Effects of Elasmopalpus lignosellus (Lepidoptera: Pyralidae)

Damage on Sugarcane Yield

Author(s): Hardev S. Sandhu, Gregg S. Nuessly, Ronald H. Cherry, Robert A.
Gilbert, and Susan E. Webb
Source: Journal of Economic Entomology, 104(2):474-483.
Published By: Entomological Society of America
DOI: http://dx.doi.org/10.1603/EC10172
URL: http://www.bioone.org/doi/full/10.1603/EC10172

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 FIELD AND FORAGE CROPS

Effects of Elasmopalpus lignosellus (Lepidoptera: Pyralidae) Damage

on Sugarcane Yield
HARDEV S. SANDHU,1,2 GREGG S. NUESSLY,1 RONALD H. CHERRY,1 ROBERT A. GILBERT,1
3
AND SUSAN E. WEBB

J. Econ. Entomol. 104(2): 474Ð483 (2011); DOI: 10.1603/EC10172

ABSTRACT Feeding by lesser cornstalk borer, Elasmopalpus lignosellus (Zeller) (Lepidoptera:
Pyralidae), larvae on sugarcane (a complex hybrid of Saccharum spp.) causes leaf damage, dead hearts,
and dead plants that can result in stand and yield loss. A 2-yr greenhouse experiment was conducted
to examine sugarcane variety and plant age-speciÞc feeding responses to E. lignosellus. Plants growing
from single-eye setts of three varieties were exposed to a single generation of E. lignosellus larvae
beginning at the three-, Þve-, and seven-leaf stages. Results indicated that the physical damage and
resulting yield loss of plants attacked by E. lignosellus larvae were dependent on the variety and leaf
stage at which they were infested. SigniÞcantly more plant damage was observed in all three varieties
when infested at the three- than at the seven-leaf stage. Larvae caused signiÞcantly more plant damage
and reduced yield in CP 89-2143 than in CP 78-1628. Tiller production increased in CP78 Ð1628 and
CP 88-1762 when infested at the three-leaf stage, whereas tiller production, biomass and sugar yield
decreased in CP 89-2143 when infested at all leaf stages, compared with the untreated control. There
was no reduction in yield when CP 78-1628 was infested at the three- or Þve-leaf stages. Biomass was
reduced in CP 88-1762 when plants were infested at any of the leaf stages, but sugar yield was reduced
only when infested at the seven-leaf stage. These results indicate that compensation in response to
E. lignosellus damage was variety dependent and declined with the delay in infestation time.

KEY WORDS CP 78-1628, CP 88-1762, CP 89-2143, compensation, lesser cornstalk borer

Florida is the leading sugarcane (a complex hybrid of
Saccharum spp.)-producing state in the United States,
with 162,348 ha of sugarcane valued at US$398.9 mil-
lion in 2008 (USDA 2008). The majority of the sug-
arcane acreage is grown in Palm Beach, Martin, Hen-
dry, and Lee counties in southern Florida. Sugarcane
is vegetatively propagated by planting mature stalks.
New primary shoots grow from lateral buds at the
nodes soon after planting (Dillewijn 1952). Primary
shoots have many small internodes each with a lateral
bud. These lateral buds develop into secondary shoots
that in turn may produce tertiary shoots.
Lesser cornstalk borer, Elasmopalpus lignosellus
(Zeller) (Lepidoptera: Pyralidae), is a pest of many
crops, including sugarcane (Falloon 1974). Larval
feeding on primary, secondary, and tertiary sugarcane
shoots results in dead heart symptoms and dead plants
that can translate into reduced sugarcane and sugar
yield at harvest (Sandhu et al. 2010). Nonlethal dam-
age is caused when larvae only chew a few millimeters
into the shoot and becomes evident when the leaves
1 Everglades Research & Education Center, Institute of Food and
Agricultural Services, University of Florida, 3200 E. Palm Beach Rd.,
Belle Glade, FL 33430.
2 Corresponding author, e-mail: hardy@uß.edu.
3 Department of Entomology and Nematology, Institute of Food
and Agricultural Services, University of Florida, P.O. Box 110620,
Gainesville, FL 32611.
push out to reveal one to several symmetrical rows of
holes (Schaaf 1974, Carbonell 1978). Leaves usually
break off above these points within a few weeks of
expansion resulting in a loss of photosynthetic surface
and nutrients.
“Compensation” enables plants respond positively
to an injury (Bardner and Fletcher 1974) and de-
creases the negative effect on yield (Pedigo 1991).
The ability of plants to withstand or recover from
insect damage also is deÞned as “tolerance” by Painter
(1951). Sugarcane has a great capacity to compensate
for mechanical damage to young shoots (up to 100%),
but this capacity diminishes with plant age (Dillewijn
1952). However, mechanical damage to 7-mo-old sug-
arcane shoots also showed 50% compensatory growth
(Demandt 1929). He showed that this compensation
was partly due to the production of new tillers and
partly due to survival of the stalks that otherwise
would have died due to lack of nutrients in the pres-
ence of the primary shoot. Loh et al. (1948) later
showed that mechanical removal (topping) of the
primary shoots increased the number of millable stalks
(primary shoots and tillers ⱖ1.5 m in height) by 11.3%
and biomass yield by 25.9%.
Shoots that die in response to E. lignosellus feeding
may not produce tillers; however, initial feeding dam-
age does not always result in stand or yield loss. Car-
bonell (1978) reported 27.8% recovery in plant canes

