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Armyworm Symposium 2002: Adamczyk et al.


SPATIAL AND TEMPORAL OCCURRENCE OF BEET ARMYWORM
(LEPIDOPTERA: NOCTUIDAE) MOTHS IN MISSISSIPPI

J. J. ADAMCZYK, JR.1, M. R. WILLIAMS2, J. T. REED2, D. W. HUBBARD1 AND D. D. HARDEE1
'USDA, ARS, Southern Insect Management Research Unit
P.O. Box 346, Stoneville, MS 38776

2Mississippi State University, Department of Entomology and Plant Pathology
Clay Lyle Building, Mississippi State, MS 39762

ABSTRACT

Throughout 1994-2000, adult beet armyworm, Spodoptera exigua (Hiibner) populations
were monitored in the delta and hill regions of Mississippi using pheromone traps. Signifi-
cant differences in the mean number of moths trapped were found among different geo-
graphical areas of the state. A trend was observed where the greatest number of moths was
found in the Mississippi Delta, located in the western region of the state. The lowest number
of moths was found in the hills located in the eastern region of the state. An annual profile
of beet armyworm populations in the western section of the Mississippi Delta also revealed
that wide-scale immigration of this pest typically begins at 200 Julian days (mid-July). This
date could be used as a benchmark to determine when and if population levels are high
enough to have the potential to cause economic damage to crops in the Mississippi Delta.

Key Words: Spodoptera, migration, movement

RESUME

A trav6z de los aios 1994 a 2000, se realizaron un monitoreo de las poblaciones de adults
del gusano trozador de la remolacha, Spodoptera exigua (Hiibner) en las regions de la Delta
y las colinas del Estado de Mississippi usando trampas de feronomas. Se encontraron dife-
rencias significativas en el numero promedio de la polillas atrapada entire las areas geogra-
ficas diferentes del Estado. Se observe un patron donde se encontr6 el numero mas alto de las
polillas en la Delta del Mississippi, ubicada en la region occidental del Estado. Se encontr6
el numero de polillas mas bajo en las colinas ubicadas en la region oriental del Estado. Un
perfil annual de la poblaci6n del gusano trozador de la remolacha en la secci6n occidental del
Delta del Mississippi tambien revel6 que una inmigraci6n de amplia escala de esta plaga ti-
picamente empieza a los 200 dias Julianos (en medio dejulio). Esta fecha puede ser utilizada
como un estandar o norma para determinar cuando y si el nivel de la poblaci6n es suficiente
alto para tener el potential de causar dano econ6mico a los cultivos del Delta del Mississippi.


The beet armyworm, Spodoptera exigua (Hiib-
ner), is an occasional but serious pest of various
vegetable and row crops in the mid-southern
United States of America. This Old-World species
was first documented in the state of Mississippi in
1920 (Mitchell 1979). Compared to other North
American armyworm species (e.g., the fall army-
worm, Spodoptera frugiperda (J. E. Smith)),
knowledge of the ecology of this pest in the Mid-
South is limited. Although this pest has no known
photoperiod or temperature induced diapause
mechanism (Kim & Kim 1997), it is able to over-
winter by continuous generations in southern
Florida and Texas. Therefore, initial populations
of beet armyworms found throughout the state of
Mississippi are believed to be the result of immi-
gration from those areas. Hendricks et al. (1995)
profiled populations of beet armyworms in the
lower Mississippi Delta and noted that moths
were found in all months, but the greatest num-


bers were found in the fall months (September
and October). However, in that study, populations
were only monitored for one season and conse-
quently conclusions concerning population struc-
ture were limited. The purpose of this study was
to examine the occurrence of beet armyworm
moths across different geographical regions of
Mississippi and to profile yearly moth popula-
tions to better understand the ecology of this pest
in the Mid-South.

MATERIALS AND METHODS

Adult populations of beet armyworms were
monitored throughout agricultural areas of Mis-
sissippi using pheromone traps. Reusable bucket
style traps (Gempler'sTM) were baited routinely
with synthetic pheromones and traps were
checked weekly as described by Hendricks et al.
(1995).











The primary objective of this study was to ex-
amine the population structure of beet army-
worms across different geographical regions of
Mississippi. An extensive trap line was conducted
from 1995-1996, and 1998-2000. Traps were lo-
cated in 51 counties across the state. For each
year, traps were typically run between 100 and
300 Julian days. Geographical regions of the state
were separated into 5 groups (W. Delta, 5 coun-
ties; C. Delta, 10 counties; E. Delta, 13 counties;
C. Hills, 9 counties; NE. Hills, 14 counties) (Fig.
1). For comparisons among groups, Julian dates
were separated into periods of 30 d, with the ex-
ception being the first time period which was in-
creased to 60 d to increase numerical entries for
analysis. Across multiple years and counties, cu-
mulative mean numbers of moths trapped per
time period were generated for each group. Differ-
ences among the groups were analyzed using
PROC MIXED (SAS Institute 2001). Further-
more, cumulative means for the entire time period
(117-326 d) were separated using the LSMEANS
option of PROC MIXED (Littrell et al. 1996).
A secondary objective was to examine the sea-
sonal profile of beet armyworms in the Missis-
sippi Delta by using an additional data set. A
continuation of a one-year survey described in
Hendricks et al. (1995) was conducted. Traps (15)






II
J c3 d NE. Hills





Washington C
CO. (W. Delta) ( J


September 2003


were run continuously (i.e., 365 d, 12 mo, 7 y)
from 1994-2000 in Washington Co., W. Delta, Mis-
sissippi. A scatter plot of the data was generated
using the graphics option of SAS Analyst (SAS In-
stitute 2001), and a 2nd-order polynomial equa-
tion that described the majority of the data (190-
350 d) also was generated using simple regression
(SAS Institute 2001).

RESULTS AND DISCUSSION

Throughout the agricultural areas of Missis-
sippi, a strong trend existed where the highest
population density of beet armyworm moths was
located in the West Delta near the Mississippi
River (Fig. 2). Between 117 and 206 Julian days,
there were no significant differences (P > 0.05) in
the cumulative mean numbers of moths trapped
between the 5 geographical regions (117-176 d: F
= 2.16, P = 0.089; 117-206 d: F = 1.69, P = 0.169).
The fact that the beet armyworm has no known
obligatory diapause (Mitchell 1979) could explain
why low numbers of moths were trapped so early
in the year (i.e., before wide-scale migration from
southern latitudes). However, as the growing sea-
son progressed, there were significant differences
(P < 0.05) in the cumulative mean numbers of
moths trapped between the 5 geographical re-
gions (117-236 d: F = 3.44, P = 0.015; 117-266 d: F
= 9.49, P < 0.001; 117-296 d: F = 13.28, P < 0.001;
117-326 d: F = 12.78, P < 0.001). In addition,
mean comparison between the 5 geographical re-
gions showed that the population of moths found
in the West Delta (i.e., the area closest to the Mis-
sissippi River) was significantly higher than pop-
ulations found in all other areas (Table 1). The
beet armyworm overwinters by continuous gener-
ations in southern latitudes (e.g., Florida, Texas,
Caribbean, and Central America) (Mitchell 1979)
and these populations probably invade Missis-


700-

600-

500
0
I
400

E 300

200-

100-


Fig. 1. Geographical regions of Mississippi moni-
tored or BAW moth populations.


117-176 117-206 117-236 117-266 117-296 117-326
Julian Dates

Fig. 2. Number of beet armyworm moths found in
different geographical regions of Mississippi.


Florida Entomologist 86(3)


W. Delta




C. Delta



E. Delta

E. Hills
// Hill"







Armyworm Symposium 2002: Adamczyk et al.


TABLE 1. NUMBER OF BAW TRAPPED THROUGHOUT THE
1995-1996, 1998-2000 GROWING SEASON FROM
VARIOUS GEOGRAPHICAL REGIONS OF MISSIS-
SIPPI.

Group' Cumulative Mean2 SE

W. Delta 733.98 91.92 a
C. Delta 450.54 65.00 b
E. Delta 204.89 57.01 c
C. Hills 101.62 68.52 c
NE. Hills 82.62 57.01 c

df 4, 45
F value 12.78
(P > F) ANOVA <0.01

Means in a column followed by the same letter are not sig-
nificantly different (a = 0.05; LSMEANS option of PROC
MIXED, SAS Institute 2001).
See Figure 1 for geographical map of Mississippi.
'Between 117-326 Julian days.


sippi (Todd 1975; Mitchell 1979). It seems likely
that seasonal weather patterns including wind
currents and atmospheric disturbances from the
south-central U.S. (Muller 1985; Johnson 1995;


450


400


350


300


z 250


r 200


150
y = -0.024x2 + 12.947x 1609.2
R2 = 0.3564
100 p < 0.001


50


0


0 50 100 150 200 250

Julian Date


Westbrook et al. 1995) could influence the distri-
bution of migratory beet armyworm moths across
the different geographical regions of Mississippi.
Furthermore, even though this pest feeds on nu-
merous hosts (>50) (Mitchell 1979), it is possible
that differences in the population density of local
and migratory moths are due to differences in lar-
val host range and abundance among the differ-
ent geographical regions of Mississippi.
The robust seasonal distribution pattern of
beet armyworm moths in the Mississippi Delta
suggests that the general time for wide-scale mi-
gration of this highly vagile pest may be predicted
in most years (Fig. 3). Although moths were
caught in all months, numbers were very low un-
til approximately 200 Julian days. Hendricks et
al. (1995) suggested that beet armyworm pupae
could overwinter in the Mississippi Delta; how-
ever, without a photoperiod or temperature in-
duced mechanism, pupal diapause seems
unlikely. A more plausible explanation is that
during mild winters, populations of beet army-
worm larvae are able to survive and develop into
pupae by feeding on wild-hosts that survive freez-
ing temperatures in the Mississippi Delta (JJA,
unpublished). In addition, Kim & Kim (1997)


300 350 400


Fig. 3. Distribution of beet armyworm moths throughout Washington Co., W. Delta, Mississippi.











showed that all life-stages of beet armyworms are
able to survive periods of subzero temperatures
due to an efficient supercooling capacity. There-
fore, low numbers of moths caught during the
winter months are probably from local popula-
tions of larvae feeding on fall hosts (JJA, unpub-
lished). The contribution and influence of this
winter population on the summer and fall popula-
tions needs to be further investigated.
The seasonal distribution curve indicates a pre-
dictable period when beet armyworm migrants are
likely to be an economic threat to local crops in the
Mississippi Delta. Moth populations in the Missis-
sippi Delta also were monitored in 2001, but very
low levels of moths were caught in traps, and the
bell-shaped curve was not apparent (data not
shown). Consequently, infestations of larvae on lo-
cal crops and wild-hosts were virtually non-exis-
tent throughout the year in the Mississippi Delta.
Thus, it may be advantageous for consultants,
growers, and researchers to begin monitoring pop-
ulations of beet armyworms at 200 Julian days
(mid-July) to predict if this serious pest will be nu-
merous enough during the season to cause eco-
nomic damage to crops in the Mississippi Delta.

REFERENCES CITED

HENDRICKS, D. E., D. W. HUBBARD, AND D. D. HARDEE.
1995. Occurrence of beet armyworm moths in the
lower Mississippi river delta as indicated by num-
bers caught in traps in 1994. Southwestern Entomol.
20: 157-164.


September 2003


JOHNSON, S. J. 1995. Insect migration in North America
synoptic-scale transport in a highly seasonal environ-
ment, pp. 31-66. In V. A. Gatehouse and V. A. Drake
[eds.] Insect Migration: Tracking Resources through
Space and Time. Cambridge University Press. Cam-
bridge, UK.
KIM, Y., AND N. KIM. 1997. Cold hardiness in
Spodoptera exigua (Lepidoptera: Noctuidae). Envi-
ron. Entomol. 26: 1117-1123.
LITTRELL, R. C., G. A. MILLIKEN, W. W. STROUP, AND R.
D. WOLFINGER. 1996. SAS system for mixed models.
SAS Institute, Cary, NC.
MITCHELL, E. R. 1979. Migration of Spodoptera exigua
and S. frugiperda, North American style, pp. 386-
393. In R. L. Rabb and G. G. Kennedy [eds.] Move-
ment of Highly Mobile Insect: Concepts and Method-
ology in Research. NCSU.
MULLER, R. A. 1985. The potential for the atmospheric
transport of moths from the perspective of synoptic
climatology, pp. 179-202. In D. R. MacKenzie, C. S.
Barfield, G. G. Kennedy, and R. D. Berger [eds.] The
Movement and Dispersal of Agriculturally Impor-
tant Biotic Agents. Claitor's Publishing Division. Ba-
ton Rouge, LA.
SAS INSTITUTE. 2001. Proprietary Software Release 8.2,
Cary, NC, USA.
TODD, E. L. 1975. New distributional records for the
beet armyworm, Spodoptera exigua (Hiibner) (Lepi-
doptera: Noctuidae). USDA Coop. Econ. Insect Rep.
25:14.
WESTBROOK, J. K., J. R. RAULSTON, W. W. WOLF, S. D.
PAIR, R. S. EYSTER, AND P. D. LINGREN. 1995. Field
observations and simulations of atmospheric trans-
port of noctuids from northeastern Mexico and the
south-central U.S. Southwest Entomol. Suppl. 18:
25-44.


Florida Entomologist 86(3)







Armyworm Symposium 2002: Carpenter & Jewett


INFLUENCE OF HERBIVORE-DAMAGED CORN AND COTTON IN THE FIELD
RECRUITMENT OF BRACONID PARASITOIDS FROM FERAL POPULATIONS

J. E. CARPENTER1 AND D. K. JEWETT2
'USDA, ARS, Crop Protection and Management Research Unit, Tifton, GA 31793

2USDA, Forest Service, Southern Research Station, Athens, GA 30602

ABSTRACT

The potential to increase parasitism by Cotesia marginiventris through response to chemical
signals emitted by herbivore-damaged plants was investigated in corn and cotton field plots.
Recruitment of feral C. marginiventris adult females was measured by increased parasitism.
Spodoptera frugiperda larvae placed in the field plots and then recollected experienced a
mean rate of parasitism of approximately 4-6%. Mean total mortality of the collected larvae
ranged from 13 to 20%. We found no significant difference in the level of parasitism, or larval
mortality between field plots containing herbivore-damaged plants and plants that were un-
damaged. Under the conditions of this study, we found no evidence that systemic host plant
volatiles induced by herbivore feeding were used by feral C. marginiventris to improve for-
aging and parasitism at specific sites within a field of corn or cotton.

Key Words: corn, cotton, Cotesia marginiventris, Spodoptera frugiperda, parasitism

RESUME

El potential para el aumentar el parasitismo por Cotesia marginiventris por medio de su res-
puesta a seiales quimicas emitidas por plants daiadas por herbivoros fu6 investigada en
parcelas de campos de maiz y algod6n. El reclutamiento de hembras adults salvajes de
C. marginiventris fu6 medido por el aumento del parasitismo. Larvas de Spodoptera fru-
giperda puestas en parcelas del campo y despu6s recolectadas, experimentaron un promedio
de la tasa de parasitismo de aproximadamente 4-6%. El promedio de la mortalidad total de
las larvas recolectadas fue de 13 a 20%. No encontramos ningun diferencia significativa en el
nivel de parasitismo, o de la mortalidad larvaria entire las parcelas del campo que tenian
plants daiadas por herbivoras y plants no daiadas. Bajo las condiciones de este studio, no
encontramos ningdn evidencia que los volatiles sist6micos de las plants hospederas induci-
dos por la alimentaci6n de herbivoras fueron usados por los C. marginiventris salvajes para
mejorar el forraje y el parasitismo en sitios especificos dentro de un campo de maiz o algod6n.


Following the Boll Weevil Eradication Pro-
gram the beet armyworm, Spodoptera exigua
(Hiibner) (Lepidoptera: Noctuidae), emerged as
the most important threat to cotton production
over large areas of the southeastern United
States (Haney et al. 1996). Other important hosts
of beet armyworms include corn, tomatoes, al-
falfa, onions, asparagus, potatoes, and citrus as
well as numerous non-economic species (Hen-
dricks et al. 1995). Prior to 1991, repeated out-
breaks of S. exigua occurred regularly in Georgia
and elsewhere, e.g., 1977, 1980, 1981, 1988, and
1990 (Douce & McPherson 1991).
The threat of the fall armyworm, S. frugiperda
(J. E. Smith) (Lepidoptera: Noctuidae), to cotton
production also has increased (Riley et al. 1997).
Historically, fall armyworms have not been an im-
portant problem to cotton production. Instead,
they have been associated with corn, sorghum,
and coastal Bermuda grass (Metcalf et al. 1951).
In corn production alone, fall armyworms have
been responsible for losses of $30- to 60-million
annually to corn production (Sparks 1979, 1986).


Additional environmental costs also have accrued
due to management programs that emphasize in-
secticides (Riggin et al. 1994).
Often, less than 0.1% of pesticides applied to
crops reach their target (Pimentel & Levitan
1986); the remainder can affect beneficial insects
and biological control agents (Pimentel et al.
1980; Ripper 1956). For instance, Cotesia (= Ap-
anteles) marginiventris (Cresson) (Hymenoptera:
Braconidae) is an endoparasitoid native to the
southeastern United States (Ashley 1979). It has
a broad host range comprising a variety oftaxa in
the Lepidoptera (Tingle et al. 1978), and it is con-
sidered important to the management of army-
worms (Loke et al. 1983; Lewis & Nordlund 1980;
Ashley 1979). Tillman & Scott (1997) reported
susceptibility of C. marginiventris adults to com-
monly applied rates of selected insecticides, in-
cluding acephate, azinphosmethyl, bifenthrin,
cyhalothrin, cypermethrin, endosulfan, esfenval-
erate, fipronil, methomyl, methyl parathion, ox-
amyl, profenofos or thiodicarb. This vulnerability
has inspired the continued pursuit of manage-







Florida Entomologist 86(3)


ment tactics that either minimize or alternate in-
secticides in order to conserve natural enemies
like C. marginiventris (Loke et al. 1983, Lewis &
Nordlund 1980).
Several studies have demonstrated that para-
sitoids can use factors liberated from the food
plant of their hosts as a method of locating a po-
tential host habitat (Vinson 1975). Many investi-
gators (Cortesero et al. 1997; Loughrin et al.
1994; Turlings & Tumlinson 1992; Turlings et al.
1991; Turlings et al. 1990; Loke et al. 1983) have
reported that corn and cotton plants on which ar-
myworms feed are attractive to C. marginiventris
adult females in laboratory experiments. Organic
volatiles from such plants emitted approximately
18-24 h after commencement of feeding by beet
armyworms and released on a diurnal cycle have
been implicated in this attraction (Turlings et al.
1990; Turlings et al. 1991; Turlings & Tumlinson
1992; Loughrin et al. 1994; Cortesero et al. 1997).
An elicitor in the oral secretion from beet army-
worms induces systematic production and emis-
sion of the attractants from plants (Alborn et al.
1997). Tumlinson et al. (1993) suggested that bio-
logical control may benefit from the use of plant
breeding or genetic engineering to produce
strains of plants that generate greater amounts of
the herbivore-induced plant attractants.
Many aspects of this chemically-mediated
tritrophic relationship have been studied in labo-
ratory, wind tunnel experiments, including the in-
fluence of herbivore-induced plant volatiles on
different host-foraging strategies of C. margini-
ventris and on another more host-specific parasi-
toid Microplitis croceipes (Cresson) (Cortesero et
al. 1997). However, no data have been published
concerning a fundamental aspect of this chemi-
cally-mediated tritrophic relationship, which is
the attraction and increased performance of
C. marginiventris females in the field by plants
experiencing armyworm herbivory. Herein, we in-
vestigate the influence of herbivore-damaged
corn and cotton plants on the recruitment of feral
C. marginiventris adult females as measured by
increased parasitism. Implications for manage-
ment of armyworms are discussed.


MATERIALS AND METHODS

Insect Rearing

Spodoptera frugiperda and S. exigua were ob-
tained from laboratory colonies at the USDA,
ARS Crop Protection and Management Research
Unit laboratory, Tifton, GA. Larvae were reared
in plastic cups (30 ml) containing meridic diet
(Burton 1969) at a photoperiod of 14:10 (L:D) h
and temperature of 28 + 1C, respectively, accord-
ing to the methods of Perkins (1979) unless indi-
cated otherwise.


Experimental Design

Experiments were conducted at the USDA-
ARS-CPMRU research farm in Tifton, GA, using
areas (0.7-1.0 ha) planted in corn (Zea mays L., cv,
Pioneer 3167) and cotton (Gossypium hirsutum
L., cv. Deltapine 90). Three trials conducted in
corn (4-6 leaf stage) were initiated on 5/8/00, 5/22/
00, and 6/29/00, and two trials conducted in cot-
ton (50-75 cm high) were initiated on 6/12/00 and
6/19/00. The field plot design was the same for all
trials. Sixty sites were established in a field plot.
Each site consisted of a designated center plant
and four sentinel plants. Two of the sentinel
plants were positioned 1m on each side of the cen-
ter plant within the same row. The other two sen-
tinel plants were positioned on each side of the
center plant in the adjacent rows (=1 m). A ran-
domized complete block design was used for the
sites within a field plot. There were six blocks sep-
arated by =7 m with 10 sites per block. Sites in
each block were separated by =8 m. Treated and
control sites (5 of each) were randomly assigned
for each block.
For treated sites, six 2nd to 3rd instar S. ex-
igua were placed on a leaf of each center plant.
Larvae were confined on a leaf in feeding disks
(between a pair of ventilated plastic soft-drink
lids held together by three curl clips) as described
by Cortesero et al. (1997). The design was modi-
fied in the present experiment. Instead of lining
the perimeter of the inside face of the lid with
polystyrene foam, cotton batting was used to pre-
vent escape of larvae without inhibiting transpi-
ration by the leaf. Also, instead of covering holes
cut in the soft-drink lids with mesh, soft-drink
lids were stippled with a pin to permit free ex-
change of air (Fig. 1).
In trials involving corn plots, larvae were
placed on the center plants at 7:00 PM, EDST, and
removed from the center plants two days later (at
7:00 PM, EDST) at which time most of the leaf
within the feeding disks had been consumed.
Feeding disks (and larvae and larval frass) were
removed by excising the leaf with a pair of scis-
sors adjacent to the feeding disk but proximal to
the plant stalk. Feeding disks also were placed on
the center plants in the control sites and excised
in a similar fashion, however, the feeding disks
contained no larvae.
Following the removal of all feeding disks, cen-
ter plants and sentinel plants in all sites (both
treated and control) were infested with FAW neo-
nates using a'bazooka' (Wiseman et al. 1980) cal-
ibrated to deliver 20 neonates to the whorl. All
center and sentinel plants were collected after
36 h (7:00 AM, EDST) and all larvae were re-
moved and placed on meridic diet in individual
30-ml plastic cups. Larval mortality and parasit-
ism were recorded. Similar methods were used in
trials involving cotton plants with the following


September 2003







Armyworm Symposium 2002: Carpenter & Jewett


Fig. 1. Cage used to contain Spodoptera exigua lar-
vae and resulting frass on corn or cotton leaves (cotton
shown here) during herbivory trials (see text for de-
scription). Feeding damage can be observed through the
perforated side of the cage. Cage was constructed of a
pair of breathable plastic soft-drink lids, with the inside
perimeter lined with cotton batting, held together by
three curl clips.



exceptions: (1) feeding disks (with and without
larvae) were placed on the center plants at
1:00 PM and removed two days later at 6:00 AM;
(2) after removal of feeding disks, center and sen-
tinel plants were infested with FAW neonates
(6:00 AM) and were collected from the field the
same day at 6:00 PM.

Statistical Analysis

Data collected from corn and cotton field trials
were analyzed using analysis of variance, with
trial, field design block, treatment, trial/treat-
ment interaction, and block/treatment interac-
tion as sources of variation (PROC ANOVA &
PROC GLM) (SAS Institute 1989). Number of lar-
vae collected from the center plants and the sen-
tinel plants, total number of larvae collected,
number of plants with parasitized larvae, number
of center plants with parasitized larvae, total
number of parasitized larvae, and total larval
mortality were the dependant variables. When
significant (P < 0.05) interactions were detected
between trial and treatment or between block and
treatment, these interactions were tested as an
error term. When significant (P < 0.05) differences
were indicated, means were separated by the
Tukey-Kramer statistic or paired t-test at P =
0.05.


RESULTS AND DISCUSSION

Combining data from all trials, which included
field plots comprised of a center corn plant that
was undamaged (control) or damaged (treated) by
herbivore feeding prior to artificial infestation
with larvae and four adjacent (sentinel) plants, a
total of 5,921 S. frugiperda (FAW) larvae were col-
lected. These larvae represent 32.9% of all larvae
(=18,000) with which plants were artificially in-
fested. Of the 5,921 larvae recovered, 2,637 were
from field plots comprised of a center plant dam-
aged by herbivore feeding prior to artificial infes-
tation and 2,225 larvae were collected from field
plots comprised of a center plant undamaged
prior to artificial infestation. The mean (S.D.)
percentage of larvae collected from center plants
in the treated plots (19.97 10.6) was not signifi-
cantly different from the mean percentage of lar-
vae collected from center plants in the control
plots (20.10 10.5) (Table 1).
All of the parasitoids reared from S. frugiperda
larvae collected from the corn plots were C. mar-
giniventris. The mean (S.D.) percent parasitism
of all S. frugiperda larvae collected from treated
plots (4.45 11.3) and control plots (5.74 12.9)
was not significantly different. Likewise, there
was no significant difference between the mean
(S.D.) percent parasitism of S. frugiperda larvae
collected from sentinel plants in treated plots
(5.55 11.3) and control plots (4.43 7.7). The
mean (S.D.) percent parasitism of larvae col-
lected from the center plants of each plot was sig-
nificantly (F = 11.58; df= 2, 35;P = 0.0009) greater
for Trial 3 (10.80 17.7) than for Trial 1 (1.56
4.6) or Trial 2 (2.7 7.2). However, differences be-
tween treated and control field plots with respect
to the mean percent parasitism of larvae collected
from the center plants were not significant (Table
1). Number of plants from which parasitized lar-
vae were recovered, percent mortality (excluding
parasitism) of recovered larvae, and percent mor-
tality (including parasitism) of recovered larvae
(Table 1) were not significantly influenced by her-
bivore feeding on the center corn plant in the
treated plots prior to larval infestation.
Trials conducted in cotton plots yielded results
similar to those conducted in corn plots except the
number of larvae recovered from cotton was less
than the number of larvae recovered from corn. A
total of 730 S. frugiperda larvae were collected
from artificially-infested cotton plants in field
plots comprised of a center plant that was undam-
aged (control) or damaged (treated) by herbivore
feeding prior to larval infestation, and four adja-
cent (sentinel) plants. These 730 larvae represent
a recovery of 6.1% of the total number (=12,000) of
larvae used to infest the plants in the two trials.
343 larvae were collected from field plots contain-
ing a center plant that was damaged by herbivore
feeding prior to larval infestation, and 387 larvae







Florida Entomologist 86(3)


TABLE 1. NUMBER, LOCATION, PARASITISM AND MORTALITY OF SPODOPTERA FRUGIPERDA LARVAE COLLECTED FROM
ARTIFICIALLY-INFESTED CORN PLANTS IN FIELD PLOTS CONTAINING A CENTER PLANT THAT WAS UNDAMAGED
(CONTROL) OR DAMAGED BY HERBIVORE FEEDING PRIOR TO LARVAL INFESTATION, AND FOUR ADJACENT (SEN-
TINEL) PLANTS.


Mean S.D.'


Corn plots with herbivore- Corn plots with
damaged center plant undamaged center plant

Total number of larvae collected 2637 2225
% of collected larvae found on center plants 19.97 10.6 a 20.10 10.5 a
% parasitism of all larvae collected 5.66 10.8 a 4.80 8.0 a
% parasitism of larvae on sentinel plants 5.55 11.3 a 4.43 7.7 a
% parasitism of larvae on center plant in trial 1 1.87 5.0 a 1.26 4.2 a
% parasitism of larvae on center plant in trial 2 4.42 9.3 a 0.99 3.7 a
% parasitism of larvae on center plant in trial 3 6.98 16.2 a 14.50 18.4 a
% of plants with a parasitized larva 17.78 26.6 a 18.00 26.1 a
% mortality of collected larvae (excluding parasitism) 14.79 11.5 a 15.30 10.5 a
% total mortality of collected larvae (including parasitism) 20.45 15.5 a 20.11 11.4 a

1Means in each row followed by the same letter are not significantly different (Tukey-Kramer test, P < 0.05).


were collected from field plots containing a center
plant that was undamaged prior to larval infesta-
tion. Mean (S.D.) percent larvae collected from
center plants in the treated plots (21.96 15.9)
was not significantly different from mean percent
larvae collected from center plants in the control
plots (25.53 17.3) (Table 2).
Similar to the trials conducted in the corn plots,
all of the parasitoids reared from S. frugiperda
larvae collected from the cotton plots were C. mar-
giniventris. There was no significant difference be-
tween mean (S.D.) percent parasitism of all S.
frugiperda larvae collected from treated cotton
plots (3.47 5.0) and control cotton plots (5.92 +
7.5). Likewise, there was no significant difference


between mean (S.D.) percent parasitism ofS. fru-
giperda larvae collected from sentinel plants in
treated plots (3.69 5.7) and control plots (6.06
8.8). Mean (S.D.) percent parasitism of larvae
collected from the center plants of each plot was
significantly (F = 5.14; df = 1, 23; P = 0.0468)
greater for Trial 1 (6.40 10.9) than for Trial 2
(2.12 9.8). However, differences between treated
and control field plots with respect to mean per-
cent parasitism of larvae collected from the center
plants were not significant (Table 2). Herbivore
feeding on the center cotton plant in the treated
plots prior to larval infestation did not signifi-
cantly affect the number of plants from which
parasitized larvae were collected, the percent


TABLE 2. NUMBER, LOCATION, PARASITISM AND MORTALITY OF SPODOPTERA FRUGIPERDA LARVAE COLLECTED FROM
ARTIFICIALLY-INFESTED COTTON PLANTS IN FIELD PLOTS CONTAINING A CENTER PLANT THAT WAS UNDAM-
AGED (CONTROL) OR DAMAGED BY HERBIVORE FEEDING PRIOR TO LARVAL INFESTATION, AND FOUR ADJACENT
(SENTINEL) PLANTS.


Mean S.D.'


Cotton plots with herbivore- Cotton plots with
damaged center plant undamaged center plant

Total number of larvae collected 343 387
% of collected larvae found on center plants 21.96 15.9 a 25.53 17.3 a
% parasitism of all larvae collected 3.47 5.0 a 5.92 7.5 a
% parasitism of larvae on sentinel plants 3.69 5.7 a 6.06 8.8 a
% parasitism of larvae on center plant in trial 1 4.65 + 10.1 a 8.22 11.7 a
% parasitism of larvae on center plant in trial 2 0.57 2.7 a 3.30 12.8 a
% of plants with a parasitized larva 15.33 21.3 a 22.33 28.5 a
% mortality of collected larvae (excluding parasitism) 9.91 11.3 11.67 7.3
% total mortality of collected larvae (including parasitism) 13.38 12.4 a 17.60 10.6 a

Means in each row followed by the same letter are not significantly different (Tukey-Kramer test, P < 0.05)


September 2003







Armyworm Symposium 2002: Carpenter & Jewett


mortality (excluding parasitism) of collected lar-
vae, nor the percent mortality (including parasit-
ism) of collected larvae (Table 2).
This study represents the first field experiments
testing recruitment of C. marginiventris females
from feral populations by herbivore-damaged corn
and cotton plants. The approaches used in these
field experiments are congruent with those de-
scribed previously for laboratory bioassays, accom-
modating such important factors as time during
which volatiles are released from plants (Loughrin
et al. 1994; Turlings et al. 1990), time of day during
which C. marginiventris females forage most ac-
tively and during which they have been used previ-
ously in bioassays (Turlings et al. 1991; Loughrin
et al. 1994), and stage at which S. frugiperda lar-
vae are preferred hosts (Loke et al. 1983; Riggin et
al. 1994). Conclusions drawn previously from the
results of laboratory bioassays concerning recruit-
ment of C. marginiventris females by herbivore-
damaged corn and cotton plants are neither con-
firmed nor contradicted by results of the present
field study. We tested for significant difference in
the level of parasitism, or larval mortality between
field plots containing herbivore-damaged plants
and plants that were undamaged. Under the condi-
tions of this study, we found no evidence that sys-
temic host plant volatiles induced by herbivore
feeding were used by feral C. marginiventris to im-
prove foraging and parasitism at specific sites
within a field of corn or cotton.
In view of our findings, it is interesting to con-
sider the observations made by Ruberson & Whit-
field (1996) in a study of the facultative
parasitization of S. exigua eggs by C. marginiven-
tris, conducted in cotton fields where larval popu-
lations of S. exigua were very low. Because
herbivore-induced plant attractants are lacking
on plants with only egg masses present and are
limited or absent in fields with low or no larval
populations, they surmised that C. marginiven-
tris was quite successful at locating egg masses
even when foraging cues induced by larval feed-
ing were rare. They suggested that female wasps
may have been attracted to the field by the feed-
ing of the few larvae present and then attacked
the egg masses. Also, they concluded that larval
damage is not the only source of cues to which
C. marginiventris is capable of responding in
close-range host location. Our results are congru-
ent with the observations and conclusions of Ru-
berson & Whitfield (1996). Considering the
substantial body of published work on the attrac-
tiveness of herbivore-damaged plant volatiles to
foraging parasitoids and our findings that female
C. marginiventris did not parasitize more larvae
at herbivore-damaged sites than undamaged
sites, we suggest that C. marginiventris may be
attracted to the field by the herbivore-induced
plant volatiles, but then rely on additional cues to
locate the specific site of the host larvae.


Identifying alternative or supplemental tactics
to insecticides for managing armyworms in cotton
and corn continues to be a valuable pursuit.
Among these tactics are biological control and host
plant resistance. Tumlinson et al. (1993) sug-
gested that biological control and host plant resis-
tance may benefit from the use of plant breeding or
genetic engineering to produce strains of plants
that generate greater amounts of the herbivore-in-
duced plant attractants. Although this tritrophic
system is an interesting one with considerable
support from laboratory experiments, data col-
lected from our field study reveal that knowledge
of how this tritrophic system is influenced by fac-
tors in the field is incomplete. Additional field
studies are necessary before conclusions may be
drawn about the potential of exploiting these her-
bivore-induced plant attractants to serve industry
and the agricultural community.

ACKNOWLEDGMENTS

We thank R. J. Caldwell, S. A. Drawdy, and R. W. Gid-
dens (Crop Protection and Management Research Unit,
USDA, ARS, Tifton, GA) for their technical assistance.
J. J. Hamm and B. R. Wiseman (USDA, ARS, Tifton,
GA-retired) made important suggestions concerning
development of techniques for artificial infestation and
recovery of fall armyworms in the field. We appreciate
J. Sivinski, J. J. Hamm, J. W. Lewis and O. G. Marti for
commenting on earlier versions of this manuscript. We
are especially grateful for the leadership of C. E. Rogers,
former Laboratory Director of the Crop Protection and
Management Research Unit, who suggested this study.

LITERATURE CITED

ALBORN, H. T., T. C. J. TURLINGS, T. H. JONES, G. STEN-
HAGEN, J. H. LOUGHRIN, AND J. H. TUMLINSON. 1997.
An elicitor of plant volatiles from beet armyworm
oral secretion. Science 276: 945-949.
ASHLEY, T. R. 1979. Classification and distribution of
fall armyworm parasites. Florida Entomol. 62: 114-
123.
BURTON, R. L. 1969. Mass rearing the corn earworm in
the laboratory. USDA-ARS Series 33-134.
CORTESERO, A. M., C. M. DEMORAES, J. O. STAPEL, J. H.
TUMLINSON, AND W. J. LEWIS. 1997. Comparisons
and contrasts in host-foraging strategies of two lar-
val parasitoids with different degrees of host speci-
ficity J. Chem. Ecol. 23: 1589-1606.
DOUCE, G. K., AND R. M. MCPHERSON (eds.). 1991. Sum-
mary of losses from insect damage and costs of con-
trol in Georgia, 1989. Georgia Agric. Expt. Stn.
Special Pub. 70: 47.
HANEY, P. B., W. J. LEWIS, AND W. R. LAMBERT. 1996.
Cotton production and the boll weevil in Georgia:
history, cost of control, and benefits or eradication.
Univ. of Georgia, Agric. Exp. Sta. Res. Bull. No. 428.
HENDRICKS, D. E., D. W. HUBBARD, AND D. D. HARDEE.
1995. Occurrence of beet armyworm moths in the
lower Mississippi River Delta as indicated by num-
bers caught in traps in 1994. Southwestern Ent. 20:
157-164.











LEWIS, W. J., AND D. A. NORDLUND. 1980. Employment
of parasitoids and predators for fall armyworm con-
trol. Florida Entomol. 63: 433-438.
LOKE, W. H., T. R. ASHLEY, AND R. I. SAILER 1983. Influ-
ence of fall armyworm, Spodoptera frugiperda, (Lepi-
doptera: Noctuidae) larvae and corn plant damage on
host finding in Apanteles marginiventris (Hymenop-
tera: Braconidae). Environ. Entomol. 12: 911-915.
LOUGHRIN, J. H., A. MANUKIAN, R. R. HEATH, T. C. J.
TURLINGS, AND J. H. TUMLINSON. 1994. Diurnal cy-
cle of emission of induced volatile terpenoids by her-
bivore-injured cotton plants. Proc. Natl. Acad. Sci.
91: 11836-11840.
METCALF, C. L., W. P. FLINT, AND R. L. METCALF. 1951.
Destructive and useful insects. McGraw-Hill, New
York. Pp. 399-402.
PERKINS, W. D. 1979. Laboratory rearing of the fall ar-
myworm. Florida Entomol. 62: 87-91.
PIMENTEL, D., AND L. LEVITAN. 1986. Pesticides:
amounts applied and amounts reaching pests. Bio-
Science 36: 86-91.
PIMENTEL, D., A. ANDOW, R. DYSON-HUDSON, D. GALLA-
HAN, S. JACOBSON, M. IRISH, S. KROOP, A. MOSS, I.
SCHREINER, M. SHEPARD, T. THOMPSON, AND B. VIN-
ZANT. 1980. Environmental and social costs of pesti-
cides: a preliminary assessment. Oikos. 34: 127-140.
RIGGIN, T. M., B. R. WISEMAN, D. J. ISENHOUR, AND K.
E. ESPELIE. 1994. Functional response of Cotesia
marginiventris (Cresson) (Hym., Braconidae) to
Spodoptera frugiperda (J. E. Smith) (Lep., Noctu-
idae) on meridic diet containing resistant or suscep-
tible corn genotypes. J. Appl. Ent. 117: 144-150.
RILEY, D. G., G. K. DOUCE, AND R. M. MCPHERSON
(eds.). 1997. Summary of losses from insect damage
and costs of control in Georgia, 1996. Univ. of Geor-
gia, Agric. Exp. Sta. Special Pub. 91.
RIPPER, W. E. 1956. Effects of pesticides on balance of ar-
thropod populations. Annu. Rev. Entomol. 1: 403-438.
RUBERSON, J. R., AND J. B. WHITFIELD. 1996. Faculta-
tive egg-larval parasitism of the beet armyworm,
Spodoptera exigua (Lepidoptera: Noctuidae) by Cote-


September 2003


sia marginiventris (Hymenoptera: Braconidae).
Florida Entomol. 79: 296-302.
SAS INSTITUTE. 1989. SAS user's guide. SAS Institute,
Cary, NC.
SPARKS, A. N. 1979. A review of the biology of the fall ar-
myworm. Florida Entomol. 62: 82-87.
SPARKS, A. N. 1986. Fall armyworm (Lepidoptera: Noc-
tuidae): potential for area-wide management. Flor-
ida Entomol. 69: 603-614.
TILLMAN, P. G., AND W. SCOTT. 1997. Susceptibility of
Cotesia marginiventris (Cresson) (Hymenoptera:
Braconidae) to field rates of selected cotton insecti-
cides. J. Entomol. Sci. 32: 303-310.
TINGLE, F. C., T. R. ASHLEY, AND E. R. MITCHELL. 1978.
Parasites of Spodoptera exigua, S. eridania (Lep.:
Noctuidae) and Herpetogramma bipunctalis (Lep.:
Pyralidae) collected from Amaranthus hybridus in
field corn. Entomophaga 23: 343-347.
TUMLINSON, J. H., W. J. LEWIS, AND L. E. M. VET. 1993.
How parasitic wasps find their hosts. Scientific
American 268: 100-106.
TURLINGS, T. C. J., AND J. H. TUMLINSON. 1992. Sys-
temic release of chemical signals by herbivore-in-
jured corn. Proc. Natl. Acad. Sci. 89: 8399-8402.
TURLINGS, T. C. J., J. H. TUMLINSON, AND W. J. LEWIS.
1990. Exploitation of herbivore-induced plant odors
by host-seeking parasitic wasps. Science 250: 1251-
1253.
TURLINGS, T. C. J., J. H. TUMLINSON, R. R. HEATH, A. T.
PROVEAUX, AND R. E. DOLITTLE. 1991. Isolation of
Allelochemicals that attract the larval parasitoid,
Cotesia marginiventris (Cresson), to the microhabi-
tat of one of its hosts. J. Chem. Ecol. 17: 2235-2251.
VINSON, S. B. 1975. Biochemical coevolution between
parasitoids and their hosts in evolutionary strate-
gies of parasitic insects and mites, pp. 14-48. In P.
Price [ed.] Plenum Press, New York.
WISEMAN, B. R., F. M. DAVIS, AND J. E. CAMPBELL. 1980.
Mechanical infestation device used in fall army-
worm plant resistance programs. Florida Entomol.
63: 425-432.


Florida Entomologist 86(3)







Armyworm Symposium 2002: Farias-Rivera et al.


EFFECT OF LEAF EXTRACTS OF TEOSINTE, ZEA DIPLOPERENNIS L.,
AND A MEXICAN MAIZE VARIETY, CRIOLLO'URUAPENO', ON THE GROWTH
AND SURVIVAL OF THE FALL ARMYWORM (LEPIDOPTERA: NOCTUIDAE)

LUIS ALBERTO FARIAS-RIVERA1, JOSE LUIS HERNANDEZ-MENDOZA2, JAIME MOLINA-OCHOA1
AND ALFONSO PESCADOR-RUBIO3
'Facultad de Ciencias Biol6gicas y Agropecuarias, Universidad de Colima
Apartado postal no. 36, Tecoman, Colima 28100, M6xico

2Centro de Biotecnologia Gen6mica, Instituto Polit6cnico Nacional, Reynosa, Tamaulipas, M6xico

'Centro Universitario de Investigaci6n y Desarrollo Agropecuario, Universidad de Colima
Apartado postal no. 36, Tecoman, Colima 28100, M6xico

ABSTRACT

The effects of leaf extracts of teosinte, Zea diploperennis L., and a Mexican maize variety, cri-
ollo 'Uruapeno', on the growth and survival of the fall armyworm larvae were evaluated un-
der laboratory conditions. Hexane, methanol, and aqueous extractions were made and the
extracts and residual fiber were separately incorporated into a modified Poitout & Bues me-
ridic diet. The hexanic and methanolic extracts were concentrated in a rotary evaporator
and 30 ml of each were mixed to 20 g of cellulose and then incorporated to a meridic diet.
Aqueous leaf extract was not concentrated. Larval cumulative mortality, larval weight, days
to pupation, pupae weight, pupae length and width, days to adult emergence were evalu-
ated. The hexanic extract of both plants enhanced most of the FAW growth parameters.
Methanolic extract and residual fiber of both plants negatively affected the pupae length
and width. The aqueous extract caused 100% of larval cumulative mortality. Larvae fed on
diets containing residual fiber of both plants exhibited antibiotic effects.

Key Words: Spodoptera frugiperda, host plant resistance, maize, corn, antibiosis

RESUME

Fueron evaluados los efectos de los extractos foliares del teosinte, Zea diploperennis L. y los
de una variedad de maiz mexicano, criollo 'Uruapeno', sobre el crecimiento y sobrevivencia
de larvas neonatas del gusano cogollero, bajo condiciones de laboratorio. Las extracciones se
llevaron a cabo con hexano, metanol y agua, y 6stas y la fibra residual s6lida fueron mezcla-
dos dentro de dieta modificada de Poitout y Bues para gusano cogollero, por separado. Los
extractos con hexano y metanol fueron concentrados en un rotaevaporador y 30 ml de cada
uno fueron incorporados a 20 g de celulosa y luego incorporados a la dieta. La fase acuosa no
fue concentrada. La mortalidad total larvaria acumulada, el peso larvario, los dias a pu-
paci6n, el peso de las pupas, la anchura y longitud de las pupas, y dias necesitados por las
pupas para alcanzar el estado adult fueron determinados. La mayoria de los parametros de
crecimiento del gusano cogollero fueron favorecidos por la dietas con extract hexanico de
cada una de las plants, el extract metan6lico y la fibra residual de ambas plants afectaron
negativamente la longitud y anchura de las pupas. Las dietas con el extract acuoso tuvieron
100% de mortalidad larvaria acumulada. Las larvas alimentadas con dietas con fibra resid-
ual de ambas plants mostraron efectos antibi6ticos sobre las larvas del gusano cogollero.

Translation provided by author.


The fall armyworm (FAW), Spodoptera fru-
giperda (J. E. Smith), has been recognized as a
polyphagous insect and important pest of many
crops, particularly maize (corn), Zea mays L., in
the southeastern region of the United States
(Luginbill 1928) and the rest of the Americas (An-
drews 1988).
Over the past five decades, relatively effective
and inexpensive synthetic pesticides have been ex-
tensively used to protect maize from losses caused


by FAW. There is a renewed interest in discovering
new sources of plant resistance to the pests and in
developing plant-derived insecticides (McMillian
et al. 1967; Meisner et al. 1977; Smith & Fischer
1983; Binder & Waiss 1984; Benner 1993; Snook et
al. 1997). Natural plant products can be of benefit
in managing pest populations as well as providing
leads for developing synthetic products with
modes of action against the pests (Balandrin et al.
1985; Shapiro 1991; Benner 1993).







Florida Entomologist 86(3)


Resistance in maize to damage by the FAW has
been extensively investigated (Wiseman 1985;
Wiseman & Widstrom 1986; Wiseman et al. 1992;
Wiseman et al. 1996; Davis et al. 1998; Williams
et al. 2000). Some degree of natural resistance to
FAW larvae is exhibited in some species of corn
for example teosinte, Zea diploperennis, a wild
perennial relative of corn, and centipede grass Er-
emochloa ophiuroides. The defense chemistry of
maize and its relatives is associated with maysin,
chlorogenic acid, caffeoylquinic acids, and other
luteolin derivates (Wiseman et al. 1990; Gueldner
et al. 1991; Gueldner et al. 1992).
We bioassayed leaf extracts from teosinte and
a Mexican maize variety, criollo'Uruapeno', to de-
termine their effects on the growth and survival
of the fall armyworm.

MATERIALS AND METHODS

Insects

Experimental insects were from a colony de-
rived from feral FAW larvae collected in corn-
fields during outbreaks in the State of Colima,
Mexico. Insects were reared on modified meridic
diet for Spodoptera frugiperda (Poitout & Bues
1974). Neonate FAW larvae were used for all bio-
assays.

Teosinte and Maize Variety

Teosinte was collected in the "Reserva de la
Biosfera de Manantlan", near El Terrero, Colima,
Mexico, and the Mexican maize variety, criollo
'Uruapeho', was collected in Cuauhtemoc,
Colima. Plants were grown in the facilities of the
Centro Universitario de Investigaci6n y Desar-
rollo Agropecuario, Universidad de Colima in
Tecoman, Colima using small seedbeds under
nursery conditions, of 24 + 3C, with a photope-
riod of 16:8 (L:D), and 60-75% RH.
Stage 1 plants, exhibiting a collar of 4th leaf
(Hanway 1963) were selected for the extracts.
Portions of leaves, about 12 cm from the whorl,
were cut, and about 50 g of leaves were macerated
and used as samples for the extractions. Then to
each sample, 200 ml of hexane was added, and the
sample was finely ground and homogenized using
a Moulinex turbomix. Samples were incubated
at 25C for approximately 2 h with stirring at 400
rpm. The solution was filtered under vacuum us-
ing Whatman No. 1 filter paper, and the residual
material was washed with 100 ml of hexane. Both
filtrates were combined. Methanol (200 ml) was
then used to extract the residual material using
the same conditions mentioned above. Finally, the
residual material was extracted with 100 ml of
distilled water following the procedures men-
tioned above and the residual material was re-
tained. The hexane and methanol extracts were


concentrated under reduced pressure on a rotary
evaporator (Caframo model OB2000) to a final
volume of 30 ml. Each was incorporated into 20 g
of cellulose and air-dried. Water extracts were not
concentrated but were directly added to the diet.
The hexane, methanol, and water extracts as
well as the residual material (fiber) of both plants
were added separately with 250 g of diet, and 10
ml of diet were dispensed into individual polysty-
rene cups (30 ml) (n = 25 cups per treatment per
trial). Cups of diet were allowed to air-dry for 12
h before each cup was infested with one first in-
star FAW. Diets were arranged in a randomized
complete block design and replicated three times.
Means were separated by Student-Newman-
Keuls test, P < 0.05. Criteria used to assess the ef-
fects of the extracts included: cumulative larval
mortality, larval weight, days to prepupation and
pupation, pupae weight, pupae length and width,
and days required post-pupation to achieve the
adult stage.

