Journal of Economic Entomology, 110(6), 2017, 2727–2730

doi: 10.1093/jee/tox276
Advance Access Publication Date: 6 November 2017
Short Communication Short Communication

A Survey of the Species of Squash Bug (Hemiptera:

Coreidae) Egg Parasitoids in Virginia and Their
Distribution
James M. Wilson1 and Thomas P. Kuhar
Department of Entomology, Virginia Tech, Blacksburg, VA, and 1Correspondence author, e-mail: keepbees@vt.edu

Subject Editor: Eric Riddick

Received 6 July 2017; Editorial decision 30 August 2017

Abstract
Squash bug, Anasa tristis DeGeer (Hemiptera: Coreidae), is a major pest of squash and pumpkins in the United
States. In order to better understand the importance of natural egg parasitism of this species in Virginia, we
conducted a 2-yr statewide survey. In total, 1,127 squash bug egg masses (~20,000 total eggs) were sampled from
squash and pumpkins from 43 counties in Virginia from 2014 to 2015. Egg masses were brought back to the lab
to record levels of squash bug nymphal emergence or adult parasitoid eclosion and identification. Over 50% of
the total squash bug eggs collected statewide were parasitized. Gryon pennsylvanicum Ashmead (Hymenoptera:
Scelionidae) was the predominant egg parasitoid accounting for over 98% of all parasitoid adults recovered. The
only other species emerging from squash bug eggs was Anastatus reduvii Howard (Hymenoptera: Eupelmidae),
which is a generalist parasitoid. G. pennsylvanicum was found in 75% of the counties surveyed with the highest
levels of parasitism occurring in the Northern, Southwestern Mountain, and Western Piedmont regions of the state
and the lowest levels of parasitism occurring in the Tidewater region in the southeastern portion of the state. Based
on this 2-yr survey, G. pennsylvanicum was determined to be a major natural enemy of squash bug, significantly
reducing the number of nymphs that emerge from deposited eggs. Conservation of this natural enemy should
therefore be a priority for integrated pest management programs in cucurbits.

Key words: Anasa tristis, biological control, parasitoid, natural control

The squash bug, Anasa tristis DeGeer (Hemiptera: Coreidae), is an
important pest of pumpkin (Cucurbita maxima), squash (Cucurbita
pepo), and other cucurbits in the United States causing wilt in plants
with its piercing sucking mouthparts and by transmitting a plant
pathogenic strain of the bacteria Serratia marcescens, which causes
Cucurbit Yellow Vine Decline (Bruton et  al. 2003, Doughty et  al.
2016). With hopes of developing a pest management strategy for
this pest that is more pollinator friendly (US EPA 2015)  and less
dependent on applications of broad-spectrum insecticides such as
pyrethroids or neonicotinoids (Doughty et al. 2016), a better under-
standing of the biological control of the insect is needed. In Virginia,
the natural enemy complex of squash bugs has not been well studied.
Egg parasitoids appear to be the most important natural ene-
mies of this pest in other regions of the United States (Nechols et al.
1989, Doughty et  al. 2016). Among the relatively few parasitoids
of squash bug, the egg parasitoid Gryon pennsylvanicum Ashmead
(Hymenoptera: Scelionidae), has been shown to be one of the most
effective (Nechols et  al. 1989) with natural parasitism levels of
31% and 55% being recorded in Kentucky and Mayland, respec-
tively (Decker and Yeargan 2008, Cornelius et  al. 2016). In 2013,
we collected 81 squash bug egg masses from two counties in south-
western Virginia and observed parasitism levels greater than 80%.
Given the high rate of parasitism recorded in 2013, we embarked
on a more extensive and robust survey of squash bug egg parasit-
ism in Virginia, which, to our knowledge, has not been conducted
before. Herein, we report the results of a 2-yr survey from 43 coun-
ties throughout Virginia, which will enable us to quantify the poten-
tial effect that this parasitoid is having on squash bug population
dynamics in the state.
Materials and Methods
Throughout the summer months of 2014 and 2015, wild squash bug
egg masses were collected from summer squash, zucchini (C. pepo
L.), and pumpkin (C. maxima Duchesne) plants located throughout
Virginia. After finding very few squash bug eggs in commercial con-
ventional fields presumably due to the insecticide spray programs
used in those fields, we focused most of our sampling in untreated
squash or pumpkins located on Virginia Tech research farms, organic
farms, and community gardens throughout Virginia. In 2014, a total