0022-0493/11/0474Ð0483$04.00/0 䉷 2011 Entomological Society of America

April 2011 SANDHU ET AL.: SUGARCANE RESPONSE TO E. lignosellus
and 48.1% recovery in stubble canes in response to E.
lignoselus damage. Information on variety speciÞc sug-
arcane recovery to E. lignosellus damage would be
useful for the industry in their variety selection pro-
gram. This information is also important for develop-
ing damage thresholds for integrated management of
this pest. The objective of this study was to document
variety and age speciÞc feeding damage in sugarcane
by lesser cornstalk borer larvae and the potential for
damaged plants to compensate for early season dam-
age.
Materials and Methods
Effects of damage caused by lesser cornstalk borer
on sugarcane growth and yield were evaluated in two,
11-mo greenhouse studies during 2008 (JanuaryÐ
November) and 2009 (November 2008 ÐSeptember
2009) conducted at the Everglades Research and Ed-
ucation Center (EREC), Belle Glade, FL. There are
multiple varieties of sugarcane grown in this area
(Rice et al. 2009) that are attacked by lesser cornstalk
borer at different growth stages. The sugarcane vari-
eties CP 78-1628, CP 89-2143, and CP 88-1762 were
selected for this study. These varieties occupy the
greatest acreage grown on Immokalee Þne sand
(sandy soil) where lesser cornstalk borer is considered
to be a major perennial problem (Rice et al. 2009). CP
89-2143 and CP 88-1762 also were ranked as Þrst and
second in total Florida sugarcane acreage. Three early
growth stages (three-, Þve-, and seven-leaf stage)
were selected for infestation with lesser cornstalk
borer larvae based on damage reports during the Þrst
2Ð3 mo of sugarcane growth (Carbonell 1978). These
three selected growth stages were present ⬇3, 5, and
8 wk after emergence of primary shoots.
Production of Sugarcane Plants. Mature stalks of
each variety were harvested from Þelds at the EREC
to obtain viable buds for planting. Stalks were cut into
10-cm-long seed pieces each with one lateral bud (i.e.,
single eye setts) and planted in plastic trays (50 by 36
by 9.5 cm) Þlled with Immokalee Þne sand to germi-
nate the buds and produce primary shoots. Immokalee
Þne sand was used as a medium for plant growth
throughout the experiment. The lesser cornstalk borer
causes more damage in sandy soil than muck soil, and
Immokalee Þne sand is one of the major sandy soils in
the sugarcane-growing area around Lake Okeechobee
in south central Florida. Two days after the emergence
of primary shoots, uniform-sized seedlings were se-
lected and transplanted to 19.0-liter (5-gal) buckets
(two seedlings per bucket) Þlled with Immokalee Þnd
sand. Plants were fertilized by adding 50 g of ammo-
nium sulfate and 20 g of a balanced granular fertilizer
(14 Ð14 Ð14) to the soil of each bucket at planting time
and again every 3 mo until harvest. Irrigation was
applied every 2 d.
Insect Rearing. Insects used in this study were ob-
tained from a laboratory colony of lesser cornstalk
borer maintained at EREC. The colony was started 4
mo before the start of the experiment by using larvae
and adults of E. lignosellus collected from sugarcane
475
Þelds at Belle Glade and Moore Haven, FL. Larvae and
adult E. lignosellus were collected from sugarcane
Þelds and added to the colony each month to improve