RESULTS

The weights of sixth instar and pupae were af-
fected by the leaf extracts of Z. diploperennis and
maize. Heavier larvae and pupae were obtained
when diets containing the hexane and methanol
extracts of both plants and regular diet were
used. Lighter FAW larvae and pupae were ob-
tained when fiber (residual material) diets were
used (Table 1). The 6th instars that fed on diets
with hexane and methanol extracts ofZ. diplope-
rennis were 1.86-fold and 1.56-fold heavier, re-
spectively, than the larvae that fed on diets
containing Z. diploperennis fiber. The weights of
6th instars and pupae reared on diets with hex-
ane and methanol extracts of Z. diploperennis
and the regular diet were statistically similar.
Larvae that developed on diets with hexane and
methanol extracts of 'Uruapeno' were 1.58 and
1.55 fold, respectively, heavier than those larvae
that developed on 'Uruapeno' fiber diets. Weights
of 6th instars and pupae reared on regular diet
were statistically similar to those from diets sup-
plemented with hexane and methanol extracts
(Table 1). Similar effects were observed on
weights of pupae from larvae reared on diets con-
taining hexane and methanol extracts. The mean
weights ranged from 191.0 mg to 211.9 mg for
Z. diploperennis, and from 187.5 mg to 207.4 mg
for 'Uruapeno' (Table 1).
Overall, FAW growth and developmental crite-
ria were affected by leaf extracts and residual fi-
ber. The criteria evaluated from insects reared on
residual fiber diets showed extended developmen-
tal times, diminished pupae length and width,
and increased cumulative larval mortality, com-
pared with those from insects reared on diets con-
taining hexane and methanol extracts of both
plants and regular diet (Table 2).


September 2003







Armyworm Symposium 2002: Farias-Rivera et al.


TABLE 1. WEIGHTS OF SPODOPTERA FRUGIPERDA LARVAE AND PUPAE FED ON A MERIDIC DIET SUPPLEMENTED WITH
LEAF EXTRACTS OF ZEA DIPLOPERENNIS AND A MEXICAN Z. MAIZE VARIETY, 'URUAPENO'.

Extracts and diets

Strain or variety Hexane (n) Methanol (n) Water (n) Fiber (n) Regular (n)

6th instar weights (mg)
Z. diploperennis 407.5 a (25) 343.3 ab (21) 219.0 c (16) 373.0 a (24)
Z. maize var. 'Uruapeno' 322.0 a (19) 315.0 ab (24) -202.8 c (14) 373.0 a (24)
Pupae weights (mg)
Z. diploperennis 211.9 a (16) 191.0 a (14) -142.5 b (12) 198.6 a (22)
Z. maize var. 'Uruapeio' 207.4 a (12) 187.5 a (15) -149.4 b (6) 198.6 a (22)

Means in the same row followed by the same letter are not significantly different (P < 0.05; Student-Newman-Keuls test).


One hundred percent larval mortality was ob-
tained when larvae were fed diets containing wa-
ter extracts. Higher larval survival was obtained
when larvae were fed diets containing hexane ex-
tracts of leaves from Z. diploperennis (100%).
Lower larval mortality was obtained with the
methanol extract of 'Uruapeio' and regular diet
(8.3%) (Table 2). A significant reduction was ob-
served in the FAW survival of larvae fed on fiber
diets of both plants when compared with the FAW
survival of larvae fed the regular diet and diets
containing hexane and methanol extracts of both
plants (Table 1).

DISCUSSION

Research on maize resistance to fall army-
worm and corn earworm (Helicoverpa zea Boddie)
has attempted to correlate resistance to chemistry
of the plant, plant parts, or tissues. The mecha-
nisms of non-preference, antixenosis or tolerance,
and antibiosis are involved in some or all of the
manifestations of plant resistance. Some effects of
antibiosis in resistant crop genotypes on insects,
including extended development, decreased pupal
weight, and increased mortality compared with
individuals reared on susceptible hosts, were de-
scribed by Painter (1951). The presence of maysin,
chlorogenic acid, apimaysin and 3'-methoxymay-
sin in leaves or silks in corn, as possible factors of
resistance to the fall armyworm and corn ear-
worm, was reported by Gueldner et al. (1992).
Maysin in corn, teosinte and centipede grass also
occurs naturally and is considered as a pest bio-
regulator (Gueldner et al. 1991).
The allelochemical activity observed among
the Z. diploperennis and corn extracts in this
study suggests that a variety of plant chemicals
may have biological activities on FAW larvae,
prepupae and pupae. FAW larvae reared on diets
containing water extract and residual fiber were
killed or were significantly lighter than those
reared on diets with hexane and methanol ex-
tracts and regular diet. Differences in larval


growth and development could have caused by an-
tibiosis as a result of the presence of maysin and
its analogues (Wiseman et al. 1992; Molina-Ochoa
et al. 1996), non-preference, or a combination of
the two mechanisms (Williams et al. 1987). Our
results are in accord with those manifestations of
antibiosis observed in resistant genotypes, but we
did not discount the non-preference mechanism,
because it was not evaluated in this study.
We suggest that one or more non-polar com-
pounds that enhanced the growth and develop-
ment of the FAW and diminished the cumulative
larval mortality may be present in the hexane
fraction of both plants. In the methanol fraction of
Z. diploperennis, the presence of polar or polar
compounds affected the pupal length and width.
Diets containing the aqueous fraction and fiber
contained dissolved antibiotic polar compounds
that negatively affected most of the growth and
developmental criteria, and increased the cumu-
lative larval mortality. The survival of larvae
reared on diets containing fiber was reduced 1.84-
fold, 1.46-fold, and 1.69-fold compared to survival
of larvae reared on diets containing hexane and
methanol extracts of Z. diploperennis, and regu-
lar diet, respectively. FAW survival was reduced
2.0-fold and 2.44-fold in larvae reared on fiber di-
ets compared with larvae reared on diets contain-
ing hexane and methanol extracts of'Uruapeio',
and was reduced 2.44-fold compared with larvae
reared on the regular diet, respectively. The aque-
ous extracts of both plants contained the antibi-
otic factors and reduced FAW larval survival to
0%. Larval mortality indicates that the concen-
tration of toxic leaf factors) is greatly increased
in extract-supplemented diets, presumably due to
the removal of phago-stimulants and/or growth-
promoting substances in water extracts (Smith &
Fischer 1983). The higher mortality exhibited in
the diets with water extracts and residual fiber of
both plants indicated that both plants contained
chemicals that severely reduced survival and
negatively affected the growth and development
of FAW larvae. Once these chemicals are removed







Florida Entomologist 86(3)


September 2003


TABLE 2. GROWTH AND DEVELOPMENTAL CRITERIA OF SPODOPTERA FRUGIPERDA LARVAE FED ON MERIDIC DIET SUP-
PLEMENTED WITH LEAF EXTRACTS OF ZEA DIPLOPERENNIS AND A MEXICAN Z. MAIZE VARIETY, 'URUAPENO'.

Extracts and diets

Strain or variety Hexane (n) Methanol (n) Water (n) Fiber (n) Regular (n)

Days to prepupation
Z. diploperennis 18.81 a (16) 19.80 a (15) 29.31 b (13) 18.77 a (22)
Z. maize var. 'Uruapeno' 18.92 a (19) 21.63 ab (24) -27.67 c (14) 18.77 a (22)
Days to pupation
Z. diploperennis 20.94 a (16) 21.14 a (14) 31.33 b (12) 20.77 a (22)
Z. maize var. 'Uruapeio' 20.67 a (12) 23.29 a (14) -28.0 b (6) 20.77 a (22)
Pupal length (cm)
Z. diploperennis 1.61 a (16) 1.51 b (14) 1.46 b (12) 1.64 a (22)
Z. maize var. 'Uruapeio' 1.61 a (12) 1.61 a (14) -1.48 b (6) 1.64 a (22)
Pupal width (cm)
Z. diploperennis 0.53 a (16) 0.47 ab (14) 0.43 c (12) 0.51 a (22)
Z. maize var. 'Uruapeno' 0.49 ab (12) 0.50 a (14) -0.46 c (6) 0.51 a (22)
Days required by pupae to reach the adult stage
Z. diploperennis 10.40 a (16) 11.18 a (11) -10.20 a (5) 11.18 a (22)
Z. maize var. 'Uruapeno' 11.17 a (12) 13.00 a (9) -11.50 a (4) 11.18 a (22)
Cumulative larval mortality
Z. diploperennis 0.0 d (24) 20.8 c(24) 100 a (24) 45.8 b (24) 8.3 c (24)
Z. maize var. 'Uruapeio' 25.0 c(24) 8.3 c (24) 100 a (24) 62.5 b (24) 8.3 c (24)

Means in the same row followed by the same letter are not significantly different (P < 0.05; Student-Newman-Keuls test).


through extraction, the larvae were able to use
the nutrients contained in the leaves. The avail-
ability of these nutrients increased the survivor-
ship to adulthood above those on the residual
fiber and regular diet. Similar results were ob-
tained by Bosio et al. (1990). Our results differ
from those obtained by Quisenberry et al. (1988),
because they did not find differences in mortality
caused by diets containing water extracts of dif-
ferent varieties of Bermudagrass, Cynodon dacty-
lon (L.). Their results suggested that the water
extractable factors did not contribute to FAW re-
sistance in the grasses evaluated.
The possibility of FAW management by foliar
application of the plant tissue extracts of insect
resistant corn varieties or its wild relatives could
also be explored in greenhouse and field condi-
tions. A knowledge of the compounds occurring in
corn "criollo", improved varieties and wild rela-
tives of corn would also facilitate the selection
and aid the development of selective breeding
procedures for varieties with an inherent resis-
tance to FAW in tropical conditions.


ACKNOWLEDGMENTS

The authors express their gratitude to Dr. Carlos
Salazar Silva, Rector of the Universidad de Colima, for
supporting this research. We also appreciate Dr. John E.


Foster, Dr. E. A. Heinrichs (Department of Entomology,
University of Nebraska Lincoln, Lincoln, NE) and Dr.
Steven R. Skoda (USDA-ARS, Midwest Livestock In-
sects Research Laboratory, Lincoln, NE for review of the
manuscript.

REFERENCES CITED

ANDREWS, K. L. 1988. Latin American research on
Spodoptera frugiperda (Lepidoptera: Noctuidae).
Florida Entomol. 71: 630-653.
BALANDRIN, M. F., J. A. KLOCKE, E. S. WURTELE, AND
W. H. BOLLINGER. 1985. Natural plant chemicals:
sources of industrial and medicinal material. Sci-
ence 228: 1154-1160.
BENNER, J. P. 1993. Pesticidal compounds from higher
plants. Pesticide Sci. 39: 95-102.
BINDER, R. G., AND A. C. WAISS, JR. 1984. Effects of soy-
bean leaf extracts on growth and mortality of boll-
worm (Lepidoptera: Noctuidae) larvae. J. Econ.
Entomol. 77: 1585-1588.
BoSIO, C. F., K. D. MCCREA, J. K. NITAO, AND W. G.
ABRAHAMSON. 1990. Defense chemistry of Solidago
altissima: effects on the generalist herbivore Tri-
choplusia ni (Lepidoptera: Noctuidae). Environ. En-
tomol. 19: 465-468.
DAVIS, F. M., W. P. WILLIAMS, AND P. M. BUCKLEY.
1998. Growth responses of southwestern corn borer
(Lepidoptera: Crambidae) and fall armyworm (Lepi-
doptera: Noctuidae) larvae fed combinations of
whorl leaf tissue from a resistant and susceptible
maize hybrid. J. Econ. Entomol. 91: 1213-1218.







Armyworm Symposium 2002: Farias-Rivera et al.


GUELDNER, R. C., M. E. SNOOK, B. R. WISEMAN, N. W.
WIDSTROM, D. S. HIMMELSBACH, AND C. E. COS-
TELLO. 1991. Maysin in corn, teosinte, and centipede
grass, pp. 251-263. In P. A. Hedin (ed.) Naturally oc-
curring pest bioregulators. ACS Symposium Series
No. 449, Washington.
GUELDNER, R. C., M. E. SNOOK, N. W. WIDSTROM, AND
B. R. WISEMAN. 1992. TLC screen for maysin, chloro-
genic acid, and other possible resistance factors to
the fall armyworm and the corn earworm in Zea
mays. J. Agric. Food. Chem. 40: 1211-1213.
HANWAY, J. J. 1963. Growth stages of maize, Zea mays.
Agron. J. 55: 487-492.
LUGINBILL, P. 1928. The fall armyworm. USDA Tech.
Bull. 34: 1-92.
MCMILLIAN, W. W., K. J. STARKS, AND C. M. BOWMAN.
1967. Resistance in corn to the corn earworm, Helio-
this zea, and the fall armyworm, Spodoptera fru-
giperda (Lepidoptera: Noctuidae). Part I. Larval
feeding responses to corn plant extracts. Ann. Ento-
mol. Soc. Am. 60: 871-873.
MEISNER, J., K. R. S. ASCHER, AND M. ZUR. 1977. Pha-
godeterrency induced by pure gossypol and leaf ex-
tracts of a cotton strain with high gossypol content in
the larva of Spodoptera littoralis. J. Econ. Entomol.
70: 149-150.
MOLINA-OCHOA, J., B. R. WISEMAN, R. LEZAMA-GUTIER-
REZ, J. J. HAMM, O. REBOLLEDO-DOMINGUEZ, M.
GONZALEZ-RAMIREZ, AND M. ARENAS-VARGAS. 1996.
Impact of resistant "Zapalote Chico" corn silks on
Spodoptera frugiperda (Lepidoptera: Noctuidae)
growth and development. Vedalia 4: 31-34.
PAINTER, R. H. 1951. Insect resistance in crop plants.
The MacMillan Co. New York, USA.
POITOUT, S., ET R. BUES. 1974. Elevage des chenilles de
vingt-huit especes de Lepidopteres Noctuidae et de
deux especes d' Arctidae sur milieu artificial simple.
Ann. Zool. Ecol. Anim. 6: 431-441.
QUISENBERRY, S. S., P. CABALLERO, AND C. M. SMITH.
1988. Influence of Bermudagrass leaf extracts develop-
ment and survivorship of fall armyworm (Lepidoptera:
Noctuidae) larvae. J. Econ. Entomol. 81: 910-913.


SHAPIRO, J. P. 1991. Phytochemicals at the plant-insect
interface. Arch. Insect Biochem. Biophysiol. 17: 191-
200.
SMITH, M. C., AND N. H. FISCHER. 1983. Chemical fac-
tors of an insect resistant soybean genotype affecting
growth and survival of the soybean looper. Entomol.
Exp. Appl. 33: 343-345.
SNOOK, M. E., A. W. JOHNSON, R. F. SEVERSON, Q. TENG,
R. A. WHITE, JR., V. A. SISSON, AND D. M. JACKSON.
1997. Hydroxygeranyllinalool glycosides from tobacco
exhibit antibiosis activity in the tobacco budworm
(Heliothis virescens (F.)). J. Agric. Food Chem. 45:
2299-2308.
WILLIAMS, W. P., P. M. BUCKLEY, AND F. M. DAVIS.
1987. Feeding response of corn earworm (Lepidop-
tera: Noctuidae) to callus and extracts of corn in the
laboratory. Environ. Entomol. 16: 532-534.
WILLIAMS, W. P., P. M. BUCKLEY, AND F. M. DAVIS.
2000. Vegetative phase change in maize and its as-
sociation with resistance to fall armyworm. Maydica
45: 215-219.
WISEMAN, B. R. 1985. Types and mechanisms of host
plant resistance to insect attack. Insect Sci. Appl. 6:
239-242.
WISEMAN, B. R., AND N. W. WIDSTROM. 1986. Mecha-
nisms of resistance in "Zapalote Chico" corn silks to
fall armyworm (Lepidoptera: Noctuidae) larvae. J.
Econ. Entomol. 79: 1390-1393.
WISEMAN, B. R., R. C. GUELDNER, R. E. LYNCH, AND R.
F. SEVERSON. 1990. Biochemical activity of centi-
pedegrass against fall armyworm larvae. J. Chem.
Ecol. 16: 2677-2690.
WISEMAN, B. R., M. E. SNOOK, D. J. ISENHOUR, J. A.
MIHM, AND N. W. WIDSTROM. 1992. Relationship be-
tween growth of corn earworm and fall armyworm
larvae (Lepidoptera: Noctuidae) and maysin concen-
tration in corn silks. J. Econ. Entomol. 85: 2473-
2477.
WISEMAN, B. R., F. M. DAVIS, W. P. WILLIAMS, AND N.
W. WIDSTROM. 1996. Resistance of a maize popula-
tion, FAWCC(C5), to fall armyworm larvae (Lepi-
doptera: Noctuidae). Florida Entomol. 79: 329-336.







Florida Entomologist 86(3)


September 2003


PATHOGENS AND PARASITIC NEMATODES ASSOCIATED
WITH POPULATIONS OF FALL ARMYWORM
(LEPIDOPTERA: NOCTUIDAE) LARVAE IN MEXICO

JAIME MOLINA-OCHOA1, ROBERTO LEZAMA-GUTIERREZ1, MARTIN GONZALEZ-RAMIREZ1,
MARILU LOPEZ-EDWARDS1, MANUEL A. RODRIGUEZ-VEGA' AND FRANCISCO ARCEO-PALACIOS1
'Facultad de Ciencias Biol6gicas y Agropecuarias, Universidad de Colima
Apartado Postal No. 36, Tecoman, Colima 28100, M6xico

ABSTRACT
Larvae of fall armyworm (FAW), Spodoptera frugiperda (J. E. Smith) and soil samples were
collected in six Mexican states. Larvae were collected from whorl-stage corn, grain sorghum,
forage sorghum, and Sudan grass fields in 64 locations during the summer of 2000, to deter-
mine the occurrence of entomopathogens and parasitic nematodes. A total of 5591 FAW lar-
vae from 64 locations were examined for indigenous FAW biological control agents. Overall
total larval mortality was 3.935%. The larval mortality percent due to entomopathogens and
parasitic nematodes was 3.524%, other causes reached 0.411% of total mortality. Three spe-
cies of entomopathogenic fungi representing two classes, Hyphomycetes (Nomuraea rileyi,
and Hirsutella sp.) and Zygomycetes (Entomophthora sp.) were recovered from FAW larvae,
and two species of Hyphomycetes (Metarhizium anisopliae and Beauueria bassiana) were
isolated from soil samples. An unidentified microsporidian was recovered from four locations
in the State of Jalisco, three from Michoacan, three from Nayarit, and one from Veracruz and
Colima, respectively. Mermithid nematodes were recovered from 24 FAW larvae at three lo-
cations in Nayarit and three larvae were recovered from two locations in Veracruz. Six lar-
vae showing symptoms of viral disease were collected from Sinaloa (2), Jalisco (2),
Michoacan (1), and Nayarit (1). Entomopathogenic nematodes from the genus Heterorhab-
ditis sp. and Steinernema sp. were isolated from soil samples from Colima in one and two lo-
cations, respectively. Steinernema sp., and Heterorhabditis sp. were isolated from soil in one
location in Michoacan. Steinernema sp. was recovered from two locations of Jalisco. In this
survey, N. rileyi, mermithid nematodes, and microsporidia were the most frequent patho-
gens and parasites.

Key Words: Spodoptera frugiperda, biological control, occurrence, survey, maize, mermithid
nematodes, entomopathogenic microorganisms

RESUME
Larvas de gusano cogollero, Spodoptera frugiperda (J. E. Smith) (FAW), y muestras de suelo
se colectaron de 64 localidades en seis estados mexicanos. Las larvas fueron recogidas de
campos cultivados con maiz, sorgo para grano, sorgo forrajero y past Sudan, en estado de
cogollo o verticilio, con la finalidad de determinar la presencia de entomopat6genos y nema-
todos parasitos, durante el verano de 2000. 5591 larvas se colectaron para buscar agents de
control biol6gico indigenous de esta plaga. En general la mortalidad total de larvas fue de
3.935%, la mortalidad larvaria provocada por patagenos y parasitos fue de 3.524%, otras
causes provocaron 0.411%. Tres species de hongos entomopat6genos pertenecientes a dos
classes, los Hyphomycetes (Nomuraea rileyi e Hirsutella sp.) y los Zygomycetes
(Entomophthora sp.) fueron recuperados de larvas de gusano cogollero, y dos species de
Hyphomycetes (Metarhizium anisopliae y Beauveria bassiana) fueron aislados de muestras
de suelos. Un microsporidio no identificado fue recuperado en cuatro localidades de Jalisco,
tres de Michoacan, tres de Nayarit, y una en Veracruz y Colima, respectivamente. Los ne-
matodos mermitidos parasitaron a veinticuatro larvas en tres localidades de Nayarit, asi
mismo, a tres larvas en dos localidades de Veracruz. Las larvas con sintomas de virosis se
presentaron en dos sitios de Sinaloa, dos en Jalisco, una en Michoacan y una en Nayarit. Los
nematodos entomopat6genos de los g6neros Steinernema sp. y Heterorhabditis sp. fueron
aislados de muestras de suelo, presentandose en Colima, en una y dos localidades, respecti-
vamente; en Michoacan ocurrieron en una localidad, respectivamente, pero Steinernema sp.
solo se present en dos localidades de Jalisco. En este inventario, N. rileyi, los nematodos
mermitidos y el microsporidio fueron los pat6genos y parasitos mas frecuentes.

Translation provided by author


The fall armyworm (FAW), Spodoptera frugi- beans and occasionally other crops, in most of the
perda (J. E. Smith), causes considerable economic countries of the Western Hemisphere (Sparks
losses in maize, sorghum, peanuts, cotton, soy- 1986). Control of this pest is usually achieved







Armyworm Symposium 2002: Molina-Ochoa et al.


through the application of synthetic insecticides
(Hruska & Gould 1997), but their high cost, envi-
ronmental contamination, development of resis-
tance to chemicals, and pest resurgence (Colborn
1995; Crowe & Booty 1995) have encouraged the
search for alternatives more compatible with the
environment. Microbial control is an environmen-
tally sound and a valuable alternative to the use
of chemicals for controlling this pest.
Interactions between insect host, environ-
ment, insect host age (Fuxa et al. 1988; Molina-
Ochoa et al. 1996), pathogens and plant to be pro-
tected (Bergman & Tingey 1979; Hamm & Wise-
man 1986; Barbercheck 1993; Wiseman & Hamm
1993; Molina-Ochoa et al. 1997, 1999) determine
the strategies for using pathogens in microbial
control (Hamm 1984). FAW larvae are susceptible
to entomopathogenic bacteria, fungi, nematodes,
protozoa, and viruses (Gardner & Fuxa 1980;
Agudelo-Silva 1986; Hamm et al. 1986; Patel &
Habib 1988; Richter & Fuxa 1990; Lezama-Guti-
errez et al. 1996; Molina-Ochoa et al. 1996; Mo-
lina-Ochoa et al. 1999). The insect host age,
habitat and soil type, pesticide use, agricultural
practices, and location, influence the natural dis-
tribution of biological control organisms (Croft &
Brown 1975; Fuxa 1982; Agudelo-Silva 1986;
Hamm et al. 1986; Sosa-Gomez & Moscardi 1994;
Vanninen 1996; Chandler et al. 1997; Mietkiew-
ski et al. 1997; Molina-Ochoa et al. 2001).
As a result of economic and environmental
concerns, surveys for natural enemies of the FAW
occurring in Mexico have been conducted to de-
velop a better understanding of the pathogen
complex, parasitic nematodes and parasitoids
(Lezama-Gutierrez 2001, Molina-Ochoa 2001,
Molina-Ochoa et al. unpublished). These two sur-
veys conducted in four Mexican states reported
the occurrence of the fungi Beauveria bassiana,
Nomuraea rileyi, and Hirsutella sp., an unidenti-
fied microsporidian, mermithid nematodes, and
an ascovirus affecting FAW larvae. The occur-
rence of the bacterium Bacillus thuringensis and
steinernematid and heterorhabditid nematodes
are reported from soil samples.
This paper reports on the presence of ento-
mopathogens and parasitic nematodes in FAW
larval populations and recovered from soil sam-
ples of corn, grain sorghum, forage sorghum, and
Sudan grass fields from six Mexican states, dur-
ing the summer of 2000.

MATERIALS AND METHODS

Isolation of Entomopathogens from FAW Larvae

During August and September of 2000, FAW
larvae were collected from whorl-stage corn,
grain and forage sorghum, and Sudan grass fields
in 64 locations in the Mexican states of Sinaloa,
Nayarit, Jalisco, Colima, Michoacan, and Ver-


acruz. Concurrently, four soil samples were ob-
tained from each location in all of the states.
Location 43 comprised a combination of collec-
tions from adjacent corn and grain sorghum field
in whorl-stage. Sample size ranged from 33 to 119
FAW larvae per field, but most often sample size
was about 90. The number collected was corrected
by subtracting the number that died from injury
or unknown causes during the first days after col-
lection. Collection data and percent infection by
entomopathogens and parasitic nematodes is pre-
sented in Table 1. Larval mortality due to insect
parasitoids is reported elsewhere (Molina-Ochoa
et al., in press).
Larvae were placed individually in 30 cc plas-
tic cups with regular pinto bean diet (Burton &
Perkins 1989) and maintained in the laboratory
to record the larvae infected by entomopathogens
and parasitic nematodes. Mermithid nematodes
that emerged from larvae were collected and
placed in crystal vials containing 2 ml of 70% eth-
anol. Dead FAW larvae showing signs of fungal
infection were placed in plastic Petri dishes (60 x
10 mm) lined with a piece of 5.5 cm-diameter fil-
ter paper (Whatman No. 1) moistened with sterile
distilled water until the fungus sporulated on the
insect surface. A medium composed of 200 ml of
V8 vegetable juice, 5 g glucose, 2 g yeast extract,
3 g CaCO,, 15 g agar, and 800 ml distilled water
(Fargues & Rodriguez-Rueda 1980) for isolating
the fungus Nomuraea rileyi was used. A Sabou-
raud-dextrosa-agar medium enriched with 1% (w/
v) yeast extract (SDAY), and 500 ppm chloram-
phenicol (Lezama et al. 1996) for growing other
fungus species was used. Entomophthorales were
not isolated.

Isolation of Entomopathogenic Fungi and Nematodes
from Soil

In each location of the six surveyed states, a 2
kg combined soil sample was collected. A soil sub-
sample, about 500 g from four different points a
few meters apart, was obtained by digging to a
depth of 10-15 cm with a small shovel. Soil sam-
ples were deposited into double plastic bags,
tagged, stored in a plastic cooler, and taken to the
laboratory where they were kept at 25C until
processing. The storage time ranged from a few
days to three weeks. Soil was thoroughly mixed
and passed through a 0.4 mm mesh sieve, break-
ing soil lumps and separating any litter.
For isolating entomopathogenic nematodes
and fungi, greater wax moth (GWM) larvae, Gal-
leria mellonella L., were used as bait (Bedding &
Akhurst 1975; Tarasco et al. 1997). From the 2 kg
combined soil sample from each location, two
samples were placed in 1000 ml capacity plastic
pots and five GWM last instar larvae were re-
leased into each pot. Pots were incubated at 25C
in the dark for a 10-day period (Woodring & Kaya








Florida Entomologist 86(3)


September 2003


TABLE 1. GEOGRAPHIC LOCATION, DATE, ALTITUDE, CROP (*), SAMPLE SIZE (N), AND TOTAL PERCENT FAW LARVAE IN-
FECTED BY ENTOMOPATHOGENS AND PARASITIC NEMATODES IN SIX MEXICAN STATES (**) DURING 2000.


Code

C1

C2

C3

C4

C5

C6

C7

C8

C9

C10

Cll

J1

J2

J3

J4

J5

J6

J7

J8

J9

J10

Jll

J12

J13

M1

M2


'Corn (c), Forage Sorghum (fs), Grain Sorghum (gs), and Sudan Grass (sg).
*Colima (C), Jalisco (J), Michoacan (M), Nayarit (N), Sinaloa (S), and Veracruz (V).


Date

08/04

08/04

08/04

08/06

08/06

08/06

08/06

08/06

08/06

08/07

08/07

08/08

08/15

08/15

08/15

08/17

08/17

08/17

08/17

08/17

08/17

08/18

08/18

08/23

08/09

08/09


Location

El poblado,
Coquimatlin
Pueblo Juarez,
Coquimatlin
Amachico,
Coquimatlin
Los mezcales, Comala

El remate, Comala

Carrizalillo, Queseria

Queseria

Villa de Alvarez

Juluapan,
Villa de Alvarez
Tepames, Colima

Estapilla, Colima

Ciudad Guzman

Los pinitos, Tonila

Pialla, Tuxpan

Atenquique, Tuxpan

Canoas, Zapotiltic

Apastepe

Teocuitatlin

Zacoalco de Torres

Acatlan de Juarez

Tlajomulco de Zuifiga

Zapopan

Magdalena

Crucero de Magdalena

Totolin

Santa In6s Tocumbo


Coordinates

19 13.698'N
103 47.722'W
19 10.752'N
103 54.634'W
19 10.667'N
10356.351'W
19 20.811'N
103 47.639'N
19 24.825'N
103 47.639'W
19 25.389'N
103 41.000'W
19 23.362'N
103 34.882'W
19 17.201'N
103 47.030'W
19 18.890'N
103 49.611'W
19 08.231'N
103 37.996'W
19 59.549'N
103 31.140'W
19 40.11'N
103 28.830'W
19 25.343'N
103 32.447W
19 27.293'N
103 28.514'W
19 31.778'N
103 27.851'W
19 34.073'N
103 27.324'W
19 38.060'N
103 30.950'W
20 07.035'N
103 32.704'W
2011.988'N
103 33.806'W
20 25.362'N
103 33.406'W
20 29.396'N
103 28.298'W
20 43.129'N
103 29.041'W
20 53.008'N
103 02.509'W
20 56.300'N
104 02.509'W
19 58.890'N
102 40.183'W
19 44.502'N
102 34.967W


Altitude
(m)

422

279

328

608

817

1550

1304

515

539

519

304
1557


1326

1079

1338

1391

1709

1369

1425

1575

1607

1670

1496

1386

1590

1630


Crop n

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 90

c 96

c 92

c 90

c 93

c 92

c 90

c 90


Infected
larvae (%)

0.000

2.222

0.000

1.111

0.000

0.000

0.000

0.000

1.111

2.222

1.111

2.222

11.111

44.444

11.111

12.111

3.333

1.111

1.111

4.166

4.347

10.000

2.150

9.782

0.000

0.000








Armyworm Symposium 2002: Molina-Ochoa et al.


TABLE 1. (CONTINUED) GEOGRAPHIC LOCATION, DATE, ALTITUDE, CROP (*), SAMPLE SIZE (N), AND TOTAL PERCENT
FAW LARVAE INFECTED BY ENTOMOPATHOGENS AND PARASITIC NEMATODES IN SIX MEXICAN STATES (**)
DURING 2000.


Code

M3

M4

M5

M6

M7

M8

M9

M10

M1l

M12

N1

N2

N3

N4

N5

N6

N7

N8

N9

N10

S1

S2

S3

S4

S5

S6


Date

08/09

08/10

08/10

08/11

08/11

08/11

08/11

08/12

08/12

08/12

08/18

08/18

08/18

08/19

08/19

08/19

08/19

08/20

08/23

08/23

08/21

08/21

08/21

08/21

08/22

08/22


Location

Periban

Cointzio

Cerro "La Esperanza"

Tejaban

Carretera a
Nueva Italia
Presa de Zicuiran

Elcenidor,
Nueva Italia
La Guadalupe
Paracuaro
Las yeguas Paracuaro

El cirian, Nueva Italia

Santa Maria del Oro

El rinc6n, Tepic

El pich6n, Tepic

Xalisco

El refili6n, Xalisco

Compostela

La presa, Compostela

Las lumbres, Acaponeta

Seboruco

Ahuacatlan

Bacurimi, Culiacan

La campana, Culiacan

Pericos, Mocorito

Rancho viejo Mocorito

Aguapepito Mocorito

Comanito Mocorito


Coordinates

1933.106'N
10226.586'W
1941.609'N
10116.398'W
1941.233'N
10118.890'W
1913.342'N
10153.714'W
1903.290'N
10202.458'W
1856.191'N
10154.650'W
1859.651'N
10211.577'W
1907.472'N
10212.519'W
1857.308'N
10216.733'W
1853.661'N
10207.483'W
21 20.121'N
10440.174'W
2132.472'N
10456.123'W
2133.479'N
10456.937W
2119.601'N
10455.060'W
2119.407'N
10455.323'W
2117.858'N
10454.044'W
2113.714'N
10452.162'W
2220.795'N
10518.141'W
2120.850'N
10440.749'W
2106.331'N
10427.427W
2451.688'N
10729.478'W
2458.415'N
10733.517W
2503.574'N
10739.547W
2506.033'N
10743.165'W
2503.861'N
10739.547W
2509.006'N
10739.645'W


Altitude
(m)

1475

1932

1998

587

442

292

350

540

359

255

1160

849

774

1042

964

920

928

48

1134

1120

70

143

80

89

68

91


'Corn (c), Forage Sorghum (fs), Grain Sorghum (gs), and Sudan Grass (sg).
*Colima (C), Jalisco (J), Michoacan (M), Nayarit (N), Sinaloa (S), and Veracruz (V).


Crop

c

c

c

c

c

c

c

fs

fs

c

c

c

c

c

c

c

c

C&gs

c

c

gs

gs

gs

gs

sg

gs
gs


Infected
larvae (%)

0.000

2.222

2.222

0.000

2.222

12.698

1.754

2.222

2.222

0.000

2.222

2.083

4.210

7.476

2.222

21.505

2.222

13.333

2.222

1.111

0.000

2.000

1.052

1.020

1.052

0.000







Florida Entomologist 86(3)


September 2003


TABLE 1. (CONTINUED) GEOGRAPHIC LOCATION, DATE, ALTITUDE, CROP (*), SAMPLE SIZE (N), AND TOTAL PERCENT
FAW LARVAE INFECTED BY ENTOMOPATHOGENS AND PARASITIC NEMATODES IN SIX MEXICAN STATES (**)
DURING 2000.

Altitude Infected
Code Date Location Coordinates (m) Crop n larvae (%)

S7 08/22 La poma Badiraguato 2515.749'N 157 c 100 0.000
10740.739'W c
S8 08/22 La majada Badiraguato 2514.076'N 145 c 92 0.000
10739.781'W
V1 09/02 Seis de Enero, Xalapa 1934.115'N 950 c 91 0.000
9650.207'W
V2 09/02 Altolucero, Almolonga 1935.063'N c 33 6.060
9647.384'W
V3 09/02 Actopan 1934.623'N c 64 1.562
9648.589'W
V4 09/02 Los Gonzalez, Actopan 1931.894'N 432 c 113 0.884
9641.294'W
V5 09/02 Bocana, Actopan 1924.416'N 311 c 119 0.000
9636.731'W
V6 09/03 El volador, Coatepec 1921.594'N 709 c 90 3.333
9651.037'W
V7 09/03 Palmillas 1912.293'N 702 c 59 0.000
9646.221'W
V8 09/03 Tierra Colorada 1913.255'N 46 c 45 4.444
9621.916'W
V9 09/04 Cerro gordo 1925.252'N 443 c 45 0.000
9639.566'W
V10 09/04 Lacumbre 1923.320'N 366 c 66 0.000
9638.807'W

*Corn (c), Forage Sorghum (fs), Grain Sorghum (gs), and Sudan Grass (sg).
**Colima (C), Jalisco (J), Michoacan (M), Nayarit (N), Sinaloa (S), and Veracruz (V).


1988, Bidochka et al. 1998). Larval cadavers were
removed and surface-sterilized with 1% Sodium
hypochlorite for a 3 minute-period, then washed
three times with sterile distilled water and placed
on damp filter paper in a 60 mm diameter sealed
Petri dish, and incubated at 25C for 12 days
(Chandler et al. 1997). Entomopathogenic fungi
from the larvae were isolated using SDAY, with
500 ppm of chloramphenicol (Lezama-Gutierrez
et al. 1996). Fungi were identified by microscopic
inspection of morphological characteristics in situ
or after isolation in SDAY according to the crite-
ria by Brady (1979) and Samson et al. (1988).
The entomopathogenic nematodes were sepa-
rated to genera by identifying coloration of Galle-
ria cadavers according to Woodring & Kaya (1988).
Entomopathogenic viruses and bacteria from
FAW larvae and soil, respectively, have not been
yet isolated or identified.

Geographical Coordinates and Collection Data

A Garmin GPS III PlusTM was used for obtaining
the coordinates and altitude data. Location, date,
place, coordinates, altitude, crop, sample size, and
percentage of infected larvae are shown in Table 1.


RESULTS

In this survey, out of 5591 FAW larvae col-
lected from 64 locations in six Mexican states, the
entomopathogens and parasitic nematodes killed
197 larvae. Overall larval mortality percentage
due to these organisms was 3.524%. Mortality
percentage per location ranged from 0.000 to
44.444% (Table 1). Considering the total mortal-
ity due to entomopathogens and parasitic nema-
todes, 137 (69.54%) larvae were killed by
entomopathogenic fungi, 26 (13.19%) larvae were
killed by microsporidia, six (3.04%) larvae were
killed by viruses, and 28 (14.21%) larvae were
killed by mermithid nematodes. Two classes of
entomopathogenic fungi were collected. The class
Zygomycetes was represented by Entomophthora
sp. which infected a larva from Colima and a
larva from Veracruz. The class Hyphomycetes
was represented by Nomuraea rileyi and Hirsu-
tella sp. which infected 134 and one larvae, re-
spectively. N. rileyi was responsible for 68.020% of
total mortality due to entomopathogens and par-
asitic nematodes, and was the most abundant and
widely distributed, occurring in all the states.
Hirsutella sp. occurred in Sinaloa, only. Mermith-







Armyworm Symposium 2002: Molina-Ochoa et al.


ids were collected from the states of Nayarit (23),
and Veracruz (5), only, and accounted for approx-
imately 14.21% of total mortality of FAW larvae.
The 26 FAW larvae infected by microsporidia
were collected in five of six states sampled (Mi-
choacan (10), Nayarit (9), Jalisco (5), Colima (1),
Veracruz (1) and Sinaloa (0)). These entomo-
pathogens accounted for 13.19% of the total FAW
larval mortality. The symptoms of larvae infected
with these entomopathogens were similar to the
unidentified microsporidia reported by Lezama-
Gutierrez et al. (2001). They often were dry and
fragile when dead, resembling cigarette ashes.
Few FAW larvae showed symptoms of ascovirosis
infection (Hamm et al. 1986). Two were detected
in Jalisco in one location, two in two separate lo-
cations in the same municipality in Sinaloa, one
in Michoacan, and one in Nayarit. The viruses
were not identified by electron microscopy. The
percentage of fall armyworm larvae infected by
pathogens and parasitic nematodes at each loca-
tion is shown in Table 2.

Entomopathogens Isolated from Soil

Two species of entomopathogenic fungi, Metar-
hizium anisopliae and Beauveria bassiana, were
recovered from 10 of 64 soil samples M. anisopliae
was recovered from four of eleven locations in
Colima, and in one location in each of Nayarit,
Jalisco, and Michoacan. Beauveria bassiana was
recovered from two locations in Veracruz and one
in Michoacan.
Two genera of entomopathogenic nematodes,
Steinernema sp., and Heterorhabditis sp. were
collected from soil samples. Steinernematid nem-
atodes were recovered from five of 64 locations
(two locations in Colima, two in Jalisco, and in
one in Michoacan). Heterorhabditid nematodes
were recovered in two locations, one in Colima
and one in Michoacan.

DISCUSSION

Current research efforts are focused on select-
ing native and exotic entomopathogens, which
are highly virulent to arthropod pests, for devel-
oping efficient and environmentally-sound bio-
insecticides. The high susceptibility of fall army-
worm larvae and other lepidopterous pests to
strains ofN. rileyi, B. bassiana and M. anisopliae
has been demonstrated (Bustillo & Posada 1986;
Habib & Patel 1990; Lecuona & Lanteri 1999).
N. rileyi has been reported infecting FAW larvae
naturally in Brasil (Valicente 1989), Venezuela
(Agudelo-Silva 1986), Puerto Rico (Pantoja et al.
1985), Colombia (Vargas & Sanchez 1983), United
States (Fuxa 1982), Mexico (Lezama-Gutierrez
et al. 2001), and other countries. In this survey,
N. rileyi caused 68.020% of the total FAW larval
mortality due to pathogens and parasitic nema-


todes, and was the most abundant and widely dis-
tributed entomopathogen, occurring in each of the
six states surveyed. Similar results were reported
by Lezama-Gutierrez et al. (2001) from a survey
conducted in Colima, Jalisco, and Michoacan.
Entomophthora aulicae was reported attack-
ing FAW larvae on grain sorghum in Georgia
(Hamm 1980; Schwehr & Gardner 1982), and Ar-
gentina (Vera et al. 1995). Entomophthora sp.
and Hirsutella sp. were reported attacking larvae
of this pest with parasitism rates that ranged
from 0.6% to 1.1%, respectively (Lezama-Gutier-
rez et al. 2001). In this study, parasitism rates for
E. sp., and H. sp. were 3.030 and 1.520%, respec-
tively. Total FAW mortality caused by all the
pathogens was 3.524% (197 larvae killed).
An unidentified microsporidian was the third
cause of FAW total larval mortality with 13.19%,
and was similar to that reported by Lezama-Guti-
errez et al. (2001). Most larvae infected with the
microsporidian were collected from Michoacan,
Nayarit, and Jalisco, with 10, 8, and 5 infected
larvae, respectively. The arrangement of the
spores of this microsporidian suggested that this
entomopathogen was neither Nosema nor Vairi-
morpha as previously reported by Gardner &
Fuxa (1980).
A few larvae showed symptoms and signs sim-
ilar to those from ascoviruses, but the identity of
these viruses was not verified by electron micros-
copy. Occurrence of entomopathogenic viruses
has been reported in Latin America, in Puerto
Rico, Argentina, Brasil, and Mexico (Valicente
1989; Pantoja & Fuxa 1992; Vera et al. 1995;
Lezama-Gutierrez et al. 2001).
In this survey, the mermithid nematodes were
important natural enemies of FAW larvae. They
were the second most important mortality factor,
causing 14.21% of total mortality. Nematodes
from the genus Hexamermis have been reported
attacking FAW larvae in Honduras, Brasil, Nica-
ragua, and Argentina (Van Huis 1981; Valicente
1989; Wheeler et al. 1989; Vera et al. 1995).
Mermithids attacking FAW larvae in Mexico were
reported by Alcocer-G6mez & Mendez-Villa (1965).
They found parasitism ranging from 8 to 100%
during a 3-year study. An association between the
pest density and percent of parasitism was deter-
mined. Rainfall also was cited as an important
factor in influencing percent parasitism. In a re-
cent survey conducted in Mexico, mermithid nem-
atodes caused larval mortality ranging from 0.0
to 14.9% in Colima (Lezama-Gutierrez et al.
2001). But in our survey, mermithids were not re-
covered in Colima. However, similar percentages
of mortalities were recorded (0.000 to 15.054%)
from other locations, with the highest rate of par-
asitism from Nayarit and Veracruz. A possible
reason for the difference between the findings of
Lezama-Gutierrez et al. (2001) and those we re-
port is that most of the locations in Colima were








Florida Entomologist 86(3)


September 2003


TABLE 2. PERCENTAGE OF FAW LARVAE INFECTED BY ENTOMOPATHOGENS AND MERMITHIDS AT EACH LOCATION.

Code* N. rileyi Hirsut. Entomoph. Mermithid Microspo. Viruses


0
2.222
0
0
0
0
0
0
1.111
1.111
1.111
0
0
0
2.222
2.222
0
2.222
0
0
1.111
1.111
0
2.222
8.888
44.444
11.111
12.222
2.222
1.111
0
4.166
4.347
7.777
2.150
8.696
0
1.041
3.157
2.803
2.222
6.451
1.111
0
2.222
1.111
0
2.000
0
0
0
0
0


0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.052
0
0


0
0
0
1.111
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0


0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
15.054
1.111
13.333
0
0
0
0
0
0
0
0
0
0


0
0
0
0
1.111
0
0
0
0
0
0
0
0
12.698
1.754
1.111
0
0
0
0
0
0
0
1.111
0
1.111
0
0
2.222
0
1.086
2.222
0
1.053
4.673
0
0
0
0
0
0
0
0
0
0
0
0
0


0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1.111
0
0
2.222
0
0
0
0
0
0
0
0
0
0
0
0
1.042
0
0
0
0
0
0
0
0
0
0
1.052
1.020
0
0
0


*Locations are described in Table 1.
N. rileyi = Nomuraea rileyi, Hirsut. = Hirsutella sp., Entomoph.
crospo. = Microsporidia, Viruses = with virosis signs.


Entomophthora sp., Mermithid = mermithid nematode, Mi-







Armyworm Symposium 2002: Molina-Ochoa et al.


TABLE 2. (CONTINUED) PERCENTAGE OF FAW LARVAE INFECTED BY ENTOMOPATHOGENS AND MERMITHIDS AT EACH
LOCATION.

Code* N. rileyi Hirsut. Entomoph. Mermithid Microspo. Viruses

V1 0 0 0 0 0 0
V2 3.030 0 3.030 0 0 0
V3 0 0 0 1.562 0 0
V4 0 0 0 0 0.884 0
V5 0 0 0 0 0 0
V6 0 0 0 3.333 0 0
V7 0 0 0 0 0 0
V8 2.222 0 0 2.222 0 0
V9 0 0 0 0 0 0
V10 0 0 0 0 0 0

*Locations are described in Table 1.
N. rileyi = Nomuraea rileyi, Hirsut. = Hirsutella sp., Entomoph. = Entomophthora sp., Mermithid = mermithid nematode, Mi-
crospo. = Microsporidia, Viruses = with virosis signs.


different than those sampled by Lezama-Gutier-
rez et al. (2001) during the summer of 1998.
The entomopathogenic fungi and nematodes
were recovered in 26.5% of the soil samples (17 of
64 locations). M. anisopliae and B. bassiana were
isolated in 15.6% of the samples. They occurred in
Colima, Michoacan, Nayarit, and Veracruz, but
were not found in Jalisco and Sinaloa.M. anisopliae
was recovered from five locations in Colima, and
one location each in Michoacan and Nayarit.
B. bassiana was recovered in two locations of Ve-
racruz and one location in Michoacan. In a study
conducted in Szczecin, Poland using soils col-
lected from forests during the spring and autumn,
the entomopathogenic fungi M. anisopliae and
B. bassiana infected wax moth larvae (Mietkiew-


ski et al. 1998), and M. anispoliae was the domi-
nant species. Recently, Lezama-Gutierrez et al.
(2001) reported three species of entomopatho-
genic fungi recovered from soil samples using the
Galleria technique; M. anisopliae, B. bassiana, and
Paecilomyces fumosoroseus, with M. anisopliae be-
ing the most dominant species. In our soil samples,
M. anisopliae was also the most dominant, occur-
ring in 10.9% of the locations, while B. bassiana
occurred only in 4.7% of the locations (Table 3).
Steinerematid and Heterorhabditid nematodes
were found in seven of 64 locations (10.9%). Stein-
ernematid nematodes were recovered from Colima
in two locations, one in Michoacan, and two in
Jalisco. Heterorhabditids occurred in Colima and
Michoacan, in one location, respectively. Low rates


TABLE 3. ENTOMOPATHOGENIC FUNGI (HYPHOMYCETES) AND NEMATODES (RHABDITIDA: STEINERNEMATIDAE AND
HETERORHABDITIDAE) RECOVERED FROM SOIL SAMPLES IN DIFFERENT MEXICAN LOCATIONS.

Code* Location Entomopathogen

C2 Pueblo Juarez, Coquimatlan Heterorhabditis sp.
C5 El remate Metarhizium anisopliae
C5 El remate Steinernema sp.
C6 Carrizalillo Steinernema sp.
C6 Carrizalillo Metarhizium anisopliae
C7 Queseria Metarhizium anisopliae
C10 Tepames Metarhizium anisopliae
J1 Ciudad Guzman Metarhizium anisopliae
M2 Santa In6s, tocumbo Metarhizium anisopliae
M4 Cointzio Beauveria bassiana
M6 Tejaban Steinernema sp.
M7 Carretera a Nueva Italia Heterorhabditis sp.
J2 Los pinitos, Tonila Steinernema sp.
J3 Pialla, Tuxpan Steinernema sp.
N6 Compostela Metarhizium anisopliae
V8 Tierra Colorada Beauveria bassiana
V9 Cerro Gordo Beauveria bassiana

*Locations are described in Table 1.











of entomopathogenic nematode recovery have been
reported in different regions around the world, and
range from 3.9% to 21.4% (Constant et al. 1998;
Tangchitsomkid et al. 1998; Griffin et al. 2000; Rosa
et al. 2000; Lezama-Gutierrez et al. 2001). Soil pH
and type, altitude, habitat, soil temperature, crop-
lands, orchards, pastures, and proximity to coastal
lands were discussed as possible factors affecting
the occurrence of these entomopathogens.
The diversity and distribution of entomopatho-
gens and parasitic nematodes occurring in Mexico
could play an important role in regulating the
FAW larval populations. Additional research is
needed on the identification, biology, and poten-
tial of the microsporidia frequently recovered in
the surveys conducted during 1998 and 2000.
There is also a need to identify the role of
mermithid nematodes as potential biological con-
trol agents. Additional research has already been
conducted at the Universidad de Colima, Mexico,
to identify the steinernematid and heterorhab-
ditid nematodes isolated, and to determine their
potential for biological control of fall armyworm
larvae and other lepidopterous pests.

