cm; Tri-State
Molded Plastic, Dixson, Ky.) by cutting two holes
(9 cm diam) in opposite sides and covering each
hole with an inwardly projecting screen funnel. In
one experiment a 20% (wt/wt) formulation of (+)-
a-terpineol-blend in plasticized polyvinyl chloride
(Tenneco, Piscataway, N.].) was prepared (Fitz-
gerald et al. 1973) and four traps were rebaited
every 4 days with 350 mg of the material. The
baited traps were hung from trees as above, alter-
nating with unbaited control traps, and were mon-
itored daily from mid-March to October 1983. In
a second experiment, ( + )-a-terpineol-blend and the
two (±)-a-terpineol-blends were mixed with Poly-
trap (Recon Associates Inc., Huguenot, N.Y.) (60%
wt/wt) and three traps were rebaited every 4 or 5
days with 75 mg of the material. Traps were hung
as above, alternating treatments and three unbait-
August 1984 ALDRICH ET AL.: PHEROMONE OF SPINED SOLDIER BUG 1033

5 e. macullventris A ~. lJlaculiventris
j: both sexes ~A 50
,\ v! male
j:
,\
j: ~ . .IJAXA
i 40 (0
:! \
,I'\
female \ 1\ i "
'
e. maculiventris
i:, \ i 30 i\ ' \
i : female
' l
j:
1i. .lIIIWll
female
\ i\ i \
, \
j \ \
20 i \ i' \\
, \
i \ i j \;\ i' \ ~
: (:
N \ \ i \\ : \: \
I:.' \i'. I I V \\ .I'
'
to'' " \ .. ! \ :
v'
': \ "

e. macullventris
male
Fig. 1. Field-attraction of P. maculiventris adults en 60
and females of the tachinid parasitoids, E. fiava and H. ~
aurata, to traps baited with (+ )-a-terpineol-blend dur- frl 50 e. maculiventris
ing 1982 and 1983. Each point is the sum of the catches female
for 5 days from seven sticky traps. ~ 40
u.
o
a: 30
w
ed control traps, and monitored daily from 24 May !Xl 20
to 18 August 1983. Only bugs or parasitoids caught
inside traps were counted. ~

Results /" \
:
~I '\
Optical Rotation. The optical rotation of a-ter- 1\,
pineol isolated from laboratory reared male P. 60 : Ij \,
I
maculiventris DAG was calculated to be [aJ B = : i
+ 115° ± 19.8 SEM. The rotation of ( + )-a-terpine- 50 Telenomus sp. I \
I \ r-,
01 according to Colonge and Crabalona (1959) is
[aJ f) = + 1000. These data suggest that the insect
40
/'" :
i V :\
produces only the plus enantiomer of a-terpineol, 30 I ' i..
although this method does not exclude the possi-
bility that a small amount of the antipode is also
20
.
" j
I
'
: \
\
\
,
produced by the insect. 10 ! \: \
Field Tests. In 1982, 1,416 P. maculiventris ) ",..
adults were caught on or nearby 30 sticky traps. o APR JUN JUL
In addition, 8,585 individuals of two parasitic flies,
Hemyda aurata Robineau-Desvoidy and Euclytia Fig. 2. (A) P. maculiventris adults caught on or near
seven sticky traps baited with (+ )-a-terpineol-blend
!lava (Townsend) (Diptera: Tachninidae), were
during 1983. (8 and C) P. maculiventris adults and the
caught on the traps. No bugs or parasitoids were parasitoids, E. fiava and Telenomus sp., caught inside
caught on or near control traps. Table 1 gives the five plastic traps baited with (+ )-a-terpineol-slow-re-
results of 1982 experiments I and II. Only traps lease-blend during 1983. Each point is the sum of trap
baited with a live male bug or artificial blends catches for 5 days.
were attractive to P. maculiventris and its tach-
inid parasitoids (Experiment I), The number of
male and female spined soldier bugs trapped was
not significantly different (Experiments I and II). males selectively release pheromone. This field-
Males and females of E. !lava were equally sus- observation agrees with observations of airborne-
ceptible to capture in traps baited with (+ )-a-ter- trapping in the laboratory that indicated males re-
pineol-blend, otherwise more female than male lease pheromone only during daylight and some
tachinids were caught in traps (Experiments I and days remain "silent."
II). The blend made with ( + )-a-terpineol was much Fig. 1 shows the spined soldier bug and female
more attractive to bugs and tachinid parasitoids tachinid parasitoid catches for 1982 and 1983, us-
than (- )-a-terpineol-blend (Experiment 11). Cap- ing seven sheet metal sticky traps baited with
tures of bugs and parasitoids in sticky traps baited (+ )-a-terpineol-blend. By starting field-trapping 6
with live males was much more sporadic than for weeks earlier in 1983, we discovered that the
traps baited with artificial blends, suggesting that emergence of overwintered bugs had been com-
1034 ENVIRONMENTAL ENTOMOLOGY Vol. 13, no, 4