heterogeneity and fecundity in the laboratory popu-
lation. The colony was maintained on a wheat germ-
and soy ßour-based artiÞcial diet as described in
Sandhu et al. (2010). Third-instar larvae were used to
infest sugarcane plants. To produce larvae for the trial,
Þrst- and second-instar larvae were removed from the
artiÞcial diet and reared on three- to four-leaf stage
shoots of respective sugarcane varieties to avoid the
effect of host change on larval feeding. The choice to
infest plants by using third-instar larvae was based on
preliminary trials on greenhouse plants, where it was
observed that Þrst and second instars had high mor-
tality and their feeding on leaves did not cause major
damage. Third-instar larvae move from leaves to the
soil to feed on shoots and tillers.
Experiment Design. A randomized complete block
design with a 3 by 4 factorial arrangement was used
during both experiment years to evaluate sugarcane
response to E. lignosellus feeding damage. The factors
were three sugarcane varieties (CP 78-1628, CP 88-
1762, and CP 89-2143) and three leaf stages infested
plus one control (i.e., no infestation and infestation at
the three-, Þve-, and seven-leaf stages). The combi-
nations of these factors were applied randomly to the
12 buckets in each block. The buckets in each block
were arranged in two rows of six buckets each in a fan-
and pad-cooled greenhouse at the EREC. Tempera-
ture within the greenhouse was maintained within 3⬚C
of the ambient temperature outside the greenhouses
for the duration of each trial. Each block was repli-
cated 12⫻ in 2008 and 15⫻ in 2009. Four, third-instar
(6 Ð7-d-old) larvae conditioned on sugarcane as above
were released near the base of shoots in each bucket
with the aid of a camelÕs-hair brush at the respective
leaf stages. The number of larvae per bucket was
selected based on preliminary trials on greenhouse
plants. Sugarcane in the buckets was exposed to a
single generation of lesser cornstalk borer. After com-
pletion of the larval stage, pupae were located in silken
tunnels in the soil surrounding the plants and removed
by straining the top 6 cm of soil through 15 mesh
screen. Timing for the completion of the larval stage
was estimated based on the results of temperature-
dependent developmental studies on sugarcane seed-
lings in the laboratory (Sandhu et al. 2010).
Damage Assessment. Feeding damage was recorded
for primary shoots and tillers in each bucket. Second-
ary and tertiary shoots are referred to as tillers in this
report. The number of dead hearts, number of shoots
with symmetrical rows of holes in leaves, and number
of dead plants per bucket were recorded weekly for 3
wk starting 1 wk after infestation at each leaf stage.
The last sample date for each leaf stage was within a
week after the insects completed larval development.
Plants were counted as dead hearts if feeding lead to
chlorosis and necrosis of the unfurled leaves in the
whorl of primary shoots or tillers. A plant with damage
that led to necrosis of all the aboveground plant tissues
in both primary shoot and tillers was counted as a dead
476 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 2

Table 1. Summary of test statistics with effects of sampling date, variety, leaf stage and their interaction on plant damage by E.
lignosellus to sugarcane in 2008 and 2009