ACKNOWLEDGMENTS
This work was supported by funds from Universidad
de Colima, Mexico, we thank Dr. Carlos Salazar Silva,
Rector of this institution. The authors express their
gratitude to Dr. J. J. Hamm, and Dr. R. E. Lynch (USDA-
ARS, Crop Protection and Management Research Unit,
Tifton, GA) for their assistance and identification of the
pathogens. We also thank Dr. J. E. Foster, Dr. E. A. Hein-
richs (University of Nebraska Lincoln, Lincoln, NE),
and Dr. S. R. Skoda (USDA-ARS, Midwest Livestock In-
sects Research Laboratory, Lincoln, NE) for review of
the manuscript. We recognize J. J. Molina-Cardenas,
O.F. Aguilar-Meza, G. May-Mora, and M. A. Reyes-
Hernandez for their technical assistance.

REFERENCES CITED

AGUDELO-SILVA, F. 1986. Naturally occurring patho-
gens of Spodoptera frugiperda (Lepidoptera: Noctu-
idae) larvae collected in Venezuela. Florida Entomol.
69: 768-769.
ALCOCER-GOMEZ, L., AND M. MENDEZ-VILLA. 1965.
Estudios preliminares sobre parasitismo en larvas
de Laphigma frugiperda Smith y Abbot, por un nem-
atodo de la familiar mermitidae. Fit6filo 48:5-20.
BARBERCHECK, M. E. 1993. Tritrophic level effects on
entomopathogenic nematodes. Environ. Entomol.
22: 1166-1171.
BEDDING, R. A., AND R. A. AKHURST. 1975. A simple
technique for the detection of insect parasitic rhab-
ditid nematodes in soil. Nematologica 21: 109-110.
BERGMAN, J. M., AND W. M. TINGEY. 1979. Aspects of in-
teractions between plant genotypes and biological
controls. Bull. Entomol. Soc. Am. 25: 275-279.
BIDOCHKA, M. J., J. E. KASPERSKI, AND G. A. M. WILD.
1998. Occurrence of the entomopathogenic fungi
Metarhizium anisopliae and Beauveria bassiana in
soils from temperate and near-northern habitats.
Can. J. Bot. 76: 1198-1204.


September 2003


BRADY, B. L. K. 1979. CMI description of pathogenic
fungi and bacteria, Sets 61 and 62 (Nos. 601-620),
Commonwealth Mycological Institute, England, UK.
BURTON, R. L., AND W. D. PERKINS. 1989. Rearing the
corn earworm and fall armyworm for maize resis-
tance studies, pp. 37-45. In CIMMYT. 1989. Toward
insect resistant maize for the third world: Proceed-
ings of the International Symposium on methodolo-
gies for developing host plant resistance to maize
insects. Mexico, D.F.: CIMMYT.
BUSTILLO-P. A. E., AND F. J. POSADA-F. 1986. Pathoge-
nicity of Nomuraea rileyi isolate to larvae of the
maize borer Spodoptera frugiperda. Revista Colom-
biana de Entomologia 12: 5-15.
COLBORN, T. 1995. Pesticides-how research has suc-
ceeded and failed to translate Science into Policy:
Endocrinological effects on wildlife. Environ. Health
Perspect. 103: 81-86.
CONSTANT, P., L. MARCHAY, M. FISCHER-LE SAUX, S.
BRIAND-PANOMA, AND H. MAULEON. 1998. Natural
occurrence of entomopathogenic nematodes (Rhab-
ditida: Steinernematidae and Heterorhabditidae) in
Guadelopupe islands. Fund. Appl. Nematol. 21: 667-
672.
CROFT, B. A., AND A. W. BROWN. 1975. Responses of ar-
thropod natural enemies to insecticides. Annu. Rev.
Entomol. 20: 285-335.
CROWE, A. S., AND W. G. BOOTY. 1995. A multi-level as-
sessment methodology for determining the potential
for groundwater contamination by pesticides. Envi-
ron. Monitor. Assess. 35: 239-261.
CHANDLER, D., D. HAY, AND A. P. REID. 1997. Sampling
and occurrence of entomopathogenic fungi and nem-
atodes in UK soils. Applied Soil Ecol. 5: 133-141.
FARGUES, J., AND D. RODRIGUEZ-RUEDA. 1980. Sensibil-
it6 des larves de Spodoptera littoralis (Lepidoptera:
Noctuidae) aux hyphomycetesentomopathogenes
Nomuraea rileyi et Paecilomyces fumosoroseus. En-
tomophaga 25: 43-54.
FUXA, J. R. 1982. Prevalence of viral infection in popu-
lations of fall armyworm, Spodoptera frugiperda, in
Southeastern Louisiana. Environ. Entomol. 11: 239-
242.
FUXA, J. R., A. R. RICHTER, AND F. AGUDELO-SILVA.
1988. Effect of host age and nematode strain on sus-
ceptibility of Spodoptera frugiperda to Steinernema
feltiae. J. Nematol. 20: 91-95.
GARDNER, W. A., AND J. R. FUXA. 1980. Pathogens for
the suppression of the fall armyworm. Florida Ento-
mol. 63: 439-447.
GRIFFIN, C. T., R. CHAERANI, D. FALLON, A. P. REID,
AND M. J. DOWNES. 2000. Occurrence and distribu-
tion of the entomopathogenic nematodes Stein-
ernema spp., and Heterorhaditis indica in Indonesia.
J. Helminthol. 74: 143-150.
HABIB, M. E. M., AND P. N. PATEL. 1990. Patogenicidade
de Nomuraea rileyi (Farlow) Samson em larvas de
Spodoptera frugiperda (J. E. Smith, 1797), praga de
milho. Revista de Agricultura Piracicaba 65: 83-90.
HAMM, J. J. 1980. Epizootics of Entomophthora aulicae
in lepidopterous pests of sorghum. J. Invertebr.
Pathol. 36: 60-63.
HAMM, J. J. 1984. Invertebrate pathology and biological
control. J. Georgia Entomol. Soc. 19: 6-13.
HAMM, J. J., AND B. R. WISEMAN. 1986. Plant resistance
and nuclear polyhedrosis virus for suppression of the
fall armyworm (Lepidoptera: Noctuidae). Florida
Entomol. 69: 541-549.


Florida Entomologist 86(3)







Armyworm Symposium 2002: Molina-Ochoa et al.


HAMM, J. J., S. D. PAIR, AND O. G. MARTI. 1986. Inci-
dence of host range of a new ascovirus isolated from
fall armyworm, Spodoptera frugiperda (Lepidop-
tera: Noctuidae). Florida Entomol. 69: 524-531.
HRUSKA, A. J., AND F. GOULD. 1997. Fall armyworm
(Lepidoptera: Noctuidae)and Diatraea lineolata
(Lepidoptera: Pyralidae): Impact of larval popula-
tion level and temporal occurrence on maize yield in
Nicaragua. J. Econ. Entomol. 90: 611-622.
LECUONA, R. E., AND A. A. LANTERI. 1999. Control mi-
crobiano con hongos entomopat6genos en la Argen-
tina. Revista de la Sociedad Entomol6gica Argentina
58: 301-306.
LEZAMA-GUTIERREZ, R., R. ALATORRE-ROSAS, L. F. Bo-
JALIL-JABER, J. MOLINA-OCHOA, M. ARENAS-VAR-
GAS, M. GONZALEZ-RAMIREZ, AND O. REBOLLEDO-
DOMINGUEZ. 1996. Virulence of five entomopatho-
genic fungi (Hyphomycetes) against Spodoptera fru-
giperda (Lepidoptera: Noctuidae) eggs and neonate
larvae. Vedalia 3: 35-39.
LEZAMA-GUTIERREZ, R., J. J. HAMM, J. MOLINA-OCHOA,
M. LOPEZ-EDWARDS, A. PESCADOR-RUBIO, M.
GONZALEZ-RAMIREZ, AND E. STYER. 2001. Occur-
rence of entomopathogens of Spodoptera frugiperda
(Lepidoptera: Noctuidae) in the mexican states of
Michoacan, Colima, Jalisco and Tamaulipas. Florida
Entomol. 84: 23-30.
MIETKIEWSKI, R. T., J. K. PELL, AND S. J. CLARK. 1997.
Influence of pesticide use on the natural occurrence
of entomopathogenic fungi in arable soils in the UK,
field and laboratory comparisons. Biocontrol Sci.
Technol. 7: 565-575.
MIETKIEWSKI, R., M. DZIEGIELEWSKA, AND K. JANOW-
ICZ. 1998. Entomopathogenic fungi isolated in the vi-
cinity of Szczecin. Acta Mycologica 33: 123-130.
MOLINA-OCHOA, J., J. J. HAMM, R. LEZAMA-GUTIERREZ,
L. F. BOJALIL-JABER, M. ARENAS-VARGAS, AND M.
GONZALEZ-RAMIREZ. 1996. Virulence of six ento-
mopathogenic nematodes (Steinernematidae and
Heterorhabditidae) on immature stages of
Spodoptera frugiperda (Lepidoptera: Noctuidae).
Vedalia 3: 25-29.
MOLINA-OCHOA, J., B. R. WISEMAN, R. LEZAMA-GUTIER-
REZ, J. J. HAMM, O. REBOLLEDO-DOMINGUEZ, M.
GONZALEZ-RAMIREZ, AND M. ARENAS-VARGAS. 1997.
Impact of resistant "Zapalote Chico" corn silks on
Spodoptera frugiperda (Lepidoptera: Noctuidae)
growth and development. Vedalia 4: 31-34.
MOLINA-OCHOA, J., R. LEZAMA-GUTIERREZ, J. J. HAMM,
B. R. WISEMAN, AND M. LOPEZ-EDWARDS. 1999. Inte-
grated control of fall armyworm (Lepidoptera: Noctu-
idae) using resistant plants and entomopathogenic
nematodes (Rhabditida: Steinernematidae). Florida
Entomol. 82: 263-271.
MOLINA-OCHOA, J., J. J. HAMM, R. LEZAMA-GUTIERREZ,
M. LOPEZ-EDWARDS, M. GONZALEZ-RAMIREZ, AND A.
PESCADOR-RUBIO. 2001. A survey of fall armyworm
(Lepidoptera: Noctuidae) parasitoids in the Mexican
states of Michoacan, Colima, Jalisco, and Tamauli-
pas. Florida Entomol. 84: 31-36.
PANTOJA, A., C. M. SMITH, AND J. F. ROBINSON. 1985.
Natural control agents affecting Spodoptera fru-
giperda (Lepidoptera: Noctuidae) infesting rice in
Puerto Rico. Florida Entomol. 68: 488-490.
PANTOJA, A., AND J. R. FUXA. 1992. Prevalence of biotic
control agents in the fall armyworm Spodoptera fru-
giperda (J. E. Smith) (Lepidoptera: Noctuidae). Fo-
lia. Entomol. Mex. 84: 79-84.


PATEL, P. N., AND M. E. M. HABIB. 1988. Protozoosis
caused by Vairimorpha necatrix microsporidia Nose-
matidae in larvae of Spodoptera frugiperda Lepi-
doptera Noctuidae. Rev. Bras. Zool. 5: 593-598.
RICHTER, A. R., AND J. R. FUXA. 1990. Effect of Stein-
ernema feltiae on Spodoptera frugiperda and Helio-
this zea (Lepidoptera: Noctuidae) in corn. J. Econ.
Entomol. 83: 1286-1291.
ROSA, J. S., E. BONIFAASSI, J. AAMARL, L. A. LACEY, N.
SIMOES, AND C. LAUMOND. 2000. Natural occurrence
of entomopathogenic nematodes (Rhabditida: Stein-
ernema, Heterorhabditis) in the Azores. J. Nematol.
32: 215-222.
SAMSON, R. A., H. C. EVANS, AND J. P. LATGE. 1988. At-
las of entomopathogenic fungi. 187 pp. Springer-
Verlag, Berlin, Germany.
SCHWEHR, R. D., AND W. A. GARDNER. 1982. Disease in-
cidence in fall armyworm and corn earworm popula-
tions attacking grain sorghum. J. Georgia Entomol.
Soc. 17: 38-46.
SOSA-GOMEZ, D. R., AND F. MOSCARDI. 1994. Effect of
till and no-till soybean cultivation on dynamics of
entomopathogenic fungi in the soil. Florida Entomol.
77: 284-287.
SPARKS, A. N. 1986. Fall armyworm (Lepidoptera: Noc-
tuidae) potential for area-wide management. Florida
Entomol. 69:603-614.
TANGCHITSOMKID, N., S. SONTIRAT, NUCHANART-TANG-
CHITSOMKID, AND SUEBSAK-SONTIRAT. 1998. Occur-
rence of entomopathogenic nematodes in Thailand.
Kasetsart Journal Natural Sciences 32: 347-354.
TARASCO, E., C. DE BIEVRE, B. PAPIEROK, M. POLISENO,
O. TRIGGIANI, AND C. DE BIEVRE. 1997. Occurrence
of entomopathogenic fungi in soils in Southern Italy.
Entomologica 31: 157-166.
VALICENTE, F. H. 1989. Levantamento dos inimigos
naturals de Spodoptera frugiperda (J. E. Smith,
1797) (Lepidoptera: Noctuidae) em diferentes re-
gioes de Minas Gerais. Annais da Sociedade Ento-
mologica do Brasil 18: 119-130.
VAN HUIS, A. 1981. Integrated pest management in the
small farmer's maize crop in Nicaragua. Med. Land.
Wageningen 81: 93-100.
VANNINEN, I. 1996. Distribution and occurrence of four
entomopathogenic fungi in Finland: Effect of geo-
graphical location, habitat type and soil type. Mycol.
Res. 100: 93-101.
VARGAS, M. L., AND G. SANCHEZ-G. 1983. Natural con-
trol of some pests of the rice varieties IR-22 and CICA-
6. Revista Colombiana de Entomologia 9: 50-54.
VERA, M. L., L. VALVERDE, S. B. POPICH, AND Z. D. AJMAT-
DE TOLEDO. 1995. Preliminary evaluation of natural
enemies ofSpodoptera frugiperda (J. E. Smith) (Lep-
idoptera: Noctuidae) in Tucuman, Argentina. Acta
Entomologica Chilena 19: 135-141.
WHEELER, G. S., T. R. ASHLEY, AND K. L. ANDREWS.
1989. Larval parasitoids and pathogens of the army-
worm in Honduran maize. Entomophaga 34: 331-340.
WISEMAN, B. R., AND J. J. HAMM. 1993. Nuclear polyhe-
drosis virus and resistant corn silks enhance mortal-
ity of corn earworm (Lepidoptera: Noctuidae) larvae.
Biological Control 3: 337-342.
WOODRING, J. L., AND H. K. KAYA. 1988. Steinernema-
tid and Heterorhabditid nematodes: A handbook of
techniques. Southern Cooperative Series Bulletin 331.
30 pp. Arkansas Agricultural Experiment Station,
Fayetteville.







Florida Entomologist 86(3)


September 2003


PARASITOIDS AND PARASITES OF SPODOPTERA FRUGIPERDA
(LEPIDOPTERA: NOCTUIDAE) IN THE AMERICAS
AND CARIBBEAN BASIN: AN INVENTORY

JAIME MOLINA-OCHOA1, JAMES E. CARPENTER2 E. A. HEINRICHS3 AND JOHN E. FOSTER3
'Facultad de Ciencias Biol6gicas y Agropecuarias, Universidad de Colima
Apartado postal no. 36, Tecoman, Colima 28100, M6xico

2United States Department of Agriculture, Agricultural Research Service
Crop Protection & Management Research Laboratory, P.O. Box 748, Tifton, GA 31793-0748, USA

3University of Nebraska Lincoln, Department of Entomology
312F Plant Industry Building, Lincoln, NE 68583-0816, USA


ABSTRACT

An inventory of parasitoids and parasites of fall armyworm (FAW), Spodoptera frugiperda
(J. E. Smith), was conducted using references describing parasitized FAW eggs, larvae, pu-
pae and adults collected from different crops or habitats throughout the Americas and the
Caribbean Basin. The crops and countries where these parasites were reported occurring in
the Americas is also inventoried. Maize was the crop where the FAW was more frequently
collected followed by rice. Overall, Chelonus insularis (Cresson) had the broadest natural
distribution in the Americas. For the North American region C. insulares, Chelonus sp., and
Euplectrus platyhypenae (Howard) were the most relevant parasitoids. In Central America,
C. insularis was the most prevalent parasitoid, and in the South American region the most
prevalent parasites were Archytas incertus (Macq.),A. marmoratus (Tns.), C. insularis, and
Meteorus laphygmae (Viereck). Diapetimorpha introita (Cresson) is the most important pu-
pal parasitoid of FAW occurring mainly in North America. An acugutturid, Noctuidonema
guyanense (Remillet & Silvain), is the most important ectoparasitic nematode attacking
adults of FAW and other noctuid moths in South and Southeastern US, and Mexico in North
America, Caribbean Basin, Central America, and Northern South America.

Key Words: parasitoids, fall armyworm, Chelonus insularis, Diapetimorpha introita, Noctu-
idonema guyanense, maize, natural distribution, biological control

RESUME

Un inventario de los parasitoides y parasitos del gusano cogollero, Spodoptera frugiperda (J.
E. Smith) se llev6 a cabo usando referencias relacionadas con parasitos de huevos, larvas,
pupas y adults del insecto plaga colectados de diferentes cultivos en su ambito de distribu-
ci6n. Ademas, un inventario se realize de los cultivos y pauses donde estos parasitos atacaron
al gusano cogollero en Am6rica. La plaga fue colectada principalmente en el maiz, seguido
por el arroz. Chelonus insularis (Cresson) fu6 el parasitoide distribuido mas ampliamente en
todo el ambito de distribuci6n del. C. insulares, Chelonus sp. y Euplectrus platyhypenae
(Howard) fueron los parasitoides mas prevalecientes en Norteam6rica. En Centroam6rica,
C. insularis fue el parasitoide mas prevalente, y en la region Sudamericana lo fueronArchy-
tas incertus (Macq.),A. marmoratus (Tns.), C. insularis y Meteorus laphygmae (Viereck). Di-
apetimorpha introita (Cresson) fue el parasitoide de pupas mas important y 6ste habit en
Norteam6rica, principalmente. Un acugguturido, Noctuidonema guyanense (Remillet & Sil-
vain) fue el nematodo ectoparasito mas important atacando adults de gusano cogollero y
otros noctuidos en el Sur y Sureste de los Estados Unidos de Am6rica y M6xico en Nortea-
m6rica, Cuenca del Caribe, Centroam6rica y Norte de Sudam6rica.

Descriptores: parasitoides, gusano cogollero, Chelonus insularis, Diapetimorpha introita,
Noctuidonema guyanense, maiz, ambito de distribuci6n, control biol6gico


Biodiversity in agro-ecosystems can be as var- biodiversity can be used for improved pest man-
ied as the crops, weeds, arthropods, and microor- agement (Altieri 1991). A major problem in all ar-
ganisms themselves, and may differ according to eas of agriculture is the lack of basic research on
geographical location, climate, soil, and human taxonomy of insect pests and their natural ene-
factors. Experimental evidence suggests that mies. This problem is greatest in tropical coun-







Armyworm Symposium 2002: Molina-Ochoa et al.


tries where the needs are also greater (Claridge
1991). Waage (1991) stated that agricultural sys-
tems are generally simpler than the natural hab-
itats from which they are developed. These
systems have fewer plant species, fewer primary
consumers, and generally fewer natural enemy
species. In tropical systems, despite their great
biodiversity, natural enemies could be particu-
larly susceptible to local extinction as a result of
habitat destruction and unfavorable cropping
practices such as indiscriminate use of pesticides
(Claridge 1991; Waage 1991).
More than 200 years ago, the fall armyworm
(FAW), Spodoptera frugiperda (J. E. Smith), was
recognized as a destructive pest of many agricul-
tural crops (Luginbill 1928). In the continental
United States the costs for chemical control and
losses due to this pest exceeded $300,000,000 dur-
ing 1977 (Gross & Pair 1986). The biological con-
trol of FAW in areas of overwintering and
throughout its annual geographical distribution
is a highly desirable alternative to conventional
control methods (Gross & Pair 1986). Luginbill
(1928) and Vickery (1929) recognized the value of
parasitoids in reducing larval populations of
FAW. Parasitoids and parasites can be highly ef-
fective at little or no cost, serve as biotic insecti-
cides in place of chemicals, provide long-term
control without the target pest developing signif-
icant resistance to them, and impose minimal or
no harm to humans or the environment (Wilson &
Huffaker 1976; Stary & Pike 1999).
Sivasubramaniam et al. (1997), and Dent
(2000) advocated that the first step in any inves-
tigation of the role of natural enemies in pest con-
trol should involve a field survey to determine
which species are present and how their numbers
vary in relation to those of the pest insects. Sur-
veys on the FAW parasitoids and other natural
enemies in different parts of its range have been
conducted because of increasing economic and en-
vironmental concerns (Carrillo 1980; Ashley
1986; Castro et al. 1989; Gross & Pair 1991; Cave
1993; Lezama-Gutierrez et al. 2001; Molina-
Ochoa et al. 2001). However, information about
distributions and host plants of the FAW and the
accompanying parasites and parasitoids are scat-
tered throughout the published literature, and
most reviews of FAW parasitoids have empha-
sized those attacking the egg and larval stages
(Ashley 1986). The aim of this paper is to summa-
rize the information and provide an inventory of
the known FAW parasites and parasitoids occur-
ring in the Americas and the Caribbean, indicat-
ing the host stage attacked, the crops from which
parasitized fall armyworm were collected, and
the country of collection. In addition, because par-
asites and parasitoids of the pupal and adult
stages of FAW have received little attention in
most previous reviews on this subject, we provide
an expanded discussion of these natural enemies.


MATERIALS AND METHODS

Sources of Information

Research was conducted to obtain information,
papers, and bibliographic references reporting
the collection of parasitized FAW from the field.
In the US, we used the Agricola and CAB Ab-
stracts database at the University of Nebraska-
Lincoln in Lincoln, Nebraska, and the University
of Georgia, Coastal Plain Experiment Station in
Tifton, Georgia. We collected references cited in
reviews and catalogs of FAW parasites (Guima-
raes 1971; Marsh 1978; Ashley 1979; Ashley
1986; Andrews 1988) to verify information con-
cerning host plant of parasitized FAW, location of
the collected FAW, and stage of FAW attacked. We
also used various internet search engines to iden-
tify published reports of parasitized FAW col-
lected from the field. In Mexico, we collected
papers from Latin American journals and pro-
ceedings from meetings of International and Mex-
ican Entomological and Biological Control
Societies. Information was also obtained from the
libraries in the Facultad de Ciencias Biol6gicas y
Agropecuarias (School of Biological, Agricultural
& Animal Sciences) of the Universidad de Colima,
and in the Centro Nacional de Referencia de Con-
trol Biol6gico (CNRCB)-Comisi6n Nacional de
Sanidad Agropecuaria-SAGARPA (National Cen-
ter for Reference on Biological Control) in
Tecoman, Colima, Mexico, during 2001 and 2002.

Organization of the Information

The classification (order, family, genus and
species) of each parasite and parasitoid collected
from FAW is presented in Table 1. The crop from
which parasitized FAW were collected, the FAW
stage attacked, the country from which parasit-
ized FAW were collected, and the bibliographic
references for each record were included in Table
1 whenever the information was available. We
preferred to list only original references that re-
port the collection of FAW from the field. There-
fore, review articles that contain lists of insects
collected by other authors (Guimaraes 1971;
Marsh 1978; Ashley 1979; Ashley 1986; Andrews
1988) usually are omitted for the entries in Table
1. Because of the quantity involved, all references
listed in Table 1 were not cited in the text. Data
on FAW parasitoids and parasites from Table 1
have been summarized as the number of species
in each taxon reported from different geographi-
cal regions (Table 2). Number of species and FAW
stage attacked reported from different countries
(Table 3), and number of species in each taxon re-
ported from FAW collected from different host
plants (Table 4). Omission of pertinent literature
from this paper is the authors' responsibility and
was unintentional.









TABLE 1. FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIBBEAN BASIN WITH
ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUNTRIES OF COLLEC-
TION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Diptera: Bombyliidae
Poecilanthrax (Anthrax) lucifer (Fabricius)
Diptera: Phoridae
Megaselia sp.

Diptera: Sarcophagidae
Helicobia morionella (Aldrich)
Syn: Sarcophaga morionella Aldrich
Rivinia assidua (Walker)
Syn: Sarcophaga assidua (Walker)
Sarcophaga georgiana (Weideman)
Sarcophaga lambens (Weideman)
Sarcophaga sp.

Sarcodexia sternodontis (Townsend)
Diptera: Tachinidae
Tachinidae sp.

Acroglossa vetula (Reinhard)
Syn: Spallanzania vetula (Reinhard)


Admontia degeerioides (Coquillett)
Archytas analis Fabricius






Archytas apicifer (Walker)
Archytas incasana Townsend
Syn:Archytas divisus (Walker)


L US


L Honduras
Nicaragua

L Honduras
Nicaragua
L US


US
Lesser Antilles
US
Venezuela
Honduras


L Mexico
Nicaragua
L Brazil
Honduras
Venezuela

L US
L Argentina
Barbados
Honduras
Mexico
Nicaragua
US
Venezuela
L US
L Argentina
Brazil
Chile
Venezuela


(N/G)
M

(N/G)
M
(N/G)

M
M
(N/G)
(N/G)
M

M,S
M
(N/G)
(N/G)
M
(N/G)
(N/G)
M
M
M,R,O,T
M
M
(N/G)
(N/G)
(N/G)
M
(N/G)
M
(N/G)


Allen 1921

Cave 1993
Maes 1989

Cave 1993
Maes 1989
Luginbill 1928

Dew 1913
Fennah 1947
Enkerlin 1975
Teran 1974
Maes 1989; Cave 1993

Lacayo 1977
Ryder & Pulgar 1969; Ashley 1986
Guimaraes 1977
Cave 1993
Notz 1972
Teran 1974
Luginbill 1928
Virla et al. 1999
Alam 1979
Cave 1993
Ravlin & Stehr 1984
Maes 1989
Luginbill 1928
Teran 1974
Ravlin & Stehr 1984
Virla et al. 1999
Guimaraes 1977
Etcheverry 1957
Teran 1974








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Archytas incertus (Macquart)


L Argentina


Barbados
Brazil


Chile
Mexico
Puerto Rico

Suriname
Trinidad
US


Archytas marmoratus (Townsend)


Uruguay
L Argentina


Barbados
Brazil

Chile
Cuba
Ecuador
Guadeloupe
Honduras

Lesser Antilles
Mexico
Nicaragua
Peru
Puerto Rico
Suriname
Trinidad
US


M
SY
(N/G)
M
M


(N/G)

M
SC
SC
SC,M
M,PN
M
M,PN
(N/G)
(N/G)
M
SY
(N/G)
M
M
(N/G)
(N/G)
(N/G)
Ve
M
M
P,S,M
M
M,S
M
M
R
N/G)
M
A


Virla et al. 1999
Molinari & Avalos 1997
Parker et al. 1953
Alam 1979
Lucchini & Almeida 1980; Patel & Habib 1982, 1984,
1986; Silveira et al. 1987; Valicente 1989; Silva et al.
1997
Parker et al. 1953; Guimaraes 1977; Milward et al.
1991a,b,c,d,e
Etcheverry 1957
Curran 1927
Van Dine 1913
Jones 1913
Segeren & Sharma 1979
Hynes 1942
Vickery 1929
Luginbill 1928
Parker et al. 1953; Silveira & Ruffinelli 1956
Valicente & Barreto 1999; Virla et al. 1999
Molinari & Avalos 1997
Avalos 1988
Alam 1979
Valicente 1989
Guimaraes 1977
Valencia & Valdivia 1973
Bruner et al. 1975
Benzing et al. 2000
Malausa 1981, 1983
Canas & O'Neil 1998
Cave 1993
Fennah 1947
Pair et al. 1986
Huis 1981; Maes 1989; Lacayo 1977
Sarmiento & Razuri 1978
Pantoja et al. 1985; Pantoja & Fuxa 1992
Van Dither 1960
Yaseen 1979
Soteres et al. 1984








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Archytas marmoratus (Townsend) L (M


Venezuela


Archytas plangens Curran


Archytas sp.


Chetogena sp.
Cuphocerini sp.
Eucelatoria armigera (Coquillett)


Eucelatoria australis (Townsend)
Eucelatoria bryani Sabrosky


Eucelatoria guimaraesi
Eucelatoria rubentis (Coquillet)


L Argentina
Brazil
Honduras

Nicaragua
Trinidad
L Argentina

Brazil
Chile
Honduras
Mexico
Nicaragua
US
L Honduras
L Argentina
L Cuba
US
Venezuela
L Peru
L Honduras
Nicaragua
US
L Brazil
L US


M,C
M,S
C,A,M,S
M,BG,Mi
(N/G)
M
(N/G)
M
(N/G)
M
(N/G)
M
M
M
SY
(N/G)
PN
M
M
M
PN
M
(N/G)
(N/G)
PN
(N/G)
(N/G)
S
M
S,PN
S,PN
M
M,C
S,PN


Campos 1965; Enkerlin 1975; Hogg et al. 1982; Gross
& Pair 1986, 1991; Riggin et al. 1992, 1993
Tingle et al. 1994
Rohlfs & Mack 1985; Pair et al. 1986; McCutcheon 1991
Butler 1958a
Reed 1980
Ravlin & Stehr 1984
Notz 1972; Fernandez & Clavijo 1984
Teran 1974
Virla et al. 1999
Guimaraes 1977
King & Saunders 1984
Cave 1993
Estrada 1960
Hynes 1942
Vera et al. 1995
Molinari & Avalos 1997
Guimaraes 1977
Enkerlin 1975
Wheeler et al. 1989
Carrillo 1980
Estrada 1960
Nickle 1976
Cave 1993
Parker et al. 1953
Bruner et al. 1975
Wall & Berberet 1975; Nickle 1976
Teran 1974
Enkerlin 1975
Cave 1993
Maes 1989
Sabrosky 1981
Sabrosky 1981
Ashley et al. 1980
Tingle et al. 1994
Sabrosky 1981








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Eucelatoria sp.


Euphorocera floridensis Townsend
Euphorocera tachinomoides (Townsend)
Syn: Chetogena tachinomoides
Euphorocera sp.


Exorista mella (Walker)
Gonia capitata DeGeer
Gonia crassicornis (Fabricius)






Gonia (Reaumuria) pacifica Townsend

Gonia texensis Reinhard
Gonia sp.

Hyphantrophaga hyphantriae (Townsend)
Syn: Exorista ceratomiae (Coquillet)
Hyphantrophaga collina (Reinhard) Syn: Zenillia blanda
Incamyia chilensis (Aldrich)




Lespesia affinis (Townsend)


L Barbados
Brazil

Chile
Nicaragua
Venezuela

L Honduras
L US

L Brazil


US
US
US
Brazil
Honduras
Lesser Antilles
Puerto Rico


US
Venezuela
L Brazil
Peru
L Cuba
L Chile
Nicaragua
L US


Cuba
Argentina
Brazil
Chile


Uruguay
L Brazil


M
M
(N/G)
PN
M
M
(N/G)
(N/G)
PN

M
(N/G)
(N/G)
PN
(N/G)
(N/G)
S
M
SC
SC,M
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)

(N/G)
(N/G)
(N/G)
M
(N/G)
(N/G)
M


Alam 1979
Silveira et al. 1987; Valicente 1989
Guimaraes 1977
Enkerlin 1975
Maes 1989; Gladstone 1991
Fernandez & Clavijo 1984
Teran 1974
Cave 1993
Wall & Berberet 1975

Goncalves & Goncalves 1973; Silva et al. 1997
Guimaraes 1977
Nickle 1976
Wall & Berberet 1975
Luginbill 1928
Goncalves & Goncalves 1973; Guimaraes 1977
Cave 1993
Fennah 1947
Van Dine 1913
Jones 1913
Luginbill 1928
Teran 1974
Guimaraes 1977
Enkerlin 1975
Bruner et al. 1975
Enkerlin 1975
Maes 1989
Luginbill 1928

Bruner et al. 1975
Blanchard 1963
Guimaraes 1977
Etcheverry 1957
Enkerlin 1975
Parker et al. 1953; Silveira & Ruffinelli 1956
Guimaraes 1983; Silva et al. 1997








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Lespesia aletiae (Riley) L Honduras (N/G)


Lespesia archippivora (Riley)
Syn:Achaetoneura archippivora (Wile)
Syn: Frontina archippivora (Scudder)


Lespesia frenchi (Williston)
Lespesia grioti (Blanchard)


US


L Argentina
Brazil


Chile
Cuba

Guadeloupe
Guatemala
Honduras

Lesser Antilles
Mexico
Nicaragua

Puerto Rico

US








Uruguay

Venezuela
L US
L Argentina

Brazil


PN
M,S
M,BG,Mi
M
M

(N/G)
M
(N/G)

M
M,Te
M
M,S
M
M
M

SC
SC,M
A
C
M
PN
PN, M
M,S
C,A,M,S
M,BG,Mi,S
(N/G)
M

(N/G)
(N/G)
M
(N/G)
(N/G)


Cave 1993
Wall & Berberet 1975
Pair et al. 1986
Reed 1980
Virla et al. 1999
Patel & Habib 1984, 1986; Guimaraes 1983; Valicente
1989
Guimaraes 1977
Etcheverry 1957
Parker et al. 1953; Ryder & Piedra 1968; Bruner et al.
1975
Malausa 1981, 1983
Painter 1955
Canas & O'Neil 1998
Cave 1993
Fennah 1947
Carrillo 1980
Estrada 1960; Huis 1981; Maes 1989; Gladstone 1991;
Lacayo 1977
Van Dine 1913
Jones 1913
Soteres et al. 1984
Tingle et al. 1994
Gross & Pair 1986; Riggin et al. 1992, 1993
Wall & Berberet 1975; Nickle 1976
Vickery 1929
Pair et al. 1986
Butler 1958a
Reed 1980
Luginbill 1928
Silveira & Ruffinelli 1956
Notz 1972; Fernandez & Clavijo 1984
Teran 1974, 1977
Luginbill 1928
Virla et al. 1999
Blanchard 1963
Guimaraes 1977








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


L Argentina

Brazil

Colombia
Cuba
Honduras
Nicaragua
Puerto Rico
US

Uruguay
Venezuela


Linnaemya annalis (Townsend)
Linnaemya comta (Fallen)
Linnaemya sp.
Nemorilla pyste (Walker)
Syn: Exorista pyste (Walker)

Parasetigena sp.

Patelloa similis (Townsend)

Patelloa sp.


Peleteria robusta Wiedeman
Phorocera claripennis (Macquart)
Syn: Chetogena claripennis (Macquart)

Phorocera floridensis (Townsend)
Syn: Chetogena floridensis (Townsend)


Venezuela
Honduras
Ecuador
US


L Brazil
Uruguay
L Brazil

L Argentina

Uruguay
L Chile
L US

Venezuela
L Brazil
Honduras
Nicaragua
US

Venezuela


Lespesia sp.


M
(N/G)
M
(N/G)
R
(N/G)
M
M
R
M
M,S
(N/G)
M
(N/G)
M
(N/G)
M
A
C
(N/G)
(N/G)
(N/G)
M
(N/G)
M
(N/G)
(N/G)
(N/G)
A
(N/G)
(N/G)
M
(N/G)
(N/G)
PN
(N/G)
(N/G)


Virla et al. 1999
Parker et al. 1953
Lucchini & Almeida 1980
Goncalves & Goncalves 1973; Guimaraes 1977
Vargas & Sanchez 1983
Ryder & Pulgar 1969
Wheeler et al. 1989
Maes 1989
Pantoja et al. 1985; Pantoja & Fuxa 1992
Ashley et al. 1980
McCutcheon 1991
Parker et al. 1953
Notz 1972
Teran 1974
Teran 1977
Cave 1993
Benzing et al. 2000
Soteres et al. 1984
Wilson 1923
Luginbill 1928
Guimaraes 1977
Parker et al. 1953; Silveira & Ruffinelli 1956
Patel & Habib 1984, 1986
Guimaraes 1977
Virla et al. 1999
Parker et al. 1953
Parker et al. 1953; Silveira & Ruffinelli 1956
Etcheverry 1957
Soteres et al. 1984
Luginbill 1928
Teran 1974
Valicente 1989
Cave 1993
Maes 1989
Enkerlin 1975
Luginbill 1928
Teran 1974








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference

Pronemorilla mima Townsend L Venezuela M Notz 1972


L Brazil
Chile
L Argentina


Brazil


Winthemia leucanae (Kirkpatrick)
Syn: Nemorea leucanae (Kirkpatrick)
Winthemia mima (Reinhard)

Winthemia quadripustulata Fabricius


Winthemia reliqua
Winthemia roblesi
Winthemia rufopicta (Bigot)








Winthemia sinuata (Reinhard)
Winthemia trinitatis (Thompson)


Winthemia sp.


US
Uruguay
L US

L Argentina
Brazil
L Chile
US
Venezuela
L Chile
L Chile
L US








L US
L Argentina
Brazil

L Argentina
Brazil


Pseudokea sp.


Teran 1974
Costa-Lima 1949
Etcheverry 1957
Virla et al. 1999
Molinari & Avalos 1997
Parker et al. 1953
Guimaraes 1971
Guimaraes 1977
Butler 1958a
Silveira & Ruffinelli 1956
Dew 1913


(N/G)
(N/G)
M
M
SY
(N/G)
M
(N/G)
C,A,M,S
M
M,S,BG,SY,C

M
(N/G)
M
(N/G)
(N/G)
(N/G)
(N/G)
A
C
M

PN
M,S
PN,BG
C,A,M,S
M,BG,Mi,S
PN
M
M
(N/G)
M
M

(N/G)


Virla et al. 1999
Guimaraes 1977
Etcheverry 1957
Walton 1913; Luginbill 1928
Teran 1974
Valencia & Valdivia 1973
Valencia & Valdivia 1973
Soteres et al. 1984
Tingle et al. 1994
Hofmaster & Greenwood 1949;Ashley et al. 1980;Riggin
et al. 1992, 1993
Wall & Berberet 1975
Rohlfs & Mack 1985; Pair et al. 1986
Enkerlin 1975
Danks 1975
Reed 1980
Wall & Berberet 1975
Virla et al. 1999
Valicente 1989
Guimaraes 1977
Vera et al. 1995; Virla et al. 1999
Escalante 1974; Patel & Habib 1984, 1986; Silva et al.
1997
Guimaraes 1977


Voria ruralis (Fallen)








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference

Winthemia sp. L Chile M Etcheverry 1957; Campos 1965


Colombia
Honduras
Lesser Antilles
Mexico
Peru
Trinidad
US


Venezuela


Hymenoptera: Bethylidae
Perisierola sp.
Hymenoptera: Braconidae
Agathis stigmatera (Cresson)


Aleiodes caphimal
Syn: Rogas caphimal
Alieodes laphygmae (Viereck)
Syn: Rogas laphygmae (Viereck)














Aleiodes terminalis (Cresson)
Syn: Rogas terminalis (Cresson)


E Argentina


L Nicaragua

L Brazil
Chile
Honduras

Mexico

Nicaragua
Puerto Rico
US







L Canada
US


M,PN
R
M,S
(N/G)
M
M
(N/G)
PN
M,S
(N/G)


M

(N/G)
(N/G)

M
A,M,C
M
M,S
M
M,S
M
R
M


S
M,S

M,BG,PG
(N/G)
A,M,C
A,M,C
(N/G)


Enkerlin 1975
Vargas & Sanchez 1983
Cave 1993
Fennah 1974
Guevara et al. 1979
Sarmiento & Razuri 1978
Yaseen 1979; Hynes 1942
Wall & Berberet 1975
McCutcheon 1991
Teran 1974


Bianchi 1944

De Santis 1967; De Santis & Esquivel 1966;Virla et al.
1999
Parker et al. 1953
Andrews 1988

Cruz et al. 1997b
Etcheverry 1957
Wheeler et al. 1989; Canas & O'Neil 1998
Cave 1993
Molina-Ochoa et al. 2001
Pair et al. 1986
Estrada 1960; Huis 1981; Maes 1989; Gladstone 1991
Pantoja & Fuxa 1992
Vickery 1929; Bianchi 1944; Ashley et al. 1982; Mitch-
ell et al. 1984; Gross & Pair 1986; Riggin et al. 1992,
1993
Rohlfs & Mack 1985
Reed 1980; Pair et al. 1986; Isenhour 1988; McCutch-
eon 1991
Ashley et al. 1983
Luginbill 1928; Muesebeck et al. 1951
Muesebeck et al. 1951
Muesebeck et al. 1951
Luginbill 1928; Marsh & Shaw 2001


L US








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Aleiodes vaughani (Muesebeck)
Syn: Rogas vaughani Muesebeck

Aleiodes sp.
Syn: Rogas sp.



Bassus sp.

Bracon kirkpatricki Wilkinson
Cardiochiles nigriceps (Viereck)
Chelonus antillarum (Marshall)

Chelonus cautus Cresson
Syn: Microchelonus cautus (Cresson)

Chelonus formosanus (Sonan)

Chelonus insularis (Cresson)
Syn: Chelonus texanus (Cresson)


L Honduras

Nicaragua
L Argentina
Cuba
Honduras
Nicaragua
US
L Argentina
Honduras
L Mexico
L US
E Barbados
Nicaragua
E Honduras
Mexico
Nicaragua
E Barbados
Trinidad
E Argentina

Barbados
Brazil



Chile
Colombia

Cuba


Haiti
Honduras

Lesser Antilles


M

M
M
M
M
M
A
(N/G)
(N/G)
(N/G)
C
M
M
M
M,S
M
(N/G)
M
M
(N/G)
(N/G)
M


M,C
A,M,C
M
R
BG
M
(N/G)
BG
M
M,S
M


Cave 1993; Passoa 1983;Wheeler et al. 1989; Canas &
O'Neil 1998
Huis 1981; Maes 1989
Virla et al. 1999
Ryder & Pulgar 1969; Ryder & Piedra 1968
Passoa 1983
Estrada 1960; Maes 1989; Lacayo 1977
Soteres et al. 1984
Parker et al 1953
Cave 1993
Moya 1980; Pena 1980
Tingle et a. 1994
Alam 1979
Ryder & Pulgar 1969
Cave 1993; Canas & O'Neil 1998
Molina-Ochoa et al. 2001
Huis 1981
Alam 1979
Yaseen 1979
Virla et al. 1999
Parker et al. 1953
Alam 1979
Patel & Habib 1982, 1984, 1986; Valicente 1989;
Rezende et al. 1995a,b; Cruz et al. 1997a,b; Silva et al.
1997
Rezende et al. 1994
Etcheverry 1957
Medina et al. 1988
Vargas & Sanchez 1983
Myers 1932
Ryder & Pulgar 1969
Ryder & Piedra 1968; Bruner et al. 1975;
Myers 1932
Wheeler et al. 1989; Canas & O'Neil 1998
Castro et al. 1989; Cave 1993
Fennah 1947








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Chelonus insularis (Cresson)
Syn: Chelonus texanus (Cresson)


E Mexico


Nicaragua


Puerto Rico
Trinidad

US


Uruguay
Venezuela
E Brazil
Mexico


Chelonus sp.


Cotesia (Apanteles) congregate (Say)

Cotesia (Apanteles) glomeratus (Linnaeus)
Cotesia (Apanteles) marginiventris (Cresson)
Syn: Apanteles grenadensis (Ashmead)
Syn: Protapanteles harnedi (Viereck)
Syn:Apanteles laphygmae (Ashmead)


Peru
Honduras
Nicaragua
Barbados
Argentina

Brazil
Chile
Honduras
Lesser Antilles


M,S
M

M,S
R
M
(N/G)
A
C
M



PN
M,BG
M,PN
M,S

C,A,M,S
(N/G)
(N/G)
M
M
M

M,S
C
(N/G)
M
M
M

M
A,M,C
M
M,S


Bahena & Garcia 1991; Nava & Castro 1991; Arce &
Garcia 1995; Molina-Ochoa et al. 2001
Pair et al. 1986
Estrada 1960; Huis 1981; Ashley 1986; Maes 1989;
Gladstone 1991
Lacayo 1977
Pantoja & Fuxa 1992
Yaseen 1979
Yaseen et al. 1981
Soteres et al. 1984
Tingle et al. 1994
Pierce & Holloway 1912; Vickery 1929; Bianchi 1944;
Muesebeck et al. 1951; Ashley et al. 1980, 1982; Ash-
ley 1983, 1986; Mitchell et al. 1984; Gross & Pair 1986;
Riggin et al. 1992
Wall & Berberet 1975
Ashley et al. 1983
Enkerlin 1975
Waddill & Whitcomb 1982; Rohlfs & Mack 1985; Pair
et al. 1986; McCutcheon 1991
Butler 1958b
Luginbill 1928
Parker et al. 1953; Silveira & Ruffinelli 1956
Notz 1972; Fernandez & Clavijo 1984
Valicente 1989
Loya-Ramirez 1978; Coronado & Ruiz 1991; Cortez &
Trujillo 1994; Sanchez-Garcia et al. 1998; Carrillo 1980
Molina-Ochoa et al. 2001
Herrera-Aranguena 1998
Cave 1993
Maes 1989
Alam 1979
Wheeler et al. 1989; Virla et al. 1999
Lucchini & Almeida 1980; Patel & Habib 1982, 1984,
1986; Cruz et al. 1997b
Etcheverry 1957
Cave 1993
Fennah 1947








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Cotesia (Apanteles) marginiventris (Cresson)
Syn: Apanteles grenadensis (Ashmead)
Syn: Protapanteles harnedi (Viereck)
Syn:Apanteles laphygmae (Ashmead)


Cotesia (Apanteles) ruficrus (Haliday)
Syn: Microplitis manilee (Ashmead)
Cotesia (Apanteles) sp.


Distatrix sp.
Glyptapanteles militaris (Walsh)


E,L3 Mexico

Nicaragua
Puerto Rico
Suriname

US











Uruguay

Venezuela

E,L3 Trinidad & Tobago
US
E,L3 Barbados
Brazil
Colombia

Guyana
Nicaragua

Peru
Trinidad
US
L Honduras
L Honduras
US


M
M,S
M
R
M,PN
(N/G)
A
C
M





M,S
PN
M,BG
M,S,C,Mi
(N/G)
(N/G)
M
SY,To
M
(N/G)
M
M
R
M
M
M
M,S
C
M
M
M
M
S
(N/G)


Marsh 1978
Pair et al. 1986; Molina-Ochoa et al. 2001
Estrada 1960; Huis 1981; Maes 1989; Gladstone 1991
Pantoja & Fuxa 1992
Segeren & Sharma 1979
Van Dither 1960
Soteres et al. 1984
Tingle et al. 1994
Muesebeck 1921; Vickery 1929; Bianchi 1944;Hofmas-
ter & Greenwood 1949; Muesebeck et al. 1951; Marsh
1978; Ashley et al. 1980, 1982; Hogg et al. 1982; Ash-
ley 1983, 1986; Mitchell et al. 1984; Gross & Pair 1986;
Riggin et al. 1992, 1993, 1994; Hamm et al. 1994; Ru-
berson & Whitfield 1996
Rohlfs & Mack 1985; Pair et al. 1986; McCutcheon 1991
Wall & Berberet 1975; Nickle 1976
Ashley et al. 1983
Reed 1980
Luginbill 1928
Parker et al. 1953; Silveira & Ruffinelli 1956
Notz 1972; Fernandez & Clavijo 1984
Teran 1980
Yaseen 1979*
McCutcheon et al. 1983**; Rajapakse et al. 1985***
Alam 1979
Patel & Habib 1984, 1986; Silva et al. 1997
Vargas & Sanchez 1983
Medina et al. 1988
Sinha 1982
Maes 1989
Lacayo 1977
Herrera-Aranguena 1998
Yaseen 1979
Hofmaster & Greenwood 1949
Cave 1993
Cave 1993
Rohlfs & Mack 1985
Muesebeck et al. 1951








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Gnathopleura sp.
Homolobus truncator (Say)
Syn: Zele mellea (Cresson)






Macrocentrus sp.

Meteorus arizonensis (Muesebeck)

Meteorus autographae (Muesebeck)










Meteorus laphygmae (Viereck)


Honduras
Honduras
Nicaragua
US


L Barbados
Brazil
L Honduras
Nicaragua
L Mexico
US









L Chile
Colombia


Honduras
Mexico

Nicaragua
Suriname

US


(N/G)
M,S
M
A
M

M,S
M,S,C
(N/G)
(N/G)
M
(N/G)
M
M,S
A
C
M

PN
S
M,S
M,BG,Mi
M,BG,PG
(N/G)
M
M
R
C,S
M,S
M
S
(N/G)
M,PN
(N/G)
M
M,BG
(N/G)


Cave 1993
Cave 1993
Huis 1981; Maes 1989
Soteres et al. 1984
Vickery 1929; Wall & Berberet 1975; Riggin et al.
1992, 1993
Pair et al. 1986; McCutcheon 1991
Reed 1980
Luginbill 1928
Alam 1979
Silva et al. 1997
Cave 1993
Maes 1989
Pair et al. 1986
Soteres et al. 1984
Tingle et al. 1994
Ashley et al. 1980, 1982; Mitchell et al. 1984; Gross &
Pair 1986; Riggin et al. 1992, 1993
Nickle 1976
Rohlfs & Mack 1985
Pair et al. 1986; McCutcheon 1991
Reed 1980
Ashley et al. 1983
Luginbill 1928; Muesebeck et al. 1951
Etcheverry 1957
Medina et al. 1988
Vargas & Sanchez 1983
Orteg6n et al. 1988
Cave 1993
Molina-Ochoa et al. 2001
Ciceros et al. 1995
Gladstone 1991
Segeren & Sharma 1979
Van Dither 1960
Vickery 1929; Bianchi 1944
Enkerlin 1975
Luginbill 1928








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Meteorus laphygmae (Viereck)

Meteorus vulgaris (Cresson)
Meteorus sp.