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2728 Journal of Economic Entomology, 2017, Vol. 110, No. 6

of 722 squash bug egg masses were collected from 34 counties, and
in 2015, 405 squash bug egg masses were collected from 24 Virginia
counties (Table 1).
Egg masses of A. tristis were removed from leaves in the field by
cutting around the egg mass and placing it individually into 60 × 15
mm Petri dishes, which were kept at room temperature (~27℃) until
all nymphs and adult parasitoids had emerged. Once all nymphs and
parasitoids had perished, samples were processed and parasitism and
squash bug hatch rates were recorded. Data from egg masses col-
lected in Virginia were grouped by NOAA (2017) climatic regions
and proportion parasitism data were transformed using an arcsine-
square root transformation to normalize the variances (Sokal and
Rohlf 1995) and then analyzed using analysis of variance, JMP
version 10.0 (SAS Institute, Cary, NC). Means were separated using
Fisher’s Protected least significant difference at the P < 0.05 level of
significance. Data are presented as original means.
Results
In 2014, squash bug egg parasitism was found in 75% (30 of the
37) counties surveyed, ranging from 0% in some counties to 100%
in others, with an overall statewide average of 39.4% of egg masses
parasitized (Table 1). G. pennsylvanicum was the predominant egg
parasitoid accounting for over 98% of all parasitoid adults recov-
ered; the only other species emerging from squash bug eggs was
Anastatus reduvii Howard (Hymenoptera: Eupelmidae), which is a

Table 1.  Numbers of squash bug egg masses collected and percentage parasitized by Gryon pennsylvanicum by region and county in
Virginia 2014 and 2015

2014 2015

NOAA climatic division Location (Co.) Egg masses (EM) % Parasitism EM (eggs) Egg masses (EM) % Parasitism EM (eggs)