Source of 2008 2009

Feeding damage
variation df F P df F P
Dead hearts Sampling date 2, 264 4.02 0.0191 2, 313 5.94 0.0029
Variety (V) 2, 264 1.77 0.1730 2, 18.7 2.00 0.1636
Leaf stage (LS) 3, 264 270.08 ⬍0.0001 3, 24.5 531.85 ⬍0.0001
V ⫻ LS 6, 264 2.79 0.0120 6, 24.5 1.04 0.4216
Rows of holes in leaves Sampling date 2, 264 4.66 0.0102 2, 313 11.22 ⬍0.0001
Variety (V) 2, 264 6.97 0.0011 2, 17.9 1.23 0.3148
Leaf stage (LS) 3, 264 86.83 ⬍0.0001 3, 24.7 261.73 ⬍0.0001
V ⫻ LS 6, 264 1.38 0.2209 6, 24.7 2.58 0.0444
Dead plants Sampling date 2, 264 1.02 0.3634 2, 312 6.31 0.0021
Variety (V) 2, 264 118.93 0.0001 2, 19.1 5.82 0.0106
Leaf stage (LS) 3, 264 132.50 ⬍0.0001 3, 24.2 77.17 ⬍0.0001
V ⫻ LS 6, 264 24.57 ⬍0.0001 6, 24.2 1.58 0.1956

Analysis of variance (PROC MIXED, SAS Institute 2008).

plant. Buckets were Þrst observed for dead plants,
then dead hearts and then holes in the leaves. Plants
counted as dead were not counted as dead hearts, and
plants with dead hearts were not counted as plants
with holes in leaves. The number of tillers per bucket
was counted 4 mo after emergence to account for
varietal and leaf stage differences in tiller production.
Sugarcane Yield Assessment. Sugarcane yield was
determined using the number and weight of millable
stalks (biomass yield), and the sucrose concentration
(sugar yield) of juice squeezed from those stalks. Mill-
able stalks are primary shoots and tillers ⬎1.5 m in
height and are traditionally counted 8 mo after the Þrst
emergence of sugarcane shoots. Millable stalks are
counted 3 mo before harvesting, because the possible
lodging in sugarcane at harvest time interferes with
determining the height and exact number of millable
stalks. Millable stalks from each bucket were har-
vested 11 mo after sugarcane emergence. Individual
stalks were weighed separately, and biomass yield
(kilograms per bucket) was calculated as the product
of the number of millable stalks and the mean stalk
weight in each bucket. To determine sugar concen-
tration, two randomly selected millable stalks per
bucket from each block were milled and crusher juice-
analyzed for brix and pol values as described by Gil-
bert et al. (2008). These values were used to calculate
sugar yield (kilograms per bucket) according to the
theoretically recoverable sugar method (Glaz et al.
2002).
Analysis. The data on response variables (dead
hearts, holes in the leaves, dead plants, number of
tillers, millabe stalk count, biomass yield, and sugar
yield) were recorded for each bucket and analyzed for
the effect of year, varieties, infested leaf stages, and
their interactions. The PROC MIXED procedure (SAS
Institute 2008) with the repeated measures statement
was used to analyze the damage data due to the po-
tential covariance structure associated with repeated
damage assessments over time in the same locations.
Degrees of freedom were calculated in PROC MIXED
by using the KenwardÐRoger correction. Normality of
the data were tested with the Shapiro-Wilk normality
test (Shapiro and Wilk 1965). Sampling dates were
used as the repeated variable and were nested under
treatment combinations in the repeated measures
statement. Several covariance structures were Þtted to
the data. The compound symmetry type (Littell et al.
1998) provided the best (lowest) Þt statistic and was
used for the analysis. Percentage data were arcsine
transformed before analysis and retransformed for
presentation purposes. Sugarcane variety, leaf stage at
infestation, and their interaction were Þxed effects in
the analysis of variance (ANOVA) model. Replicates
and their interaction with the main plots were used as
random effects in the model.
Yield traits were recorded at the end of each ex-
periment (i.e., no repeated measures); therefore, the
data were analyzed using the PROC GLM procedure
for a factorial design (SAS Institute 2008). Sugarcane
variety, leaf stage at infestation, and their interactions
were used as Þxed effects in the ANOVA model. Rep-
licates and their interaction with the main plots were
used as random effects in the model. Orthogonal con-
trasts (SAS Institute 2008) were used for mean sepa-
ration among treatments where ANOVA F values were
signiÞcant at the P ⬍ 0.10 level. Orthogonal contrasts
were used due to the incomplete factorial experimen-
tal design and for better comparisons between the
untreated control and treatments.
Results
Damage. The study year affected the percentage of
dead plants at P ⫽ 0.0001 (F ⫽ 301.99; df ⫽ 1, 883; P ⬍
0.0001) and the percentage of plants with dead hearts
at P ⫽ 0.06 (F ⫽ 3.73; df ⫽ 1, 884; P ⫽ 0.0538);
therefore, the data on all damage types were analyzed
separately for 2008 and 2009 (Table 1). The study year
was not a signiÞcant source of variation in the model
for percentage of plants with holes in the leaves (F ⫽
0.01; df ⫽ 1, 879; P ⫽ 0.9401). Sampling date also
signiÞcantly affected the measured damage parame-
ters (Table 1); therefore, the mean percentage of dead
hearts, leaves with holes, and dead plants were pre-
sented by date (Fig. 1AÐF). The data for untreated
controls were not included in the Þgures because
April 2011 SANDHU ET AL.: SUGARCANE RESPONSE TO E. lignosellus 477