Microplitis sp.
Palinzele sp.
Stantonia sp.

Hymenoptera: Chalcididae
Brachymeria ovata (Say)


Brachymeria robusta (Cresson)

Conura (Spilochalcis) femorata (Fabricius)



Conura (Ceratosmicra) hirtifemora (Ashmead)
'.i. I .......... *..... hirfitemora (Ashmead)
Conura (Ceratosmicra) immaculate (Cresson)
Syn: Conura fulvomaculata (Cameron)
Conura (Ceratosmicra) meteori (Burks)
Conura igneoides (Kirby)
.. .'....... i..... . uittata (Ashmead)
Hymenoptera: Eulophidae
Euplectrus comstockii Howard

Euplectrus furnicus Walker


Euplectrus hircinus (Say)
Euplectrus insularis Howard


L Venezuela

L US
L Mexico

Peru
L Uruguay
L Trinidad
L Honduras
Nicaragua

P Argentina

US
P US

L Honduras
Nicaragua
US

L US

L Honduras
Nicaragua
L US
L US


L Nicaragua
US
L Argentina

Puerto Rico
L Panama
L Honduras
Nicaragua


M

(N/G)
M
M,S
C
(N/G)
M
(N/G)
M

M
(N/G)
M
C
(N/G)
(N/G)
(N/G)
C
(N/G)
M

(N/G)
M
M
C
(N/G)

M
(N/G)
M

R
(N/G)
M
(N/G)


Notz 1972; Teran 1980; Fernandez & Clavijo 1984;
Fernandez & Teran 1990a,b
Luginbill 1928; Muesebeck et al. 1951
Peraza-Lizarraga 1982; Coronado & Ruiz 1991
Molina-Ochoa et al. 2001
Herrera-Aranguena 1998
Parker et al. 1953; Silveira & Ruffinelli 1956
Yaseen 1979
Cave 1993
Maes 1989

Virla et al. 1999
Parker et al. 1953
Ashley 1979
Wilson 1923
Luginbill 1928
Cave 1993
Maes 1989
Wilson 1923
Luginbill 1928
Riggin et al. 1992, 1993

Cave 1993
Maes 1989
Hofmaster & Greenwood 1949
Wilson 1923
Luginbill 1928; Muesebeck et al. 1951

Maes 1989
Luginbill 1928
De Santis 1979; De Santis 1989; De Santis & Fidalgo
1994; Virla et al. 1999
Pantoja & Fuxa 1992
Andrews 1988
Wheeler et al. 1989
Huis 1981








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Euplectrus marginatus Ashmead
Euplectrus plathypenae Howard


Euplectrus ronnai (Brethes)
Euplectrus sp.


Trichodischia caerulea
Trichodischia soror (Bigot)


Trichospilus pupivora (Ferriere)
Trichospilus sp.


L Nicaragua
L Barbados
Brazil
Chile
Colombia
Cuba

Guyana
Lesser Antilles
Mexico

Nicaragua
Puerto Rico
Trinidad
US



Venezuela

L Brazil
L Brazil
Cuba
Honduras
Mexico

Nicaragua

US



L Brazil
L Argentina

Brazil
P Barbados
L Argentina


M
M
(N/G)
M
M
Pm&Pb
(N/G)
M
M
M


M
R
(N/G)
M

PN
(N/G)
M
S,C
M
M
BG
M
M
M,S
M
M,S
BG
M
M,S
(N/G)
(N/G)
M
(N/G)
(N/G)
M
(N/G)


Maes 1989
Alam 1979
Costa-Lima 1962
Etcheverry 1957
Vargas & Sanchez 1983
Myers 1932
Ryder & Piedra 1968; Bruner et al. 1975
Sinha 1982
Fennah 1947
Guevara et al. 1979; Montoya-Burgos 1980; Guti6rrez-
Rodriguez 1982; Molina-Ochoa et al. 2001
Ryder & Pulgar 1969
Pantoja & Fuxa 1992
Yaseen 1979
Vickery 1929; Muesebeck et al. 1951; Ashley et al.
1980, 1982; Hogg et al. 1982; Riggin et al. 1992, 1993
Wall & Berberet 1974, 1975
Luginbill 1928
Marin-Acosta 1966
Guagliami 1962
De Santis 1980
Lucchini & Almeida 1980
Myers 1932
Cave 1993
Montoya-Burgos 1979, 1980; Cortez & Trujillo 1994
Pair et al. 1986
Huis 1981
Lacayo 1977
Reed 1980
Keller 1980
Pair et al. 1986
Enkerlin 1975
Guimaraes 1977
Virla et al. 1999
Parker et al. 1953; Blanchard 1963
Cortes 1980
Alam 1979
Parker et al. 1953








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Hymenoptera: Ichneumonidae
Amblyteles sp.

Ancyloneura sp.
Anomalon ejuncidum (Say)
Campoletis chloridae (Vierech)
Campoletis curvicauda (Blanchard)
Campoletis flavicincta (Ashmead)







Campoletis grioti (Blanchard)




Campoletis oxylus (Cresson)
Syn: Sagaritis oxylus (Cresson)

Campoletis sonorensis (Cresson)


Campoletis sp.


L Brazil
Chile
L Argentina
L US
L Barbados
L Peru
L Brazil
Honduras
Mexico
Nicaragua
US


Uruguay
L Argentina
Brazil

US
Uruguay
L US


L Brazil
Chile
Honduras

Mexico
US


L Argentina
Brazil


Nicaragua
US


(N/G)
M
(N/G)
M,S
M
M
M
(N/G)
M,S
M
A
M
PN
(N/G)
M
M

M
M
M
M,S
(N/G)
M
M
M
M,S
M,S
A
M
M,S
M
M

(N/G)
M
M,S


Costa-Lima 1949
Etcheverry 1957
Parker et al. 1953
Pair et al. 1986
Alam 1979
Ayqui-Vilca 1993
Patel & Habib 1982, 1984, 1986; Cruz et al. 1997a,b
Cave 1993
Molina-Ochoa et al. 2001
Huis 1981; Maes 1989
Soteres et al. 1984
Hogg et al. 1982
Wall & Berberet 1975
Parker et al. 1953;Yaseen et al. 1981
Virla et al. 1999
Lucchini & Almeida 1980; Silveira et al. 1987; Cruz et
al. 1997b
Ashley 1983
Morey 1971
Muesebeck et al. 1951
Pair et al. 1986
Luginbill 1928
Cruz et al. 1997b
Machuca et al. 1989
Canas & O'Neil 1998
Cave 1993
Pair et al 1986
Soteres et al. 1984
Gross & Pair 1986; Riggin et al. 1992, 1993
Pair et al. 1986; Isenhour 1988
Vera et al. 1995
Silva et al. 1997; Silveira et al. 1987; Valicente & Bar-
reto 1999
Cruz et al. 1999
Maes 1989
McCutcheon 1991








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Campoplex sp.
Cryptus albitarsis (Cresson)
Diadegma sp.



Diapetimorpha introita (Cresson)


Eiphosoma annulatum (Cresson)
Eiphosoma vitticole (Cresson)


Eiphosoma sp.


Enicospilus flavus (Fabricius)
Syn: Enicospilus concolor (Cresson)
Enicospilus merdarius (Gravenhorst)
Syn: Enicospilus purgatus (Say)


Goryphina sp.
Hyposoter sp.


L Brazil
P US
L Argentina
Brazil
Cuba
Mexico
P Honduras
US

L Venezuela
L Bolivia

Brazil

Colombia
Honduras

Mexico
Nicaragua
US
Venezuela
L Brazil
Mexico
Trinidad

Venezuela
L US

L Argentina
Cuba
Honduras
Nicaragua
US

L Brazil
L Honduras
Uruguay


M
M
M
M
M
M
M
M
M,S
M
M
(N/G)
M

M
M
M,S
M,S
M
M
(N/G)
M
M
M
(N/G)
M
C
(N/G)
M
M
(N/G)
M
C,M
(N/G)
M
M
(N/G)


Silva et al. 1997
Pair & Gross 1989
Porter 1998; Virla et al. 1999
Silva et al. 1997
Ayala-Sifontes et al. 1978
Flores-Davila et al. 1991
Cave 1993
Pair & Gross 1984, 1989
Pair et al. 1986
Notz 1972
Ashley 1979
Yaseen et al. 1981
Patel & Habib 1982, 1984, 1986; Valicente 1989; Sil-
veira et al. 1987; Cruz et al. 1997b
Medina et al. 1988
Wheeler et al. 1989; Canas & O'Neil 1998
Cave 1993
Pair et al. 1986
Huis 1981
Ashley 1983
Giraldo-Vanegas & Garcia-R. 1992, 1994a,b, 1995
Cruz et al. 1997a; Silva et al. 1997
Cortez & Trujillo 1994
Hynes 1942
Yaseen et al. 1981
Fernandez & Clavijo 1984
Wilson 1923
Luginbill 1928
Muesebeck et al. 1951
Bruner et al. 1975
Cave 1993
Maes 1989
Dew 1913
Luginbill 1928
Silva et al. 1997
Cave 1993
Parker et al. 1953; Silveira & Ruffinelli 1956








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Ichneumon ambulatorius Fabricius
Ichneumon promissorius (Cresson)
Isdromas lycaenae (Howard)
Mesochorus disceitergus (Say)


Microcharops anticarsiae (Gupta)

Netelia sayi (Cushman)
Netelia sp.

Parania (Atrometus) tricolor I MI..1. i
Pristomerus spinator (Fabricius)
Syn: Neopristomerus appalachianus (Viereck)











Ophion ancyloneura (Wichsee)


Ophion bilineatus (Say)


Ophion flavidus (Brulle)


L Honduras

L US
L Peru
US
L Uruguay
L Brazil
Honduras

Mexico

Nicaragua

US





L Argentina

Uruguay
L Chile
US


L Argentina
Brazil

Honduras


M
S
C
M
A
(N/G)
M
M
M,S
M
M,S
M

A
C
M
PN
M,S
(N/G)
M
(N/G)
(N/G)
M
M
PN
(N/G)
M
M

M
M,S
(N/G)


New record (JEC)
New record (JEC)
Riggin et al. 1992, 1993
Hofmaster & Greenwood 1949; Riggin et al. 1992,
1993
Wheeler et al. 1989
Cave 1993
Tingle et al. 1994
Escalante 1974
Soteres et al. 1984
Parker et al. 1953; Silveira & Ruffinelli 1956
Patel & Habib 1984, 1986
Wheeler et al. 1989; Canas & O'Neil 1998
Cave 1993
Carrillo 1980
Pair et al. 1986; Molina-Ochoa et al. 2001
Estrada 1960; Huis 1981; Gladstone 1991; Maes 1989;
Lacayo 1977
Soteres et al. 1984
Tingle et al. 1994
Vickery 1929; Bianchi 1944
Wall & Berberet 1975
Pair et al. 1986
Luginbill 1928
De Santis 1967; Virla et al. 1999
Parker et al. 1953
Silveira & Ruffinelli 1956
Etcheverry 1957
Vickery 1929
Enkerlin 1975
Luginbill 1928
Virla et al. 1999
Goncalves 1973; Patel & Habib 1982, 1984, 1986; Sil-
veira et al. 1987
Wheeler et al. 1989; Canas & O'Neil 1998
Cave 1993
Passoa 1983










TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Ophion flavidus (Brulle)


Ophion merdarius (Gravenhorst)
Syn: Enicospilus purgatus (Gravenhorst)
Ophion sp.









Sagaritis dubitatus (Cresson)

Trachysphyrus cleonis (Viereck)
Temelucha difficilis (Dasch.)


Temelucha grapholithae (Cushman)


Temelucha sp.


Vulgichneumon brevicintor (Say)
Hymenoptera: Perilampidae
Perilampus hyalinus (Say)
(Reported as hiperparasitoid)


L Mexico

Nicargua
US



L Nicaragua
US
L Argentina
Brazil
Mexico
Nicaragua
Peru
US



Uruguay
L US

L Peru
L US


L Honduras

Nicaragua
L Honduras
Nicaragua
US
P US

L Honduras
Venezuela


M
M,S
M
M

M,S
M,BG,Mi,S
M
(N/G)
M
M
M
M
M
C
M
PN
M,PG
(N/G)
M
(N/G)
M
M


Molina-Ochoa et al. 2001
Pair et al. 1986
Maes 1989; Gladstone 1991
Hogg et al. 1982; Gross & Pair 1986, 1991; Riggin et al.
1992, 1993
Rohlfs & Mack 1985; Pair et al. 1986
Reed 1980
Maes 1989
Luginbill 1928
Vera et al. 1995
Silva et al. 1997
Ashley 1986
Huis 1981; Maes 1989
Escalante 1974
Tingle et al. 1994
Mitchell et al. 1984
Nickle 1976
Ashley et al. 1983
Parker et al. 1953
Vickery 1929
Luginbill 1928
Escalante 1974
Mitchell et al. 1984; Gross & Pair 1986
Pair et al. 1986

Canas & O'Neil 1998
Cave 1993
Gladstone 1991
Wheeler et al. 1989
Maes 1989
Ashley et al. 1980, 1982, 1983
Pair & Gross 1989

Cave 1993
Notz 1972








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Hymenoptera: Pteromalidae
Catalaccus aeneoviridis (Girault)
Trichomalopsis viridescens (Walsh)
Syn: Eupteromalus viridescens
Hymenoptera: Scelionidae
Telenomus remus (Nixon)


L US
L US


E Antigua
Barbados
Brazil
Colombia
Dominican Republic
Guadeloupe
Guyana
Honduras
Nicaragua
Puerto Rico
Suriname
Trinidad
US


Venezuela


Telenomus sp.


Hymenoptera: Trichogrammatidae
Trichogramma demoraesi (Zucchi)
Trichogramma fasciatum Perkins

Trichogramma minutum Riley


E Brazil
Colombia

Cuba

Guadeloupe
Mexico


E Chile
E Barbados
Nicaragua
E Nicaragua
US


M
M
M
M
(N/G)
(N/G)
M,S
M
M
(N/G)
M
M
M,S
(N/G)
M

M
M
R
M

M
M


M
M
M
M
M
(N/G)


Hofmaster & Greenwood 1949
Hofmaster & Greenwood 1949; Riggin et al. 1992, 1993


Irving 1978
Alam 1979
Correa-Figueiredo et al. 1999
Alvarez & Roa 1995
Yaseen et al. 1981
Yaseen et al. 1981
Sinha 1982
Cave & Acosta 1999
Maes 1989
Wojcik et al. 1976
Segeren & Sharma 1979
Yaseen 1979
Waddill 1977; Waddill & Whitcomb 1982
Wojcik et al. 1976
Hernandez et al. 1989;Hernandez & Diaz 1995, 1996a,b;
Gonzalez-Narvaez & Zocco 1996; Ferrer 1998a,b
Cruz et al. 1999
Alvarez & Roa 1995
Vargas & Sanchez 1983
Armas-Garcia & Ayala-Sifontes 1987; Ayala-Sifontes
et al. 1992
Malausa 1983
Montoya-Burgos 1979; Morales-P6rez 1982; Canseco-
Roman 1988; Garcia-Lagunas 1988; Barilla-Vera 1989

Pratissoli et al. 1999
Alam 1979
Maes 1989
Maes 1989
Muesebeck et al. 1951
Luginbill 1928








TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference


Trichogramma pretiosum (Riley)

Trichogramma sp.
















Nematoda: Acugutturidae
Noctuidonema guyanense Remillet & Silvain


E Brazil
Nicaragua
E Argentina
Brazil

Colombia
Cuba

Guadeloupe

Mexico





Nicaragua

US

A Bahamas
Belize
Bermuda
Cayman Islands
Colombia
Costa Rica
Dominica
El Salvador
French Guiana


Grenada
Guadeloupe
Honduras
Martinique
Mexico
Panama
Puerto Rico


M
M
M
M
(N/G)
R
M

M
(N/G)
M




(N/G)
M

M,S

(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)


(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)
(N/G)


De Sa & Parra 1994; Zucchi et al. 1991
Huis 1981
Virla et al. 1999
De Sa & Parra 1994
Cruz et al. 1999
Vargas & Sanchez 1983
Armas-Garcia & Ayala-Sifontes 1987; Ayala-Sifontes
et al. 1988
Malausa 1983
Yaseen et al. 1981
Loya-Ramirez 1978; Montoya-Burgos 1979, 1980;
Guevara et al. 1979; Rodriguez-Luna 1982; Wong-
Arevalo 1982; Johannes-Toonders & Carrillo-Sanchez
1987
Bahena & Garcia 1991
Huis 1981; Maes 1989; Mulock et al. 1990
Waddill & Whitcomb 1982



Simmons & Rogers 1990b
Simmons & Rogers 1990b
Simmons & Rogers 1990b; Simmons et al. 1991
Simmons & Rogers 1990b
Simmons & Rogers 1990b
Simmons & Rogers 1990b
Simmons & Rogers 1990b
Simmons & Rogers 1990b
Remillet & Silvain 1988; Marti et al. 1990; Rogers et
al. 1990a, 1991, 1993; Silvain & Remillet 1993; Marti
et al. 2000
Simmons & Rogers 1990b; Simmons et al. 1991
Marti et al. 2000
Simmons & Rogers 1990b
Simmons & Rogers 1990b; Marti et al. 1990
Simmons & Rogers 1990b
Simmons & Rogers 1990b; Simmons et al. 1991
Simmons & Rogers 1990b; Simmons et al. 1991




















TABLE 1. (CONTINUED) FALL ARMYWORM, Spodoptera frugiperda (J. E. SMITH) (LEPIDOPTERA: NOCTUIDAE), PARASITES AND PARASITOIDS IN THE AMERICAS AND CARIB-
BEAN BASIN WITH ACCOMPANYING REFERENCES INDICATING FAMILY, HOST STAGE ATTACKED, CROPS FROM WHICH FALL ARMYWORM WERE COLLECTED AND COUN-
TRIES OF COLLECTION.

Host stage Country of
Classification of parasite and parasitoid attacked' collection Crop2 Reference

Noctuidonema guyanense Remillet & Silvain A Suriname (N/G) Simmons & Rogers 1990b
Trinidad (N/G) Simmons & Rogers 1990b
US (N/G) Marti et al. 1990; Simmons & Rogers 1990b; Rogers et
al. 1990b, 1991, 1993, 1996; Simmons et al. 1991; Sim-
mons & Rogers,1990a,b, 1991, 1994; Marti et al. 2000;
Rogers & Marti 1994, 1996
M Simmons & Marti 1992
M, S Rogers & Marti 1992a,b, 1993a; Marti & Rogers 2000
Virgin Island (N/G) Simmons & Rogers 1990b

*Imported from Pakistan, ** Imported from Australia, *** Imported from Thailand & Imported from India.
'Host stage attacked: E (Egg), L (Larva), P (Pupa), A (Adult).
'A = Alfalfa (Medicago sativa L.), BG = Bermudagrass (Cynodon dactylon L.), C = Cotton (Gossypium spp.), M = Maize (Zea mays L.), Mi = Millet (Panicum miliaceum L.), O = Onion
(Allium cepa L.), P = Hot pepper (Capsicum annuum L.), Pb = (Panicum barbinode L.), PG = Paragrass (Brachiarie mutica L.), Pm = (P. maximum L.), PN = Peanut (Arachis hypogaea L.),
R = Rice (Oryza sativa L.), S = Sorghum (Sorghum bicolor L. (Moench.)), SC = Sugarcane (Saccharum officinarum L.), SY = Soybean (Glycine max L.), T = Tomato (Lycopersicum esculentum
L.), Te = Teosinte (Euchlaena mexicana Schard), To = Tobacco (Nicotiana tabacum L.), Ve = Vegetable Crops, (N/G) = Not given.
'Reported attacking eggs and larva (no confirmation).







Armyworm Symposium 2002: Molina-Ochoa et al.


TABLE 2. NUMBER OF SPECIES IN EACH TAXON REPORTED FROM DIFFERENT REGIONS.

Region

North America Central America
Order Family (Mexico-US) & Caribbean Basin South America

Diptera
Bombyliidae 1
Phoridae 1
Sarcophagidae 3 3 1
Tachinidae 29 24 38
Hymenoptera
Bethylidae 1
Braconidae 18 22 11
Chalcididae 6 2 1
Eulophidae 3 8 7
Ichneumonidae 27 18 21
Perilampidae 1 1
Pteromalidae 2
Scelionidae 2 2 2
Trichogrammatidae 2 4 3
Nematoda
Aphelenchoididae 1 1 1


RESULTS AND DISCUSSION

A great diversity of parasitoids and parasites
of FAW has been reported occurring in the Amer-
icas and the Caribbean basin. In this inventory,
approximately 150 species of parasitoids and par-
asites have been recorded from 14 families, nine
in Hymenoptera, four in Diptera and one in Nem-
atoda (Table 1). Ichneumonids and braconids
were the most diverse families in Hymenoptera,
represented by 36 and 28 species, respectively.
The most diverse family in Diptera, as well as
overall, was Tachinidae with 55 species.
Distribution among the taxa was highly vari-
able. Three of the 14 families were only reported
from one country, and five of the 14 families were
reported from two or fewer countries. Bethylid and
bombyliid parasitoids were only reported in the
United States (US) during the 1920s and 1940s
(Table 1). Of the =150 species, 74 were reported
from only one country, and 102 were reported from
only one geographical region. In contrast, nine of
the 14 families and 18 of the =150 species were re-
ported from all geographical regions (North Amer-
ica, Central America and the Caribbean, and
South America), and eight species were reported
from 10 or more countries (Tables 1 and 2).
The number of parasitoid and parasite species
reported from each FAW stage attacked in each
country is summarized in Table 3. The highest
number of parasitoid species attacking FAW lar-
vae was reported from the US (75), followed by
Brazil (45), Honduras (43) and Nicaragua (42).


Parasitoid species that attack FAW eggs were
more prevalent in Nicaragua (11) Brazil (8), Bar-
bados (7) and Mexico (6). Parasitoid species that
attack FAW pupae were reported from only 4
countries, with the greatest number of these spe-
cies collected from the US (6).
Parasitized FAW were collected from 19 differ-
ent host plants (Table 4). FAW collected from corn
were parasitized by species from all families ex-
cept Bombyliidae. The number of parasite and
parasitoid species attacking FAW was greatest
for FAW collected from corn (134), sorghum (40),
cotton (28), peanut (24), alfalfa (17), rice (13), and
Bermuda grass (11). Six of the 14 parasitoids and
parasite families were recovered from FAW that
had been collected from a single host plant spe-
cies. Ten of the 14 parasitoids and parasite fami-
lies were recovered from FAW that had been
collected from three or fewer host plant species.
The most diverse parasitoid families also were re-
covered from more FAW host plant species. Ta-
chinids, braconids, eulophids, and ichneumonids
were recovered from FAW collected from 15, 11, 8
and 8 FAW host plant species, respectively.
A considerable number of parasitoid species
(=33%) were reported only from one geographical
region (Table 1). These findings emphasize the
need for more surveys and taxonomical studies of
the natural enemies in the different habitats of
the natural distribution of the pest (Molina-Ochoa
et al. 2001). It is important to consider the differ-
ent developmental stages of the pest and the
growing season of the crop to determine if differ-







Florida Entomologist 86(3)


TABLE 3. NUMBER OF SPECIES OF PARASITOIDS AND STAGE HOST ATTACKED REPORTED FROM DIFFERENT COUNTRIES.

Number of parsitoid species

Country Egg Larval Pupae Adult

Antigua 1
Argentina 4 28 1
Bahamas 1
Barbados 7 9 1
Belize 1
Bermuda 1
Bolivia 1
Brazil 8 45
Cayman Is. 1
Canada 1
Chile 3 19
Colombia 5 6 1
Costa Rica
Cuba 3 11
Dominica 1
Dominican Republic 1
Ecuador 2
El Salvador 1
French Guinea 1
Grenada 1
Guadeloupe 3 2 1
Guatemala 1
Guyana 2 2
Haiti 1
Honduras 5 43 1 1
Lesser Antilles 2 8
Martinique 1
Mexico 6 22
Nicaragua 11 42
Panama 1
Peru 2 10
Puerto Rico 3 9 1
Suriname 2 4 1
Tobago 1 1
Trinidad 4 9 1
United States 3 75 7 1
Uruguay 1 16
Venezuela 2 29
Virgin Is. 1


ences in natural distribution of the parasitoids ex-
ist (Ashley 1979; Molina-Ochoa et al. 2001) or if
the records related to unique occurrence are due
to inadequate surveys. It also is important to de-
termine if the occurrence of FAW parasitoids is as-
sociated with the developmental stage of the pest
and the host plant. Studies related to tritrophic
interactions are needed to elucidate the role of the
plant metabolites on the susceptibility of the pest
to parasitoids and pathogens, as well as their suit-
ability (Molina-Ochoa et al. 1999). A high diver-
sity of FAW parasitoids has been reported in its
natural distribution exerting significant mortal-


ity on egg and larval populations. An understand-
ing of induced parasitization of the complex of
parasitoids is needed to determine the species
with higher capability to attack each stage of the
FAW. It is necessary to unite institutional efforts
to establish programs of release of FAW parasi-
toids in overwintering areas and to reduce the mi-
gration of the pest northward (Gross & Pair 1986).
Because much of the published work does not in-
clude environmental data with the collections, we
encourage authors to include this information in
the future. These data would help select candi-
date parasitoids for a specific or broad region.


September 2003

















TABLE 4. NUMBER OF PARASITOID SPECIES IN EACH TAXON REPORTED FROM FAW COLLECTED FROM DIFFERENT HOST PLANTS.

FAW host plants

Order Family A BG C M Mi O P Pb PG Pm PN R S SC SY T Te To Ve

Diptera
Bombyliidae 1
Phoridae 1
Sarcophagidae 4
Tachinidae 6 4 7 34 4 1 1 15 4 13 3 5 1 1 1

Hymenoptera
Bethylidae 1
Braconidae 7 4 12 24 2 2 4 5 11 1 1
Chalcididae 3 4
Eulophidae 1 1 9 1 1 1 2 2
Ichneumonidae 4 1 5 36 1 1 4 11
Perilampidae 1
Pteromalidae 2
Scelionidae 2 1 1
Trichogrammatidae 5 1 1

Nematoda
Acuguttuidae 1 1

A = Alfalfa, BG = Bermuda grass, C = Cotton, M = Maize, Mi = Millet, O = Onion, P = Hot Pepper, Pb = Panicum barbinode, PG = Para grass, Pm = P. maximum, PN = Peanut, R = Rice,
S = Sorghum, SC = Sugarcane, SY = Soybean, T = Tomato, Te = Teosinte, To = Tobacco, Ve = Vegetable crops







Florida Entomologist 86(3)


Parasites and Parasitoids of Spodoptera frugiperda
Pupae and Adults

Spodoptera frugiperda pupae and adults are
attacked by several parasite and parasitoid spe-
cies. Five species of Ichneumonidae: Diapetimor-
pha introita (Cresson), Cryptus albitarsis
(Cresson), Ichneumon promissorius (Erichson),
Ichneumon ambulatorius and Vulgichneumon
brevicinctor (Say) have been reported attacking
pupae of S. frugiperda and other noctuids
(Bechinski & Pedigo 1983; Pair & Gross 1984;
Wilson 1983; Pair & Gross 1989; Fitt & Daly
1990; Pavuk & Stinner 1991). Two generalist pu-
pal parasitoids of Chalcididae, Brachymeria
ovata (Say) and B. robusta (Cresson), have been
collected from S. frugiperda (Wilson 1923; Lugin-
bill 1928; Parker et al. 1953; Ashley 1979; Virla et
al. 1999). Only one eulophid species, Trichospilus
pupivora (Ferriere), has been reported as a gener-
alist parasitoid of Lepidopteran pupae, including
S. frugiperda (Alam 1979, Anantanarayanan
1934). Also, only one species (the ectoparasitic
nematode, Noctuidonema guyanense Remillet &
Silvain (Remillet & Silvain 1988)), has been ob-
served parasitizing S. frugiperda adults.

Studies on Diapetimorpha introita

Diapetimorpha introita, a parasitoid which at-
tacks FAW in the pupal stage, was reported for
first time by Pair & Gross (1984) in Tifton, Geor-
gia. The levels of parasitization in six trials ranged
from 0.0 to 23.7%, and the percent of intact pupae
that were parasitized during that study averaged
13.5%. Later, Gross & Pair (1986) emphasized the
need for more efforts to explore the role of species
of parasitoids that employ similar strategies. Pair
& Gross (1989) reported the seasonal incidence of
D. introita with rates of parasitism that averaged
5.2% (range 0-23.7%) and 8.4% (range 0-50.0%),
respectively, during 1983 and 1984, with the high-
est rate occurring during September to November
of each year. Because male D. introita are at-
tracted to chemicals emitted by the female wasps
(Jewett & Carpenter 1998) sticky traps baited
with live female wasps were used to study the sea-
sonal abundance of D. introita (Jewett & Carpen-
ter 2001). The highest number of adult males was
caught during early autumn which corresponded
to the rates of parasitism reported by Pair and
Gross (1989). As the investigation of their impor-
tance to biological control of Spodoptera spp. pro-
ceeds, more convenient methods of monitoring D.
introita in the field are needed.
Pair (1995) studied the biology and rearing of
D. introita on host and non-host noctuid pupae,
Spodoptera spp., Helicoverpa zea (Boddie), and
Heliothis virescens (F.) to identify factors that in-
fluence the reproduction and developmental rate
of D. introita. This parasitoid was successfully


reared in the laboratory on S. frugiperda pupae.
Carpenter & Greany (1998) compared the devel-
opmental time, weight, fecundity, longevity, and
ability to parasitize hosts forD. introita wasps de-
veloping on artificial diet and wasps reared on
S. frugiperda pupae. They conclude that the abil-
ity to rear D. introita on an inexpensive, artificial
diet significantly enhances the potential of mass
rearing this parasitoid for inundative releases
against species in the genus Spodoptera.

Studies on Ichneumon promissorius
and Collection of Ichneumon ambulatorius

I. promissorius was collected in Australia from
Helicoverpa armigera (Hfbner), and H. punctig-
era (Wallengren) pupae (Chadwick & Nikitin
1976; Wilson 1983; Fitt & Daly 1990). The parasi-
toid was imported into the US and released in Ar-
kansas, Georgia, Oklahoma, and Texas from 1992
to 1997 (J. E. C. et al., unpublished data). Follow-
ing releases of promissorius in ear-stage corn in
the lower Rio Grande valley in Texas during 1993
and 1994, feral noctuid pupae, including FAW,
were removed from the soil. During 1993, 575
FAW pupae were collected, 8 of which were para-
sitized by I. promissorius. Of the 13 FAW pupae
collected in 1994, none were parasitized by I.
promissorius. A similar study was conducted in a
cornfield in Rabun County, Georgia. During 1993,
141 of the 300 FAW pupae collected were parasit-
ized by I. promissorius and 24 were parasitized by
I. ambulatorius, a new host record. Only 5 FAW
pupae collected in 1994. Two of these FAW pupae
were parasitized by I. promissorius and one was
parasitized by I. ambulatorius.
Carpenter et al. (1994) compared several in-
digenous lepidopteran species in Tifton, GA, for I.
promissorius acceptance and development. Pupae
of H. zea, H. virescens (F.), H. subflexa (Guenee),
S. frugiperda, S. exigua (Hfbner), Trichoplusia ni
(Hfbner), Agrotis ipsilon (Hufnagel), and Antic-
arsia gemmatalis (Hfbner) were used as hosts.
Ninety pupae/species were tested, resulting in 74,
72, 68, 66, and 62 wasps emerging from T. ni, A.
ipsilon, H. suflexa, S. exigua, and S. frugiperda,
respectively. Lowest emergence of wasps was ob-
tained on A. gemmatalis (6). The most acceptable
hosts of promissorius were H. zea and H. vire-
scens, as expected because I. promissorius was
collected in Australia from heliothid species (Fitt
& Daly 1990). Carpenter et al. (1994) considered
that H. zea should be the primary host species of
this pupal parasitoid in the US.
Carpenter (1995) examined the influence of
host species, host availability, and mating status
on I. promissorius fecundity and oviposition. The
results from this study suggest that the females
budget their energy expenditures and regulate
oogenesis to maximize their reproductive poten-
tial. Host size, as well as the host species, may


September 2003







Armyworm Symposium 2002: Molina-Ochoa et al.


have contributed to differences in weight of the
wasps, and contributed to differences in fecun-
dity, longevity, and oviposition between wasps
reared on S. exigua and H. zea pupae. Virgin fe-
males reared on S. exigua pupae laid fewer eggs
than virgin females reared on H. zea pupae.

Studies on Vulgicheumon brevicinctor
and Cryptus albitarsis

Two V brevicinctor were recovered from FAW
pupae collected in 1982 from a cornfield near Tif-
ton, GA (Pair & Gross 1989). Five V. brevicinctor
were recovered from a sample of 300 FAW pupae
collected from a corn field in Rabun County, GA,
in 1993, and one V. brevicinctor was recovered
from a sample of 3 FAW pupae collected from the
same field in 1994 (J. E. C., unpublished data). V.
brevicinctor also has been reported from other
noctuids and species of other lepidopteran fami-
lies (Carlson 1979). For example, Bechinski &
Pedigo (1983) studied the population dynamics of
the green cloverworm (Plathypena scabra F.) in
soybeans in Iowa during 1979 and 1980. They and
found that V. brevicinctor acted in a delayed den-
sity-dependent manner on green cloverworm pu-
pal mortality. Pavuk & Stinner (1991) reported
that V. brevicinctor was reared from pupae of an
arctiid, Cisseps fulvicollis (Hiibner), and pupae
from a pyralid, Ostrinia nubilalis (Htibner).
Pair & Gross (1989) collected C. albitarsis in
Tifton, GA, from a single sample taken November
2, 1984. Four C. albitarsis also were recovered
from a sample of 300 FAW pupae collected from a
corn field in Rabun County, GA, in 1993 (J. E. C.,
unpublished data). Pair & Gross (1989) reported
that C. albitarsis had been established in a labo-
ratory colony in Tifton, GA. C. albitarsis cohorts
from this laboratory colony were successfully
reared on an artificial diet devoid of any host com-
ponents (Greany and Carpenter 1998).

Studies on Ectopasitic Nematodes Attacking Adults
of Spodoptera frugiperda

Remillet & Silvain in 1982 discovered and re-
ported an ectoparasitic nematode, Noctuidonema
guyanense Remillet & Silvain infecting Spodoptera
androgea (Cramer) in French Guiana. It was sub-
sequently described as a new genus and species
(Remillet & Silvain 1988) most commonly found on
moths of FAW, Spodoptera latifascia (Walker), S.
marima (Schaus), Anicla infecta (Ochsenheimer),
and Leucania spp. (Remillet & Silvain 1988).
Rogers et al. (1990a) determined the life cycle
and host range for N. guyanense in French Gui-
ana. They collected moths using a white sheet illu-
minated by UV light, by pheromone traps, and
sweeping vegetation from a variety of habitats in
Northeastern French Guiana. Moths in five fami-
lies, Lasiocampidae, Noctuidae, Notodontidae,


Pyralidae, and Sphingidae were naturally infested
with this nematode species. Twenty-five species of
Noctuidae were infected by N. guyanense, the
hosts most commonly infected were Lesmone for-
mularis (Htibner), S. dolichos (F.), S. frugiperda,
and Xanthopastis timais (Cramer) (Rogers et al.
1990a). Using the same methodologies, Rogers et
al. (1990b) determined that multiple species of
Mocis and Spodoptera were parasitized by the
nematode in Florida and Georgia, and it was the
first record of this parasite in North America. Sim-
mons & Rogers (1990a) determined the distribu-
tion and prevalence of the nematode in tropical
Americas, occurring in northern South America,
Mexico, Texas, Florida, Bermuda, most of the Car-
ibbean basin countries, and Central America.
Since 1990, numerous studies have been con-
ducted on this nematode including biology (Sim-
mons & Rogers 1994; Marti & Rogers 2000),
pathological effects on the host (Marti et al. 1990;
Rogers et al. 1993), infestation dynamics (Rogers
& Marti 1992a; Silvain & Remillet 1993; Rogers
& Marti 1993b), geographical distribution (Rog-
ers et al. 1993; Rogers et al. 1991; Simmons et al.
1991; Rogers et al. 1997; Marti et al. 2000), ecol-
ogy (Silvain & Remillet 1993), population profiles
(Rogers & Marti 1992a, 1994), mating behavior
(Simmons & Marti 1992), prevalence (Simmons &
Rogers 1990b), maintenance of colonies (Rogers &
Marti 1993a, b), host range (Simmons & Rogers
1996; Rogers & Marti 1996; Marti et al. 2000),
and the bionomics of host insects ofN. guyanense
(Rogers et al. 1996; Marti et al. 2000). Subse-
quently however, a study on the speciation in
Acugutturidae shows thatN. guyanense is limited
to the lepidopteran genera Spodoptera and Pseu-
daletia (Marti et al. 2002).
Simmons et al. (1991) studied the seasonal
chronology of the nematode in the tropical and
subtropical America, and determined that host
parasitism and nematode population density var-
ied among locations and over time. Examples in-
clude higher parasitism on male moths in Grenada
(77%) than in Texas (1%), and higher nematode
populations at lower latitudes than higher lati-
tudes. Simmons & Rogers (1991) also studied the
dispersal and seasonal occurrence ofN. guyanense
on FAW adults in the US. They found that nema-
tode populations and parasitism of FAW males
were higher in Eastern States than in the Plains,
Midwestern, and Central states, and that the per-
cent parasitism and the number of nematodes per
infested FAW changed over time at each location.

ACKNOWLEDGMENTS

The authors thank J. J. Hamm and 0. G. Marti
(USDA-ARS-Crop Protection & Management Research
Laboratory, Tifton, GA), for their critical review of ear-
lier versions of this manuscript, and thank O. G. Marti
for providing information on ectoparasitic nematodes.











The authors appreciate the assistance of Duncan Mc-
Clusky (University of Georgia Libraries, Tifton, GA),
and Marco Antonio Mellin-Rosas (CNRCB, Tecoman,
Colima, M6xico) in obtaining scientific literature. The
authors gratefully acknowledge and thank Susan
Drawdy (USDA-ARS-Crop Protection & Management
Research Laboratory, Tifton, GA) for her exceptional
technical assistance and dedication to this project. In-
sect identifications and verification of insect classifica-
tions were provided by several scientists associated
with the USDA/ARS Systematic Entomology Labora-
tory, including B. Carlson, E. E. Grissell, P. Marsh, N. E.
Woodley, and N. W. Gates. This paper is a contribution of
the University of Nebraska Agricultural Research Divi-
sion, Lincoln, NE 68583. Journal Series No. 13681, De-
partment of Entomology, University of Nebraska-
Lincoln. The authors thank Universidad de Colima, and
CONACYT-Mexico for supporting the senior author.


REFERENCES CITED

ALAM, M. M. 1979. Attempts at the biological control of
major insect pests of maize in Barbados, W. I. Symp.
On maize and peanut. Paramaribo, Suriname, Nov. 13-
18, 1978. Proc. Caribbean Food Crops Soc. 15: 127-135.
ALLEN, H. W. 1921. Notes on a bombylid parasite and a
polyhedral disease of the southern grass worm, La-
phygma frugiperda. J. Econ. Entomol. 14: 510-511.
ALTIERI, M. A. 1991. Increasing biodiversity to improve
insect pest management in agro-ecosystems, Chap-
ter XIV, pp. 165-182. In D. L. Hawksworth [ed.] The
Biodiversity of Microorganisms and Invertebrates:
Its Role in Sustainable Agriculture. CAB Interna-
tional, Wallingford, UK.
ALVAREZ, L. R., Y F. G. ROA. 1995. Comportamiento
parasitico de Telenomus sp. en Spodoptera fru-
giperda. Rev. Col. Entomol. 21: 191-197.
ANANTANARAYANAN, K. P. 1934. On the bionomics of a
eulophid Trichospilus pupivora Ferr.), a natural en-
emy of the coconut caterpillar (Nephantis serinopa,
Meyr.) in South India. Bull. Ent. Res. 25:55-61.
ANDREWS, K. L. 1988. Latin American research on
Spodoptera frugiperda (Lepidoptera: Noctuidae).
Florida Entomol. 71: 630-653.
ARCE, G. F., Y J. GARCIA G. 1995. Parasitismo natural y
biologia de Chelonus insularis Cresson (Hymenop-
tera: Braconidae) sobre Spodoptera frugiperda (J.E.
Smith) Lepidoptera: Noctuidae, en los valles cen-
trales de Oaxaca, pp. 48-49. En XVIII Congreso Na-
cional de Control Biol6gico. Soc. Mex. Contr. Biol.
Noviembre de 1995. Tapachula, Chiapas, M6xico.
ARMAS-GARCIA, J. L., Y J. L. AYALA-SIFONTES. 1987.
Parasitos de huevos de la palomilla del maiz,
Spodoptera frugiperda (J.E. Smith) (Lepidoptera:
Noctuidae). Centro-Agricola 14: 88-89.
ASHLEY, T. R. 1979. Classification and distribution of fall
armyworm parasites. Florida Entomol. 62: 114-123.
ASHLEY, T. R., E. R. MITCHELL, N. C. LEPPLA, AND E. E.
GRISSELL. 1980. Parasites attacking fall armyworm
larvae Spodoptera frugiperda in late planted field
corn. Florida Entomol. 63: 136-142.
ASHLEY, T. R., V. H. WADDILL, E. R. MITCHELL, AND J.
RYE. 1982. Impact of native parasites of the fall ar-
myworm, Spodoptera frugiperda (Lepidoptera: Noc-
tuidae), in South Florida and release of the exotic
parasite, Eiphosoma vitticole (Hymenoptera: Ichneu-
monidae). Environ. Entomol. 11: 833-837.


September 2003


ASHLEY, T. R. 1983. Growth pattern alterations in fall
armyworm, Spodoptera frugiperda larvae, after par-
asitization by Apanteles marginiventris, Campoletis
grioti, Chelonus insularis, and Eiphosoma vitticole.
Florida Entomol. 66: 260-266.
ASHLEY, T. R., C. S. BARFIELD, V. H. WADDILL, AND E. R.
MITCHELL. 1983. Parasitization of fall armyworm
larvae on volunteer corn, bermuda grass, and para-
grass. Florida Entomol. 66: 267-271.
ASHLEY, T. R. 1986. Geographical distributions and par-
asitization levels for parasitoids of the fall army-
worm, Spodoptera frugiperda. Florida Entomol.
69:516-524.
AVALOS, D. S. 1988. Moscas tachinidae de la provincia de
C6rdoba (Argentina). Rev. Peruana Entomol. 31:48-50.
AYALA-SIFONTES, J. L., M. GONZALEZ-ALVAREZ, H.
GRILLO-RAVELO, Y S. CABALLERO-FIGUEROA. 1978.
Observaciones preliminares sobre el ciclo biol6gico
de Diadegma sp., parasito de Heliothis virescens
(Fabricius). Centro Agricola 5: 1-13.
AYALA-SIFONTES, J. L., J. RAMIREZ-OBREG6N, S. ROD-
RIGUEZ-CUELLAR, Y J. L. ARMAS-GARCIA. 1988.
Habitos oviposicionales de Spodoptera frugiperda
(J.E. Smith) (Lepidoptera: Noctuidae) en plants de
arroz. Rev. Centro-Agricola 15: 11-15.
AYALA-SIFONTES, J. L. ARMAS-GARCIA, T. RODRIGUEZ, Y
M. BORGES-SOTO. 1992. Parasitismo de las puestas
de Spodoptera exigua (Hubner) por Telenomus sp. en
condiciones de laboratorio y campo. Rev. Centro-
Agricola 19: 42-46.
AYQUI-VILCA, S. 1993. Morfologia y biologia de Campo-
letis curvicauda (Hymenoptera: Ichneumonidae)
parasitoide de Spodoptera frugiperda. Rev. Peruana
Entomol. 35: 31-36.
BAHENA, J. F., Y O. GARCIA M. 1991. Enemigos naturales
del gusano cogollero del maiz Spodoptera frugiperda
(J. E. Smith) (Lepidoptera: Noctuidae) en Morelos,
M6xico, pp. 54-61 En XIV Congreso Nacional de Con-
trol Biol6gico. Soc. Mex. Contr. Biol. Octubre 10-11,
1991. Buenavista, Saltillo, Coahuila, M6xico.
BARILLA-VERA, M. 1989. T6cnica de cria del gusano
cogollero y sus parasitoids, pp. 59-62. En XII Re-
uni6n Nacional de Control Biol6gico. Secretaria de
Agriculture y Recursos Hidrailicos-DGSV, Noviem-
bre 22-24, 1989, Torre6n, Coahuila, M6xico.
BECHINSKI, E. J., AND L. P. PEDIGO. 1983. Green clover-
worm (Lepidoptera: Noctuidae) population dynam-
ics: Pupal life table studies in Iowa soybeans.
Environ. Entomol. 12: 656-661.
BENZING, A., R. G. KLEESPIES, Y F. PONCE. 2000. Mor-
talidad natural de larvas de Noctuidae (Lepidoptera)
en los Andes ecuatorianos: un primer acercamiento.
Rev. Colombiana Entomol. 26: 57-60.
BIANCHI, F. A. 1944. The recent introduction of army-
worm (S. exempta, S. exigua, S. frugiperda) para-
sites from Texas. Hawaii Planter's Rec. 48: 203-212.
BLANCHARD, E. E. 1963. Dipteros parasitos de Noctu-
idae argentinos. Rev. Inv. Agrop. INTA 17: 129-254.
BRUNER, S. C., L. C. SCARAMUZZA, AND A. R. OTERO.
1975. Catalogo de los insects que atacan a las plan-
tas econ6micas de Cuba. 2nda. Edici6n. Academia de
Ciencias de Cuba. Institute de Zoologia. 401 pp.
BUTLER, G. D., JR. 1958a. Tachinid flies reared from
lepidopterous larvae in Arizona, 1957. J. Econ. Ento-
mol. 51: 561-62.
BUTLER, G. D., JR. 1958b. Braconid wasps reared from
lepidopterous larvae in Arizona, 1957. Pan-Pac. Ent.
34: 221-223.


Florida Entomologist 86(3)







Armyworm Symposium 2002: Molina-Ochoa et al.


CAMPOS, L. 1965. Investigaciones sobre el control bi-
ol6gico del "cogollero" del maiz, Spodoptera fru-
giperda (J.E. Smith) y otros noctuideos. Rev.
Peruana Entomol. 8:126-131.
CANAS, L. A., AND R. J. O'NEIL. 1998. Applications of
sugar solutions to maize, and the impact of natural
enemies on fall armyworm. Int. J. Pest Management
44: 59-64.
CANSECO-ROMAN, 0. 1988. M6todo para la cria de
Spodoptera frugiperda (Smith) y sus parasitoides
Telenomus spp. En XI Reuni6n Nacional de Control
Biol6gico. Secretaria de Agricultura y Recursos
Hidraulicos, Agosto 15-17, 1988. Hermosillo, So-
nora. 11 pp.
CARLSON, R. W. 1979. Ichneumonidae, pp. 315-741 In
Krombein et al., Catalog of Hymenoptera in America
north of Mexico. Smithsonian Institution Press,
Washington.
CARPENTER, J. E., S. D. PAIR, AND G. P. FITT. 1994. Ich-
neumon promissorius (Hymenoptera: Ichneumoni-
dae): Development on North American hosts. J. Econ.
Entomol. 87: 929-932.
CARPENTER, J. E. 1995. Ichneumonpromissorius (Erich-
son) (Hymenoptera: Ichneumonidae): Factors affect-
ing fecundity, oviposition, and longevity. J. Entomol.
Sci. 30: 279-286.
CARPENTER, J. E., AND P. D. GREANY. 1998. Comparative
development and performance of artificially reared
versus host-reared Diapetimorpha introita (Cres-
son) (Hymenoptera: Ichneumonidae) wasps. Biologi-
cal Control 11: 203-208.
CARRILLO, H. 1980. Determinaci6n del parasitismo nat-
ural de gusano cogollero, Spodoptera frugiperda
(J.E. Smith) en Quintana Roo. Folia Entomol. M6x.
45: 111-112.
CASTRO, M. T., H. N. PITRE, AND D. H. MECKENSTOCK.
1989. Populations of fall armyworm, Spodoptera fru-
giperda (J.E. Smith), larvae and associated natural
enemies in sorghum and maize cropping systems in
Southern Honduras. Trop. Agric. 66: 259-264.
CAVE, R. D. 1993. Parasitoides larvales y pupales de
Spodoptera frugiperda (Smith) (Lepidoptera: Noctu-
idae) en Centro Am6rica con una clave para las espe-
cies encontradas en Honduras. Ceiba 34: 33-56.
CAVE, R. D., Y N. M. ACOSTA. 1999. Telenomus remus
Nixon: un parasitoide en el control biol6gico del gu-
sano cogollero, Spodoptera frugiperda (Smith).
Ceiba 40: 215-227.
CHADWICK, C. E., AND M. I. NIKITIN. 1976. Records of
parasitism in the families Ichneumonidae, Braconidae
and Aulacidae. J. Entomol. Soc. Aust. (N.S.W.) 9: 28-38.
CICEROS, O. J., J. M. MORALES, W. H. KING C., Y G.
SANCHEZ, G. 1995. Meteorus laphygmae (Viereck)
endoparasito del gusano cogollero en sorgo, pp. 56.
En XVIII Congreso Nacional de Control Biol6gico.
Soc. Mex. Contr. Biol. Noviembre de 1995. Tapa-
chula, Chiapas, M6xico.
CLARIDGE, M. F. 1991. Genetic and biological diversity
of insect pests and their natural enemies. Chapter
XV, pp. 183-194 In D. L. Hawksworth [ed.]. The
Biodiversity of Microorganisms and Invertebrates:
Its role in Sustainable Agriculture. CAB Interna-
tional, Wallingford, UK.
CORONADO, B. J. M., YE. RUIZ, C. 1991. Brac6nidos (Hy-
menoptera) de la Reserva "El Cielo" de Tamaulipas,
pp.72-76. En XIV Congreso Nacional de Control Bi-
ol6gico. Soc. M6x. Contr. Biol. Octubre 10-11, 1991.
Buenavista, Saltillo, Coahuila, M6xico.