Table 2. P. maculiventri. and parasitoids caught inside field-traps baited with (+ )-a-terpineol-blend or (± )-a-
terpineol-blend

Parasitoids
P. maculi-
Lure Telenomus
ventrisa E. flava F. crinita
n. sp.
I: Blend with
I volume of
(+ )-a-terpineol 165a U8a 36a 1I3a
II: Blend with
2 volumes of
(±)-a-terpineol 138ab 48b 15b 142a
III: Blend with
I volume of
(±)-a-terpineol 1I3b 92a 30a 1I8a

• Numbers in a column are significantly different if followed by different letters (P < 0.05: x2 test).

pletely missed in 1982, Many bugs were caught in
April 1983, before the first tachinid parasitoids (E.
!lava) were caught on 29 April. The captures of
bugs and tachinids for May through September
were similar for 1982 and 1983, The most bugs
were caught during June, more tachinids were
caught than bugs, and the capture-curves for the
tach inids were out of phase with each other. There
was no indication that the 1982 field-trapping had
any effect on the local populations of P. maculi-
ventris, H. aurata, or E. jlava in 1983.
Comparison of the number of p, maculiventris
adults caught on or near sticky traps baited with
the ( + )-a-terpineol-blend from April through Sep-
tember 1983 (Fig, 2A), to the number of adult
bugs caught in the new plastic traps baited with
the 20% (+ )-a-terpineol-slow-release-blend during
the same period (Fig. 2B), shows that both trap-
ping methods detected emerging overwintered
bugs. However, after early May, relatively few bugs
were caught in the plastic traps. E. jlava and fe-
males of the phoretic egg parasitoid, Telenomus
n. sp, (Buschman and Whitcomb 1980) were caught
in large numbers in the plastic traps baited with
the (+ )-a-terpineol-blend (Fig, 2C) when relative-
ly few bugs entered traps. Females of an ectopar-
asitic biting midge, Forcipomyia crinita Saunders,
were also common in these traps and were often
observed feeding on adult bugs inside traps. H.
aurata was commonly observed around the baited
plastic traps but none were caught in these traps.
Individuals of E. jlava were frequently seen rest-
ing on the screen funnel entrances to baited plastic
traps and approaching bugs appeared to be wary
of entering these traps,
The results of the May to August 1983 experi-
ment comparing the attractiveness of (+ )-a-ter-
pineol-blend with blends made with one or two
volumes of (±)-a-terpineol are presented in Table
2. Only those parasitoids caught in a plastic trap
on days when no P. maculiventris were in a trap
are included in Table 2 to eliminate parasitoids
that may have been attracted to live bugs rather
than to artificial blends. (- )-a- Terpineol did not
inhibit the response of P. maculiventris to the at-
tractant; it appears simply to act as a diluent. Sim-
ilarly, (- )-a-terpineol showed no inhibition of the
response of Telenomus n. sp. to attractant. How-
ever, significantly fewer individuals of E. jlava and
F, crinita were caught in traps baited with blend
II than blends I or III, indicating that the excess
(- )-a-terpineol in blend II is inhibitory to these
parasitoids.
Table 3 gives the numbers of bugs and parasit-
oids caught in plastic traps on successive days after
baiting for the May to August 1983 experiment,
using 60% (wt/wt) formulations of (+ )-a-terpine-
01- and (±)-a-terpineol-blends. Day 4 catches of
p, maculiventris were 34.4% of day 1 catches,
compared with 34.1% for Telenomus sp., 17.0%
for F. crinita, and 9.0% for E. jlava.
Tachinid Parasitization. During 1982, the

Table 3. P. maculiventris and parasitoids caught in slow-release attractant-baited traps from 1 to 4 days after
baiting

Parasitoids
Days after P. macult-
baiting ventris· Telenomus
E. flava F. crinita
n. sp.
I 163a 277a 36a 232a
2 94b lOOb 53a 210a
3 95b 45c 13b 130b
4 56c 25d 9b 79c

• Numbers in a column are significantly different if followed by different letters (P < 0.05; X2 test).
August 1984 ALDRICH ET AL.: PHEROMONE OF SPINED SOLDIER BUG 1035