Fig. 1. Percentage of sugarcane plants damaged by E. lignosellus by variety, leaf stage, and sample date: dead hearts in
2008 (A) and 2009 (B), holes in leaves in 2008 (C) and 2009 (D), and dead plants in 2008 (E) and 2009 (F).

there was no feeding damage observed in the un-
treated controls.
Dead hearts were the most commonly observed
result of E. lignosellus feeding damage to sugarcane.
The change in the percentage of dead hearts with
sampling dates varied with the variety and the leaf
stage at which the plants were infested (Fig. 1A and
B). Leaf stage was a signiÞcant source of variation
(Table 1) in the model; the earlier the plants were
infested, the greater the resulting dead heart symp-
toms were produced. The percentage of plants with
dead hearts was greater in the sequence three- ⬎
Þve- ⬎ seven-leaf stage (Fig. 1A and B). Although
variety alone was not a signiÞcant source of variation
in the model for dead hearts in either year, the sig-
niÞcant affect of sample dates and leaf stages com-
478 JOURNAL OF ECONOMIC ENTOMOLOGY
bined to produce too much variation to separate out
the varieties when the data were pooled across time
and leaf stage. The variety ⫻ leaf stage interaction was
also a signiÞcant source of variation (Table 1) in the
model for dead hearts in 2008. In all three varieties,
plants infested at the three-leaf stage had a greater
percentage of plants with dead hearts than those in-
fested at the seven-leaf stage.
Symmetrical rows of holes in the leaves were the
second most commonly observed damage by E. ligno-
sellus feeding on sugarcane. The change in the per-
centage of leaves with holes among sampling dates also
varied with the variety and the leaf stage at which the
plants were infested (Fig. 1C and D). Leaf stage was
a signiÞcant source of variation in the model for per-
centage of leaves with holes in both years (Table 1).
In contrast to dead hearts, mean percentage of plants
with holes in the leaves was greater in late-infested
(seven-leaf stage) than early-infested (three-leaf
stage) plants (Fig. 1C and D). The mean percentage
of plants with holes in the leaves at the seven-leaf stage
was signiÞcantly greater than at the Þve-leaf stage, and
more were found on plants infested at the Þve-leaf
stage than at the three-leaf stage in all three varieties.
However, variety was signiÞcant only in 2008 and the
variety ⫻ leaf stage interaction was signiÞcant only in
2009. Among varieties in 2008, CP 88-1762 had a
greater percentage of plants with holes in leaves than
CP 89-2143.
Dead plants were the third type of observed damage
caused by E. lignosellus feeding on sugarcane. The
variation in percentages of dead plants among sam-
pling dates was dependent upon the variety and the
infested leaf stage (Fig. 1E and F). Variety and leaf
stage were signiÞcant sources of variation in the model
for dead plants in both years, but the variety ⫻ leaf
stage interaction was signiÞcant only in 2008 (Table
1). Among varieties, CP 89-2143 had a signiÞcantly
greater percentage of dead plants than the other two
varieties (Fig. 1E and F). The mean percentages of
plants that died when infested at the three- and Þve-
leaf stages were signiÞcantly greater than when in-
fested at the seven-leaf stage. Infestation at the three-
and Þve-leaf stages in CP 78-1628 and CP 88-1762, and
at the three-leaf stage in CP 89-2143 produced greater
percentages of dead plants than when infested at the
seven-leaf stage in each variety. No CP 78-1628 plants
died when infested at the seven-leaf stage; however,
early infestation of CP 89-2143 caused the greatest
percentage of plant deaths.