CORREA-FIGUEIREDO, M. DE L. C., I. CRUZ, E T. M. C. D.
LUCIA. 1999. Control integrado de Spodoptera fru-
giperda (Smith & Abbott) utilizando-se o parasitoide
Telenomus remus Nixon. Pesq. Agrop. Bras. 34:
1975-1982.
CORTES, R. 1980. Neotropical Tachinidae (Diptera) I.
Notes, records, distribution and descriptions. Rev.
Bras. Entomol. 24: 105-110.
CORTEZ, M. H., Y J. TRUJILLO A. 1994. Incidencia de gu-
sano cogollero y sus enemigos naturales en tres agro-
sistemas de maiz. Turrialba 44: 1-9.
COSTA-LIMA, A. 1949. Insetos do Brasil, 6to. Tomo. Es-
cola Nacional de Agronomia. Seria Didatica No. 8:
184-185.
COSTA-LIMA, A. 1962. Insectos do Brasil. XII. Himenop-
teros. Rio de Janeiro, Escola Nac. Agr., 1962. 393 p.
(Ser. No. 14).
CRESSON, E. T. 1869. List of the North American species
of the genusAleiodes Wesmael. Trans. Am. Entomol.
Soc. 2: 377-382.
CRUZ, I., M. L. C. FIGUEIREDO, E. P. GONCALVES, D. A.
N. LIMA, E E. E. DINIZ. 1997a. Efeito da idade de
lagartas de Spodoptera frugiperda (Smith) (Lepi-
doptera: Noctuidae) no desempenho do parasitoide
Campoletis flavicincta (Ashmead) (Hymenoptera:
Ichneumonidae) e consume foliar por lagartas para-
sitadas e nao-parasitadas. An. Soc. Entomol. Brasil
26: 229-234.
CRUZ, I., M. L. C. FIGUEREIDO, F. H. VALICENTE, E A. C.
OLIVEIRA. 1997b. Application rate trials with a nu-
clear polyhedrosis virus to control Spodoptera fru-
giperda (Smith) on maize. An. Soc. Entomol. Brasil
26: 145-152.
CRUZ, I., M. L. C. FIGUEIREDO, E M. J. MATOSO. 1999.
Control biol6gico de Spodoptera frugiperda utili-
zando o parasitoide de ovos Trichogramma. Circular
T6cnica No. 30, Centro Nacional de Pesquisa de
Milho e Sorgo (CNPMS), Sete Lagoes, Brazil, 40 pp.
CURRAN, C. H. 1927. A new tachinid parasitic on army-
worm in Mexico. Proc. Hawaii Ent. Soc. 6: 497-498.
DANKS, H. V. 1975. Factors determining levels of parasit-
ism by Winthemia rufopicta (Diptera: Tachinidae),
with particular reference to Heliothis spp. (Lepi-
doptera: Noctuidae) as hosts. Can. Ent. 107: 655-684.
DE SA, L. A. N., AND J. R. P. PARRA. 1994. Natural par-
asitism of Spodoptera frugiperda and Helicoverpa
zea (Lepidoptera: Noctuidae) eggs in corn by Tri-
chogramma pretiosum (Hymenoptera: Trichogram-
matidae) in Brazil. Florida Entomol. 77: 185-188.
DE SANTIS, L., Y L. ESQUIVEL. 1966. Tercera lista de hi-
men6pteros parasitos de los insects de la Republica
Argentina. Rev. Mus. La Plata 9: 47-217.
DE SANTIS, L. 1967. Catalogo de los himen6pteros ar-
gentinos de la series parasitica, incluyendo Bethy-
loidea. Com. Invest. Cient. Prov. De Buenos Aires,
Gobernaci6n. La Plata, 337 p.
DE SANTIS, L. 1979. Catalogo de los himen6pteros Calc-
idoideos de Am6rica al Sur de los Estados Unidos.
Com. Invest. Cient. Prov. De Buenos Aires, Gober-
naci6n, 488 p.
DE SANTIS, L. 1980. Nueva sinonimia, nueva combi-
naci6n y nuevas citas de Himen6pteros Calcicoideos
para la Republica Argentina. Neotr6pica 26:153-154.
DE SANTIS, L. 1989. Catalogo de los Himen6pteros Calc-
idoideos (Hymenoptera) al Sur de los Estados Unidos.
Segundo Suplemento. Acta Ent. Chilena 15: 9-90.
DE SANTIS, L., Y P. FIDALGO. 1994. Catalogo de los Hi-
men6pteros Calcicoideos de Am6rica al Sur de los











Estados Unidos. Tercer Suplemento. Acad. Nac.
Agron. Vet. (Buenos Aires, Argentina) 13: 154 pp.
DENT, D. 2000. Insect pest management. 2nd Ed. CABI
Publishing, University Press, Cambridge, UK, 410 pp.
DEW, J. A. 1913. Fall armyworm Laphygma frugiperda
(S&A). J. Econ. Entomol. 6: 361-366.
ENKERLIN, D. 1975. Review of Spodoptera in Lation
America. Summary of research at Monterrey Tech.
Inst., N.L. Mexico. 13 pp.
ESCALANTE G., J. A. 1974. Contribuci6n al conocimiento
de la biologia de Heliothis zea y Spodoptera frugiperda,
en el Cusco. Rev. Peruana Entomol. 17: 121-122.
ESTRADA R., F. A. 1960. Lista preliminary de insects aso-
ciados al maiz en Nicaragua. Turrialba, 10: 68-73.
ETCHEVERRY, M. 1957. Laphygma frugiperda (Abbot
and Smith) en Chile (Lepidoptera: Noctuidae). Rev.
Chilena Entomol. 5: 183-192.
FENNAH, R. G. 1947. The insect pests of food-crops in
the Lesser Antilles. Dept. Agric. Windward IS., St.
George's Grenada, B.W.I. pp. 77-79.
FERNANDEZ, B. R. I., Y A. S. CLAVIJO. 1984. Efectos de
dos insecticides (uno quimico y otro biol6gico) sobre
el parasitismo observado en larvas de Spodoptera
frugiperda (S.) provenientes de parcelas experimen-
tales de maiz. Rev. Fac. Agron. Univ. Central
Venezuela 13: 101-109.
FERNANDEZ, B. R. I., Y B. J. B. TERAN. 1990a. Biologia
de Meteorus laphygmae Viereck (Hymenoptera: Bra-
conidae). Parte I. Fases de huevo, larva y pupa. Rev.
Fac. Agron. Univ. Central Venezuela 16: 177-198.
FERNANDEZ, B. R. I., Y B. J. B. TERAN. 1990b. Biologia
de Meteorus laphygmae Viereck (Hymenoptera: Bra-
conidae). Parte II. Adulto, partenogenesis e interrel-
aciones parasito-hospedero. Rev. Fac. Agron. Univ.
Central Venezuela 16: 199-206.
FERRER, W. F. 1998a. Alcances sobre el control biol6gico
en la region andina de Sudam6rica, pp. 25-26 En
Aportes del control biol6gico en la agriculture sos-
tenible. I Congr. Latinoam. Sec. Reg. Neotrop. Org.
Inter. Lucha Biol. Mayo 18-22, 1998, Lima, Peru.
FERRER, W. F. 1998b. El control biol6gico en el control de
plagas de importancia econ6mica en maiz y cafia, pp.
48-49. En Aportes del control biol6gico en la agricul-
tura sostenible. I Congr. Latinoam. Sec. Reg. Neotrop.
Org. Inter. Lucha Biol. Mayo 18-22, 1998, Lima, Peru.
FITT, G. P., AND J. C. DALY. 1990. Abundance of over-
wintering pupae and the spring generation of Heli-
coverpa spp. (Lepidoptera: Noctuidae) in Northern
New South Wales, Australia: implications for pest
management. J. Econ. Entomol. 83: 1827-1836.
FLORES-DAVILA, M., L. A. AGUIRRE-URIBE, Y E. RUIZ-
CANCINO. 1991. Ichenumonidae (Hymenoptera) del
Sureste de Coahuila, pp. 82-87 En XIV Congreso Na-
cional de Control Biol6gico. Soc. Mex. Contr. Biol.
Octubre 10-11, 1991. Buenavista, Saltillo, Coahuila,
M6xico.
GARCIA-LAGUNAS, E. D. 1988. Evaluaci6n del parasitoide
Telenomus solitus y Telenomus remus (Hymenoptera:
Scelionidae) sobre huevecillo de Spodoptera frugi-
perda (Smith) en condiciones de campo en Emiliano
Zapata, Mor. En XI Reuni6n Nacional de Control
Biol6gico. Secretaria de Agricultura y Recursos
Hidraulicos, Agosto 15-17, 1988. Hermosillo, So-
nora. 12 pp.
GIRALDO-VANEGAS, H., Y J. L. GARCIA R. 1992. Determi-
naci6n del numero de instares de Eiphosoma vitti-
cole Cresson (Hymenoptera: Ichneumonidae). Bol.
Entomol. Venezolana 7: 133-137.


September 2003


GIRALDO-VANEGAS, H., Y J. L. GARCIA R. 1994a. Com-
portamiento, descripci6n y tiempo de desarrollo de
los estadios inmaduros de Eiphosoma vitticolle Cres-
son (Hymenoptera: Ichneumonidae), parasito de
Spodoptera frugiperda (J.E. Smith) (Lepidoptera:
Noctuidae). Agron. Trop. Maracay 44: 645-665.
GIRALDO-VANEGAS, H., Y J. L. GARCIA R. 1994b. Com-
portamiento de los adults de Eiphosoma vitticole
Cresson (Hymenoptera: Ichneumonidae), parasito
de Spodoptera frugiperda (Smith) (Lepidoptera:
Noctuidae). Bol. Entomol. Venezolana 9: 15-20.
GIRALDO-VARGAS, H., Y J. L. GARCIA R. 1995. Influencia
de la alimentaci6n sobre la capacidad reproductive
de Eiphosoma vitticole Cresson (Hymenoptera: Ich-
neumonidae), parasito de Spodoptera frugiperda
(J.E. Smith). Agron. Trop. Maracay 45: 159-170.
GLADSTONE, S. H. 1991. Parasitos del cogollero,
Spodoptera frugiperda Smith (Lepidoptera: Noctu-
idae) en maiz sembrado en la 6poca seca en Nicara-
gua. Ceiba 32: 201-206.
GONCALVES, C. R. 1973. Observacoes sobre hospedeiros
de himenopteros da familiar Ichneumonidae, no Bra-
sil. An. Soc. Entomol. Brasil. 2: 31-36.
GONCALVES, C. R., AND A. J. L. GONCALVES. 1973. Novas
observacoes sobre insetos hospedeiros de moscas da
familiar Tachinidae (Diptera). Agronomia 31: 9-15.
GONZALEZ-NARVAEZ, C. E., YJ. L. ZOCCO. 1996. Control
integrado de Spodoptera frugiperda (Smith) utili-
zando Telenomus remus (Nixon) en Zea mays L. Rev.
Invest. Agric. DANAC 1: 1-5.
GREANY, P. D., AND J. E. CARPENTER. 1998. Culture me-
dium for parasitic and predaceous insects. U.S.
Patent No. 5799607.
GROSS, H. R., JR., AND S. D. PAIR 1986. The fall army-
worm: Status and expectations of biological control
with parasitoids and predators. Florida Entomol.
69:502-515.
GROSS, H. R., JR., AND S. D. PAIR. 1991. Seasonal distri-
bution, response and host developmental stage, and
screened-cage performance ofArchytas marmoratus
(Diptera: Tachinidae) and Ophion flavidus
(Hymenoptera: Ichneumonidae) on Spodoptera fru-
giperda (Lepidoptera: Noctuidae). Florida Entomol.
74: 237-245.
GUAGLIAMI, P. 1962. Las plagas de la cafia de azucar en
Venezuela. Maracay, Centro de Investigaciones
Agron6micas. 1962. V.2, pp. 520-521. (Monografia 2).
GUEVARA, M. A., C. G. PEREZ, Y F. J. GUTIERREZ. 1979.
Dinamica de poblaciones de insects ben6ficos en el
cultivo del maiz. VII Reuni6n Nacional de Control
Biol6gico. Secretaria de Agricultura y Recursos
Hidraulicos, Sanidad Vegetal. 24-26 de Abril de
1979, Veracruz, Veracruz, M6xico, 7 pp.
GUIMARAES, J. H. 1971. "Fam. Tachinidae (Larvaevori-
cidae)", In A catalogue of the Diptera of the America
South of the United States. Mus. Zool. Univ. Sao
Paulo 104: 1-333.
GUIMARAES, J. H. 1977. Host-parasite and parasite-host
catalogue of South American Tachinidae (Diptera).
Arq. Zool. Mus. Zool. Sao Paulo 28: 1-131.
GUIMARAES, J. H. 1983. Taxonomy of Brazilian flies of
the genus Lespesia (Robineau-Desvoidy) (Diptera,
Tachinidae). Papeis Avulsos de Zoologia 35: 11-30.
GUTIERREZ-RODRIGUEZ, F. J. 1982. T6cnica para cria
masiva de Euplectrus plathypenae (Howard) (Hy-
menoptera: Eulophidae) y su hospedero Spodoptera
frugiperda (Smith) (Lepidoptera: Noctuidae), pp. 17-
24. En X Reuni6n Nacional de Control Biol6gico.


Florida Entomologist 86(3)







Armyworm Symposium 2002: Molina-Ochoa et al.


Grupo Sectorial Agropecuario y Forestal. Secretaria
de Agriculture y Recursos Hidraulicos. Abril 1982,
Durango, Durango, M6xico.
HAMM, J. J., L. D. CHANDLER, AND H. R. SUMNER. 1994.
Field tests with a fluorescent brightener to enhance
infectivity of fall armyworm (Lepidoptera: Noctu-
idae) nuclear polyhedrosis virus. Florida Entomol.
77: 425-437.
HERNANDEZ, D., F. FERRER, Y B. LINARES. 1989. Intro-
ducci6n de Telenomus remus Nixon (Hym: Scelion-
idae) para controlar Spodoptera frugiperda (Lep.:
Noctuidae) en Yaritagua-Venezuela. Agron. Trop.
Maracay 39: 199-205.
HERNANDEZ, D., Y F. DIAZ. 1995. Efecto de la edad del
parasitoide Telenomus remus Nixon (Hymenoptera:
Scelionidae) sobre su capacidad de ovipostura y pro-
porci6n sexual de la descendencia. Bol Entomol.
Venezolana 10: 167-175.
HERNANDEZ, D., Y F. DIAZ. 1996a. Efecto de la edad del
hospedero Spodoptera frugiperda (Smith) (Lepi-
doptera: Noctuidae) sobre el parasitismo y la propor-
ci6n sexual de la descendencia (psd) de Telenomus
remus Nixon (Hymenoptera: Scelionidae). Bol. Ento-
mol. Venezolana 11: 27-32.
HERNANDEZ, D., Y F. DIAZ. 1996b. Efecto de la temper-
atura sobre el desarrollo de Telenomus remus Nixon
(Hymenoptera: Scelionidae) parasitoide de
Spodoptera frugiperda (Smith) (Lepidoptera: Noctu-
idae). Bol. Entomol. Venezolana 11: 149-153.
HERRERA-ARANGUENA, J. M. 1998. El control biol6gico
en la represi6n de plagas de importancia econ6mica
en el algodonero, pp. 40-44. En Aportes del control
biol6gico en la agriculture sostenible. I Congr. Lati-
noam. Sec. Reg. Neotrop. Org. Inter. Lucha Biol.
Mayo 18-22, 1998, Lima, Peru.
HOFMASTER, R. N., AND D. E. GREENWOOD. 1949. Fall
armyworm control on forage and truck crops. J.
Econ. Ent. 42: 502-506.
HOGG, D. B., R. E. ANDERSON, AND H. N. PITRE. 1982.
Early-season parasitization of fall armyworm (Lepi-
doptera: Noctuidae) larvae in Mississippi. Florida
Entomol. 65: 584-585.
HUIS, A. V. 1981. Integrated pest management in the
small farmer's maize crop in Nicaragua. Med.
Landb. Wageningen 81: 221 pp.
HYNES, H. B. N. 1942. Lepidopterous pests of maize in
Trinidad. Tropical Agric. 29: 194-202.
IRVING, N. S. 1978. Report on the Antigua, Monserrat
and St. Kitts-Nevis-Anguilla entomology pro-
gramme. Centre for Overseas pest Research, Lon-
don, United Kingdom, 55 pp.
ISENHOUR, D. J. 1988. Interactions between two hy-
menopterous parasitoids of the fall armyworm (Lepi-
doptera: Noctuidae). Environ. Entomol. 17: 616-620.
JEWETT, D. K., AND J. E. CARPENTER. 1998. Chemical
communication between males and females of a pupal
parsitoid Diapetimorpha introita (Hymenoptera: Ich-
neumonidae). Ann. Entomol. Soc. Am. 91: 748-753.
JEWETT, D. K., AND J. E. CARPENTER. 2001. Seasonal
abundance of pupal parasitoid, Diapetimorpha in-
troita (Hymenoptera: Ichneumonidae). Florida Ento-
mol. 84: 50-54.
JOHANNES-TOONDERS, T., Y J. L. CARRILLO-SANCHEZ.
1987. Evaluaci6n de la efectividad de Trichogramma
spp. (Hymenoptera: Trichogrammatidae) en el com-
bate de Spodoptera frugiperda (J. E. Smith) (Lepi-
doptera: Noctuidae). Recomendaciones para su uso.
Agrociencia 67: 75-84.


JONES, T. H. 1913. Some notes on Laphygma frugiperda
S. and A. in Porto Rico. J. Econ. Entomol. 6: 230-236.
KELLER, M. S. 1980. Effects of temperature and corn
phenology on Fall Armyworm Biology. Masters The-
sis, University of Florida, 83 pp.
KING, A. B. S., AND J. L. SAUNDERS. 1984. Las plagas in-
vertebredas de cultivos anuales alimenticios en
Am6rica Central. Overseas Development Adminis-
tration, London, 62 pp.
LACAYO, L. 1977. Especies parasiticas de Spodoptera
frugiperda Smith, Diatraea lineolata (Wer.) y Tri-
choplusia ni (Hbn) en zonas de Managua y Ma-
satepe. XXIII Reuni6n Annual del PCCMCA,
Panama, Panama 1977, 28 pp.
LEZAMA-GUTIERREZ, R., J. J. HAMM, J. MOLINA-OCHOA,
M. LOPEZ-EDWARDS, A. PESCADOR-RUBIO, M.
GONZALEZ-RAMIREZ, AND E. L. STYER. 2001. Occur-
rence of entomopathogens of Spodoptera frugiperda
(Lepidoptera: Noctuidae) in the Mexican states of
Michoacan, Colima, Jalisco and Tamaulipas. Florida
Entomol. 84: 23-30.
LOYA-RAMIREZ, J. 1978. Estudio sobre el gusano cogol-
lero, Spodoptera frugiperda (J.E. Smith), en More-
los. Institute Nacional de Investigaciones Agricolas
(INIA), M6xico. Informe T6cnico de la Coordinaci6n
del Apoyo Entomol6gico 3: 45-55.
LUCCHINI, F., E A. A. ALMEIDA. 1980. Parasitas da
Spodoptera frugiperda (Smith & Abbot, 1797) (Lep.,
Noctuidae), lagarta do cartucho do milho, encon-
trado em Ponta Grossa, PR. An. Soc. Entomol. Bra-
sil. 9: 115-121.
LUGINBILL, P. 1928. The fall armyworm. U.S. Dept. Ag-
ric. Tech. Bull. 34, 91 pp.
MACHUCA, J. R., P. ARRETZ, Y J. ARAYA. 1989. Presencia
de Campoletis sonorensis (Cameron, 1986) (Hy-
menoptera: Ichneumonidae) en Chile. Acta Entomol.
Chilena 15: 269-270.
MAES, J. M. 1989. Catalogo de los insects controladores
biol6gicos en Nicaragua. Vol. III. Insectos Parasi-
toides. Rev. Nicar. Entomol. 10: 1-138.
MALAUSA, J. C. 1981. Etude de la dynamique des popu-
lations des chenilles de Spodoptera frugiperda Abbot
& Smith et d'Heliothis zea Boddie (Lepidoptera: Noc-
tuidae) sur deux varieties de mais en Guadeloupe
(Antilles Francaises). Agronomie 1: 701-705.
MALAUSA, J. 1983. Etude bioecologique des noctuelles
(Lepidoptera: Noctuidae) des Antilles Francaises.
Bol. Soc. Portuguesa Entomol. 2: 49-67.
MARIN-ACOSTA, J. C. 1966. Nota sobre Euplectrus plat-
hypenae HOW., ectoparasito de Laphygma frugi-
perda S. y A. y Prodenia eridania (Cramer). Agron.
Trop. 16: 155-159.
MARSH, P. M. 1978. The braconid parasites (Hy-
menoptera) of Heliothis species (Lepidoptera: Noctu-
idae). Proc. Entomol. Soc. Wash. 80: 15-36.
MARSH, P. M., AND S. R. SHAW. 2001. Revision of North
AmericanAleiodes Wesmael (Part 6): the Gasterator
(Jurine) and Unipunctator (Thunberg) species-
groups (Hymenoptera: Braconidae: Rogadinae).
Proc. Entomol. Soc. Wash. 103: 291-307.
MARTI, O. G., C. E. ROGERS, J. F. SILVAIN, AND A. M.
SIMMONS. 1990. Pathological effects of an ectopara-
sitic nematode Noctuidonema guyanense (Nema-
toda: Aphelenchoididae) on adults of the fall
armyworm (Lepidoptera: Noctuidae). Ann. Entomol.
Soc. Am. 83: 956-960.
MARTI, O. G. JR., AND C. E. ROGERS. 2000. Effect ofNoc-
tuidonema guyanense (Nematoda: Acugutturidae)











on the longevity of feral male Spodoptera frugiperda
(Lepidoptera: Noctuidae) moths. J. Entomol. Sci. 35:
259-266.
MARTI, O. G. JR., B. LALANNE-CASSOU, J. F. SILVAIN, A.
KERMARREC, AND A. M. SIMMONS. 2000. Ectopara-
sitic nematodes (Aphelenchoididae: Acugutturidae)
of Lepidoptera and Blattodea in Guadeloupe. Nema-
tology 2: 669-684.
MARTI, O. G., JR., J. F. SILVAIN, AND B. J. ADAMS. 2002.
Speciation in the Acugutturidae (Nematoda: Aphe-
lenchida). Nematology 4: 489-504.
MCCUTCHEON, G. S., W. Z. SALLEY, JR., AND S. G. TUR-
NIPSEED. 1983. Biology of Apanteles ruficrus, an im-
ported parasitoid of Pseudoplusia includes,
Trichoplusia ni, and Spodoptera frugiperda (Lepi-
doptera: Noctuidae). Environ. Entomol. 12: 1055-
1058.
MCCUTCHEON, G. S. 1991. Late-season parasitoids of
the fall armyworm in South Carolina. J. Agric. Ento-
mol. 8: 219-221.
MEDINA, T. M. C., D. P. CAMACHO, Z. J. E. LUQUE, Y P.
A. SIABATTO. 1988. Ciclo de vida y descripci6n de
Chelonus insularis Cresson (Hymenoptera: Bra-
conidae), parasito de Spodoptera spp. Rev. Colombi-
ana Ent. 14: 13-21.
MILWARD-DE AZEVEDO, E. M. V., J. R. POSTAL PARRA,
J. H. GUIMARAES, E R. DE PAULA-ALMEIDA. 1991a.
Aspectos da biologia de Archytas incertus (Diptera:
Tachinidae) e de suas inter-relacoes com Spodoptera
frugiperda (Lepidoptera: Noctuidae). 1. Metodologia
de criacao e determinacao do instar mais adequado
para a producao do parasitoide. Rev. Bras. Entomol.
35: 485-497.
MILWARD-DE AZEVEDO, E. M. V., J. H. GUIMARAES, E J.
R. POSTALI-PARRA. 1991b. Aspectos da biologia de
Archytas incertus (Diptera: Tachinidae) e de suas in-
ter-relacoes com Spodoptera frugiperda (Lepi-
doptera: Noctuidae). 2. Carga parasitariae de regiao
inoculacao no corpo do hospedero. Rev. Bras. Ento-
mol. 35: 499-507.
MILWARD-DE AZEVEDO, E. M. V., E J. R. POSTALLI-
PARRA. 1991c. Aspectos da biologia de Archytas in-
certus (Diptera: Tachinidae) e de suas inter-relacoes
com Spodoptera frugiperda (Lepidoptera: Noctu-
idae). 3. Influencia da temperature na ontogenia do
parasitoide. Rev. Bras. Entomol. 35: 509-516.
MILWARD-DE AZEVEDO, E. M. V., E J. R. POSTALI-PARRA.
1991d. Aspectos da biologia de Archytas incertus
(Diptera: Tachinidae) e de suas inter-relacoes com
Spodoptera frugiperda (Lepidoptera: Noctuidae). 4.
Sincronismo. Rev. Bras. Entomol. 35: 517-520.
MILWARD-DE AZEVEDO, E. M. V., J. R. POSTAL PARRA, E
J. H. GUIMARAES. 199 le. Aspectos da biologia de Ar-
chytas incertus (Diptera: Tachinidae) e de suas inter-
relacoes com Spodoptera frugiperda (Lepidoptera:
Noctuidae). 5. Fase reproductive e longevidade. Rev.
Bras. Entomol. 35: 521-530.
MITCHELL, E. R., V. H. WADDILL, AND T. R. ASHLEY.
1984. Population dynamics of the fall armyworm
(Lepidoptera: Noctuidae) and its larval parasites on
the whorl stage corn in pheromone-permeated field
environments. Environ. Entomol. 13: 1618-1623.
MOLINA-OCHOA, J., R. LEZAMA-GUTIERREZ, J. J. HAMM,
B. R. WISEMAN, AND M. LOPEZ-EDWARDS. 1999. Inte-
grated control of fall armyworm (Lepidoptera: Noctu-
idae) using resistant plants and entomopathogenic
nematodes (Rhabditida: Steinernematidae). Florida
Entomol. 82: 263-271.


September 2003


MOLINA-OCHOA, J., J. J. HAMM, R. LEZAMA-GUTIERREZ,
M. LOPEZ-EDWARDS, M. GONZALEZ-RAMIREZ, AND A.
PESCADOR-RUBIO. 2001. A survey of fall armyworm
(Lepidoptera: Noctuidae) parasitoids in the Mexican
states of Michoacan, Colima, Jalisco, and Tamauli-
pas. Florida Entomol. 84:31-36.
MOLINARI, A. M., Y S. D. AVALOS. 1997. Contribuci6n al
conocimiento de taquinidos (Diptera) parasitoides de
defoliadores (Lepidoptera) del cultivo de la soja. Rev.
Soc. Entomol. Argentina 56: 131-136.
MONTOYA-BURGOS, J. A. 1979. Estudio preliminary del
ectoparasito del g6nero Euplectrus del gusano cogol-
lero Spodoptera frugiperda. VII Reuni6n Nacional
de Control Biol6gico. Secretaria de Agricultura y Re-
cursos Hidraulicos, Sanidad Vegetal. 24-26 de Abril
de 1979, Veracruz, Veracruz, M6xico, 4 pp.
MONTOYA-BURGOS, J. A. 1980. Observaciones sobre la
importancia de Euplectrus sp. en la integraci6n de
los m6todos de control de plagas del maiz. VIII Re-
uni6n Nacional de Control Biol6gico. Secretaria de
Agriculture y Recursos Hidraulicos, Sanidad Vege-
tal. 22-25 de Abril de 1980, Manzanillo, Colima,
M6xico, 7 pp.
MORALES-PEREZ, A. 1982. Cria masiva del parasito Te-
lenomus sp. usando como huesped huevecillos de
Spodoptera exigua, pp. 13-16. En X Reuni6n Nacio-
nal de Control Biol6gico. Grupo Sectorial Agropec-
uario y Forestal. Secretaria de Agricultura y
Recursos Hidraulicos. Abril 1982, Durango, Du-
rango, M6xico.
MOREY, C. S. 1971. Biologia de Campoletis grioti (Blan-
chard) (Hymen.: Ichneumonidae) parasito de la
"lagarta cogollera del maiz" Spodoptera frugiperda
(J.E. Smith). Sociedad Entomol6gica del Peru, pp.
263-271. En Anales del Primer Congreso Latino-
americano de Entomologia, Cusco Peru, 12-18 de
Abril de 1971.
MOYA, M. 1980. Parasitismo de Bracon Kirkpatricki
Wilkinson (Hymenoptera-Braconidae) en larvas de
Spodoptera frugiperda Smith. Ibid. 14 pp.
MUESEBECK, C. F. W. 1921. A revision of the North
American species of Ichneumon-flies belonging to the
genus Apanteles. U.S. Nat. Mus., Proc. 58: 483-576.
MUESEBECK, C. F. W., K. V. KROMBEIN, AND H. K.
TOWNES (Eds.). 1951. Hymenoptera of America
North of Mexico. Synoptic Catalog. USDA Agric.
Monogr. No. 2 1420 pp.
MULOCK, B. S., S. S. SWEZEY, C. NARVAEZ, P. CASTILLO,
AND C. M. RIZO. 1990. Development ofbaculoviruses
as a contribution to biological control of lepidopterous
pests of basic grains in Nicaragua, pp. 179-183. In
Proc. Vth. Inter. Colloq. Invertebr. Pathol. Microb.
Control. Adelaide, Australia, 20-24 August 1990.
MYERS, J. G. 1932. Biological observations on some neo-
tropical parasitic hymenoptera. Trans. Ent. Soc.
Lond. 80: 121-136.
NAVA, R. A., Y N. S. CASTRO. 1991. Parasitismo del gu-
sano cogollero Spodoptera frugiperda (J.E. Smith)
en la region central del Estado de Tamaulipas., pp.
62-64 En XIV Congreso Nacional de Control Bi-
ol6gico. Soc. Mex. Contr. Biol. Octubre 10-11, 1991.
Buenavista, Saltillo, Coahuila, M6xico.
NICKLE, D. A. 1976. The peanut agroecosystem in Cen-
tral Florida. Ph.D. dissertation, University of Flor-
ida, Gainesville, 131 pp.
NOTZ, P. A. 1972. Parasitismo de Diptera e Hy-
menoptera sobre larvas de Spodoptera frugiperda
(Smith) (Lepidoptera: Noctuidae) recolectadas en


Florida Entomologist 86(3)







Armyworm Symposium 2002: Molina-Ochoa et al.


maiz, Maracay, Venezuela. Rev. Fac. Agron. Univ.
Central. Venezuela 6: 5-16.
ORTEG6N, E. J., N. C. TORRES, E. LUQUE, YA. SIABATTO.
1988. Estudio sobre la longevidad, habitos, progenie
y evaluaci6n preliminary de Meteorus laphygmae (Vi-
ereck), parasito de Spodoptera spp. Rev. Colombiana
Entomol. 14: 7-12.
PAINTER, R. H. 1955. Insects on corn and teosinte in
Guatemala. J. Econ. Ent. 48: 36-42.
PAIR, S. D. 1995. Biology and rearing of Diapetimorpha
introita (Cresson) (Hymenoptera: Ichneumonidae)
on host and non-host noctuid pupae. J. Entomol. Sci.
30: 468-480.
PAIR, S. D., AND H. R. GROSS, JR. 1984. Field mortality
of pupae of the fall armyworm, Spodoptera fru-
giperda (J.E. Smith), by predators and a newly dis-
covered parasitoid, Diapetimorpha introita. J.
Georgia Entomol. Soc. 19:22-26.
PAIR, S. D., J. R. RAULSTON, A. N. SPARKS, AND P. B.
MARTIN. 1986. Fall armyworm (Lepidoptera: Noctu-
idae) parasitoids: Differential spring distribution
and incidence on corn and sorghum in the southern
United States and Northeastern Mexico. Environ.
Entomol. 15: 342-348.
PAIR, S. D., AND H. R. GROSS, JR. 1989. Seasonal inci-
dence of fall armyworm (Lepidoptera: Noctuidae)
pupal parasitism in corn by Diapetimorpha introita
and Cryptus albitarsis (Hymenoptera: Ichneu-
monidae). J. Entomol. Sci. 24: 339-343.
PANTOJA, A., C. M. SMITH, AND J. F. ROBINSON. 1985.
Natural control agents affecting Spodoptera fru-
giperda (Lepidoptera: Noctuidae) infesting rice in
Puerto Rico. Florida Entomol. 68: 488-490.
PANTOJA, A., AND J. R. FUXA. 1992. Prevalence of biotic
control agents in the fall armyworm Spodoptera fru-
giperda (J. E. Smith) (Lepidoptera: Noctuidae). Folia
Entomol. Mex. 84: 79-84.
PARKER, H., P. BERRY, AND A. S. GUIDO. 1953. Host-par-
asite and parasite-host lists of insects reared in the
South American parasite laboratory. Montevideo,
Republica Oriental del Uruguay, 100 p.
PASSOA, S. 1983. Lista de los insects asociados con los
granos basicos y otros cultivos selectos en Honduras.
CEIBA 25. 97 pp.
PATEL, P. N., E M. E. M. HABIB. 1982. Ocorrencia natu-
ral de Aspergillus parasiticus em populacoes de
Spodoptera frugiperda (Abbot & Smith, 1797) (Lepi-
doptera: Noctuidae) e sua trasmissao por insects
parasites. Rev. Agric. Piracicaba 57: 223-232.
PATEL, P. N., E M. E. M. HABIB. 1984. Levantamento e
eficacia de insects parasites de Spodoptera fru-
giperda (Abbot & Smith, 1797) (Lepidoptera: Noctu-
idae). Rev. Agric. Brasil 59: 229-237.
PATEL, P. N., E M. E. M. HABIB. 1986. Levantamento e
eficacia de insects parasites de Spodoptera fru-
giperda (Abbot & Smith, 1797) (Lepidoptera: Noctu-
idae). Rev. Agric. Piracicaba 61: 93-100.
PAVUK, D. M., AND B. R. STINNER. 1991. New lepi-
doptera-parasitoid associations in weedy corn plant-
ings: a potential alternate host for Ostrinia nubilalis
(Lepidoptera: Pyralidae) parasitoids. Great Lakes
Entomol. 24:219-223.
PENA, G. 1980. Estudio de Bracon Kirkpatricki Wilkin-
son en diferentes huespedes, con fines de control bi-
ol6gico. VIII Reuni6n Nacional de Control Biol6gico,
Manzanillo, Col., Mex. 17 pp.
PERAZA-LIZARRAGA, J. C. 1982. Dinamica de poblaciones
de insects ben6ficos en cultivo de maiz en Durango,


Durango, pp. 91-95. En X Reuni6n Nacional de Con-
trol Biol6gico. Grupo Sectorial Agropecuario y For-
estal. Secretaria de Agricultura y Recursos
Hidraulicos. Abril 1982, Durango, Durango, M6xico.
PIERCE, W. D., AND T. E. HOLLOWAY. 1912. Notes on the
biology of Chelonus texanus Cress. J. Econ. Entomol.
5: 425-428.
PORTER, C. C. 1998. Guia de los g6neros de Ichneu-
monidae en la region Neantartica del Sur de
Sudam6rica. Opera Lilloana 42: 234 pp.
PRATISSOLI, D., J. B. TORRES, Y J. C. ZANUNCIO. 1999.
Selectividad de Trichogramma demoraesi (Hy-
menoptera: Trichogrammatidae) en huevos de tres
lepid6pteros inviabilizados. Agro-Ciencia 15: 75-80.
RAJAPAKSE, R. H. S., T. R. ASHLEY, AND V. H. WADDILL.
1985. Biology and host acceptance of Microplitis ma-
nilae (Hymenoptera: Braconidae) raised on fall ar-
myworm larvae Spodoptera frugiperda. Florida
Entomol. 68: 653-657.
RAVLIN, F. W., AND F. W. STEHR. 1984. Revision of the ge-
nus Archytas (Diptera: Tachinidae) for America North
of Mexico. Misc. Pub. Entomol. Soc. Am. 58, 59 pp.
REED, D. J. 1980. Toxicological and parasitological stud-
ies on the fall armyworm, Spodoptera frugiperda
(J.E. Smith), in Alabama. M.S. Thesis, Auburn Uni-
versity, Auburn, Alabama, 55 pp.
REMILLET, M., AND J. F. SILVAIN. 1988. Noctuidonema
guyanense n.g., n. sp. (Nematoda: Aphelenchoididae)
ectoparasite de noctuelles du genre Spodoptera
(Lepidoptera: Noctuidae). Rev. Nematol. 11: 21-24.
REZENDE, M. A. A., I. CRUZ, E T. M. C. DELLA LUCIA.
1994. Consumo foliar de milho e desenvolvimento de
lagartas de Spodoptera frugiperda (Smith) parasita-
das por Chelonus insularis (Cresson) (Hymenoptera:
Braconidae). An. Soc. Entomol. Brasil 23: 473-478.
REZENDE, M. A. A., I. CRUZ, E T. M. C. DELLA LUCIA.
1995a. Aspectos biologicos do parasitoide Chelonus
insularis (Cresson) (Hymenoptera, Braconidae) cria-
dos em ovos de Spodoptera frugiperda (Smith) (Lep-
idoptera: Noctuidae). Rev. Bras. Zool. 12: 779-784.
REZENDE, M. A. A., T. M. C. DELLA LUCIA, E I. CRUZ.
1995b. Comportamento de lagartas de Spodoptera
frugiperda (Lepidoptera: Noctuidae) parasitadas por
Chelonus insularis (Hymenoptera, Braconidae), sobre
plants de milho. Rev. Bras. Entomol. 39: 675-681.
RIGGIN, T. M., B. R. WISEMAN, D. J. ISENHOUR, AND K.
E. ESPELIE. 1992. Incidence of fall armyworm (Lepi-
doptera: Noctuidae) parasitoids on resistant and
susceptible corn genotypes. Environ. Entomol. 21:
888-895.
RIGGIN, T. M., K. E. ESPELIE, B. R. WISEMAN, AND D. J.
ISENHOUR. 1993. Distribution of fall armyworm
(Lepidoptera: Noctuidae) parasitoids on five corn
genotypes in South Georgia. Florida Entomol. 76:
292-302.
RIGGIN, T. M., B. R. WISEMAN, D. J. ISENHOUR, AND K.
E. ESPELIE. 1994. Functional response of Cotesia
marginiventris (Cresson) (Hym., Braconidae) to
Spodoptera frugiperda (J.E. Smith) (Lep., Noctu-
idae) on meridic diet containing resistant or suscep-
tible corn genotypes. J. Appl. Ent. 117: 144-150.
RODRIGUEZ-LUNA, S. J. 1982. Parasitismo natural de
Trichogramma sp. sobre huevecillos de Spodoptera
frugiperda (Smith) en maiz de temporal de Jiutepec,
Emiliano Zapata y Xotitepec, Mor., pp. 22-33. En X
Reuni6n Nacional de Control Biol6gico. Secretaria
de Agriculture y Recursos Hidraulicos, Abril 1982,
Durango, Durango, M6xico.











ROGERS, C. E., O. G. MARTI, A. M. SIMMONS, AND J. F.
SILVAIN. 1990a. Host range of Noctuidonema guyan-
ense (Nematoda: Aphelenchoididae): An ectoparasite
of moths in French Guiana. Environ. Entomol. 19:
795-798.
ROGERS, C. E., A. M. SIMMONS, AND O. G. MARTI. 1990b.
Parasitism of Lepidoptera adults by Noctuidonema
guyanense Remillet and Silvain (Nematoda: Aphe-
lenchoididae) in Southeastern United States. J. Ag-
ric. Entomol. 7: 241-245.
ROGERS, C. E., A. M. SIMMONS, AND O. G. MARTI. 1991.
Noctuidonema guyanense: an ectoparasitic nema-
tode of fall armyworm adults in the tropical Ameri-
cas. Florida Entomol. 74: 246-257.
ROGERS, C. E., AND O. G. MARTI, JR. 1992a. Noctui-
donema guyanense (Nematoda: Aphelenchoididae):
population profiles on male and female fall army-
worm moths. J. Entomol. Sci. 27: 354-360.
ROGERS, C. E., AND O. G. MARTI, JR. 1992b. Infestation
and dispersal of Noctuidonema guyanense (Nema-
toda: Aphelenchoididae) on Spodoptera frugiperda
(Lepidoptera: Noctuidae). Environ. Entomol. 21:
417-421.
ROGERS, C. E., AND O. G. MARTI, JR. 1993a. Infestation
dynamics and distribution of Noctuidonema guyan-
ense (Nematoda: Aphelenchoididae) on adults of
Spodoptera frugiperda and Mocis latipes (Lepi-
doptera: Noctuidae). Florida Entomol. 76: 326-333.
ROGERS, C. E., AND O. G. MARTI, JR. 1993b. Mainte-
nance of an ectoparasitic nematode, Noctuidonema
guyanense (Nematoda: Aphelenchoididae), on
Spodoptera frugiperda (Lepidoptera: Noctuidae)
moths. J. Entomol. Sci. 28: 335-337.
ROGERS, C. E., O. G. MARTI, JR., AND A. M. SIMMONS.
1993. Noctuidonema guyanense (Nematoda: Aph-
elenchoididae): Host range and pathogenicity to the
fall armyworm, Spodoptera frugiperda (Lepi-
doptera: Noctuidae), pp. 27-32. In Nematodes and
the biological control of insect pests. Bedding, R., R.
Akhurst & H. Kaya [eds.] CSIRO, Australia.
ROGERS, C. E., AND O. G. MARTI, JR. 1994. Population
structure and transfer success of Noctuidonema guy-
anense (Nematoda: Aphelenchoididae) on moths of
Spodoptera frugiperda (Lepidoptera: Noctuidae).
Ann. Entomol. Soc. Am. 87: 327-330.
ROGERS, C. E., AND O. G. MARTI, JR. 1996. Beet army-
worm (Spodoptera exigua) as a host for the ectopar-
asitic nematode, Noctuidonema guyanense. J. Agric.
Entomol. 13: 81-86.
ROGERS, C. E., O. G. MARTI, JR., L. D. CHANDLER, AND
A. M. SIMMONS. 1996. Bionomics of the fall army-
worm (Lepidoptera: Noctuidae) and its ectoparasitic
nematode, Noctuidonema guyanense, in South Geor-
gia. J. Entomol. Sci. 31: 209-217.
ROGERS, C. E., O. G. MARTI, JR., AND J. A. CLAYTON.
1997. Report of the ectoparasitic nematode Noctui-
donema guyanense (Acugutturidae) infesting Lepi-
doptera in the Fiji Islands. Nematologica 43: 505-506.
ROHLFS, W. M., AND T. P. MACK. 1985. Seasonal para-
sitism rates, host size, and adult emergence pattern
of parasitoids of the fall armyworm, Spodoptera fru-
giperda (J.E. Smith), with emphasis on Ophion fla-
vidus Brulle (Hymenoptera: Ichneumonidae). Ann.
Entomol. Soc. Am. 78: 217-220.
RYDER, W. D., AND N. PIEDRA. 1968. Field investiga-
tions into the control for some insects pests of maize
in Cuba with high-volume sprays of DDT, diazinon,
malathion, and phosdrin. Ibid 2: 295-303.


September 2003


RYDER, W. D., AND N. PULGAR 1969. A note on parasit-
ism of the fall armyworm, Spodoptera frugiperda, on
maize. Rev. cubana Cienc. Agric. (English ed.) 3: 267-
271.
SABROSKY, C. W. 1981. A partial revision of the genus
Eucelatoria (Diptera: Tachinidae), including impor-
tant parasites of Heliothis. Tech. Bull. Sci. & Educ.
Adm. USDA, No. 1635, 18 pp.
SANCHEZ-GARCIA, J. A., J. ROMERO-NAPOLES, S. RAMA-
RIZ-ALARCON, S. ANAYA-ROSALES, Y J. L. CARILLO-
SANCHEZ. 1998. G6neros de Braconidae del estado de
Guanajuato (Insecta: Hymenoptera). Acta Zool.
Mex. 74: 59-137.
SARMIENTO, M. J., Y R. V. RAZURI. 1978. Bacillus thur-
ingiensis en el control de Spodoptera frugiperda y de
Diatraea saccharalis en maiz. Rev. Peruana Ento-
mol. 21: 121-124.
SEGEREN, P. A., AND S. R. SHARMA. 1979. Insect control
on maize in Suriname. Proc. Caribbean Food Crops
Soc. 15: 142-155.
SILVA, F. M. A., H. G. FOWLER, E R. N. S. LEMOS. 1997.
Parasitismo em lagarta-do-cartucho, Spodoptera
frugiperda (Smith), na regiao do Triangulo Mineiro,
MG. An. Soc. Entomol. Brasil 26: 235-241.
SILVAIN, J. F., AND M. REMILLET. 1993. Ecology and bi-
ology of Noctuidonema guyanense (Nematoda: Aph-
elenchoididae), an ectoparasite of Spodoptera
frugiperda (Lep., Noctuidae), in French Guiana. En-
tomophaga 38: 465-474.
SILVEIRA, G. A., Y A. RUFFINELLI. 1956. Primer catalogo
de los parasitos y predatores encontrados en el Uru-
guay. Bol. Fac. Agron. Montev. 32: 80 pp.
SILVEIRA, J. C. F. DA, E. T. SASAKI, M. A. FORMER, M. S.
HONDA, E M. H. CALAFIORI. 1987. Ocorrencia de par-
asitoides de Spodoptera frugiperda (J.E. Smith,
1797) em cultural de milho, em Espirito Santo do Pin-
hal, SP. Ecossistema 12: 41-44.
SIMMONS, A. M., AND C. E. ROGERS. 1990a. Tempera-
ture and humidity effects on Noctuidonema (Nema-
toda: Aphelenchoididae), an ectoparasite of adult
Spodoptera frugiperda (Lepidoptera: Noctuidae),
and transfer success during host mating. Ann. Ento-
mol. Soc. Am. 83: 1084-1087.
SIMMONS, A. M., AND C. E ROGERS. 1990b. Distribution
and prevalence of an ectoparasitic nematode, Noctu-
idonema guyanense, on moths of the fall armyworm
(Lepidoptera: Noctuidae) in the tropical Americas. J.
Entomol. Sci. 25: 510-518.
SIMMONS, A. M., AND C. E. ROGERS. 1991. Dispersal and
seasonal occurrence of Noctuidonema guyanense, an
ectoparasitic nematode of adult fall armyworm (Lep-
idoptera: Noctuidae), in the United States. J. Ento-
mol. Sci. 26: 136-148.
SIMMONS, A. M., C. E. ROGERS, K. U. BUCKMIRE, B.
GRAY, K. D. MONKMAN, A. PANTOJA, J. R. RAULSTON,
AND V. H. WADDILL. 1991. Seasonal chronology of
Noctuidonema, an ectoprasitic nematode of adult
moths, in tropical and subtropical America. Florida
Entomol. 74: 311-319.
SIMMONS, A. M., AND O. G. MARTI, JR. 1992. Mating by
the fall armyworm (Lepidoptera: Noctuidae): fre-
quency, duration, and effect of temperature. Envi-
ron. Entomol. 21: 371-375.
SIMMONS, A. M., AND C. E. ROGERS. 1994. Effect of an
ectoparasitic nematode, Noctuidonema guyanense,
on adult longevity and egg fertility in Spodoptera
frugiperda (Lepidoptera: Noctuidae). Biological Con-
trol 4: 285-289.


Florida Entomologist 86(3)







Armyworm Symposium 2002: Molina-Ochoa et al.