Tidewater Accomack 20 20.0 (2.2) 0.0 0.0

Greensville 5 0.0 – –
Northampton 14 0.0 0.0 0.0
Virginia Beach – – 6 0.0
Eastern Piedmont Amelia 22 9.1 (2.0) – –
Buckingham 25 4.0 (0.7) 16 31.3 (10.6)
Caroline 4 0.0 – –
Cumberland 20 30.0 (14.8) 12 41.7 (27.8)
Fluvanna 13 15.4 (10.3) 8 37.5 (11.6)
Goochland 46 23.9 (51.7) 14 28.6 (18.4)
Hanover – – 15 86.7 (44.5)
Henrico - – 15 93.3 (43.1)
Louisa 11 18.2 (1.4) 0.0 0.0
Mecklenburg 16 18.8 (50.0) – –
Nottaway – – 15 100.0 (82.8)
Powhatan 28 3.6 (4.8) 15 6.7 (7.4)
Prince Edward – – 5 0.0
Spottsylvania – – 14 71.4 (33.2)
Western Piedmont Albemarle 9 88.9 (92.0) 19 63.2 (30.6)
Appomattox – – 10 90 (57.5)
Bedford 14 71.4 (76.9) – –
Campbell – – 15 46.7 (14.1)
Franklin 32 9.4 (4.1) – –
Henry 8 0.0 – –
Nelson – – 15 93.3 (38.8)
Pittsylvania 9 88.9 (53.0) – –
Northern Virginia Clarke 2 50.0 (16.7) – –
Fauquier 10 100.0 (98.5) – –
Fredrick 26 96.2 (64.4) – –
Loudoun 6 83.3 (35.6) – –
Madison – – 15 0.0
Orange – – 17 29.4 (54.5)
Page 6 0.0 – –
Rappahannock 10 60.0 (37.4) – –
Shenandoah 21 85.7 (71.7) – –
Central Mountain Augusta 15 13.3 (12.6) 25 100 (99.7)
Botetourt 3 0.0 – –
Roanoke 16 56.3 (47.7) – –
Rockbridge 4 50.0 (50.0) – –
Rockingham 22 100.0 (100.0) 36 66.7 (47.7)
Southwestern Mountain Floyd 4 0.0 – –
Montgomery 235 68.5 (37.0) 45 100.0 (95.6)
Pulaski 6 83.3 (81.6) 5 100.0 (97.3)
Washington 21 28.6 (13.6) – –
Wythe 19 63.0 (40.0) – –
Journal of Economic Entomology, 2017, Vol. 110, No. 6
Fig. 1.  Mean ± SE percentage parasitism of squash bug eggs by Gryon pennsylvanicum by region and year of in Virginia (means in columns with a letter in
common are not significantly different, Fisher’s Protected least significant difference α = 0.05.)
generalist parasitoid that attacks eggs from multiple insect orders
(Burks 1967). In 2015, 18 of the 24 sampled counties had parasitism
by G.  pennsylvanicum, with 49.4% of collected egg masses para-
sitized (Table 1).
There was a significant effect of climatic region on parasitism
rates (Fig.  1). In 2014, there were significant differences among
regions in egg mass parasitism (F = 2.5515; df = 5, 33; P < 0.0505)
but not in total egg parasitism (F = 1.7182; df = 5, 33; P > 0.1631);
however, numeric trends were the same with the highest parasitism
occurring in the Northern Region (67.9% of egg masses and 46.3%
of eggs) and the lowest occurring in the Tidewater Region (6.7%
of egg masses and 0.8% of eggs) and Eastern Piedmont Regions
(13.7% of egg masses and 15.1% of eggs) (Fig. 1). In 2015, there
was also a significant effect of region on parasitism of egg masses
(F = 4.8651; df = 5, 23; P < 0.0055) and eggs (F = 5.9854; df = 5,
23; P < 0.002). The highest parasitism occurred in the Southwestern
Mountains (100.0% of egg masses and 96.3% of eggs) and the low-
est again occurring in the Tidewater Region (0.0% of egg masses)
(Fig. 1).
Discussion
Sampling squash bug eggs throughout Virginia in 2014 and 2015
showed widespread occurrence of the egg parasitoid G. pennsylvani-
cum, which was recovered from 32 out of 43 counties, resulting in
significant egg mortality in the state, with 100% parasitism of egg
masses recorded from several samples each year. G.  pennsylvani-
cum has been previously reported from squash bug eggs in North
Carolina, Kansas, and Kentucky but at lower parasitism levels than
were found in our survey of Virginia (Schell 1943, Olson et al. 1996,
Decker and Yeargan 2008). In the Southwestern Mountain region of
Virginia, parasitism averaged 63.3% based on a sample of 335 egg
masses collected over 2 yr. This level is consistent with that reported
recently by Cornelius et  al. (2016), who observed G.  pennsylvani-
cum parasitism levels averaging 55.7% from wild squash bug eggs
collected in Beltsville, MD. The physical removal of egg masses of
varying age from the field likely prevented rates of parasitization
from reaching their natural level and so underestimated parasitiza-
tion rates. Thus, G. pennsylvanicum appears to be a significant natu-
ral enemy of squash bugs in Virginia, and the rest of the Mid-Atlantic
region of the United States may be similarly impacted. However,
2729
there appear to be additional factors that can greatly impact the level
of parasitism that occurs.
Low levels of parasitism were found in the Tidewater Region of
Virginia, which has the warmest climate in the state. Climate can
impact survival of parasitoids as well as synchrony with the popu-
lation dynamics of its host. Determining the ecological reasons for
low parasitism in the Tidewater Region should be a focus of addi-
tional research. Studies have shown that G. pennsylvanicum has a
host range that appears to be limited to the leaf-footed bugs (family
Coreidae) and has been recently introduced as a classical biological
control agent for the western conifer seed bug, Leptoglossus occi-
dentalis Heidemann (Hemiptera: Coreidae) in Italy (Peverieri et al.
2013, Roversi et al. 2014). Adult G. pennsylvanicum do not feed on
the host eggs (Vogt and Nechols 1993) but instead feed on the exu-
date from cucurbit leaf trichomes (extra-floral nectaries) that serve
as sources of basic sugars and protein (Olson et al. 1996). G. penn-
sylvanicum has been considered for augmentative biological con-
trol of squash bug in the past (Olson et al. 1996) but was not cost
efficient when compared to conventional squash bug management.
In addition to regional differences, very few squash bug eggs
or parasitoids were found in conventional commercial squash and
pumpkin farms in Virginia. Presumably the frequent use of broad-
spectrum insecticides like pyrethroids is a major limiting factor. Given
the obvious biological importance of this parasitoid, conservation of
this natural enemy should therefore be a priority for integrated pest
management programs in cucurbits, especially in crops under reduced
spray management. In Virginia, organic and or reduced spray-man-
aged squash and pumpkin fields will likely benefit from the conserva-
tion of G. pennsylvanicum. Using an integrated approach to squash
bug management as suggested by Doughty et al. (2016) may be the
most sound and sustainable way to control squash bug.
Acknowledgments
This work was funded in part by the Southern Sustainable Agriculture Research
and Education Graduate Student Project grant GS14-131. Additional support
was generously provided by the Virginia Agricultural Council.
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