E. maculiventris H. aurata larvae emerged from 10.1% (n = 297)

A
80 male of these overwintered adult bugs. No E. jlava
emerged from these bugs, although this was the
70 first tachinid to be attracted to traps in the spring;
60 Tachinid eggs H. aurata was not caught in attractant-baited sticky
traps until 18 May 1983. Bugs caught in plastic
50 traps after the appearance of E. jlava often had
(f) 40 many tachinid eggs on them and several times these
Cl multiply-parasitized bugs were observed feeding
Cl 30 on E. jlava flies inside a trap. Thus, parasitization
..•.
w
of bugs around traps baited with artificial blends
(f)
I-
20
0 is probably unnaturally high. Overestimation of
w 10 tachinid parasitization is less of a problem for at-
(f)
~ 0 tractant-baited sticky traps than plastic traps be-
I.L. B E. maculiventris cause the flies are unable to escape from the sticky
0 80 female
a: material.
w
7
Tachinid eggs Discussion
~ 60
50 P. maculiventris adults periodically migrate in
search of food and shelter (Evans 1982), as do many
40
hemipteran species, but the details of the recolo-
30 nization process remain obscure. The attraction of
20 1\ spined soldier bugs to traps baited with male bugs
/ \ or mixtures of compounds blended to mimic the
10 / ''---, male DAG secretion unequivocally demonstrates
0
../ \ that this secretion constitutes the long-range at-
JUN JUL AUG SEP tractant pheromone of P. maculiventris. It ap-
pears that males often search for food first, then
Fig. 3. Tachinid eggs on 1982 field-collected male
(A) and female (B) P. maculiventris adults.
call females with the pheromone from their DAG.
The intensity of parasitization on P. maculiven-
tris is both surprising and interesting. The major
number of tachinid eggs on field-collected bugs tachinid parasitoids of spined soldier bugs in
was recorded (Fig. 3) and parasitized adult bugs Maryland have population cycles out of synchrony
collected around traps were placed in Petri dishes with each other, minimizing direct competition
in the insectary for parasitoid emergence. There for hosts. E. jlava flies are first to parasitize P.
were 254 tachinid eggs on the 666 field-collected maculiventris in the spring although they appar-
adult male P. maculiventris, and 69 eggs on the ently overwinter as larvae inside adults of other
750 adult female bugs collected in the field in 1982. pentatomid species (Eger and Ables 1981). Our
Male bugs often had more than one tachinid egg records of F. crinita are the first report of a hemip-
on them (up to nine on a single male), while par- teran host for a biting midge and the first time a
asitized female bugs usually had only one egg on ceratopogonid has been shown to be attracted to
them. Adults of 44 E. jlava were reared from P. a host pheromone. While phoresy is common in
maculiventris adults parasitized in the field; 32 of the Scelionidae (Clausen 1976), host-finding in fe-
these were reared from male bugs and 12 from male Telenomus n. sp. is unusual because it entails
female bugs. Adults of seven H. aurata were reared orienting to a male-produced aggregation phero-
from field-parasitized spined soldier bugs; three mone. Female wasps can then become phoretic on
from male bugs and four from female bugs. Many mated female bugs or search directly for eggs in
bugs died without a parasitoid emerging. Parasit- an area assured to contain mated female bugs. Be-
ized bugs usually died within a day after emer- cause calling males are more vulnerable to para-
gence of the maggot, but on several occasions a fly sitoids than noncalling males, it may be advanta-
maggot emerged from its P. maculiventris host geous for a male to remain silent sometimes and
and the bug fed on the maggot before it could try to usurp females attracted to calling males.
form a puparium. P. maculiventris females col- Artificial pheromone of P. maculiventris may
lected near attractant-baited traps and brought into be useful in luring predators to pest infestations or
the laboratory in order to rear their parasitoids out of infested fields before an insecticide appli-
often laid viable eggs, demonstrating that mated cation, in suppressing parasitism, in establishing
females are attracted to the pheromone. the predator in exotic regions, and in monitoring
In 1983, all the adult bugs caught alive in the population levels of the predator.
plastic traps during April (before the appearance Predaceous hemipterans are instrumental in
of tachinids in the field) were placed in dishes with suppressing countless agricultural pests. For ex-
wire mesh, false bottoms for parasitoid emergence. ample, in the United States the most abundant
1036 ENVIRONMENTAL
predaceous insects in soybean are hemipterans:
Nabis spp. (Nabidae), Geocoris spp. (Lygaeidae),
Grius insidiosus (Say) (Anthocoridae), and P.
maculiventris (Turnipseed and Kogan 1983). Mor-
phological and behavioral data suggests that at least
some nabids and anthocorids rely on male-pro-
duced aggregation pheromones to congregate
(Stoner 1973, Carayon 1954, Peet 1979). Synthetic
aggregation pheromones for a few key species of
these predaceous Hemiptera could be invaluable
in combating a wide variety of pests.
Acknowledgment
We thank the following scientists of the Systematic
Entomology Laboratory (USDA) for identifications of
specimens: C. w. Sabrosky (Tachinidae), W. E. Wirth
(Ceratopogonidae), P. M. Marsh (Scelionidae), and T. J.
Henry (Pentatomidae). We also thank Norman Johnson,
Dept. of Entomology, Ohio State Univ., for examining
the scelionids, and Richard Chromecek, Recon Associ-
ates Inc., Huguenot, N.Y., for preparing the pheromone
polymer. We also thank Joe Sexton for technical assis-
tance.
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Received for publication 20 January 1984; accepted
10 April 1984.