SIMMONS, A. M., AND C. E. ROGERS. 1996. Ectoparasitic
acugutturid nematodes of adult Lepidoptera. J.
Nematol. 28: 1-7.
SINHA, A. K. 1982. Integrated pest control programme
in intermediate savannahs of Guyana. Caribbean
Plant Protec. Newsl. 1: 2.
SIVASUBRAMANIAM, W., S. D. WRITTEN, AND J. KLIMA-
SZEWSKI. 1997. Species composition, abundance, and
activity of predatory arthropods in carrot fields,
Canterburry, New Zealand. New Zealand J. Zool. 24:
205-212.
SOTERES, K. M., R. C. BERBERET, AND R. W. MCNEW.
1984. Parasitic insects associated with lepidopterous
herbivores on alfalfa in Oklahoma. Environ. Ento-
mol. 13: 787-793.
STARY, P., AND K. S. PIKE. 1999. Uses of beneficial insect
diversity in agroecosystem management. Chapter 4,
pp. 49-67 In Biodiversity in Agroecosystems. W. W.
Collins & C. O. Qualset [eds.]. CRC Press, Boca Ra-
ton, Florida, USA.
TERAN, B. J. 1974. Lista preliminary de dipteros parasiti-
cos de otros insects en Venezuela. Rev. Fac. Agron.
Alcance (Maracay) 23: 64-65.
TERAN, B. J. 1977. Diptros parasiticos de otros insects
en Venezuela. Rev. Fac. Agron. (Maracay) 9: 134-135.
TERAN, B. J. 1980. Lista preliminary de Hymenoptera
parasitos de otros insects en Venezuela. Rev. Fac.
Agron. 11: 283-389.
TINGLE, F. C., E. R. MITCHELL, AND J. R. MCLAUGHLIN.
1994. Lepidopterous pest of cotton and their parasi-
toids in a double-cropping environment. Florida En-
tomol. 77: 334-341.
VALENCIA, L., AND R. VALDIVIA. 1973. Noctuideos del
Valle de Ica, sus plants hospederas y enemigos nat-
urales. Rev. Peru. Entomol. 16: 94-101.
VALICENTE, F. H. 1989. Levantamento dos inimigos
naturals de Spodoptera frugiperda (J.E. Smith,
1797) (Lepidoptera: Noctuidae) em diferentes re-
gioes de Minas Gerais. An. Soc. Entomol. Brasil 18:
119-130.
VALICENTE, F. H., E M. R. BARRETO. 1999. Levanta-
mento dos inimigos naturals da lagarta do cartucho
do milho, Spodoptera frugiperda (J.E. Smith) (Lepi-
doptera: Noctuidae), na regiao de Cascavel, PR. An.
Soc. Entomol. Brasil 28: 333-337.
VAN DINE, D. L. 1913. The insects affecting sugar cane
in Porto Rico. J. Econ. Entomol. 6: 251-257.
VAN DITHER, J. B. M. 1960. Insect pests of cultivated
plants in Surinam. Bull. Landouwproefstation. Suri-
name 76. 131 pp.
VARGAS, M. L., Y G. G. SANCHEZ. 1983. Control natural
de algunas plagas de arroz en las variedades IR-22 y
CICA-6. Rev. Colombiana Entomol. 9: 50-54.
VERA, M. L., L. VALVERDE, S. B. POPICH, Y Z. D. AJMAT-
DE TOLEDO. 1995. Evaluaci6n preliminary de los ene-
migos naturales de Spodoptera frugiperda (J.E.
Smith) (Lepidoptera: Noctuidae) en Tucuman, Ar-
gentina. Acta Entomol. Chilena 19: 135-141.
VICKERY, R. A. 1929. Studies on the fall armyworm in
the Gulf Coast District of Texas. USDA Tech. Bull.
138, 63 pp.
VIRLA, E. G., M. V. COLOMO, C. BERTA, Y L. VALVERDE.
1999. El complejo de los parasitoides del "gusano
cogollero" del maiz, Spodoptera frugiperda, en la


Republica Argentina (Insecta: Lepidoptera).
Neotr6pica 45: 3-12.
WAAGE, J. K. 1991. Biodiversity as a resource for biolog-
ical control. Chapter XIII, pp. 149-163. In D. L.
Hawksworth [ed.]. The Biodiversity of Microorgan-
isms and Invertebrates: Its Role in Sustainable Ag-
riculture. CAB International, Wallingford, UK.
WADDILL, V. I. 1977. Shadow sampling: A fast, painless
method for collecting fall armyworm egg masses.
Florida Entomol. 60: 215-216.
WADDILL, V. H., AND W. H. WHITCOMB. 1982. Release of
Telenomus remus (Hym. Scelionidae) against
Spodoptera frugiperda (Lep.:Noctuidae) in Florida,
USA. Entomophaga 27: 159-162.
WALL, R., AND R. C. BERBERET. 1974. The life cycle of
Euplectrus platyhypenae a gregarious external para-
sitoid of peanut foliage feeders in Oklahoma. Envi-
ron. Entomol.3: 744-746.
WALL, R., AND R. C. BERBERET. 1975. Parasitoids asso-
ciated with lepidopterous pests on peanuts; Okla-
homa fauna. Environ.Entomol. 4: 877-882.
WALTON, W. R. 1913. Efficiency of a tachinid parasite on
the last instar of Laphygma. Proc. Ent. Soc. Wash.
15:128-131.
WHEELER, G. S., T. R. ASHLEY, AND K. L. ANDREWS.
1989. Larval parasitoids and pathogens of the fall
armyworm in Honduran maize. Entomophaga 34:
331-340.
WILSON, A. G. L. 1983. Abundance and mortality of
overwintering Heliothis spp. J. Aust. Entomol. Soc.
22: 191-199.
WILSON, C. E. 1923. Insect pests of cotton in St. Croix
and a means of combating them. Virgin Islands Ag-
ric. Exp. Stn. Bull. 3: 1-20.
WILSON, F., AND C. B. HUFFAKER. 1976. The philosophy,
scope, and importance of biological control, pp. 3-15.
In C. B. Huffaker and P. S. Messinger [eds.] Theory
and Practice of Biological Control. Academic Press,
New York, USA.
WOJCIK, B., W. H. WHITCOMB, AND D. H. HABECK. 1976.
Host range testing of Telenomus remus (Hymenoptera:
Scelionidae). Florida Entomol. 59: 195-198.
WONG-AREVALO, B. 1982. Un nuevo sistema de lib-
eraci6n del parasito Trichogramma spp en el Soco-
nusco Chiapas, M6xico, pp. 34-46 En X Reuni6n
Nacional de Control Biol6gico. Secretaria de Agricul-
tura y Recursos Hidraulicos, Abril 1982, Durango,
Durango, M6xico.
YASEEN, M. 1979. Introduction of exotic parasites for
control of Spodoptera and Heliothis in Trinidad.
Symp. on maize and peanut. Paramaribo, Suriname,
Nov. 13-18, 1978. Proc. Caribbean Food Crops Soc.
15: 136-141.
YASEEN, M., F. D. BENNETT, AND R. M. BARROW. 1981.
Introduction of exotic parasites for control of
Spodoptera frugiperda in Trinidad, the eastern Car-
ibbean and Latin America, pp. 161-171. In C. W. D.
Brathwaite and G. V. Pollard [eds.] Urgent plant pest
and disease problems in the Caribbean. IICA Misc.
Publ. 378.
ZUCCHI, R. A., J. R. P. PARRA, AND S. SILVEIRA NETO.
1991. Trichogramma species associated with some
lepidopterous pests in Brazil. Coll. De-L'INRA 56:
131-134.







Florida Entomologist 86(3)


September 2003


EVALUATION OF REMOTE SENSING TO IDENTIFY VARIABILITY IN COTTON
PLANT GROWTH AND CORRELATION WITH LARVAL DENSITIES OF BEET
ARMYWORM AND CABBAGE LOOPER (LEPIDOPTERA:NOCTUIDAE)

D. L. SUDBRINK JR.1, F. A. HARRIS', J. T. ROBBINS1, P. J. ENGLISH' AND J. L. WILLERS2
'Mississippi State University, Delta Research & Extension Center, P.O. Box 197, Stoneville, MS 38776

2USDA-ARS-CSRL-GPARU, P.O. Box 5367, Mississippi State, MS 39762

ABSTRACT

Field experiments were conducted from 2000 to 2002 in the Mississippi Delta to evaluate re-
mote sensing technologies for identifying factors in cotton growth and development related
to infestations of beet armyworm and cabbage looper. Larval defoliation of plants was mon-
itored using remote sensing techniques including aerial and hand-held sensors as well as vi-
sual measurements of damage. Percent reflectance differed for beet armyworm infested
leaves compared to uninfested leaves. In two whole field studies, more beet armyworm hits
were found in zones of less vigorous and open canopy, which corresponded to lower normal-
ized difference vegetation index (NDVI) values calculated from remotely sensed imagery.
Percent light penetration of canopy was greater for plots damaged by looper larvae than for
less damaged plots where looper larvae were controlled with insecticide, but NDVI values
were not different.

Key Words: Insect management, beet armyworm, cabbage looper, cotton defoliators, remote
sensing

RESUME

Se llevaron a cabo experiments de campo desde el ano 2000 al 2002 en el Delta del Missis-
sippi para evaluar las t6cnicas de observaci6n remota (remote sensing) para identificar los
factors en el crecimiento y desarrollo del algod6n relacionadas con las infestaciones del gu-
sano trozador de la remolacha y el gusano medidor del repollo. La defoliaci6n de plants por
las larvas fu6 monitoreada usando t6cnicas de observaci6n remota incluyendo sensors de-
reos y de mano y medidas visuales del daho. El porcentaje de la reflecci6n vari6 en las hojas
infestadas con el gusano trozador comparado con hojas no infestadas. En dos studios que
abarcaron todo el campo, se encontraron mas concentraci6n (encuentros positives) del gu-
sano trozador de la remolacha en sonas donde el dosl de las plants es abierto y vigoroso,
lo cual corresponde a valores del indice de la diferencia vegetal normalizada (NDVI en in-
gl6s), mas bajos calculados de las imagenes de observaci6n remota. El porcentaje de la pene-
traci6n de luz al dosl fue mas alto en las parcelas dahadas por larvas del medidor que en
las parcelas menos dahadas donde las larvas de medidor fueron controladas con insecticide,
pero los valores de NDVI no fueron diferentes.


Beet armyworm, Spodoptera exigua (Hubner),
is an occasional pest of cotton in the Midsouth
that can become a severe pest under some envi-
ronmental conditions (Leigh et al. 1996). Beet ar-
myworm outbreaks are typically associated with
high temperatures, drought conditions, and in-
tensive insecticide regimes that eliminate natu-
ral enemies (Stewart et al. 1996). Infestations of
beet armyworm in cotton also are associated with
canopy development and varying levels of plant
nutrients such as low levels of potassium and
high levels of zinc (Parajulee et al. 1999; Graham
& Gaylor 1997; Akey et al. 1990).
Cabbage looper, Trichoplusia ni (Hubner), is
an occasional pest of cotton that only reaches
damaging levels in late-season in Mississippi
(Jost & Pitre 2002). High plant densities and vig-


orously growing plants are typically attractive for
cabbage looper oviposition and larval densities
are usually greater under these conditions (Wil-
son et al. 1982; Greene 1984).
Remote sensing is a promising technology that
may provide early detection of localized infesta-
tions of these pests based on associated crop con-
ditions (Allen et al. 1999). Remotely sensed data
may permit reduced applications of insecticides
using variable rate technology (Dupont et al.
2000). Insect pests like tarnished plant bug, have
been found in abundance in vigorously growing
portions of cotton fields that generally have faster
fruiting rates, taller plants and/or greater canopy
closure (Willers et al. 1999). These vigorous
growth zones can be identified in remotely sensed
imagery to target site-specific insecticide applica-







Armyworm Symposium 2002: Sudbrink et al.


tions with variable rate technology (Dupont et al.
2000, Willers et al. 2000). Multi-spectral remotely
sensed imagery of cotton fields is acquired aeri-
ally and the normalized difference vegetation in-
dex (NDVI) is calculated. The NDVI is associated
with crop vigor and is a calculation of the near in-
frared (NIR) and red (R) wavelengths such that
NDVI = (NIR-R)/(NIR+R) (Willers et al. 1999). In
remotely sensed imagery, NDVI values can be
used to identify spatial variability in the cotton
canopy Insecticide savings of 20-50% in control of
tarnished plant bug, Lygus lineolaris (Palisot de
Beauvois), can be achieved through the use of pre-
scription application maps that are generated
from classed NDVI values (DuPont et al. 2000;
Sudbrink et al. 2001).
Studies to determine the utility of this technol-
ogy for managing other pests are needed. For ex-
ample, other researchers report that beet
armyworm infests stressed or open cotton canopy
zones (Stewart et al. 1996). These zones may be
treatable on a site-specific basis. More informa-
tion is needed to determine if those zones can be
identified with remotely sensed imagery. The ob-
jective of these studies was to evaluate remote
sensing technologies for identifying factors in cot-
ton growth and development related to insect pest
infestation populations including infestations of
beet armyworm and cabbage looper.

MATERIALS AND METHODS

Field experiments were conducted from 2000
to 2002 on the Delta Branch Experiment Station,
Stoneville, Mississippi, or the nearby (ca. 10 miles
distance) satellite station at Tribbett, MS, to eval-
uate potential for remotely sensed data to detect
cotton plant characteristics associated with infes-
tations of leaf feeding insects such as beet army-
worm and cabbage looper. These tests included
grid-sampled fields as well as plot studies where
varietal, insecticidal, and plant growth regulator
effects on these pests in cotton were investigated.
Statistical analyses were performed with ARM
software (Gylling Data Management, Inc., Brook-
ings, SD) and correlation analyses were per-
formed using SAS for Windows 8e (SAS Institute
1990).

Test 1 Tribbett-2000

A test was conducted at Tribbett, MS in 2000
on a 2.4 acre field (320-ft x 320-ft square) subdi-
vided into a geometrically square 8 x 8 grid. Each
grid unit was 40-ft x 40-ft square. The 64 basic
units of the grid were further subdivided diago-
nally to create 128 sub-sample units, each a 40-ft
x 40-ft x 56.6-ft right triangle.
Plots were geo-spatially mapped with a Trim-
ble (Trimble navigation, Sunnyvale, CA) Ag124
GPS unit. Plant development was monitored


weekly using the COTMAN expert system, which
includes measurements of plant height, square
shed, and nodes above white flower (NAWF)
(Cochran et al. 1998). Beet armyworm damage
was monitored by observations of hits per 80-row
ft. Data were converted to hits per 100-row ft. for
treatment decision purposes and for data analysis
and presentation. A beet armyworm hit is defined
as an area on a cotton leaf where a group of beet
armyworm larvae feed and skeletonize the lower
leaf surface, often spinning silk over the site.
Treatment decision (spray or not spray) was based
on the Mississippi State University Extension Ser-
vice Cotton Insect Control Guide (Layton, 2000).
Aerial remote sensing fly-overs were made ap-
proximately every 7-14 d. Spectral reflectance
data were acquired in the aerial fly-overs with a
Duncan MS2100, 3-Chip Progressive Scan, Digi-
tal Smart Camera. Spectro-radiometry data were
also recorded from field plots on fly-over dates
and other intervening dates with a GER 1500
spectro-radiometer (Geophysical Environmental
Research Corp., Millbrook, NY).

Test 2 Stoneville-2002

Whole field observations of beet armyworm in-
festations were made in an 8-acre cotton field at
Stoneville in late August, 2002. Imagery data (ae-
rially acquired with equipment as described for
Test 1 Tribbett 2000) were used to select paired
observation sites-one in closed canopy (higher
NDVI) cotton and one in open canopy (lower
NDVI) cotton. Paired observations were made at
fifteen locations over the field. Each observation
consisted of beet armyworm hits/100 row ft. (n =
30). Means and standard errors of beet army-
worm hits were calculated from data that were
classed into four equal-interval categories of the
NDVI values.

Test 3 Stoneville-2001

A plant growth regulator by cotton variety
trial was arranged as a factorial experiment in a
randomized-complete-block design replicated
four times. Cotton was planted on 05/21/01. Each
plot was 26.7 ft (8 rows) wide by 50 ft long. Mepi-
quat chloride treatments were applied on 07/20/
01, 07/21/01 and 08/3/01. The two factors were (1)
plant-growth-regulator treatments (PGR) (two
levels, non-treated, and treated with mepiquat
chloride {Pix, 8 oz/acre, applied 2 times}), and
(2) cotton varieties (four levels-Stoneville 474
[non-transgenic], Deltapine 5415 [non-trans-
genic], Deltapine NuCotn 33B [transgenic], and
Stoneville 4691B [transgenic]). Spectroradiome-
try readings (GER 1500 spectro-radiometer as de-
scribed for Test 1 Tribbett 2000) were taken
weekly from each plot and NDVI values were cal-
culated with these data.







Florida Entomologist 86(3)


The test was modified in late season after beet
armyworm and cabbage looper infestations be-
came established. Plots in each replicate were di-
vided (without randomization and perpendicular
to row direction) into two equal size plots. The
south end plots were untreated and north end
plots were treated with spinosad. This non-ran-
dom assignment of Factor C treatments was nec-
essary to limit potential influence of drift. The
spinosad treatment was applied on 09/19/01.
The final experimental arrangement was a fac-
torial RCB design replicated four times with
three factors, (A) PGR treatment-2 levels, (B)
cotton variety-4 levels, and (C) caterpillar insec-
ticide treatment-(2 levels, untreated and
treated with spinosad {Tracer, 0.07 lb ai/acre}).
The purpose of this experimental design was to
create plant growth differences with different va-
rieties and different PGR treatments and to cre-
ate different beet armyworm and/or cabbage
looper infestations with different insecticide
treatments, and to determine if the differences
could be detected with remotely sensed data.

RESULTS

Test 1 Tribbett-2000

A beet armyworm infestation reached treat-
ment threshold levels during August in the grid-
sampled test at Tribbett in 2000. Results from
spectroradiometry readings of individual leaves
revealed that percent reflectance patterns were
distinctive for BAW damaged leaves compared to
healthy leaves (Fig. 1). Damaged leaves had lower
near infrared values than the healthy leaves.
Beet armyworm hits were found above treatment
threshold levels in zones that had lower NDVI
values calculated from aerial remote imagery
(Fig. 2). The NDVI values in the gray zones in Fig.
2 were lower than NDVI values in the white
zones. The image was classed into four equal in-
terval NDVI classes. Average plant height was
significantly taller with each progressively higher
quartile of NDVI values. Progressively higher av-
erage numbers of BAW hits were associated with
progressively lower NDVI classes (Table 1). The
lower NDVI classes were associated with less vig-
orous plants that were shorter than those in the
higher NDVI classes.
Correlation analyses revealed that there was a
significant negative correlation between beet ar-
myworm hits and NDVI value on two dates (7 &
14 August) (Table 1).

Test 2 Stoneville-2002

In late August of 2002 a beet armyworm infes-
tation reached treatment threshold levels in the
parts of the field study site at Stoneville. On 30
August 2002, aerial image-based observations re-


80-
70
w --
soI


- healthy
BAW hits


10 t
20 I
10

300 4000 500 00 700 800 900 1000 1100
Wavelength (nm)
Fig. 1 Reflectance curves for leaf canopy with beet
armyworm damage versus healthy canopy, Tribbett,
MS, 8 August 2000.

vealed that beet armyworm hits occurred over
treatment threshold levels in the two lower NDVI
class zones sampled (which were associated with
open or nearly open canopy) (Table 2). Sub-
threshold levels of beet armyworm hits were
found in samples taken in the two higher NDVI
class zones. Similar observations have been made
in very large commercial cotton fields at Gunni-
son, MS, about 60 miles north of Stoneville. Im-
age based scouting there revealed that beet
armyworm was found only in areas of lower
NDVI, albeit in extremely low populations (<1%)
(J.L.W., unpublished).
Correlation analysis revealed a highly signifi-
cant negative relationship between beet army-
worm hits and NDVI values (Table 2).


Fig. 2. Test 1 field with low (gray) and high (white)
NDVI zones. Overlaid triangular sample units had beet
armyworm hits above treatment threshold.


September 2003







Armyworm Symposium 2002: Sudbrink et al.


TABLE 1. NDVI CLASS VALUES FROM AERIAL REMOTE SENSING, PLANT HEIGHTS, AND MEAN NUMBER OF BEET ARMY-
WORM (BAW) HITS/100 FT ROW AND CORRELATIONS BETWEEN BEET ARMYWORM HITS AND NDVI, TRIBBETT,
MS, AUGUST 2000.

Plant height (inches) BAW hits/ BAW hits/
NDVI class and range Mean SEM 100ft. row 8/7/00 100ft. row 8/14/00
8/7/00 n 7/26/00 Mean SEM Mean SEM

Class I 0.367-0.423 15 36.60 1.04 4.67 0.55 5.42 0.51
Class II 0.423-0.479 20 40.35 1.01 3.44 0.49 4.63 0.51
Class III 0.479-0.535 40 42.58 0.65 2.78 0.38 4.03 0.35
Class IV 0.535-0.591 53 43.38+ 0.71 2.19 0.21 2.97 0.23

Correlation analyses
Correlation
BAW hits vs. NDVI n Slope Intercept r P

8/7/00 128 -12.52 9.248 -0.343 <0.0001
8/14/00 128 -14.19 11.09 -0.386 <0.0001


Test 3 Stoneville-2001

Populations of beet armyworm larvae re-
mained below treatment threshold levels at the
Stoneville test in 2001. Late in the season, an in-
festation of cabbage looper approached economic
threshold levels. There were no significant differ-
ences in numbers of cabbage looper larvae among
variety or mepiquat chloride treatments and no
significant interactions.
Spinosad treatments had significantly fewer
cabbage looper larvae than untreated plots (Table
3). Larval feeding was measured by determining
percentage light penetration through the canopy
as measured by a Li-Cor@ (Li-Cor, Inc., Lincoln,
NE) light bar. Percent light penetration was sig-
nificantly lower in spinosad treated plots than in
untreated plots. This result indicates that less
feeding occurred in the spinosad treated plots.
However, there was no significant difference in
mean NDVI between spinosad treated and un-
treated plots (Table 3). Thus, despite measurable


differences in looper feeding damage in this test, a
difference in spectral reflectance was not detected.

DISCUSSION

Results from these experiments indicate that
beet armyworm infestations were associated with
lower NDVI values in remotely sensed data that
represented zones of open and/or stressed canopy.
This association may be useful in the develop-
ment of future sampling plans or site-specific
management techniques that direct insecticide
applications for beet armyworm at lower NDVI
zones in a field.
The remotely sensed spectral reflectance data
did not detect crop damage by cabbage looper lar-
vae despite measurable differences in light pene-
tration between infested (damaged) and sprayed
(less damaged) canopy. This illustrates the diffi-
culty of detection via remote sensing of insect
damage, even visually observable leaf feeding, be-
fore it is too late for corrective action. Additional


TABLE 2. NDVI CLASS VALUES FROM AERIAL REMOTE SENSING AND MEAN NUMBER OF BEET ARMYWORM (BAW) HITS/100'
ROW AND CORRELATION BETWEEN BEET ARMYWORM HITS AND NDVI, STONEVILLE, MS, 30 AUGUST, 2002.

BAW hits/ 100 ft. row
NDVI class and range n Mean SEM

Class I <-0.097>-0.024 3 5.67 0.67
Class II 0.024-0.145 7 4.71 0.42
Class III 0.145-0.266 10 2.90 0.94
Class IV 0.266-0.387 10 0.50+ 0.22

Correlation analyses

Correlation n Slope Intercept r P

BAW hits vs. NDVI 30 -16.794 6.074 -0.774 <0.0001







Florida Entomologist 86(3)


September 2003


TABLE 3. EFFECT OF SPINOSAD ON A CABBAGE LOOPER INFESTATION, PERCENT LIGHT PENETRATION IN PLANT CANOPY,
AND NDVI VALUES FROM HAND-HELD SPECTRO-RADIOMETRY, FIELD 11, STONEVILLE, MS, 2001.

Mean looper larvae/6 ft row % Light penetration NDVI value
Treatment 24-Sep-01 01-Oct-01 01-Oct-01

Control 11.7 7.85 0.7623
Spinosad 2.7 5.34 0.7788
LSD 1.9 1.42 0.0330
Prob (F) 0.0001* 0.0001* 0.488 ns

*Indicates significant difference (P = 0.05) in Factorial test.


study will be needed to determine if cabbage
looper infestation can be associated with plant
characteristics that are detectable via remote
sensing techniques.
Image-based scouting through characteriza-
tion of canopy development for beet armyworm
may be useful in site-specific management of this
cotton pest. Further research is required to eluci-
date the relationship of lower NDVI levels to beet
armyworm hits and develop it into a useful sam-
pling and site-specific management plan.

ACKNOWLEDGMENTS

We greatly appreciate the assistance of Randy Furr,
Research Associate III, and student workers, David Sul-
livan, Whit Clark, Kelly Ross, Mary Grace Dye, and
Amanda Trotter, at the Delta Research and Extension
Center, Stoneville, MS. We also thank Johnny Williams,
GPS Inc., Greenwood, MS, for assistance with remotely
sensed imagery. This research was funded in part by
grants from MSU, Remote Sensing Technologies Center,
NASA, and the USDA-Advanced Spatial Technologies
in Agriculture Project.

REFERENCES CITED

AKEY, D. H., H. M. FLINT, AND J. R. MAUNEY. 1990. In-
creased damage to cotton foliage by beet armyworm
from application of zinc chelate and ammonium sul-
phate, pp. 277-278. In Proc. Beltwide Cotton Prod.
Res. Conf., National Cotton Council of America,
Memphis, TN.
ALLEN, J. C., D. D. KOPP, C. C. BREWSTER, AND S. J.
FLEISCHER. 1999. 2011: An agricultural odyssey.
Amer. Entomol. 45: 96-104.
COCHRAN, M. J., N. P. TUGWELL, F. M. BOURLAND, D. M.
OOSTERHUIS, AND D. M. DANFORTH. 1998. COTMAN
Expert System. Version 5.0. D. M. Danforth, and P.
F. O'Leary (eds.) University of Arkansas, Agric.
Expt. Sta., Fayetteville, AR. Published Cotton Incor-
porated, Raleigh, NC, p. 198.
DUPONT, K., R. CAMPANELLA, M. R. SEAL, J. L. WILLERS,
AND K. B. HOOD. 2000. Spatially variable insecticide
applications through remote sensing, pp. 426-429. In
Proc. Beltwide Cotton Prod. Res. Conf., National
Cotton Council of America, Memphis, TN.
GRAHAM, L. C. AND M. J. GAYLOR. 1997. Effects of potas-
sium fertility on beet armyworms, pp. 1320-1324. In


Proc. Beltwide Cotton Prod. Res. Conf., National
Cotton Council of America, Memphis, TN.
GREENE, G. L. 1984. Seasonal populations of eggs and
larvae in North America. In Suppression and manage-
ment of cabbage Looper populations, P. D. Lindgren
and G. L. Greene [eds.]. USDA Tech. Bull. 1684. 150 p.
JOST, D. J., AND H. N. PITRE. 2002. Soybean looper and
cabbage looper (Lepidoptera: Noctuidae) popula-
tions in cotton and soybean cropping systems in Mis-
sissippi. J. Entomol. Sci. 37: 227-235.
LAYTON, M. B. 2000. Cotton Insect Control Guide, 2000.
Mississippi State Univ. Ext. Serv. Publ. 343. 35 pp.
LEIGH, T. F., S. H. ROACH, T. F. WATSON. 1996. Biology
and ecology of important insect and mite pests of cot-
ton, pp. 17-69. In E. G. King, J. R. Phillips, and R. J.
Coleman [eds.] Cotton Insects and Mites. The Cotton
Foundation Publisher, Memphis, TN.
PARAJULEE, M. N., J. E. SLOSSER, AND D. G. BOR-
DOVSKY. 1999. Cultural practices affecting the abun-
dance of cotton aphids and beet armyworms in
dryland cotton, pp. 1014-1016. In Proc. Beltwide
Cotton Prod. Res. Conf., National Cotton Council of
America, Memphis, TN.
SAS INSTITUTE. 1990. SAS/STAT users guide: statistics,
version 6, 4th ed. SA Institute, Cary, NC.
STEWART, S. D., M. B. LAYTON JR., AND M. R. WILLIAMS.
1996. Occurrence and control of beet armyworm out-
breaks in the cotton belt, pp. 846-848. In Proc. Belt-
wide Cotton Prod. Res. Conf., National Cotton
Council of America, Memphis, TN.
SUDBRINK, D. L., F. A. HARRIS, J. T. ROBBINS, AND P. J.
ENGLISH. 2001. Remote sensing and site-specific
management of cotton arthropods in the Mississippi
Delta, pp. 1220-1223. In Proc. Beltwide Cotton Prod.
Res. Conf., National Cotton Council of America,
Memphis, TN.
WILLERS, J. L., J. K. DUPONT, R. CAMPANELLA, M. R.
SEAL, K. B. HOOD, J. WILLIAMS, AND D. WOODARD.
2000. Employment of spatially variable insecticide
applications for tarnished plant bug control in cot-
ton, p. 1133. In Proc. Beltwide Cotton Prod. Res.
Conf., National Cotton Council of America, Mem-
phis, TN.
WILLERS, J. L., M. R. SEAL, AND R. G. LUTTRELL. 1999.
Remote sensing line intercept sampling for tar-
nished plant bugs (Heteroptera: Miridae) in Mid-
South cotton. J. Cotton Sci. 3: 160-170.
WILSON, L. T., A. P. GUTIERREZ, AND D. B. HOGG. 1982.
Within-plant distribution of cabbage looper, Trichop-
lusia ni (Hubner), on cotton: development of a sam-
pling plan for eggs. Environ. Entomol. 11: 251-254.







Armyworm Symposium 2002: Sumerford


LARVAL DEVELOPMENT OF SPODOPTERA EXIGUA (LEPIDOPTERA:
NOCTUIDAE) LARVAE ON ARTIFICIAL DIET AND COTTON LEAVES
CONTAINING A BACILLUS THURINGIENSIS TOXIN:
HERITABLE VARIATION TO TOLERATE CRYIAC

D. V. SUMERFORD
USDA-ARS, Southern Insect Management Research Unit, Stoneville, MS 38776

Present Address: USDA-ARS, Corn Insect and Crop Genetic Research Unit, Ames, IA 50011

ABSTRACT

Studies were conducted to determine if beet armyworms, Spodoptera exigua (Htibner), pos-
sess the genetic variation necessary to respond to selection for improved tolerance of the Ba-
cillus thuringiensis Berliner (Bt) toxin CrylAc. Spodoptera exigua individuals that pupated
earliest when fed the CrylAc diet (ca. the first 20% to pupate) produced offspring that de-
veloped significantly faster on the CrylAc diet than their parental-control strain. In addi-
tion, after two generations of selection, the selected population reached pupation 2 d faster
than the Parental population. The selected group also developed significantly faster on
transgenic-Bt cotton leaves (cv. NuCOTN 33B) than the Parental strain. Individuals se-
lected to more rapidly develop on media containing CrylAc developed no more rapidly on ar-
tificial diet containing Cry2Aa than the parental, control colony of S. exigua.

Key Words: Spodoptera exigua, Bacillus thuringiensis, insecticide resistance, heritability,
CrylAc

RESUME

Se llevaron a cabo studios para determinar si el gusano trozador de la remolacha,
Spodoptera exigua (Hibner), posee la variaci6n gen6tica necesaria para responder a la se-
lecci6n para la tolerancia mejorada de toxin CrylAc de Bacillus thuringiensis Berliner (Bt).
Los individuos de Spodoptera exigua que se empuparon mas temprano cuando fueron ali-
mentados de una dieta de CrylAc (ca. los primeros 20% que se empuparon) produjieron des-
cendientes que se desarollaron significativamente mas rapidos con la dieta de CrylAc que el
grupo control de la raza de su parientes. Ademas, despu6s de dos generaciones de selecci6n,
la poblaci6n seleccionada se empuparon 2 dias mas rapida que la poblaci6n pariente. El
grupo seleccionado tambien se desarroll6 significativamente mas rdpido en las hojas de al-
god6n transg6nicas-Bt (cv. NuCOTN 33B) que la raza Pariente. Los individuos seleccionados
para desarrollarse mas rdpidos en un medio contiendo CrylAc no se desarrollaron mas ra-
pidamente en la dieta artificial con Cry2Aa que en la colonia control pariente de S. exigua.


Cotton varieties containing a gene from Bacil-
lus thuringiensis (Berliner) (Bt) that expresses
the insecticidal protein CrylAc have been com-
mercially available since 1996. The CrylAc-ex-
pressing varieties of Bt cotton were developed
primarily to control the tobacco budworm, Helio-
this virescens F. and the pink bollworm, Pectino-
phora gossypiella (Saunders). The abilities of
different species of insects feeding on cotton to
tolerate Bt proteins will play an important role in
determining how Bt cotton influences the popula-
tion dynamics of these pests.
Spodoptera exigua (Hiibner), a secondary pest
of cotton, is more tolerant of CrylAc than the to-
bacco budworm (Gould & Tabashnik 1998). Al-
though S. exigua can survive initial encounters
with cotton tissue expressing CrylAc, its larval
development is delayed. Delayed larval develop-
ment not only may slow the buildup of population


size in this species, but it may have the potential
to increase its generation time. One important
step is to determine if the genetic potential exists
in S. exigua to become more tolerant of CrylAc,
i.e., develop more rapidly on tissue containing
CrylAc. The purpose of this research is to deter-
mine if there is genetic potential in this species to
more rapidly finish larval development when
feeding on food sources containing CrylAc. In ad-
dition, the performance of S. exigua on a second
Cry protein is also investigated.

MATERIALS AND METHODS

Insect Colony and Diet

The S. exigua colony used during this study is
maintained at the USDA-ARS, Southern Insect
Management Research Unit, Stoneville, MS. Lar-











vae were fed a standard artificial diet developed
for Lepidoptera (Raulston & Lingren 1972).
CrylAc diet was prepared by mixing a concen-
trated stock solution of MVPII powder into 300-
500 ml batches of artificial diet to obtain a final
concentration of 1.0 pg/ml CrylAc (approx.). A
single neonate larva was placed into a 30-ml cup
containing approx. 10 ml of CrylAc diet. This con-
centration (6.8 pg/g dry weight of diet) is compa-
rable to reported amounts of CrylAc present in
cotton tissue (6.7 pg/g dry weight of diet; Green-
plate 1999) during the latter part of the growing
season (116 d after plating). Larvae from all test
groups were also placed on non-CrylAc diet as a
control. Environmental conditions for these tests
were 27 1C, 45-60% RH, and a photoperiod of
14:10 (L:D).
For two generations, individuals that com-
pleted larval development more rapidly when
feeding on CrylAc diet were selected based on the
number of days required to reach pupation (<16 d;
Fig. 1, "Selected" colony). In addition to testing
the Selected colony on CrylAc diet, its perfor-
mance was also assessed on non-CrylAc diet. The
Parental strain was always tested on both types
of diet at the same time as the selected colony.

First and Second Generations

During the first generation, the number of
days to pupation for each tested individual from
the Parental colony was recorded. Parental-col-
ony larvae were tested on both CrylAc (N = 510)
and non-CrylAc diets (N = 60). The pupae result-
ing from individuals tested on CrylAc diet were
placed into three groups: (1) the "Selected" group
was composed of individuals that had pupated by
14-16 d on CrylAc diet; (2) an "Intermediate"
group was composed of individuals that pupated
by 17-20 d on CrylAc diet; and (3) a "Slow" group
was composed of individuals that pupated after
21 d of exposure to CrylAc diet. Adults from each


Selection for Tolerance
Generation 1: Control
Selected individuals from
CrylAc diet that pupated t16d
Generation 2: "FastG2"
Selected individualsfrom
CrylAcdiet that pupated t16d
Generation 3: "FastG,"
Increased size of population:
Used individualsfrom non-
CrylAc diet for Generation 4
Generation 4: "FastG4"

Fig. 1. Mating design for the selection
exigua individuals and colonies mo
CrylAc.


September 2003


group were mated inter se to produce their sec-
ond-generation progenies. The larval develop-
ments (days to pupation) of these offspring were
compared among the above groups and also to the
Parental colony on both CrylAc (N > 200 per
group) and non-CrylAc diets (N = 30 per group).
In addition, larvae from the two extreme groups
("Selected" and "Slow") and the Parental strain
were weighed at 7 d. The weights were used as
another measure of larval development on
CrylAc and non-CrylAc diets.
Differences between groups for the number of
days to pupation were analyzed via non-paramet-
ric statistics. Comparisons between two groups
(first generation) were made using Wilcoxon
rank-sum tests (SAS 1995). Comparisons of >2
groups (second generation) were made with
Kruskal-Wallis ANOVA (SAS 1995). When
Kruskal-Wallis results indicated significant dif-
ferences, means were separated via pairwise-
comparisons among the groups using the Wil-
coxon rank-sum tests. Bonferroni adjustments
were made in the significance levels for follow-up
tests to control the overall Type I error atP = 0.05
(n = 4 groups, therefore significance was set at P
< 0.008).

Third Generation

Third-generation individuals from the Se-
lected colony were tested on CrylAc and non-
CrylAc diets and compared with the unselected,
Parental colony. As with the previous genera-
tions, the number of days to pupation was re-
corded for each individual. Comparisons between
the Selected and Parental groups were made via
Wilcoxon rank sum tests (SAS 1995).

Fourth Generation

Most of the third-generation larvae from the
Selected group were fed non-CrylAc diet in order


to not put the colony through another bottleneck
and also to increase the size of the colony for the
fourth generation of testing on plant tissue. As a
Colonies Tested
consequence, parents of the fourth-generation
Control larvae from the Selected group underwent no se-
lection for improved tolerance of CrylAc. During
the fourth generation, larvae from the Parental
Fast, Intermediate colony and Selected colony were assayed on trans-
Slow, Control genic-Bt cotton leaves (cv. NuCOTN 33B) and
non-Bt cotton leaves (cv. DP5415). Growth of lar-
vae on both types of cotton leaves was evaluated
Fast, Control via 10-d weights.
To test for differences in larval growth be-
tween the Selected and Parental groups on leaves
of Bt and non-Bt cotton, a single leaf from the up-
Fast, Control per canopy of cotton plants was placed into a cir-
n and tests of S. cular leaf cup. Five larvae were placed into each
re tolerant of cup. To help control for environmental heteroge-
neity in the performance of the cotton, tests were


Florida Entomologist 86(3)







Armyworm Symposium 2002: Sumerford


blocked by the location where the cotton was
grown. NuCOTN 33B and DP5415 were planted
in four sets of paired plots. For each block, leaves
were collected from the same set of paired plots
and replaced every 2 d. Ten cups per insect group
per cotton variety were randomized on a tray for
each block. Two blocks were set up with the first
egg clutch from both colonies, while the remain-
ing two blocks were set up the next day with lar-
vae from the second egg clutch. As a consequence,
egg clutch and block are not independent of each
other. In our ANOVA, block effects and cups
nested within (block x cotton variety) treatments
were considered random sources of error. Insect
group was considered a fixed source of variation.
Proc Mixed was used to analyze the log-trans-
formed weights and estimated the denominator
degrees of freedom based on Satterthwaite's ap-
proximation (Littell et al. 1996).

Performance on CrylAc and Cry2Aa

The duration of larval development for larvae
from the Selected and Parental colonies was com-
pared on non-Bt, CrylAc, and Cry2Aa diets. The
duration of larval development was measured as
the number of days required to finish larval devel-
opment (days to pupation). The purpose was to
determine if the performance of colonies when
feeding on CrylAc and Cry2Aa was correlated.
CrylAc diet was prepared as described above.
Cry2Aa was supplied by B. Moar, Auburn Univer-
sity. A concentration of 10 pg per ml Cry2Aa was
chosen for investigating larval development. In
other work, the expression of Cry2Ab, a similar
protein to Cry2Aa, in BollgardII cotton was ap-
prox. 10x greater than the expression of CrylAc.
The CrylAc concentration was increased 10-fold
to determine our concentration of Cry2Aa used in
this experiment.

RESULTS

Generations 1 and 2

The development of S. exigua larvae was sig-
nificantly different on CrylAc compared with
non-CrylAc diet (Wilcoxon test P < 0.0001). Days
to pupation for the Parental colony during the
first generation ranged from 15-28 d when larvae
were reared on 1.0 pg/ml CrylAc diet. All individ-
uals had pupated by 14 d on the non-CrylAc diet.
Individuals from the Selected group produced
second-generation offspring that pupated signifi-
cantly earlier on the CrylAc diet than individuals
from the Parental, Intermediate, and Slow groups
(Fig. 2A; K-W ANOVA H = 149.94; df = 3; P <
0.0001). All groups differed significantly in their
development on the toxic diet (P < 0.0001 for all
comparisons). Individuals from the Selected group
pupated ca. 1 d earlier than the larvae from the


B.2.8
2.4
2.0.
1.6.
.0
W 1.2
M 0.8.
o
S0.4.
0.0.


FAST IMD SLOW CONTROL


Non-CrvIAc CrvlAc
Fig. 2. Second-generation tests of S. exigua groups.
(A) Days to pupation for mating groups feeding on
CrylAc diet and (B) average (SE, mg) log weight of
mating groups on non-CrylAc and CrylAc diets. Bars
with the same letter do not differ significantly as deter-
mined by Wilcoxon tests (A) and analysis of least-
squared means (B).


Parental colony. On non-CrylAc diet there were
also significant differences among groups in their
days to pupation (H = 79.93; df = 3; P < 0.0001).
Greater than 90% of the larvae from the Selected,
Intermediate, and Parental groups had pupated
by 14 d. The only group to significantly differ from
the others was the Slow group. Individuals from
the Slow group took significantly longer to pupate
(mean SE = 13.90 + 0.10, 14.00 + 0.08, 16.14
0.08, and 14.77 0.10 d, for Selected, Intermedi-
ate, Slow, and Parental, respectively).
The 7-d log weights of larvae supported the
days to pupation results during the second gener-
ation. There were significant differences among
the log weights (mg) of the second-generation col-
onies on non-CrylAc diet (F = 158.43; df = 2, 55; P
< 0.0001) and CrylAc diet (F = 155.15; df= 2,203;
P < 0.0001). On both diets the Slow group was sig-
nificantly smaller than the Parental and Selected
group (Fig. 2B). However, the Selected group grew
significantly faster on the CrylAc diet than the
Parental group, but there was no difference in
growth on the non-CrylAc diet (Fig. 2B). The ra-
tios of the average log weight on CrylAc relative
to non-CrylAc diet were 0.706, 0.607, and 0.560
for the Selected, Parental, and Slow groups, re-
spectively. In addition, the percentage differences


ST
SLOW
B CONTROL







Florida Entomologist 86(3)


for the log weight of groups on CrylAc relative to
non-CrylAc were -29.43, -39.30, and -44.02% for
the Selected, Parental, and Slow groups, respec-
tively.

Generation 3

The improved growth of the Selected group
continued into the third generation. During the
third generation, the Selected group pupated sig-
nificantly earlier than the Parental group when
larvae were fed CrylAc diet (P < 0.0001; 16.24 +
0.14 vs. 18.75 0.15 d for the Selected and Paren-
tal groups, respectively). Most individuals (>90%)
from both groups pupated by 14 d on the non-
CrylAc diet (P > 0.6).
The mean days to pupation significantly dif-
fered among generations (F = 46.65; df = 2,445; P
< 0.0001). All pair-wise comparisons of least-
squared means for generations differed from each
other (all Bonferroni-adjusted P's < 0.0001). The
percentage of the total individuals feeding on
CrylAc diet that pupated <15 d increased with
each generation of selection (2.6, 6.2, and 33.3%
for generation 1, 2, and 3, respectively). The mean
number of days to pupation also decreased after
each episode of selection (Fig. 3).

Generation 4

During the fourth generation, the Selected col-
ony was compared with the Parental colony in 10-
d tests involving leaves of NuCOTN 33B (Bt cot-
ton) and DP5415 (non-Bt cotton). Two-way
ANOVA found significant effects of cotton variety
(F = 84.90; df = 1, 77.7; P < 0.0001), BAW colony
(F = 17.06; df = 1, 647; P < 0.0001) and variety x
colony (F = 10.21; df= 1, 647; P = 0.0015) on larval
weights. Tests of least-squared means (Slice op-
tion of Ismeans statement, Proc Mixed) found no
significant and significant differences between
larval weights of Selected and Parental colonies
when they were feeding on non-Bt leaves
(DP5415 F = 0.45; df = 1, 646; P = 0.5046) and Bt
leaves (NuCOTN 33B F = 26.30; df = 1, 648; P <
0.0001), respectively. Larvae from the Selected
strain were significantly larger than individuals


from the Parental strain after 10 d of feeding on
Bt cotton leaves.
During the leaf tests, comparisons between the
Selected and Parental groups were also made on
artificial diets. There were no differences between
the two groups after 11 d of feeding on non-
CrylAc diet (P = 0.099, with slightly better
growth in Parental group). However, larvae from
the Selected group were significantly larger than
larvae from the Parental group after feeding for
11 d on CrylAc diet (average weights = 151.3 and
75.3 mg, for Selected and Parental, respectively;
F = 37.00; df = 1, 10.3; P < 0.0001).

CrylAc and Cry2Aa Tests

Larval development was influenced by larval
diet (F = 157.68; df= 1,331; P < 0.0001; Fig. 5) and
the interaction between larval diet and S. exigua
colony (F = 4.37; df = 2,331; P = 0.0134). Larvae
from the selected colony pupated significantly
earlier than larvae from the Parental colony
when exposed to CrylAc diet (F = 9.55; df = 1,331;
P = 0.0022; Fig. 5). However, there were no signif-
icant differences in the number of days to reach
pupation between the Parental and selected colo-
nies when larvae fed on non-Bt diet or Cry2Aa di-
ets (P > 0.6; Fig. 5).

DISCUSSION

Spodoptera exigua had sufficient genetic vari-
ation to respond to selection for improved toler-
ance of CrylAc. After two generations of minimal
selection, the Selected group pupated approxi-
mately 2 d earlier on CrylAc diet than the Paren-
tal strain. The better growth of the Selected group
on the leaves of Bt cotton supports that the Se-
lected colony had become more tolerant of CrylAc


SControl
DFastI


Gen. 1


Fig. 3. Mean days to pu
each generation of selection


Gen.2


Gen.3


[pation (SE) during after


Fig. 4. Mean log weight (SE, mg) of the Selected and
Parental groups when tested on Bt cotton leaves (A) and
non-Bt cotton leaves (B) during generation 4.


September 2003







Armyworm Symposium 2002: Sumerford


P = 0.6160 P = 0.0022 P = 0.6938


14 1


Non-Bt CrylAc Cry2Aa
Fig. 5. Mean days to pupation (SE) wh
elected and Parental colonies are exposed to a
ets containing the no Bt proteins, CrylAc an


relative to the parental colony. In add
rapid response to selection suggests th
little recessive gene action for develop
larvae are exposed to tissue expressing
The growth of larvae on convention
tissue was significantly more rapid
growth of selected larvae on CrylAc-e
tissue. There is a potential for assortati
of adults based on the relative growth r
vae while feeding on expressing and nor
ing cotton tissue. The degree of a
mating will be dependent on the overla]
development on the two types of tissi
population becomes more tolerant of tl
tissue, there should be greater overlap
velopment of larvae feeding on non-Ci
sue. Although there is little recessive ge
one consequence of the greater overlap
development and its reduction in positive
tive mating will be to slow the evolut
proved growth when larvae are fe
CrylAc-expressing tissue.
Unlike conventional insecticides us
trol late-season populations of S. exigua


tinuous expression of CrylAc in cotton leaves
may create unwanted selection for improved tol-
erance in field populations that are below eco-
nomically-damaging densities. However, the lack
of a significant relationship between larval devel-
oParental opment on CrylAc and Cry2Aa will delay the re-
MSelected sponse to selection for improved growth on tissue
expressing these two toxins or Cry proteins re-
lated to these proteins. Stacked varieties may
therefore play an important role in the evolution
of ecological characters that will influence popu-
lation development and generation time.

en the Se- ACKNOWLEDGMENT
artificial di-
d Cry2Aa. I thank A. Combest and M. Mullen for technical as-
sistance during this study. I also thank W. Moar for pro-
viding Cry2Aa protein.
lition, the
it there is REFERENCES CITED
ent when
CrylAc. FALCONER, D. S., AND T. F. C. MACKAY. 1996. Introduc-
nal cotton tion to quantitative genetics. 4th ed. Longman, Es-
than the sex, England.
expressing GOULD, F., AND B. TABASHNIK. 1998. Bt-cotton resis-
ve mating tance management, pp. 67-105. In M. Mellon and J.
sites oflar- Rissler [eds.] Now or never: serious new plans to
n-express- save a natural pest control. Union of Concerned Sci-
ssortative entists, Washington, DC.
ssortative GREENPLATE, J. T. 1999. Quantification of bacillus thu-
p of larval ringiensis insect control protein CrylAc over time in
te. As the Bollgard cotton fruit and terminals. J. Econ. Ento-
le CrylAc mol. 92: 1377-1383.
in the de- LITTELL, R. C., G. A. MILLIKEN, W. W. STROUP, AND R.
rylAc tis- D. WOLFINGER. 1996. SAS system for mixed models.
ne action, SAS Institute, Cary, NC.
in larval LYNCH, M., AND B. WALSH. 1998. Genetics and the anal-
e assorta- ysis of quantitative traits. Sinauer Associates, Sun-
ion of im- derland, MA.
ion of im- RAULSTON, J. R., AND P. D. LINGREN. 1972. Methods for
ending on large-scale rearing of the tobacco budworm. U.S.
Dep. Agric. Prod. Res. Rep.
ed to con- SAS INSTITUTE. 1995. SAS procedure guide for personal
, the con- computers, vers. 6th ed. SAS Institute, Cary, NC.







Florida Entomologist 86(3)


September 2003


ARTHROPODS ASSOCIATED WITH ABOVE-GROUND PORTIONS OF THE
INVASIVE TREE, MELALEUCA QUINQUENERVIA, IN SOUTH FLORIDA, USA

SHERYL L. COSTELLO, PAUL D. PRATT, MIN B. RAYAMAJHI AND TED D. CENTER
USDA-ARS, Invasive Plant Research Laboratory, 3205 College Ave., Ft. Lauderdale, FL 33314

ABSTRACT

Melaleuca quinquenervia (Cav.) S. T. Blake, the broad-leaved paperbark tree, has invaded ca.
202,000 ha in Florida, including portions of the Everglades National Park. We performed
prerelease surveys in south Florida to determine if native or accidentally introduced arthro-
pods exploit this invasive plant species and assess the potential for higher trophic levels to
interfere with the establishment and success of future biological control agents. Herein we
quantify the abundance of arthropods present on the above-ground portions of saplings and
small M. quinquenervia trees at four sites. Only eight of the 328 arthropods collected were
observed feeding on M. quinquenervia. Among the arthropods collected in the plants adven-
tive range, 19 species are agricultural or horticultural pests. The high percentage of rare
species (72.0%), presumed to be transient or merely resting on the foliage, and the paucity
of species observed feeding on the weed, suggests that future biological control agents will
face little if any competition from pre-existing plant-feeding arthropods.

Key Words: Paperbark tree, arthropod abundance, Oxyops vitiosa, weed biological control

RESUME

Melaleuca quinquenervia (Cav.) S. T Blake ha invadido ca. 202,000 ha en la Florida, inclu-
yendo unas porciones del Parque Nacional de los Everglades. Nosotros realizamos sondeos
preliminares en el sur de la Florida para determinar si los art6podos natives o accidental-
mente introducidos explotan esta especie de plant invasora y evaluar el potential de los ni-
veles tr6ficos superiores para interferir con el establecimento y 6xito de futures agents de
control biol6gico. En cuatro sitios, nosotros cuantificamos la abundancia de art6podos presen-
tes en las porciones sobre el terreno de los renuevos y pequeios arboles de M. quinquenervia.
Solamente ocho de los 328 art6podos recolectados fueron observados alimentandose en la M.
quinquenervia. Entre los art6podos colectados en las areas no nativas de la plant, 19 species
son plagas agricolas 6 de hortalizas. El alto percentage de species raras (72.0%), presumidos
de ser transeuntes o meramente descansando en el follaje, y la escasez de species observadas
alimentandose de la maleza, sujiere que los futures agents de control biol6gico enfrentaran
poca o ninguna competencia de los art6podos herbivoras ya presents en la plant.


Melaleuca quinquenervia (Cav.) S.T. Blake, the
broad-leaved paperbark tree, was introduced into
south Florida during the late 1800s (Thayer & Bo-
die 1990). Although threatened in its native range
along the east coast of Australia and a few nearby
South Pacific islands, life history characteristics
ofM. quinquenervia melaleucaa) combine with fa-
vorable ecological characteristics of Everglades
habitats to make this tree an explosive weed in
south Florida (Meskimen 1962; Myers 1983; Bal-
ciunas & Center 1991; Hofstetter 1991). Cur-
rently, melaleuca occurs on about 202,000 ha of
Florida wetlands (Bodle et al. 1994) and has his-
torically spread at a rate of about 2,850 ha/yr
(Center et al. 2000). The negative impacts of
melaleuca on native flora and public health prob-
lems have been documented (Di Stefano & Fisher
1983; Myers 1983; Molnar et al. 1991; Bodle et al.
1994). Diamond et al. (1991), for instance, deter-
mined that if unchecked, potential losses to the
Florida economy as a result of this invasive tree
could reach $169 million annually.


Melaleuca infested areas can be restored
through removal of existing trees, followed by
measures to preempt reinvasion and subsequent
recruitment. Conventional control tactics com-
bine mechanical and chemical means to eliminate
seedlings, saplings, entire stands of mature trees,
or isolated plants in sensitive areas (Stocker &
Sanders 1981; Bodle et al. 1994). However, biolog-
ical attributes of this weed necessitate repeated
mechanical and chemical treatments, which im-
pose an accumulation of negative impacts on non-
target organisms, including endangered plants.
These adverse impacts limit the frequent use of
such methods. In contrast, classical weed biologi-
cal control has been described as the most ecolog-
ically benign tactic for controlling exotic pests
(McEvoy & Coombs 1999) and has been consid-
ered a desirable addition to conventional methods
(Browder & Schroeder 1981; Bodle et al. 1994).
Development of a weed biological control pro-
gram typically proceeds in a stepwise fashion,
including: selection of a natural enemy, risk







Costello et al.: Arthropods ofMelaleuca quinquenervia


analysis, release, monitoring establishment, and
finally assessing the effectiveness and ecological
impact of the introduced biological control agent
(Harris 1975; McEvoy & Coombs 1999). An often
recommended initial phase in a classical weed bio-
logical control program includes surveys of herbi-
vores associated with the invasive weed in the new
adventivee) geographic range (Harris 1975; Olck-
ers & Hulley 1995). Such surveys are intended to
identify herbivores already exploiting the weed
and to ascertain whether niche competition could
influence agent establishment and impact (Harris
1971). Although surveys for natural enemies were
performed in Australia during 1987 to 1991 (Balci-
unas et al. 1995), surveys of arthropods associated
with melaleuca in its adventive range had never
been done. Failure to perform such surveys could
increase costs due to wasted effort associated with
selecting, screening and releasing herbivores that
may already be present, having accompanied the
invasive weed upon introduction or thereafter.
Therefore, specific objectives of this study were: 1)
assess the current abundance of arthropods asso-
ciated with melaleuca in south Florida, 2) deter-
mine if native herbivores are exploiting the
invasive plant, 3) determine if co-evolved natural
enemies from the native range inadvertently ac-
companied melaleuca into south Florida, and 4)
inventory those higher trophic levels associated
with the plant that could potentially interfere with
the establishment or impact of introduced biologi-
cal control agents.

MATERIALS AND METHODS

Arthropod surveys were performed at four loca-
tions in south Florida. Site 1 was located near
Ft. Lauderdale, Broward Co., FL (N26.05606 and
W80.25168). The site was a 0.5 ha field consisting
of 2 to 5 m tall trees occurring at a plant density of
ca. 21,560 trees/ha. In general, melaleuca trees
were growing in high organic soils typical of re-
claimed 'glades' systems. Although melaleuca was
the dominant species, other plants commonly oc-
curring in the site included Blechnum serrulatum
Rich., Ampelopsis arborea (L.) Koehne, Vitis aesti-
valis Michx., and Ludwigia peruviana (L.) H. Hara.
Site 2 was located under a power line right-of-
way near Weston, Broward Co., FL (N26.035483
and W80.43495). Prior to 1997 land managers cut
melaleuca trees near their bases, resulting in multi-
stemmed branches re-growing from the stumps.
The survey area was ca. 0.5 ha and trees were 2-5 m
tall, occurring at a density of2,517 trees/ha. The site
was swale-like with common vegetation other than
melaleuca including: Sagittaria lancifolia L., Cla-
dium jamaicensis Crantz, and Andropogon glomer-
atus (Walt.) B.S.P (Anonymous 1990).
Site 3 was located near Estero, Collier Co., FL
(N26.4255 and W81.81033) and consisted of an 8
ha area of drained wetland converted to pasture.


To suppress melaleuca growth, land managers
mowed trees at ca. 6-month intervals, resulting in
coppices 0.5-2 m in height. These coppicing
clumps formed a dense, nearly continuous canopy
of leaves with 4,406 clumps/ha. In contrast to the
previous sites, the soil type was primarily sand,
consistent with an invaded pine flatwoods habitat
type (Anonymous 1990). Other than melaleuca,
the subdominant vegetation included Ludwigia
sp., Centella asiatica (L.) Urb., Rhynchospora
globularis (Chapm.) Small, Rhynchospora eximia
(Nees) Boeck., and Rhynchospora filifolia Gray.
Site 4 consisted of a 1 ha area within histori-
cally mesic flatwoods in the Picayune Forest,
Collier Co., FL (N26.10478 and W81.63392)
(Anonymous 1990). A fire burned much of the
melaleuca dominated areas during June 1998, re-
sulting in recruitment of 129,393 trees/ha com-
posed of primarily small 1-2 m tall saplings,
interspersed with an occasional large, mature
tree. Pinus elliottii Engelm. and a parasitic (dod-
der-like) species growing on the melaleuca were
the only other common vegetation.
Surveys were conducted monthly at each site
from November 2000 through June 2001. Sites
were surveyed between 10 a.m. and 2 p.m. on
days without precipitation. To survey arthropods
associated with melaleuca canopies, we swept fo-
liage, and occasionally trunks, with a 90-cm-di-
ameter sweep net. One sample consisted of 100
sweeps in a 180 sweeping motion spaced ca. 1.0 m
apart along a randomly selected 100 m transect.
Four samples along separate transects were col-
lected each month. The contents of the net after
100 sweeps were emptied into a 3.78 liter sealable
plastic bag and frozen at -19 (+1) C until pro-
cessed. Arthropods were then separated from
plant material, sorted by morphological types,
and stored in 70% ethanol.
One limitation of our sweep sampling method
included collecting arthropods that were not
closely associated with melaleuca, but were tran-
sients, merely resting on the plant foliage or dis-
turbed from understory vegetation while
sampling. Additionally, this method was biased to-
wards those species that are poor fliers or slow to
disperse from a disturbance and, unlike previous
Australian surveys, endophages were not in-
cluded. Therefore, caution should be used when
drawing inferences from these data due to the un-
known relationships between some of these ar-
thropods and melaleuca. For this reason, a
minimum of two observers searched for direct her-
bivory on the above ground portions of melaleuca
trees for 30 min./month at each site. Arthropods
observed feeding on melaleuca are reported inde-
pendently from those collected in sweep samples.
For each species collected, species abundance
per site was calculated for the entire survey pe-
riod by first averaging the number of specimens
from the four monthly samples and then averag-







Florida Entomologist 86(3)


ing among all sample dates. Average species
abundance among all sites was determined by to-
tal specimens collected throughout the entire sur-
vey (rare = 1-5 specimens, occasional = 6-10
specimens, common = >10 specimens). Occasion-
ally, arthropods were collected by hand to facili-
tate identification. Where possible, arthropods
were identified to species. Identifications that
could not be confirmed are indicated by posss."
(possibly) before the scientific name. Some
Diptera were not sent for identification because
specialists were not available or specimens were
damaged and lacked key identifying features.
Such specimens were combined into an "unidenti-
fied spp." group and the number of morphological
types is denoted in parentheses. All morphologi-
cal types, except for immatures that could be as-
sociated with their adult forms, were included in
the total species count.
All specimens, except formicids, were submit-
ted to and deposited at the Florida State Collec-
tion of Arthropods (FSCA, Division of Plant
Industry (DPI), Gainesville, FL) for identification
and incorporated into their taxonomic database.
Most formicids were identified and retained by
L. Davis at the Fire Ant Unit, Agricultural Re-
search Service, USDA, Gainesville, FL. A few
formicids were identified by M. Deyrup at the
Archbold Biological Station, Lake Placid, FL.
Several dipteran specimens were identified at the
Systematic Entomology Laboratory, Agricultural
Research Service, USDA, Beltsville, MD.

RESULTS AND DISCUSSION

Surveys of herbivores associated with an inva-
sive plant in its adventive range are often recom-
mended as a prelude to a weed biological control
project (Harris 1975). Historically, scientists have
ignored this recommendation, possibly due to the
supposition that native herbivores are already
suppressing the weed to the greatest level pos-
sible. In contrast, native arthropods can cause
considerable damage to non-indigenous weeds
(Newman et al. 1998). The native weevil, Euhry-
chiopsis lecontei Deitz, for instance, shows promise
for control of Eurasian watermilfoil, Myriophyllum
spiacata (L.) (Newman & Beisoer 2000). In addi-
tion to natives, co-evolved herbivores and diseases
may also be accidentally introduced from the
plant's native range. The biological control agents
Megastigmus aculeatus chalcidd wasp) and Phyllo-
coptes fructiphilus (an eriophyoid mite), for exam-
ple, were collected in West Virginia during surveys
of arthropods associated with the exotic weed
Rosa multifloria (Thunb.). The eriophyoid mite,
and the virus it transmits, is considered the most
effective agent for the suppression ofR. multiflora
(Amrine 1996).
In its adventive range, however, it appears that
melaleuca has not acquired native herbivores at


sufficient densities to cause appreciable damage
to trees in south Florida. For instance, of the 18
orders, 117 families, and 328 species collected in
this study, only 54 species were classified as com-
mon and 33 species were classified as occasional
(Tables 1 and 2). Of the most commonly occurring
species, 33 (66.7%) were predators or detritivores
(Table 2), and 11 (20.4%) were herbivores (Table
1). Both adult and immature stages of H. coagu-
lata, the glassy-winged sharpshooter, were ob-
served on melaleuca, suggesting that melaleuca
may serve as an alternative host for this insect.
However, during the sampling period none of
these arthropods were directly observed feeding
on melaleuca. Furthermore, out of 409 herbivo-
rous arthropods found attacking melaleuca in
Australia, none were found on melaleuca in south
Florida indicating that no co-evolved natural ene-
mies accompanied melaleuca into south Florida
upon introduction or thereafter (Balciunas et al.
1995). The most intuitive explanation for these
findings is probably due to the fact that all known
importations of the invasive tree were in the form
of seed (F. A. Dray, pers. comm.).
In contrast, we have observed several arthro-
pod species feeding on melaleuca that were never
recovered in the sweep samples. Both early and
late instars of the polyphagous saddleback cater-
pillar, Sibile stimulea (Clem.), were observed feed-
ing on mature melaleuca leaves at Site 3. Larvae
of the caterpillar were concentrated on a single
sapling, defoliated much of the tree, and were only
present during late winter. After inspection of a
single damaged sapling (5 cm diam), larvae of the
generalist cerambycid Neoclytus cordifer (Klug)
were also collected, allowed to pupate and suc-
cessfully emerged as adults (2 males and 1 fe-
male). Two phytophagous mites, Oligonychus
coffeae (Nietner) and Brevipalpus obovatus Don-
nadieu, were observed feeding and developing
large (>100 individuals), although isolated popu-
lations. Populations of these generalist mites oc-
curred on mature leaves and were only observed
once. The Florida red scale, C'i., ...... .l,.. aoni-
dum L., the stellate scale, Vinsonis stellifera, and
an unidentified Coccus sp. often co-occurred on
mature Melaleuca leaves. Although the scale oc-
curred in surprisingly high densities (>10 per
leaf), no apparent foliar damage was visible. Two
polyphagous aphids, Aphis gossypii Glover and
Toxoptera aurantii (Boyer de Fonscolombe), were
observed feeding on stems of developing branches.
Infestations of both polyphagous aphids were
slight (<50 individuals per plant). Although these
arthropod species were observed feeding on mela-
leuca, no damage was visible. These observational
findings suggest that, unlike some invasive plants
that can be stressed by native arthropods in the
adventive range, the arthropod community cur-
rently associated with melaleuca provides little if
any suppressive effect on the exotic tree. The pau-


September 2003









TABLE 1. HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Coleoptera
Aderidae
Ganascus ventricosus LeConte
Anthicidae
Vacusus vicinus (LaFerte-Senecteere)
Anthribidae
Trignorohinus sp.
Bruchidae
Sennius fallax (Boheman)
Buprestidae
Taphrocerus puncticollis Schwarz
Cantharidae
Chauliognathus marginatus (Fabricius)
Chrysomelidae
Altica sp. A
Altica sp. B
Bassareus brunnipes (Olivier)
Chrysomela scripta Fabricius
Graphopus curtipennis Blake
Lexiphanes saponatus (Fabricius)
Ophraella notulata (Fabricius)
Paria sp.
Curculionidae
Auleutes sp.
Diaprepes abbreviatus (L.)
Listronotus cryptops (Dietz)
Pheloconus hispidus (LeConte)
Trichodirabius longulus (LeConte)
Elateridae
Drapetes rubricollis LeConte
Languriidae
Loberus sp.
Lycidae
Plateros sp.


0.03 (0.09)

0.03 (0.09)

0.03 (0.09)



0.19 (0.26)


0.44 (0.90)



0.09(0.19)



0.03 (0.09)
0.03 (0.09)


0.03 (0.09) R 5


0.03 (0.09)



0.03 (0.09)


0.03 (0.09)



0.03 (0.09)



0.06 (0.18)


R 5

R 11

R -

0 3-6


0.03 (0.09)


0.03 (0.09)


0.03 (0.09) R
R
C
R
0.03 (0.09) R
R
R
0.03 (0.09) R


0.06 (0.18) 0.06 (0.12)
0.06 (0.12)


0.06 (0.12)


12 H N

H N

H N


H N

H N

H N


11 12 H
H
H
3 H
1 H
H
H
6 H


11 11, 12
- 4, 11


0.03 (0.09) R

R

0.03 (0.09) R


3 H N


4, 12


H N


2 D/H1 N









TABLE 1. (CONTINUED) HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA,
USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Scarabaeidae
Trigonopeltastes delta (Forster)
Collembola
Sminthuridae
Sminthurus sp.
Sminthurinus sp.
Dermaptera
Forficulidae
poss.12 Doru taeniatum Dohrn
Diptera
Agromyzidae
Melangromyza sp.
Unidentified sp.
Bibionidae
Unidentified sp.
Otitidae
Chaetopsis massyla (Walker)
Euxesta juncta Coquiller
Sarcophagidae
Ravinia derelicta Walker
Sciaridae
Unidentified sp.
Syrphidae
Toxomerus boscii (Macquart)

Toxomerus politus (Say)
Tephritidae
Acinia picture (Snow)
Dioxyna picciola (Bigot)

Euaresta bella (Loew)
Trupanea actinobola (Loew)
Xanthaciura insecta (Loew)


0.09 (0.27)


0.03 (0.09)
0.03 (0.09)


0.03 (0.09)


0.09 (0.27) 0.09 (0.19)
0.03 (0.09) 0.03 (0.09)



0.03 (0.09) 0.03 (0.09)
0.03 (0.09) -


R 5


R 11
R 11


- 11


11 4, 11
12 11


0.03 (0.09)


0.03 (0.09) 0.03 (0.09) 0.38 (1.06) 0.03 (0.09)


0.03 (0.09)


0.16 (0.30)


C 4 5 5


0 5


0.13(0.13)


3,6,
11, 12
11, 12


0.09 (0.19) 0.03 (0.09)


0.06 (0.12) 0.28 (0.53)
0.69 (0.86) 0.22 (0.43)


0.22 (0.41) 0.06 (0.18)


0.03 (0.09)


0.03 (0.09)
0.50 (0.97)


0.13 (0.35)


C 5, 11 1,3, 12
C 1-5, 11, 4, 11,
12 12
R 11
0 12 -
C 1,11, 12 -


D/H N


H
H


H


H/H
H/H

H/H

H/H
H/H

12 D/H


11, 12


1 -






11









TABLE 1. (CONTINUED) HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA,
USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Therevidae
Cyclotelus picitipennis (Wiedmann)
Tipulidae
Unidentified spp. (2 morphotypes)
Hemiptera
Alydidae
Hyalymenus sp. A
Hyalymenus sp. B
Imm. sp.
Coreidae
Leptoglossus phyllopus (L.)
Issidae
Acanalonia servillei Spinola
Largidae
Largus davisi Barber
Lygaeidae
Neortholomus koreshanus (Van Duzee)
Neopamera bilobata (Say)
poss. Nysius sp.
Oedancala crassimana (Fabricius)
Oncopeltus fasciatus (Dallas)
Paromius longulus (Dallas)
Imm. spp.
Miridae
Creontiades sp.
Dagbertus semipictus (Blatchley)
Reuteroscopus ornatus (Reuter)
T .. ... .. .... pallidulus (Blanchard)
Unidentified sp. A
Unidentified sp. B
Pentatomidae
Loxa sp.
Thyanta custator (Fabricius)
Thyanta perditor (Fabricius)


0.03 (0.09)


0.06(0.18)


0.03 (0.09) 0.38 (0.35)
0.06 (0.12) 0.03 (0.09)


0.03 (0.09) 0.09 (0.13)


0.03 (0.09)




0.03 (0.09)
0.03 (0.09)

0.41 (0.60)
0.03 (0.09)
0.22 (0.41)
0.25 (0.44)





0.06 (0.12)
0.03 (0.09)
0.03 (0.09)


0.03 (0.09)

0.03 (0.09)


R 11


4 1, 2, 4-6
5, 11 12


0.03 (0.09)


R 11 1,4, 12


R 6


0.03 (0.09)


0.06 (0.12) -
0.13 (0.27) -
0.06(0.12) 0.25(0.52)

0.13(0.35) 0.03(0.09)
0.41(0.65) 0.03(0.09)

0.03(0.09) 0.03(0.09)
0.03 (0.09)
0.03 (0.09)
0.06(0.18) 0.13(0.27)


006(0.12) R
R
R
C
R
C
C


5 1, 12 -
6, 12 -
5, 6, 11 3, 6 1, 4, 12
1 -
11, 12 12 12
1, 11, 12 2, 11, 12 4

1 4
2
6
1,2 4, 11 6, 11


0.06(0.18)


(D
0

lz1-



0
o
re
o


a
co






* 5

re


H N

H

H N


2, 12 H
H
H
H
H
H
H

H
H
H
H








TABLE 1. (CONTINUED) HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA,
USA.

Abundance per site' Months collected2
Trophic3 Native/ Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Rhopalidae
Liorrhysus hydlinus (Fabricius)
Homoptera
Aphididae
Aphis spiraecola Patch
Aphis sp.
Eulachnus rileyi (Williams)
Hysteroneura setariae (Thomas)
Schizaphis sp.
Tetraneura nigriabdominalis (Sasaki)
Toxoptera aurantii (Boyer de Fonscolombe)
Cercopidae
Clastoptera xantocephala Germar
Lepyronia sp.
Cicadellidae
Balclutha sp.
Cuerna costalis (Fabricius)
Draeculacephala sp. A
Draeculacephala sp. B

poss. Empoasca sp.

Graminella nigrifrons (Forbes)
Graphocephala versuta (Say)

Gypona sp.

Homalodisca coagulata (Say)

Hortensia similis (Walker)
Oncometopia nigricans (Walker)
Paraulacizes irrorata (Fabricius)
Stragania sp.
Tropicanus costamaculatus (Van Duzee)
Imm. spp.


0.13 (0.27) 0.03 (0.09)


0.03 (0.09)


0.06(0.18) 0.03(0.09)


0.03 (0.09)


0.03 (0.09)
0.03 (0.09)
0.03 (0.09)
0.03 (0.09)
0.03 (0.09)
0.03 (0.09)


0.03 (0.09) 0.03 (0.09)


0.06 (0.12)




0.13(0.19)


0.06 (0.12) 0.13 (0.13)


1,2 3 11
- -- 11
1
5 11
-- 11
-- 11
11 2

11 4
2, 12 -


0.03 (0.09) 0.03 (0.09)
0.13(0.19)

0.59(0.80) 0.03(0.09)



0.16(0.27) 0.03(0.09)


1, 11, 12


0 1,4 1,2,
4, 12


0.19 (0.22)


0.19(0.22) 0.06(0.18) 0.09(0.18)

0.94 (0.74) 0.41(0.33) -


C 2,4, 2
11, 12
0.03(0.09) C 1-6, 1,4-6,
11, 12 11, 12


0.03 (0.09)
0.03 (0.09) -
0.03 (0.09) -
0.03(0.09) 0.09(0.19) 0.25(0.48)
0.06 (0.12) -
2.19(1.47) 1.94 (1.84) 0.72 (1.11) 0.13(0.27)


1

1
3, 12
1-6,
11, 12


1
11, 12


4
4, 6, 12

2,4,
11, 12


4, 6, 12


11, 12



12


11, 12


1-6, 3, 4,
11, 12 11, 12


11 H
H
H
12 H

H

H
H

H

11 H


01
0D









TABLE 1. (CONTINUED) HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA,
USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Cixiidae
Bothriocera sp.
Myndus crudus Van Duzee
Delphacidae
Delphacodes puella (Van Duzee)
Delphacodes sp. A
Delphacodes sp. B
Imm. sp.
Flatidae
Imm. spp.
Membracidae
Spissistilus festinus (Say)
Stictocephala lutea (Walker)

Psyllidae
Diaphorina citri Kuwayama
Hymenoptera
Agaonidae
Unidentified sp.
Anthophoridae
Exomalopsis sp.
Halictidae
Agapostemon splendens (Lepeletier)
Augochlora sp.
Lasioglossum sp.
Lepidoptera
Heliconiidae
Heliconius charitonius Tuckeri
Geometridae
Unidentified sp. A
Unidentified sp. B
Unidentified sp. C
Gracillariidae
Phyllocnistis sp.


0.19 (0.44)
0.03 (0.09) 0.06 (0.12)


- 5,6
11 4,5

- 1, 11


0.06 (0.12)


0.03 (0.09)


0.03 (0.09) 0.03 (0.09)

0.03 (0.09) 0.03 (0.09)


0.47 (0.59) 0.03 (0.09)


0.16(0.44) C 3-6,11 3


0.22 (0.53)
0.16(0.19)


0.03 (0.09)


0.03 (0.09)


H N
H N

H
12 1 H
- H
12 12 H

- 5 H


1, 12
3,4,
11, 12


R 11


0.03 (0.09)

0.06 (0.12)
0.03 (0.09)


0.03 (0.09)

0.03 (0.09)




0.03 (0.09)


R
R
0.03 (0.09) R


5, 12
5


H
H
6 H


R 12


0.03 (0.09)


R
0.03 (0.09) R
R


R 4


- H
11 H
H

H


C
0










0
5"
0
(1


H N
H N


H E


H N








TABLE 1. (CONTINUED) HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA,
USA.

Abundance per site' Months collected2
Trophic3 Native/ Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Noctuidae
Unidentified sp.
Pyralidae
Unidentified sp. A
Unidentified sp. B
Unidentified sp. C
Unidentified sp. D
Unidentified sp. E
Unidentified sp. F
Unidentified sp. G
Unidentified sp. H
Unidentified sp. I
Orthoptera
Acrididae
Leptysma marginicollis (Serville)
Schistocerca damnifica (Saussure)
Orphulella pelidna (Burmeister)
Paroxya atlantica Scudder
Imm. spp.
Gryllidae
Cyroxipha poss. columbiana Caudell
Oecanthus quadripunctatus Beutenmuller
Tetrigidae
Tettrigidea lateralis (Say)
Tettrigidea sp.
Tettigoniidae
Conocephalus sp.

Imm. sp.
Phasmatodea
Pseudophasmatidae
Anisomorpha buprestoides (Stoll)
Psocoptera
Peripsocidae
Peripsocus madescens (Walsh)


0.16 (0.44)


0.03 (0.09) R


0.03 (0.09)


0.09 (0.27)



0.06(0.18)
0.06 (0.12)

0.03 (0.09)
0.31 (0.48)

0.03 (0.09)


0.03 (0.09)


0.13(0.19)
0.03 (0.09)
0.03 (0.09)
0.03 (0.09)

0.06 (0.12)


0.03 (0.09)


0.03 (0.09) R
0.03 (0.09) R
0.03 (0.09) R
0.03 (0.09) R
0.25(0.40) 0.50(0.64) C


0.09(0.19)


0.09(0.19)


1
5, 12

5
4,5,6


0.06 (0.12) R
R

0.03 (0.09) R
R


0.53 (0.59) 0.72 (0.86) 0.03 (0.09)

0.03 (0.09) -


0.22(0.41) 0.06(0.12)


12 H


6, 11, 12
11
11
11


11 H
4 H
6 H
6 H
3,4,6 2-6 H


- 11,12 H N
3, 12 H N

- 11 H N
2,3 H N


1,2,6, 3, 4, 6,
11, 12 11, 12
1


H N


2,4 3,4 H N


1.84 (2.15) C 1,3-6 H N


1.84 (2.15) C


1, 3-6 H N




















TABLE 1. (CONTINUED) HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA,
USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status

Psocidae
Indiopsocus ceterus Mockford -- 0.03 (0.09) R 4 H N
gen. sp. -- 0.03 (0.09) R 5 H
Thysanoptera
Phlaeothripidae
Haplothrips gowdeyi (Franklin) 0.03 (0.09) R 11 H N

'Abundance per transect for each site averaged over 8 months. Each transect equals 100 sweeps with a 90-cm diameter sweep net. One sweep consists of an 180 sweeping motion.
Samples were taken from November (month 11) through June (month 6)..
'D = Detritivore (including scavengers), H=Herbivore (including pollen and nectar feeders), P = Predator, U = Undetermined.
'N = Native, E = Exotic, Blank space = Undetermined.
5An indicates that the species is a known economic pest.
'Weston, FL, Broward Co., N26.035483 and W80.43495, M. quinquenervia stand under a power line.
'University Rd. and Griffin Rd., Fort Lauderdale, FL, Broward Co., N 26.05605 and W -80.25168, vacant lot occupied byM. quinquenervia.
8Tamiami Tr. and Corkscrew Rd., Estero, FL, Collier Co., N 26.4255 and W -81.81033, Cow pasture occupied with small M. quinquenervia stumps.
'Belle Meade, FL, Collier Co., N 26.10478 and W -81.63392, M. quinquenervia stand in the Picayune Forest.
"Average abundance among all sites includes total number of specimens collected. R = Rare, 1-5 specimens; O = Occasional, 6-10 specimens; C = Common, >10 specimens.
/ indicates a difference in trophic level of larvae stage and adult stage. Trophic level data include larval then adult trophic level.
Sposs. indicates a possible identification that could not be confirmed.









TABLE 2. NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Acari
Anystidae
Anystis agilis Banks
Microtrombidiidae
Trichotrombidum muscarum (Riley)
Araneae
Anyphaenidae
Hibana sp.

Lupettiana mordax (0. P.-Cambridge)
Wulfila alba (Hentz)
Araneidae
Acacesia hamata (Hentz)

Cyclosa turbinata (Walckenaer)
Eriophora ravilla (C.L. Koch)
Gasteracantha cancriformis (L.)
Kaira alba (Hentz)
Mangora spiculata (Hentz)
Mangora imm. sp.
Neoscona arabesca (Walckenaer)
Neoscona imm. sp.

Wagneriana tauricornis (0. P.-Cambridge)
Imm. spp.

Clubionidae
Clubiona sp.
Corinnidae
Castianeira sp.
Trachelas volutus Gertsch
Linyphiidae
Eperigone bryantae Ivie & Barrows
Meioneta sp.
Unidentified sp. A


0.03 (0.09) 0.06 (0.12)


R 2 2,3


0.09 (0.27)


0.13 (0.27) 0.03 (0.09) 0.03 (0.09) 0.34 (0.52)


0.06 (0.12)
0.03 (0.09)


0.06 (0.12) R
R


0.47 (0.34) 0.44 (0.44) 0.06 (0.12) 0.06 (0.12)


0.03 (0.09) 0.13 (0.27)
0.16 (0.23) 0.03 (0.09)
0.03 (0.09)
0.03 (0.09)


0.19(0.18)
0.97 (1.26)


0.03 (0.09)


0.06 (0.18)
0.03 (0.09)


0.47 (0.77) 0.09 (0.19)


0.66(0.63) 0.06(0.12)


0.19 (0.44)

0.03 (0.09) -
0.06 (0.12) -


0.03 (0.09) -
0.13 (0.27) -


0.06 (0.18) 0.03 (0.09)
0.03 (0.09) 0.13 (0.13)


1,4 6 12 1,3
11, 12
- 3, 11 2, 5


C 2-6,
11, 12
0 4
0 5, 6, 11
R
R
R
R
0 1, 3-6
C 1-4,
11, 12
R -
C 1, 2, 4, 6,
11, 12


0.16 (0.35) C


1, 3, 6,
11, 12
5,6
3
1
3



1-3, 11


3,6


5, 11


R 5
R 5, 12


6,11 2,11


P N

Pa/P1 N


P N

P N
P N

P N


- P
11 P
-
-
12 P
6 P
-
-


4 12
4 1,6,
11, 12


P N
P N


11, 12 P N


P N
P N


0.03 (0.09)
0.03 (0.09)









TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Lycosidae
Pardosa littoralis Banks
Pardosa imm. sp.
Pirata sp.
Mimetidae
Mimetus sp.
Miturgidae
Cheiracanthium inclusum (Hentz)

Oxyopidae
Peucetia viridans (Hentz)

Pisauridae
Pisaurina mira (Walckenaer)
Pisaurina undulata (Keyserling)
Pisaurina imm. spp.

Imm. sp.
Salticidae
Eris flava (Peckham & Peckham)
Eris imm. sp.
Habronattus sp.
Hentzia palmarum (Hentz)

Lyssomanes viridis (Walckenaer)
Pelegrina galathea (Walckenaer)

Pelegrina sp.
Phidippus clarus Keyserling
Phidippus regius C.L. Koch
Phidippus sp.
Thiodina peurpera (Hentz)
Zygoballus sexpunctatus (Hentz)
Zygoballus sp.


0.03 (0.09)
0.56 (1.12)

0.03 (0.09)


0.06(0.18) -
0.50 (1.41) -
0.06(0.12) 0.84(2.39) 0.69(1.56)

0.06(0.18) 0.50(0.53)


3.00 (2.10) 4.38 (4.89) 0.31 (0.35) 0.50 (0.44)


4.28 (2.87) 0.28 (0.39) 0.34 (0.42) 0.13 (0.13)


0.03 (0.09) -
0.41 (0.77) 0.16 (0.27)


0.03 (0.09)

0.06 (0.12)


0.09(0.19) 0.03(0.09) -

0.13(0.23) 0.16(0.30) 0.50 (0.92) 0.28 (0.62)
0.03(0.09) 0.03(0.09) -
0.03(0.09) 0.06(0.12) -
2.25(0.90) 1.41(0.57) 0.31(0.42) 1.72(1.86)

0.13(0.19) -
0.38(0.48) 0.16(0.19) 0.44 (0.37) -


0.03 (0.09)
0.59 (1.68)


0.06 (0.18)
0.03 (0.09)


0.03 (0.09)


- 12
- 12
3, 12 12


C 3 -

C 1-6, 12 1-6, 12


-
-
1, 11,12 P


12 1-4, 11

1, 2, 4, 3, 5, 6,
6, 12 11, 12


C 1-6, 12 1, 3, 2-4, 6, 1, 2,
5, 12 12 6, 12


R -
R 6
C 1,2,
5, 11
R 1,12


0.03 (0.09) R
R
R
0.16 (0.35) 0
C
R
0.03 (0.09) R


6

1,4


1, 4, 11


2, 12 2, 12
11 12
2
1-6, 1-6,
11, 12 11, 12
- 3-5
1-4, 12, 1,
11, 12 2, 6



5
6


P N


P N
P N
P N

P N


1, 11, 12 1, 12

12, 4
1, 3, 6, 1, 2, 4-6,
11, 12 11, 12


1-4,
11, 12

4
12


4


P N
P N

5 P N
P N
P N
12,5 P N
P N
P N
2 P N








TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Tetragnathidae
Glenognatna foxi (McCook)
Glenognatha sp.
Leucauge argyra (Walckenaer)
Tetragnatha sp.

Theridiidae
Anelosimus studiosus (Hentz)

Chrysso pulcherrima (Mello-Leitao)
Dipoena nigra (Emerton)
Latrodectus geometricus C.L. Koch
Theridion flavonotatum Becker

Theridion glaucescens Becker
Theridion imm. sp.
Thymoites sp.
Thomisidae
Misumenoides formosipes (Walckenaer)
Misumenops bellulus (Banks)

Misumenops oblongus (Keyserling)
Misumenops imm. spp.

Tmarus sp.

Coleoptera
Coccinellidae
Brachiacantha decora Casey
Coelophora inaequalis (Fabricius)
Cycloneda sanguine (L.)

Exochomus marginipennis (LeConte)
Psyllobora parvinotata Casey
Scymnus securus J. Chapin
Scymnus sp.


0.03 (0.09)
1.19(0.61) 0.28(0.28)


2.09 (1.41) -

0.03 (0.09) -


0.75 (0.69) -

0.38 (0.57) -
0.13(0.27) 0.03(0.09)
0.06(0.18) -

0.03 (0.09)
0.81(0.53) 1.16(1.14)


0.50(0.40) 0.13(0.19)

0.06 (0.12) -


0.13 (0.35)


0.16 (0.35)


R
0.06 (0.18) R
R
0.03 (0.09) C


0.09 (0.27) -

0.03(0.09) 0.13(0.35)
0.03 (0.09)
0.03 (0.09) -
0.06 (0.12)


0.03 (0.09)


0.03 (0.09) C
-
R


0.84 (0.67) 0.47 (0.39)

0.06 (0.18) 0.03 (0.09)
1.09 (1.13) 0.19 (0.44)

0.03 (0.09) 1.63 (0.86)


0.06 (0.18) -
0.03 (0.09) -
0.03(0.09) 0.06(0.12) 0.09(0.13)


0.03 (0.09) 0.03 (0.09)

0.03 (0.09) 0.09 (0.13)


0.06 (0.12)

0.03 (0.09)


12
1-6, 12,
11, 12 2-4, 6

1-6,
11, 12
11


11, 1-3, -
3, 5,6
11, 3, 4 -
2,5 5
12


- 11
1-5, 1-6,
11, 12 11, 12

1-4, 6, 2, 11, 12
11, 12
11, 6



6
12
12 1,3


3 12

4 2, 3, 5


12 P
11 P
- P
11,12 11 P


12 P

11 11 P
5 P
4 P
11.2 P


2


1-4, 6,
11, 12
4
1-4,
11, 12
1


11, 1,
2, 4-6
1
5, 11

1-6,
11, 12


P N
P N

P N
P N

P N


1, 3,
6, 11
11, 12


N








TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Lampyridae
Pyropyga minute LeConte
Scirtidae
Cyphon sp.
Diplopoda
Spirobolidae
Chicobolus spinigerus (Wood)
Dictyoptera
Blattellidae
Chorisoneura parishi Rehn

Imm. sp.
Mantidae
Gonatista grisea (Fabricius)
Stagmomantis sp.
Thesprotia graminis (Scudder)
Imm. spp.
Diptera
Ceratopogonidae
Atrichopogon sp.
Unidentified sp. A
Unidentified sp. B
Unidentified sp. C
Chironomidae
Unidentified spp. (11 morphotypes)

Chloropidae
Apallates dissidens Tucker
Apallates neocoxendrix Sabrosky
Coniscinella sp.
Chlorops sp.
Ectecephala unicolor (Loew)
Hippelates plebejus Loew
Liohippelates pusio Loew


0.03 (0.09)

0.25 (0.53)


0.06(0.18)


R 3

0 4,6


R 12


0.19 (0.22) 0.28 (0.60)


C 1, 5, 3, 5, 12
11, 12


0.03 (0.09)


0.03 (0.09)


0.06 (0.12)

0.03 (0.09)
0.09 (0.19)


0.06 (0.12) 0.63 (1.13) -
0.03 (0.09) 0.25 (0.48) 0.38(1.06)
0.03 (0.09) -


R
R
R
0.16 (0.30) O


0.03 (0.09) C
C
0.50 (1.41) C
0.03 (0.09) R


0.53 (0.36) 1.28 (2.07) 0.84 (1.08) 0.03 (0.09)


0.13(0.19)
0.03(0.09) 0.06(0.18)
0.19(0.29) 0.03(0.09)

0.03 (0.09)


0.34 (0.97) 0.06 (0.18)
0.19 (0.53)
0.03(0.09) 0.06(0.08)
0.22 (0.41) -


11, 12

12
5, 12


2,11 1,11, 12
2 1, 11
11


C 1, 2, 4-6, 1, 2, 1, 3, 4,
11, 12 11, 12 11, 12


11, 12
5 1
1, 2, 11 2

1


0.06 (0.18)
0.03 (0.09) 0.59 (1.29)


P N

D N


D N


D E


-
-
-
3,4 P


12 D/Pa
- D/Pa
11 D/Pa
1 D/Pa

11 U/U


U/U
U/U
U/U
U/U
U/U
D/Pa
U/U









TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Unidentified sp. A
Unidentified sp. B
Unidentified sp. C
Unidentified sp. D
Unidentified sp. E
Clusiidae
Unidentified sp.
Culicidae
Unidentified spp. (2 morphotypes)
Dolichopodidae
Chrysotus sp.
Chrysotus picticornis Loew
Condylostylus sp.
Condylostylus tonsus Aldrich
Empididae
Euhybus poss. stramaticus Melander
Euhybus sp.
Syneches simplex Walker
Unidentified sp.
Ephydridae
Unidentified sp. A
Unidentified sp. B
Unidentified sp. C
Unidentified sp. D
Unidentified sp. E
Lauxaniidae
Unidentified sp. A
Unidentified sp. B
Milichiidae
Desmometopa sp.
Muscidae
Stomoxys calcitrans (L.)
Unidentified spp. (8 morphotypes)


0.03 (0.09)
0.06 (0.18)
0.03 (0.09)


0.03 (0.09)
0.06 (0.18)
0.03 (0.09)

0.03 (0.09)


0.13 (0.23) R


1.34 (1.56)
0.13 (0.35)
0.03 (0.09)


1.69 (1.99) 0.72 (1.47) 0.06 (0.12)
0.22 (0.62) -
0.03 (0.09) -
0.38(0.46) 0.16(0.44) -


0.13(0.27) 0.09(0.19)
0.03 (0.09) -
0.06 (0.12)
0.03 (0.09)

0.09 (0.19)
0.03 (0.09)

0.03 (0.09) -
0.13 (0.27) -

0.03(0.09) 0.16(0.30)


0.06 (0.18)


0.13 (0.27) 0.28 (0.28)


6, 11 D/Pa


1-6, 11
5
5


2, 5
6


0.03 (0.09)

0.03(0.09) 0.06(0.18)



0.09(0.27) 0.03(0.09)
0.06 (0.18)





0.03 (0.09)



0.03 (0.09)
0.59(0.86) 0.19(0.44)


1-6, 11 4, 11, 12
-- 11
-- 11
2, 4, 12 4


1, 12
4

11, 12
11

3
3, 5

2 1,2


2 -

6
4,5 1,2,4, 3, 4, 6,
5, 12 11, 12


4 P/P
P/P
12 12 P/P
- P/P


- U/U
12 U/U
- U/U
- U/U
- U/U

- D/U
- D/U

- D/U


11, 12


Pa/D
U/U








TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Otitidae
Herina narytia (Walker)


Sciomyzidae
Dictya sp.
Sepsidae
Palaeosepsis insularis (Williston)
Stratiomyidae
Nemotelus glaber Loew
Tabanidae
Chrysops sp.
Hemiptera
Pentatomidae
Euthyrhynchus floridanus (Pointer)
Podisus mucronatus Uhler

Podisus sagitta (Fabricius)
Sphyrocoris obliquus (Germar)
Stiretrus anchorago (Fabricius)
Imm. spp.
Phymatidae
Unidentified sp.
Reduviidae
Zelus longipes (L.)

Homoptera
Derbidae
Cedusa sp.

Hymenoptera
Aphidiidae
Lysiphebus testaceipes (Cresson)
Braconidae
Apantales sp.
Bassus sp.


2.00 (1.57)


0.03 (0.09)



0.13 (0.23)

0.06(0.18)


0.03 (0.09)
0.19(0.18)


0.03 (0.09) 0.03 (0.09)


0.03 (0.09)



0.03 (0.09) 3.06 (5.29) 0.13(0.35)


0.06 (0.18) 0.03 (0.09)
0.03(0.09) 0.13(0.27)
0.03 (0.09)


0.03 (0.09)

0.78 (0.73)



0.13(0.19) 1.22(1.28)


0.03 (0.09)
0.09 (0.27)


C 1-6,
11, 12

R 4

R -

R 4,5

R 4


R 4
C 2, 3, 5,
11, 12
R 11
0 12
R 11
R -


11 11


2 1, 3, 4,
11, 12
5
11, 12


R 11

0.03 (0.09) C 1-3, 5, 6,
11, 12


C 3, 11, 12 2-6,
11, 12


0.03 (0.09)


0.06(0.18)


0.03 (0.09)
0.03 (0.09) 0.03 (0.09)


R 1 3
R 11 11


D/U

U/U

D/Pa


P N
P N

P N
P N

P

P

P N



D


Pa N
Pa N








TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status


Brachymyrmex obscurior Forel
Camponotus floridanus (Buckley)

Camponotus planatus Roger


0.06(0.12) R 3, 11 Pa N
0.06 (0.12) R 3, 12 Pa
0.03(0.09) R 12 Pa
0.03(0.09) R 12 Pa
0.03(0.09) R 3 Pa
0.03(0.09) R 6 Pa


0.03 (0.09)


Cotesia sp.
Unidentified sp. A
Unidentified sp. B
Unidentified sp. C
Unidentified sp. D
Unidentified sp. E
Bethylidae
Unidentified sp. A
Chrysididae
Chrysis sp.
Encyrtidae
poss.Aenasioidea sp.
Anagyrus sp. A
Anagyrus sp. B
poss. Cercobelus sp.
poss. Syrphophagus sp.
Eucoilidae
Eucoila sp.
Unidentified sp.
Eulophidae
Cirrospilus poss. pictus (Nees)
Unidentified sp.
Eumenidae
Zethus slossonae (Zethusculus)
Eupelmidae
Anastatus sp.
Eupelmus sp.
Eurytomidae
Eurytoma sp.
Formicidae


0 2 5, 12


0.03 (0.09)


0.03 (0.09)


0.03 (0.09)


1


1
11 11


R 12


Pa N

Pa
Pa
Pa
Pa
Pa

Pa
Pa


P N


0.03 (0.09)
0.03 (0.09)

0.03 (0.09)


0.09 (0.13) 0.03 (0.09)
0.44 (0.55) -

2.34(1.08) 1.44 (1.23) 0.06 (0.12)


R 1,5,11
0.06(0.12) C 3,5,6,
11, 12
C 1-6,
11, 12


5


1-6,
11, 12


- P
1,5 P


P E


0.03(0.09) 0.13(0.23)

0.03 (0.09) -
0.03(0.09) 0.03(0.09)
0.03 (0.09) -
0.03 (0.09) -


0.03 (0.09)


0.03 (0.09)
0.03 (0.09) 0.03 (0.09)








TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.10 A B C D level Exotic status


Camponotus sexguttatus (Fabricius)
Cardiocondyla wroughtoni obscurior
Wheeler
Crematogaster ashmeadi Mayr
Crematogaster atkinsoni Wheeler
Cyphomyrmex rimosus (Spinola)
Dolichoderus pustulatus Mayr
Dorymyrmex bureni (Trager)
Gnamptogenys aculeaticoxae (Santschi)
Odontomachus ruginodus Smith
Paratrechina guatemalensis (Forel)

Paratrechina longicornis (Latreille)

Platythyrea punctata (Smith)
Pseudomyrmex ejectus (Smith)
Pseudomyrmex gracilis (Fabricius)

Pseudomyrmex pallidus (Smith)

Solenopsis invicta Buren

Technomyrmex albipes (Smith)
Ichneumonidae
Diadegma sp.
Megaspilinidae
Dendrocerus sp.
Mutillidae
Dasymutilla sp.
Mymaridae
Anaphes sp.
Pteromalidae
Pachyneuron sp.
poss. Pteromalus sp.
Unidentified sp. A
Unidentified sp. B


0.03 (0.09)
0.03 (0.09)


0.06 (0.12)

0.19(0.44)
0.03 (0.09)
0.03 (0.09) -
0.03 (0.09) -
0.50(0.94) 0.06(0.18)

2.66(4.86) 3.19(5.37)

0.03 (0.09) -
0.03(0.09) 0.03(0.09)
0.06(0.12) 0.03(0.09)


R 5
R 11


0.03 (0.09)

0.03 (0.09)


- R
- R
- R
-
- R


1, 11

6, 12


R 5
R 11
0.19 (0.53) C 3,4,
11, 12
0.13(0.27) C 1-4,
11, 12
R 12
R 5
0.19(0.22) 0 4,6


0.47 (0.49) 0.34 (0.68) -

0.41 (0.58) 0.50 (0.33) 0.50 (0.61) 2.00 (4.55)


C 1, 2, 4,
6, 11, 12
C 1, 2, 4,
11, 12


2


1

3, 5, 6,
11, 12

6
4

1, 3,
4, 11
1-4, 6,
11, 12


0.03 (0.09)

0.03 (0.09)


0.03 (0.09)

0.03 (0.09)



0.03 (0.09)

0.03 (0.09)
0.03 (0.09)


R 2

R 11


0.06 (0.12)


0.03 (0.09)


P E
P E


-
-
-
-
-
-
-
12 P

2, 11 P


3, 4,
6, 11


2, 11


P N


2, 4, 6,
11, 12
12


Pa

Pa N








TABLE 2. (CONTINUED) NON-HERBIVOROUS ARTHROPODS COLLECTED IN THE ABOVE-GROUND PORTIONS OF THE INVASIVE TREE, MELALEUCA QUINQUENERVIA IN SOUTH
FLORIDA, USA.

Abundance per site' Months collected2
Trophic3 Native/4 Pest5
Species A6 B7 C8 D9 Ave.O1 A B C D level Exotic status

Scelionidae
Macroteleia sp. 0.03 (0.09) R 12 Pa
Telenomus sp. 0.06(0.12) 0.03(0.09) R 1, 11 12 Pa
Trissolcus sp. 0.25 (0.33) 1, 2, Pa
5, 12
Sphecidae
Tachytes sp. 0.03 (0.09) R 11 Pa
Torymidae
Torymus sp. 0.09 (0.19) R 3, 12 Pa
Vespidae
Mischocyttarus mexicanus (Saussure) 0.09 (0.13) R 2,4,5 P N
Polistes dorsalis (Fabricius) 0.09 (0.19) R 11, 12 P N
Polistes major Beauvios 0.03 (0.09) R 1 P N
Neuroptera
Chrysopidae
Ceraeochrysa sp. 0.03 (0.09) 0.13 (0.23) O 1 11, 12 P
Chrysopa quadripunctatus Burmeister -- 0.03 (0.09) R 11 P
Chrysoperia sp. 0.06(0.18) 0.06(0.12) -R 1 1, 5 P
Odonata
Coenagrionidae
Ischnura hastata (Say) 0.03 (0.09) 0.03 (0.09) 0.06 (0.12) R 3 11 11, 12 P N
Nehalennia pallidula Calvert 0.03 (0.09) -R 5 P N
Libellulidae
Erythrodiplax minusula (Rambus) -- 0.03 (0.09) R 11 P N
Thysanoptera
Phlaeothripidae
Nesothrips latiuentris (Karny) 0.16 (0.44) 12 D N

Abundance per transect for each site averaged over 8 months. Each transect equals 100 sweeps with a 90cm diameter sweep net. One sweep consists of an 180 sweeping motion.
'Samples were taken from November (month 11) through June (month 6).
'D = Detritivore (including scavengers), Pa = Parasitiod (including secretion feeders and blood suckers), P = Predator, U = Undetermined.
'N = Native, E = Exotic, Blank space = Undetermined.
'An indicates that the species is a known economic pest.
'Weston, FL, Broward Co., N 26.035483 and W -80.43495, M. quinquenervia stand under a power line.
'University Rd. and Griffin Rd., Fort Lauderdale, FL, Broward Co., N 26.05605 and W -80.25168, vacant lot occupied byM. quinquenervia.
8Tamiami Tr. and Corkscrew Rd., Estero, FL, Collier Co., N 26.4255 W -81.81033, Cow pasture occupied with smallM. quinquenervia stumps.
'Belle Meade, FL, Collier Co., N 26.10478 W -81.63392, M. quinquenervia stand in the Picayune Forest.
"Average abundance among all sites includes total number of specimens collected. R = Rare, 1-5 specimens; O = Occasional, 6-10 specimens; C = Common, >10 specimens.
/ indicates a difference in trophic level of larvae stage and adult stage. Trophic level data include larval then adult trophic level.







Costello et al.: Arthropods ofMelaleuca quinquenervia


city of herbivores indicates that direct competi-
tion between natives and introduced biological
control agents will be minimal.
Habitats dominated by invasive plants are of-
ten assumed to be sterile environments with few
wildlife species utilizing the ecosystem (Bodle et
al. 1994). However, Mazzotti et al. (1981) deter-
mined that differences exist among invasive
plants in their ability to support native fauna, in-
dicating that habitats invaded and dominated by
non-indigenous plants are not necessarily biologi-
cal deserts. After eight months of surveying ar-
thropods in melaleuca dominated ecosystems,
rarefaction curves of both herbivorous and non-
herbivorous arthropods suggests that continued
surveying efforts would result in the collection of
additional species (Figs. 1 and 2; Magurran 1988).
The variety of arthropods, both collected (Tables 1
and 2) and predicted (Figs. 1 and 2), reported
herein indicates that melaleuca dominated habi-
tats do support an arthropod community. How-
ever, this does not necessarily imply that
melaleuca is a superior habitat for such fauna as
indicated by the paucity of basal trophic levels
(i.e., herbivores). Without the ability to compare
arthropod diversity in surrounding native habi-
tats, the probability that many species are tran-


140




120 -


sient, and considering the dearth of commonly
collected arthropods, caution should be exercised
when making conclusions concerning the func-
tional well being of melaleuca invaded ecosystems.
The role of invasive species as facilitators of
other invasive species has received little attention
in the literature (Simberloff & Von Holle 1999).
One example of this interaction may include the
ability of nonindigenous plants to modify the habi-
tat in a way that favors exotics over natives. In this
study, 20 exotic species were collected in the mela-
leuca habitat (Tables 1 and 2). Among the exotic
species, Solenopsis invicta Buren, the red imported
fire ant, was common (Table 2) and is included as
one of the most ecologically destructive invasive
species in the southeastern U.S. These ant colonies
not only cause human disturbance, but also are
known to cause 70% mortality of freshwater turtle
hatchlings (Pseudemys nelsoni Carr), can nega-
tively impact the endangered Schaus swallowtail
(Papilio aristodemus porceanus), and can dramati-
cally change arthropod communities (Porter et al.
1988; Allen et al. 2001; Forys et al. 2001). Although
native to Florida, the glassy-winged sharpshooter
is an invasive species in California, where it vec-
tors Xylella fastidiosa Wells et al., the causal agent
of Pierce's disease in vineyards. Because the


Nov Dec Jan Feb Mar Apr May June


Month


Fig. 1. Rarefaction curve for cumulative herbivorous species collected from M. quinquenervia (Nov.-June).







Florida Entomologist 86(3)


220 -


200 -


180 -


160 -


140


120


100


O U I I I I I I I I
Nov Dec Jan Feb Mar Apr May June

Month
Fig. 2. Rarefaction curve for cumulative non-herbivorous species collected from M. quinquenervia (Nov.-June).


glassy-winged sharpshooter is commonly associ-
ated with melaleuca in Florida, it may be predicted
that the plant also provides a refuge for the inva-
sive sharpshooter in California. In this manner,
melaleuca may serve as a reservoir for these and
other invasive species in Florida and beyond.
In addition to the facilitation of ecological im-
pacts by exotic species, invasive weeds may also
harbor agricultural pests. For instance, 1/3 of the
phytophagous insects associated with Salsola
kali L. var. tenuifolia Tausch (Russian thistle)
and 1/2 of the insect species on Carduus pycno-
cephalus L. (Italian thistle) proved to be pests of
agricultural importance (Goeden & Ricker 1968).
In our study, 18 arthropods collected from mela-
leuca canopies are major or minor economic pests
of agricultural crops. Three species,Aphis spirae-
cola Patch (Aphididae), T aurantii (Aphididae),
and S. invicta (Formicidae), were commonly asso-
ciated with melaleuca. Both aphid species are cos-
mopolitan, phytophagous pests of Citrus spp. and
many other plants. An infestation of these aphid
species can result in abortion of Citrus flower
buds and both aphids produce honeydew, thus fa-
voring the development of sooty molds.
Native predators, parasitoids, and pathogens
have interfered with half of the published case


histories involving insect introductions for weed
control (Goeden & Louda 1976). Parasitoids and
pathogens, for instance, caused 24% larval mor-
tality of the introduced moth, Samea multiplica-
lis Guenee (Semple & Forno 1987). Herein, we
collected several generalist predators that may
potentially impact current and future biological
control agents, including Euthyrhynchus florida-
nus (Pointer) (Pentatomidae), Podisus mucronatus
Uhler (Pentatomidae), Podisus saggita (Fabri-
cius) (Pentatomidae), Stiretrus anchorage (Fabri-
cius) (Pentatomidae), and Zelus longipes (L.)
(Reduviidae), as well as various ant and spider
species. Predation on populations of the recently
released biological control agent Boreioglycaspis
melaleucae Moore melaleucaa psyllid, Psyllidae)
by various pentatomid and coccinellid species has
been observed in the field and may be negatively
affected by generalist predators. During host
specificity testing and under mass rearing condi-
tions prior to its introduction, B. melaleucae was
attacked by multiple arachnid species. However,
the level of predation observed in the field or un-
der laboratory conditions does not appear to im-
pact colonies in a significant way (P. D. Pratt,
pers. obs.; S.A. Wineriter pers. comm.). Studies
on other psyllids, Psylla pyricola Forester (pear


1


September 2003







Costello et al.: Arthropods ofMelaleuca quinquenervia


psyllid) and Diaphorina citri Kuwayama (Asian
citrus psyllid), have shown that their populations
are reduced by generalists predators such as:
CL., ....., sp. (Chrysopidae), Anthocoris sp.
(Anthicoridae), and Olla v-nigrum (Mulsant)
(Coccinellidae) (Watson & Wilde 1963; Michaud
2001). Furthermore, Watson & Wilde (1963) and
Santas (1987) demonstrated a reduction in psyl-
lid populations by generalist predators. Never-
theless, in each study psyllid populations were
suppressed by generalist predators at different
levels, suggesting that predicting the acquisition
and impact of these predators on introduced bio-
logical control agents is tenuous.
During our study, we also collected several par-
asitic hymenopteran species associated with mela-
leuca in south Florida (Table 2). Hymenopteran
species in Australia parasitized ca. 40% of galls
formed by the potential biological control agent
Fergusonia spp. (gall fly) (Davies et al. 2001).
Davies et al. (2001) suggested the impact by Fer-
gusonia spp. as biological control agents of mela-
leuca will likely be reduced due to parasitism from
local hymenopteran species in Florida. However,
predicting which parasitoids may exploit this or
other proposed biological control agents is diffi-
cult. Initial steps may include a taxonomic com-
parison among the co-evolved parasitoids in the
agent's native and adventive ranges. For instance,
Cirrospilus sp. (Eulophidae), Eupelmus sp. (Eu-
pelmidae) and Eurytoma sp. (Eurytomidae) were
collected in Australia associated with Fergusonia
spp. and during our survey we also collected para-
sitoids belonging to these genera in south Florida
(Goolsby et al. 2001). Unfortunately, species deter-
mination was not possible for those reported
herein. Due to the diversity of both genera, geo-
graphic separation over evolutionary time, and
lack of Fergusoninidae in the New World, it is un-
likely that the species occurring in Australia and
Florida are the same. Other genera found during
our survey do not correspond to those genera
known to parasitize current and candidate biolog-
ical control agents in their native range, including
Fergusonina spp., B. melaleucae, Poliopaschia li-
thochlora (Lower) (tube-dwelling moth) and Lo-
phyrotoma zonalis (Rohwer) melaleucaa sawfly)
(Jensen 1957; Riek 1962; Burrows & Balciunas
1997; Davies et al. 2001; J. A. Goolsby, USDA/ARS,
Aust. Bio. Cont. Lab., pers. comm.). Predictions
based solely on this survey may grossly underesti-
mate parasitoid acquisition as additional species
may be recruited to the system after introduction
of the biological control agent. In the future a more
accurate assessment may be obtained by survey-
ing melaleuca for endophagic arthropods and com-
paring regional species databases or arthropod
collections in the native and adventive ranges.
Further studies may also include an evaluation of
predator and parasitoid arthropod recruitment af-
ter the release of new biological control agents.


ACKNOWLEDGMENTS
The authors thank the following specialists and as-
sociates for identification of most arthropod species:
J. Bramblia, G. B. Edwards, G. A. Evans, S. E. Halbert,
J. B. Heppner, J. M. Kingsolver, B. F. Mauffray, C. C. Por-
ter, L. A. Stange, G. J. Steck, M. C. Thomas, W. C. Wel-
bourn, J. R. Wiley of the FSCA, Division of Plant
Industry, Gainesville, FL. We also thank formicid spe-
cialists L. R. Davis of the Fire Ant Unit, Agricultural Re-
search Service, USDA, Gainesville, FL, and M. A.
Deyrup of the Archbold Biological Station, Lake Placid,
FL.; diptera specialists F. C. Thompson and N. E. Wood-
ley of the Systematic Entomology Laboratory, Agricul-
tural Research Service, USDA, Beltsville, Maryland;
Collembola specialist R. J. Snider at Michigan State
University; and Psocopotera specialist E. L. Mockford at
Illinois State University.

REFERENCES CITED
ALLEN, C. R., E. A. FORYS, K. G. RICE, AND D. P. WOJCIK.
2001. Effects of fire ants (Hymenoptera: Formicidae)
on hatching turtles and prevalence of fire ants on sea
turtle nesting beaches in Florida. Florida Entomol.
84(2): 250-253.
AMRINE, J. W. 1996. Phyllocoptes fructiphilus and bio-
logical control of multiflora rose, pp. 741-749. In
E. E. Lindquist, M. W. Sabelis and J. Bruin [eds.],
Eriophyoid Mites: Their Biology, Natural Enemies
and Control. Elsevier Science, 790 pp.
ANONYMOUS. 1990. Guide to the natural communities of
Florida. Florida Natural Areas Inventory. Florida De-
partment of Natural Resources, Tallahassee, 118 pp.
BALCIUNAS, J. K., AND T. D. CENTER. 1991. Biological
control of Melaleuca quinquenervia: prospects and
conflicts, pp. 1-22. In T. D. Center, R. F. Doren, R. L.
Hofstetter, R. L. Myers and L. D. Whiteaker [eds]
Proc. Symp. Exotic Pest Plants, Miami, Florida, 2-4
Nov. 1988, U.S. Dept. Interior, National Park Ser-
vice, Denver, CO, 387 pp.
BALCIUNAS, J. K., M. F. BURROWS, AND M. F. PURCELL.
1995. Australian insects for the biological control of
the paperbark tree, Melaleuca quinquenervia, a seri-
ous pest of Florida, USA, wetlands, pp. 247-267. In
E. S. Delfosse, and R. R. Scott [eds.], Proc. Symp. on
Biological Control of Weeds. 2-7 February 1992, Lin-
coln University, Canterbury, New Zealand, 735 pp.
BODLE, J. J., A. P. FERRITER, AND D. D. THAYER. 1994.
The biology, distribution, and ecological conse-
quences of Melaleuca quinquenervia in the Ever-
glades, pp. 341-355. In S. M. Davis and J. C. Ogden
[eds.], Everglades: The Ecosystem and Its Restora-
tion. St. Lucie Press, Delray Beach, 826 pp.
BROWDER, J. A., AND P. B. SCHROEDER 1981. Melaleuca
seed dispersal and perspectives on control, pp. 17-21.
In R. K. Geiger [ed.], Proceedings of a Melaleuca
Symposium. Florida Dept. of Agriculture and Con-
sumer Services, Division of Forestry, Tallahassee,
Florida, 140 pp.
BURROWS, D. W., AND J. K. BALCIUNAS. 1997. Biology,
distribution and host-range of the sawfly, Lophyro-
toma zonalis (Hym., Pergidae), a potential biological
control agent for the paperbark tree, Melaleuca
quinquenervia. Entomophaga 42(3): 299-313.
CENTER, T. D., T. K. VAN, M. RAYACHHETRY, G. R.
BUCKINGHAM, F. A. DRAY, S. WINERITER, M. F. PUR-
CELL, AND P. D. PRATT. 2000. Field colonization of











the melaleuca snout beetle (Oxyops vitiosa) in south
Florida. Bio. Cont. 19: 112-123.
DAVIES, K. A., J. MAKINSON, AND M. F. PURCELL. 2001.
Observations on the development and parasitoids of
Fergusonina/Fergusobia galls on Melaleuca quin-
queneruia (Myrtaceae) in Australia. Trans. Royal
Soc. S. Aust. 125: 45-50.
DI STEFANO, J. F., AND R. F. FISHER. 1983. Invasion po-
tential ofMelaleuca quinqueneruia in southern Flor-
ida, U.S.A. Forest Ecol. Manage. 7: 133-141.
DIAMOND, C., D. DAVIS, AND D. C. SCHMITZ. 1991. Eco-
nomic impact statement: The addition of Melaleuca
quinqueneruia to the Florida Prohibited Aquatic
Plant List. In T. D. Center, R. F. Doren, R. L. Hofstet-
ter, R. L. Myers and L. D. Whiteaker [eds] Proc.
Symp. Exotic Pest Plants, Miami, Florida, 2-4 Nov.
1988, U.S. Dept. Interior, National Park Service,
Denver, CO, 387 pp.
FORYS, E. A., A. QUISTORFF, AND C. R. ALLEN. 2001. Po-
tential fire ant (Hymenoptera: Formicidae) impact
on the endangered Schaus Swallowtail (Lepidop-
tera: Papilionidae). Florida Entomol. 84(2): 254-258.
GOEDEN, R. D., AND S. M. LOUDA. 1976. Biotic interfer-
ence with insects imported for weed control. Ann.
Rev. Entomol. 21: 325-342.
GOEDEN, R. D., AND D. W. RICKER. 1968. The phytopha-
gous insect fauna of Russian thistle (Salsola kali
var. tenuifolia) in southern California. Ann. Ento-
mol. Soc. Amer. 61: 67-72.
GOOLSBY, J. A., C. J. BURWELL, J. MAKINSON, AND F.
DRIVER 2001. Investigation of the Biology of Hy-
menoptera Associated with Fergusonina sp.
(Diptera: Fergusoninidae), a Gall Fly of Melaleuca
quinqueneruia, Integrating Molecular Techniques.
Journal of Hymenoptera Research 2(2): 172-2000.
HARRIS, P. 1971. Biological control of weeds. Env. Let-
ters 2: 75-88.
HARRIS, P. 1975. General approach to biocontrol of
weeds in Canada. Phytoprotection 56(3): 135-141.
HOFSTETTER, R. L. 1991. The current status of Mela-
leuca quinqueneruia in southern Florida, pp. 159-
176. In T. D. Center, R. F. Doren, R. L. Hofstetter,
R. L. Myers and L. D. Whiteaker [eds] Proc. Symp.
Exotic Pest Plants, Miami, Florida, 2-4 Nov. 1988,
U.S. Dept. Interior, National Park Service, Denver,
CO, 387 pp.
JENSEN, D. D. 1957. Parasites of the Psyllidae. Hilgar-
dia 27: 71-99.
MAGURRAN, A. E. 1988. Ecological diversity and its
measurement. Princeton University Press, NJ.
MAZZOTTI, F. J., W. OSTRENKO, AND A. T. SMITH. 1981.
Effects of the exotic plants Melaleuca quinquenervia
and Casuarina equisetifolia on small mammal popu-
lations in the eastern Florida Everglades. Florida
Sci. 44(2): 65-71.
McEVOY, P. B., AND E. M. COOMBS. 1999. Why things
bite back: Unintended consequences of biological
control of weeds, pp. 1-31. In P. A. Follett and J. J.
Duan [eds.], Nontarget Effects of Biological Control.
Kluwer Academic Publishers, Boston, MA, 316 pp.


September 2003


MESKIMEN, G.F. 1962. A silvical study of the melaleuca
tree in south Florida. University of Florida, Thesis,
Gainesville, pp. 177.
MICHAUD, J. P. 2001. Numerical Response ofOlla V-nig-
rum (Coleoptera: Coleoptera) to Infestations of
Asian Citrus Psyllid (Hemiptera: Psyllidae) in Flor-
ida. Florida Entomol. 84(4): 608-612.
MOLNAR, G., R. H. HOFSTETTER, R. F. DOREN, L. D.
WHITEAKER, AND M. T. BRENNAN. 1991. Manage-
ment of Melaleuca quinqueneruia within East Ever-
glades wetlands, pp. 237-253. In T. D. Center, R. F.
Doren, R. L. Hofstetter, R. L. Myers and L. D.
Whiteaker [eds.] Proc. Symp. Exotic Pest Plants, Mi-
ami, Florida, 2-4 Nov. 1988, U.S. Dept. Interior, Na-
tional Park Service, Denver, 387 pp.
MYERS, R. L. 1983. Site susceptibility to invasion by the
exotic tree Melaleuca quinqueneruia in southern
Florida. J. Appl. Ecol. 20: 645-658.
NEWMAN, R. M., D. C. THOMPSON, AND D. B. RICHMAN.
1998. Conservation strategies for the biological con-
trol of weeds, pp. 371-396. In P. Barbosa [ed.], Con-
servation Biological Control. Academic Press, San
Diego, 396 pp.
NEWMAN, R. M., AND D. D. BIESOER 2000. A decline of
Eurasian watermilfoil in Minnesota associated with
the milfoil weevil, Euhrychiopsis lecontei. J. Aqua.
Plant Manag. 38: 105-111.
Olckers, T, and P. E. Hulley. 1995. Importance of prein-
troduction surveys in the biological control of
Solanum weeds in South Africa. Agric. Ecosys. Envi-
ron. 52: 179-185.
PORTER, S. D., B. VAN EIMEREN, AND L. E. GILBERT.
1998. Invasion of red imported fire ants (Hymenop-
tera: Formicidae): microgeography of competitive re-
placement. Ann. Entomol. Soc. Am. 81: 913-918.
RIEK, E. F. 1962. The Australian species of Psyllaepha-
gus (Hymenoptera: Encyrtidae), Parasites of Psyl-
lids (Homoptera). Aust. J. Zool. 10: 682-757.
SANTAS, L. A. 1987. The predators' complex of pear-
feeding psyllids in unsprayed wild pear trees in
Greece. Entomophaga 32: 291-297.
SEMPLE, J. L. AND I. W. FORNO. 1987. Native parasi-
toids and pathogens attacking Samea multiplicalis
Guenee (Lepidoptera: Pyralidae) in Queensland. J.
Aust. Ent. Soc. 26: 365-366.
SIMBERLOFF, D., AND B. VON HOLLE. 1999. Positive in-
teractions of nonindigenous species: invasional melt-
down. Bio. Invasions 1: 21-32.
STOCKER, R. K., AND D. R. SANDERS. 1981. Chemical
Control of Melaleuca quinqueneruia. In R. K. Geiger
[ed.], Proceedings of Melaleuca Symposium, Sept.
23-24, 1980. pp. 129-134. Florida Department of Ag-
riculture and Consumer Services, Division of For-
estry, 140 pp.
THAYER, D. D., AND M. BODLE. 1990. Melaleuca quin-
quenervia: The paperbark tree in Florida or an
Aussie out of control. Aquatics 12(3): 4-9.
WATSON, T. K., AND W. H. A. WILDE. 1963. Laboratory
and field observations on two predators of the Pear
Psylla in British Columbia. Can. Ent. 95: 435-438.


Florida Entomologist 86(3)







Landolt et al.: Trapping Yellowjackets with Heptyl Butyrate


TRAPPING YELLOWJACKETS (HYMENOPTERA: VESPIDAE) WITH HEPTYL
BUTYRATE EMITTED FROM CONTROLLED-RELEASE DISPENSERS

P. J. LANDOLT1, H. C. REED2 AND D. J. ELLIS2
'USDA-ARS Yakima Agricultural Research Laboratory, 5230 Konnowac Pass Road, Wapato, WA 98951

2Dept. Biology, Oral Roberts University, Tulsa, OK 74171


ABSTRACT

Numbers of workers of Vespula pensylvanica (Saussure) (western yellowjacket) and V atro-
pilosa (Sladen) trapped with heptyl butyrate in Washington increased with greater release
of the attractant from vial dispensers, up to an estimated 2.3 milligrams heptyl butyrate per
hour. Vespula germanica (F.) (German yellowjacket) workers were also captured in signifi-
cant numbers, and numbers of workers captured increased with increased release of heptyl
butyrate, up to an estimated 1.4 milligrams per hour. Numbers of workers of Vespula squa-
mosa (Drury) trapped with heptyl butyrate in Oklahoma increased with increased release of
heptyl butyrate from dispensers, up to an estimated 3.3 milligrams per hour. Vial dispensers,
with holes of 6, 12, 22, and 33 mm diameter in the vial lid, lost 0.42, 1.4, 2.3, and 3.3 milli-
grams of heptyl butyrate per hour in the laboratory and these rates changed little over a pe-
riod of 4 weeks, indicating close to a zero order rate of release pattern. Rates of loss of 2 ml
heptyl butyrate applied to a cotton ball decreased with exposure time, from an initial rate of
6.0 milligrams per hour to near zero at 16 days post-treatment. Captures of wasps in traps
sold commercially might be improved with the use of a controlled-release dispenser.

Key Words: Wasp, yellowjacket, trap, attractant, lure, heptyl butyrate

RESUME

El numero de trabajadores de Vespula pensylvanica (Saussure), la avispa "chaqueta amari-
lla" occidental, y los de V atropilosa (Sladen) atrapados con el butirato heptilico en el estado
de Washington aument6 con una mayor liberaci6n del atrayente en frascos dispensadores,
hasta un estimado de 2.3 miligramos de butirato heptilico por hora, los trabajadores de Vesp-
ula germanica (F.), la avispa "chaqueta amarilla" aleman, tambien fueron capturados en nu-
meros significantes, y el numero de trabajadores capturados aument6 con la mayor
liberaci6n de butirato heptilico, hasta un estimado de 1.4 miligramos por hora. El numero de
los trabajadores de Vespula squamosa (Drury) atrapados con butirato heptilico en Oklahoma
aument6 con el aumento de la liberaci6n de butirato heptilico de los dispensadores, hasta un
estimado 3.3 miligramos por hora. Los frascos dispensadores, con hoyos de 6, 12, 22 y 33 mm
de diametro en el tap6n del frasco, perdieron 0.42, 1.37, 2.3, y 3.3 miligramos de butirato
heptilico por hora en el laboratorio y estas proporciones cambiaron poco sobre un period de
4 semanas, indicando una proporci6n del orden cerca del cero del patron de liberaci6n. La
proporci6n de perdida de 2 ml de butirato heptilico aplicado a una pelota de algod6n baj6 con
el tiempo de exposici6n, desde una proporci6n inicial de 6.0 miligramos por hora hasta cerca
de cero a los 16 dias despu6s del tratamiento. La recolecci6n de avispas en trampas para
venta commercial puede ser mejorada con el uso de un dispensador de liberaci6n controlada.


Heptyl butyrate is attractive to some species of
yellowjackets, and is sold commercially as a lure
for traps. Workers of the western yellowjacket,
Vespula pensylvanica (Saussure), and both
queens and workers of Vespula atropilosa
(Sladen), are captured in large numbers in traps
baited with heptyl butyrate (Davis et al. 1969;
MacDonald et al. 1973; Landolt 1998). Workers of
Vespula acadica (Sladen), Vespula consobrina
(Saussure) (blackjacket), Vespula germanica (F.)
(German yellowjacket), Vespula maculifrons
(Buysson) (eastern yellowjacket), Vespula squa-
mosa (Drury), Vespula sulphurea (Saussure),
Vespula vidua (Saussure), Vespula vulgaris (L.),


and other wasps have also been captured in traps
baited with heptyl butyrate (Grothaus et al. 1973;
Landolt 1998; MacDonald et al. 1974; Reed &
Landolt 2003; Reierson & Wagner 1975, 1978).
These trapping studies indicate a broad-based
but variable response of a taxonomic range of yel-
lowjacket species to heptyl butyrate. However, it
has been concluded that the chemical is effective
primarily in attracting V pensylvanica and V at-
ropilosa and is weak or ineffective as a lure for
other species of social wasps (Akre et al. 1981).
The use of heptyl butyrate as a trap lure might
be improved by providing a controlled release sys-
tem for optimizing the amount of the chemical







Florida Entomologist 86(3)


evaporated from traps. In studies evaluating
wasp responses to this chemical in traps, heptyl
butyrate has been applied directly to the bottom
of the trap (0.25 ml) (Davis et al. 1968, 1969), to
cotton swabs (0.4 ml) (Howell et al. 1974), to cot-
ton balls (one to 5 ml) (Davis et al. 1973; Mac-
Donald et al. 1973, 1974; Sharp & James 1979),
rubber stoppers (Chang 1988), and in polyethyl-
ene caps (Landolt 1998). Currently, measured
amounts of heptyl butyrate are applied to cotton
balls as a means of baiting traps sold commer-
cially for yellowjackets (Rescue Trap@, Sterling
International, Inc., Veradale, WA). Although a
dose of one to several ml on cotton has been effec-
tive as a lure for the yellowjackets V pensylvanica
and V atropilosa (Davis et al. 1973, MacDonald et
al. 1973, 1974), there are no published determina-
tions of the relationship between dosage or re-
lease rate of heptyl butyrate and captures of
wasps in traps. Thus, we do not know how to pro-
vide optimum amounts of heptyl butyrate for
maximum attraction or capture of wasps in traps.
We report here the trapping of several species
of Vespula with heptyl butyrate emitted at varied
release rates, and the first documentation of
worker V germanica and V squamosa response to
heptyl butyrate in controlled experiments. This
study was designed to test for trapping of V pen-
sylvanica in Washington and V squamosa in
Oklahoma as primary pest species. Our long term
objective is to improve the efficacy of trapping
systems for these pest wasps.

MATERIALS AND METHODS

The Dome trap (yellowjacket trap of Gem-
pler's, Bellevue, WI), was used in all field experi-
ments. This plastic trap is opaque yellow below
and clear above, with a bottom entrance through
which wasps enter the trap. Traps contained
about 200 ml of a drowning solution made of
0.01% unscented dishwashing detergent (Palmol-
ive Concentrated Dishwashing Liquid, Colgate-
Palmolive, New York, NY) in water. Polypropy-
lene vials (Nalge Nunc International, Rochester,
NY, 15 ml 2118-9050 and 30 ml 2118-0001) were
used as a means of dispensing heptyl butyrate
(Aldrich Chemical Co., Milwaukee, WI). Ten ml of
heptyl butyrate were pipetted onto three balls (ca
2.5 cm diam) of cotton wedged into the bottom
half of the vial. Chemical release was through a
hole in the lid of the vial. Vials were suspended in
the top center of the inside of traps by wire.
A range of release rates of heptyl butyrate was
provided by varying the diameter of the hole in
the lid of vial dispensers. Comparisons of cap-
tures of wasps in traps with varying heptyl bu-
tyrate release rates were conducted in both the
Yakima, Washington and Tulsa, Oklahoma areas,
in order to obtain data for V pensylvanica and
V squamosa respectively. The same chemical,


lures, traps, and methods were used for the exper-
iments in both Yakima and Tulsa. A randomized
complete block design was used for each test, with
each of a series of vial hole sizes (providing a dif-
ferent release rate) represented within each of
the five replicate blocks. Holes of different sizes
were made using sets of drill bits. The first test
compared heptyl butyrate in 15 ml vials with
holes of 0, 1.0, 1.5, 3.0, 6.0, and 12 mm diameter.
The vial with no hole (0 mm hole diameter) was
presumed to not release heptyl butyrate. This test
was followed by another experiment comparing
heptyl butyrate in 30 ml vials with holes of 1.5,
3.0, 6.0, 12, 22, and 33 mm diameter.
In Yakima, the first test was conducted from 19
to 31 July 2000. Traps were hung on fences, shrub-
bery, and low branches of trees at a golf course, at
a height of 2 m, with 30 m spacing between traps.
The second test in Yakima was conducted from 4
to 21 August 2000. Traps were positioned as de-
scribed above, with some blocks at a golf course
and others at a commercial tree nursery.
In Tulsa, Oklahoma, the first test was con-
ducted from 12 September to 6 October 2000,
with traps on vegetation on and adjacent to the
campus of Oral Roberts University. The second
test in Tulsa was conducted from 6 October to 3
November 2000, with traps again on vegetation
on and near the campus of Oral Roberts Univer-
sity. At both locations, traps were checked twice
per week, and the drowning solution of traps was
changed weekly. In Tulsa, heptyl butyrate dis-
pensers were replaced after two weeks. In both lo-
cations, trap positions were randomized initially
and each time that traps were checked.
Release rates of heptyl butyrate from vials
were determined by weighing vials that con-
tained heptyl butyrate at intervals to determine
weight loss with time. Weight loss was deter-
mined for four replicates of 30 ml vials loaded
with 10 ml of heptyl butyrate on cotton balls, and
with vial hole diameters of 0, 6, 12, 22, and 33
mm. These vial hole sizes were selected because
we sought to determine the range of release rates
of heptyl butyrate that are effective in attracting
wasps to traps. Weight loss was also determined
for cotton balls placed on aluminum foil weighing
dishes, with an initial application of two ml of
heptyl butyrate to the cotton. Vials and cotton
balls were weighed every two or three days over
the 3-week period of time following the initial
loading of the vials with heptyl butyrate. Vials
and cotton balls on weighing dishes were held in
a fume hood inside the USDA-ARS Yakima Agri-
cultural Research Laboratory, for the 21 day du-
ration of the experiment. Temperature inside of
the fume hood was 22.5 + 1C, with the tempera-
ture variance likely due to changes in building air
handling rates and heating/cooling which could
alter both the air flow rate through the fume hood
and the ambient temperature.


September 2003







Landolt et al.: Trapping Yellowjackets with Heptyl Butyrate


For each of the four trapping experiments, trap
catch data were subjected to a regression analysis
(DataMost 1995) to determine if there was a sig-
nificant positive relationship between numbers of
wasps captured and vial hole diameter. Also, data
from the first experiment for V germanica in
Yakima, Washington, and for V squamosa in
Tulsa, Oklahoma were subjected to an ANOVA,
with treatment means compared using a paired t
test (DataMost 1995), to determine if workers of
these two species were captured in significant
numbers in traps baited with heptyl butyrate.
Weight loss data for heptyl butyrate from cotton
balls and from vials were subjected to a regres-
sion analysis (DataMost 1995) to determine if
there was a significant negative relationship be-
tween dispenser age and rate of loss.

RESULTS

In the first field test conducted in Yakima, there
was a significant positive regression of numbers of
wasps trapped in relation to vial hole diameter,
and the greatest numbers of V pensylvanica were
captured in traps baited with vials with the largest
hole size (12 mm diam) (Table 1). Although num-
bers of V atropilosa and V germanica captured in
this test were small, there was a significant posi-
tive relationship between vial hole diameter and
numbers of workers trapped for both species (Ta-
ble 1). Numbers of V germanica workers in traps
with vials releasing heptyl butyrate were statisti-
cally significant compared to traps with vials that
did not release heptyl butyrate (0 mm hole), for the
1.5 mm (t = 2.33, p = 0.02), 3 mm (t = 1.79, p =
0.05), 6 mm (t = 1.92, p = 0.04), and 12 mm (t =
3.87, p = 0.001) diameter holes in vial lids. In this
test, totals of 695 V pensylvanica, 58 V atropilosa,
and 106 V germanica workers were captured.
In the second field test conducted in Yakima,
there was a significant linear regression of num-


bers of V pensylvanica trapped with all vial hole
diameters tested (Table 1). Greatest numbers of
V pensylvanica and V atropilosa were captured
in traps baited with vials with 22 mm diameter
holes, with no increase in catches of these wasps
with the largest diameter hole in vials, 33 mm
(Table 1). The regression of wasps captured ver-
sus hole diameter was more significant for vial
hole diameters of 1.5 to 22 mm (r2 = 0.98, p =
0.002, df = 5), compared to 1.5 to 33 mm. Simi-
larly, there was a significant regression of num-
bers of V atropilosa trapped with all vial hole
diameters tested (Table 1), which was stronger for
1.5 to 22 mm (r2 = 0.98, p = 0.0015, df = 5), com-
pared to 1.5 to 33 mm. There was not a significant
linear relationship between vial hole diameter
and numbers of V germanica captured in this test
(Table 1). Totals of 5075 V pensylvanica, 172 V at-
ropilosa, 114 V germanica, 18 Dolichovespula
maculata (L.), 4 Dolichovespula arenaria (F.), 2
Polistes aurifer Saussure, and 1 Polistes dominu-
lus (Christ) were captured in this second test in
Yakima.
In the first dose-response test conducted in
Tulsa, Oklahoma, there was a significant rela-
tionship between vial hole diameter and numbers
of wasps trapped and greatest numbers of V
squamosa workers were captured in traps baited
with vials with the largest hole size (12 mm diam)
(Table 2). Also, in this test, numbers of V squa-
mosa workers captured in traps with vials releas-
ing heptyl butyrate were significantly greater
than numbers of workers in traps with vials that
did not release heptyl butyrate, for vial hole di-
ameters of 1.5 mm (t = 2.03, p = 0.02, df = 29), 3
mm (t = 2.95, p = 0.003), 6 mm (t = 4.90, p = 1.7 x
10-5), and 12 mm (t = 6.2, p = 4.4 x 10-7). No other
vespine wasps, but six female Polistes perplexus
(Cresson) and nine female Polistes fuscatus (F.)
were captured in traps in this test. These num-
bers were not sufficient for statistical analyses.


TABLE 1. MEAN (SE) NUMBERS OF WASPS CAPTURED PER TRAP PER TRAP-CHECK. TRAPS BAITED WITH HEPTYL BU-
TYRATE IN POLYPROPYLENE VIALS WITH VARIED HOLE DIAMETERS. YAKIMA, WASHINGTON, 2000.

Test 1, July 19-31 Vial hole diameter (mm) Regression statistics

Wasp species 0 1 1.5 3 6 12 r2 p

V atropilosa 0.0 0.0 0.1 0.1 0.2 0.1 0.2 0.1 0.5 0.1 1.9 0.4 0.93 0.002
V germanica 0.0 0.0 0.2 0.2 0.4 0.2 0.6 0.3 1.2 0.6 1.6 0.4 0.94 0.001
V pensylvanica 0.0 0.0 1.3 0.3 2.4 0.7 2.5 0.7 6.8 2.4 22.0 7.1 0.95 0.001

Test 2, August 4-21 Vial hole diameter (mm) Regression statistics

Wasp species 1.5 3 6 12 22 33 r2 p

V atropilosa 0.1 0.1 0.3 0.1 0.6 0.2 1.0 0.3 2.6 0.5 2.3 0.4 0.84 0.01
V germanica 0.3 0.2 0.9 0.3 0.4 0.1 1.2 0.4 1.1 0.4 1.0 0.3 0.37 0.20
V pensylvanica 5.3 1.4 12.7 2.3 27.8 5.5 34.9 4.0 63.2 5.1 58.3 8.2 0.84 0.01






Florida Entomologist 86(3)


TABLE 2. MEAN (SE) NUMBERS OF WASPS CAPTURED PER TRAP PER TRAP-CHECK. TRAPS BAITED WITH HEPTYL BU-
TYRATE IN POLYPROPYLENE VIALS WITH VARIED HOLE DIAMETERS. TULSA, OKLAHOMA, 2000.
Test 1, September 12-October 6 Vial hole diameter (mm) Regression statistics
Wasp species 0 1.0 1.5 3 6 12 r2 p
V. squamosa 0.2 0.2 0.3 0.1 0.9 0.3 1.3 0.3 5.5 1.0 11.2 1.7 0.99 <0.001
Test 2. October 6-November 3 Vial hole diameter (mm) Regression statistics
Wasp species 1.5 3 6 12 22 33 r2 p
V squamosa 0.4 0.1 1.1 0.2 3.1 0.7 6.3 1.3 12.3 2.1 17.8 3.0 0.99 <0.001


In the second dose-response test conducted in
Tulsa, there was a significant positive regression
of numbers of wasps trapped with vial hole diam-
eter, up through the largest hole tested, 33 mm,
and the greatest numbers of V squamosa cap-
tured were in traps baited with vials with 33 mm
diameter holes (Table 2). A total of 582 V squa-
mosa workers were captured in the first test in
Tulsa, while 1641 V squamosa workers were cap-
tured in the second test in Tulsa. No other
vespines, but six female P. fuscatus, were cap-
tured in these traps; not numerous enough for a
statistical analysis.
For all vial hole diameters, weight losses from
vials were steady over the 28 day duration of the


study (Fig. 1). Y-intercepts of best fit line equa-
tions indicated initial release rates (weight losses)
of 0, 0.42, 1.37, 2.30, and 3.30 milligrams per hour
for vials with holes of 0, 6, 12,22, and 33 mm in di-
ameter. For vials with holes of 6, 12, 22, and 33
mm in diameter, there was no significant slope (for
6 mm diam holes, r2 = 0.05, df = 10, p = 0.49; for 12
mm diam holes, r2 = 0.02, df = 10, p = 0.65; for 22
mm diam holes, r2 = 0.00, df = 10, p = 0.89; for 33
mm diam holes, r2 = 0.25, df = 10, p = 0.12), indi-
cating a near zero order release rate function.
Heptyl butyrate on cotton balls was released at a
higher rate initially, but decreased over time (Fig.
2). Initial weight losses were near 6 milligrams
per hour, decreasing to 5 mg/hr by day 9 and near


* 6nun A 12nun 22nnn o 33nun


45
4
35-
3
2.5
2-
15-
1-
05
0


DAYS IN AGE


Fig. 1. Mean (SE) milligrams of heptyl butyrate lost per hour from 30 ml polypropylene vials in a laboratory
fume hood. Vials had lid holes with diameters of 6, 12, 22, and 33 mm.


is I>
*P *


~I A 4 A
h o 67
. 0 S .'


September 2003







Landolt et al.: Trapping Yellowjackets with Heptyl Butyrate


U- ui EW i
0 5 10 15 20 25

DAYS IN AGE
Fig. 2. Mean (SE) milligrams of heptyl butyrate lost per hour from cotton balls in a laboratory fume hood. Cot-
ton balls were soaked with 2 ml heptyl butyrate, and then were placed on aluminum foil weighing dishes.


zero mg/hr by day 16. Rate of loss of heptyl bu-
tyrate from exposed cotton balls was related to
dispenser age (r2 = 0.93, df = 9, p < 0.01).

DISCUSSION
These results provide conclusive experimental
evidence of worker response to heptyl butyrate for
the southern yellowjacket V squamosa and the
German yellowjacket V germanica. Vespula squa-
mosa is widely distributed in eastern and central
North America from the mid Atlantic states to the
Gulf of Mexico (Akre et al 1981), and in areas of
Central America as far south as Honduras (Hunt
et al. 2001). Vespula germanica is native to Eur-
asia, and has become widely distributed and
abundant throughout much of temperate North
America (MacDonald & Akre 1984; Akre et al.
1989; Vetter et al. 1995). Both species are signifi-
cant pests. Consistent and significant numbers of
V squamosa were captured in this study in traps
baited with heptyl butyrate, demonstrating at-
traction to this chemical. The studies of Grothaus
et al. (1973) suggested attractiveness of heptyl
butyrate and other chemicals to V squamosa, but
did not include unbaited traps or traps that did
not release heptyl butyrate as experimental con-
trols, leaving open the question of whether or not
V squamosa workers are attracted. Sharp and
James (1979) used a mixture of heptyl butyrate
and octyl butyrate in their studies of V squamosa,
which did not include control traps for compari-
son. They did not address the question of V squa-
mosa worker attraction to heptyl butyrate, but
rather sought to evaluate the attractiveness of
trap colors.


A response of V germanica workers to heptyl
butyrate was also undocumented prior to this
study. Small numbers of V germanica workers
were previously captured in traps baited with hep-
tyl butyrate dispensed from a polyethylene cap
(Landolt 1998), but unbaited traps were not in-
cluded in that study as a control. Thus, conclusions
could not be drawn regarding the attractiveness of
the chemical to V germanica. The experimental
demonstration of attractiveness of these wasps to
heptyl butyrate with their capture in baited traps
does not address the question of the utility of the
chemical as a practical means of trapping either
species. However, comparisons of heptyl butyrate
with lures that release acetic acid and isobutanol
indicate a much stronger response by V germanica
to the latter attractant (Landolt 1998).
It is of interest, but not known, why numbers
of V germanica workers captured in traps in the
first test in Yakima increased with vial hole diam-
eter, and yet did not increase with vial hole diam-
eter in the second test in Yakima. Possible
variables to consider in future studies are the
temperature effects on heptyl butyrate release
rates in the field, competitive interactions with
other social wasps at or in traps, and changes in
responsiveness of V germanica wasps to heptyl
butyrate with the advance of the season, or vari-
ance in the availability of competing odor sources
and food materials.
Greatest numbers of V squamosa workers
were captured in traps with heptyl butyrate in vi-
als with the largest hole size tested (33 mm),
which gave an estimated rate of loss of about 3.3
milligrams of heptyl butyrate per hour at 22.5C
in the laboratory. It is possible that more V squa-


.4











mosa wasps might be caught at even higher hep-
tyl butyrate release rates.
It has been thoroughly established that very
large numbers of V pensylvanica workers can be
captured in traps baited with heptyl butyrate
(Davis et al. 1969; MacDonald et al. 1973), and re-
sults of this work are consistent with those reports.
Significant numbers ofV. atropilosa and V german-
ica workers were captured in heptyl butyrate-
baited traps here, indicating attraction to the
chemical. Vespula atropilosa has been trapped ex-
tensively with heptyl butyrate (Davis et al. 1969;
MacDonald et al. 1973). The numbers of both V.
atropilosa and V germanica captured were dramat-
ically less than the numbers ofV. pensylvanica cap-
tured. The relative differences in numbers of
workers captured for different species may reflect
both differences in population density at trapping
sites and differences in responsiveness to the at-
tractant (MacDonald et al. 1973).
These results provide information that can be
used to optimize captures ofyellowjackets in traps
baited with heptyl butyrate and to provide a
means of long-term sustained release of heptyl bu-
tyrate at an optimized release rate. An optimum
release rate was determined for heptyl butyrate as
an attractant for trapping V pensylvanica as well
as V atropilosa. Laboratory data indicated release
rates (weight losses) of 2.3 and 3.3 milligrams per
hour respectively from vials with those hole diam-
eters at laboratory temperatures. The amounts of
heptyl butyrate emitted from these vials in the
field will change with temperature, wind, and
other variable environmental parameters, but
these results provide a benchmark for comparison
to other types of controlled release technologies
and materials. The weight losses of vial dispens-
ers loaded with heptyl butyrate were steady for 30
days for all hole sizes evaluated. Application of
heptyl butyrate to a cotton ball (a presently-used
commercial application) provided a release rate
that was high and likely to be a strong attractant
for 6 to 9 days, but changed with time and was re-
duced to near zero after two weeks, suggesting a
relatively short period of attractiveness. The dis-
pensing of heptyl butyrate from a suitable device
has the potential of increasing both the attractive-
ness of the lure to wasps by stabilizing the release
rate at a targeted amount and the longevity of the
lure at that desired release rate.

ACKNOWLEDGMENTS

Technical assistance was provided by J. Beauchene,
J. Brumley, P. Chapman, D. Green, and L. Tom.

REFERENCES CITED

AKRE, R. D., A. GREENE, J. F. MACDONALD, P. J.
LANDOLT, AND H. G. DAVIS. 1981. Yellowjackets of
America North of Mexico. U.S. Department of Agri-
culture, Agriculture Handbook No. 552, 102 pp.


September 2003


AKRE, R. D., C. RAMSAY, A. GRABLE, C. BAIRD, AND A.
STANFORD. 1989. Additional range extension by the
German yellowjacket, Paravespula germanica (Fab-
ricius) in North America (Hymenoptera: Vespidae).
Pan-Pacific Entomol. 65: 79-88.
CHANG, V. 1988. Toxic baiting of the western yellow-
jacket (Hymenoptera: Vespidae) in Hawaii. J. Econ.
Entomol. 81: 228-235.
DATAMOST. 1995. StatMost statistical analysis and
graphics. DataMost, Salt Lake City, UT.
DAVIS, H. G., T. P. MCGOVERN, G. W. EDDY, T. E. NEL-
SON, K. M. R. BERTUN, M. BEROZA, AND J. C. INGA-
NGI. 1968. New chemical attractants for yellow
jackets (Vespula spp.). J. Econ. Entomol. 61:459-462.
DAVIS, H. G., G. W. EDDY, T. P. MCGOVERN, AND M.
BEROZA. 1969. Heptyl butyrate, a new synthetic at-
tractant for yellow jackets. J. Econ. Entomol. 62: 1245.
DAVIS, H. G., R. W. ZWICK, W. M. ROGOFF, T. P. MCGov-
ERN, AND M. BEROZA. 1973. Perimeter traps baited
with synthetic lures for suppression ofyellowjackets
in fruit orchards. Environ. Entomol. 2: 569-571.
GROTHAUS, R. H., H. G. DAVIS, W. M. ROGOFF, J. A.
FLUNO, AND J. M. HIRST. 1973. Baits and attractants
for east coast yellowjackets, Vespula spp. Environ.
Entomol. 2: 717-718.
HOWELL, J. 0., T. P. MCGOVERN, AND M. BEROZA. 1974.
Attractiveness of synthetic compounds to some east-
ern Vespula species. J. Econ. Entomol. 67: 629-630.
HUNT, J. H., R. H. CAVE, AND G. R. BORJAS. 2001. First
records from Honduras of a yellowjacket wasp, Ves-
pula squamosa (Drury) (Hymenoptera: Vespidae:
Vespinae). J. Kansas Entomol. Soc. 74: 119-119.
LANDOLT, P. J. 1998. Chemical attractants for trapping
yellowjackets Vespula germanica and Vespula pen-
sylvanica (Hymenoptera: Vespidae). Environ. Ento-
mology 27: 1229-1234.
MACDONALD, J. F., R. D. AKRE, AND W. B. HILL. 1973.
Attraction of yellowjackets (Vespula spp.) to heptyl
butyrate in Washington State (Hymenoptera: Vespi-
dae). Environ. Entomol. 2: 375-379.
MACDONALD, J. F., R. D. AKRE, AND W. B. HILL. 1974.
Comparative biology and behavior of Vespula atropi-
losa and V. pensylvanica (Hymenoptera: Vespidae).
Melanderia 18: 1-93.
MACDONALD, J. F., R. D. AKRE, AND R. W. MATTHEWS.
1976. Evaluation of yellowjacket abatement in the
United States. Bull. Entomol. Soc. America 22: 397-
401.
MACDONALD, J. F. AND R. D. AKRE. 1984. Range exten-
sion and emergence of subterranean nesting by the
German yellowjacket Vespula germanica, in North
America. Entomol. News 95: 5-8.
REED, H. C., AND P. J. LANDOLT. 2003. Michigan social
wasps (Hymenoptera: Vespidae) captured in traps
baited with heptyl butyrate, acetic acid, and isobu-
tanol. Great Lakes Entomol. 35: 71-77.
REIERSON, D. A., AND R. E. WAGNER 1975. Trapping yel-
lowjackets with a new standard plastic wet trap. J.
Econ. Entomol. 68: 395-398.
REIERSON, D. A., AND R. E. WAGNER 1978. Trapping to
determine the sympatry and seasonal abundance of
various yellowjackets. Environ. Entomol. 7: 418-422.
SHARP, J. L., AND J. JAMES. 1979. Color preference of
Vespula squamosa. Environ. Entomol. 8: 708-710.
BETTER, R. S., P. K. VISSCHER, AND D. A. REIERSON.
1995. Vespula germanica (Fabr.) in southern Califor-
nia (Hymenoptera: Vespidae). Pan-Pac. Entomol. 71:
246-248.


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