Tiller Production. Experiment year signiÞcantly af-
fected tiller production (F ⫽ 34.85; df ⫽ 1, 322; P ⬍
0.0001) with approximately one additional tiller pro-
duced per bucket in 2009 than in 2008. Due to the
signiÞcant year affect, the data were analyzed sepa-
rately for 2008 and 2009. Variety and leaf stage in 2008
and only variety in 2009 were the signiÞcant sources
of variation in the model for number of tillers per
bucket in both years (Tables 2 and 3). CP 78-1628
produced the greatest number of tillers followed by
CP 88-1762 and CP 89-2143 in 2008 (Table 2). In 2009,
both CP 78-1628 and CP 88-1762 produced more tillers
Vol. 104, no. 2
than CP 89-2143 (Table 3). Buckets infested at the
three- and Þve-leaf stages produced signiÞcantly more
tillers than those infested at the seven-leaf stage in
2008, but there was no difference among the leaf stages
for tiller production in 2009. The variety ⫻ leaf stage
interaction was not a signiÞcant source (at P ⫽ 0.05)
of variation in either year. However, leaf stage signif-
icantly (P ⬍ 0.10) affected the number of tillers pro-
duced by CP 78-1628 and CP 88-1762 in 2008 with
more tillers produced when infested at the three-leaf
than at the seven-leaf stage or in the untreated control
(Table 2). In 2009, damage at all the leaf stages re-
sulted in fewer tillers produced CP 89-2143 than in the
untreated control (Table 3).
Sugarcane Yield Traits. Yield traits refer to the num-
ber of millable stalks, biomass yield, and sugar yield.
The experiment year (df ⫽ 1, 322) was again a sig-
niÞcant source of variation in the models for numbers
of millable stalks (F ⫽ 88.28, P ⬍ 0.0001), biomass yield
(F ⫽ 49.78, P ⬍ 0.0001), and sugar yield (F ⫽ 11.83, P ⫽
0.0007). Therefore, the data were analyzed separately
for 2008 and 2009. The signiÞcant year affect resulted
from greater yields in 2009 than in 2008. Overall there
was approximately one extra stalk produced per
bucket in 2009 than in 2008, which lead to a 0.75-kg
increase in biomass yield and a 0.5-kg increase in sugar
yield per bucket in 2009 than in 2008. Mean ⫾ SEM
values of yield traits for 2008 and 2009 are presented
in Tables 2 and 3, respectively.
The number of millable stalks per bucket was sig-
niÞcantly affected by variety and variety ⫻ leaf stage
in 2008 (Table 2). In 2009, both main effects and their
interaction were signiÞcant sources of variation in the
model (Table 3). More millable stalks were produced
by CP 78-1628 followed by CP 88-1762 and then by CP
89-2143 in both years (Tables 2 and 3). Plants infested
at the three-leaf stage produced more millable stalks
than when infested at the seven-leaf stage in 2009
(Table 3). E. lignosellus damage to CP 78-1628 at the
three-leaf stage in 2008 (Table 2) and at the three- and
Þve-leaf stages in 2009 (Table 3) resulted in increased
millable stalk production over the untreated controls
and those damaged at the seven-leaf stage. In CP
88-1762, plants infested at the three-leaf stage pro-
duced more millable stalks than plants infested at the
seven-leaf stage in 2009. SigniÞcantly fewer millable
stalks were produced by CP 89-2143 when plants were
infested at all three-leaf stages compared with the
untreated control in 2008 (Table 2).
Biomass yield was signiÞcantly affected by variety
and leaf stage in both study years (Tables 2 and 3).
Biomass yield was the greatest in CP 78-1628 followed
by CP 88-1762 and then by CP 89-2143 in 2008 (Table
2), whereas CP 78-1628 and CP 88-1762 produced
similar biomass yields that were both greater than CP
89-2143 (Table 3). In 2008 and 2009, untreated control
plants produced greater biomass yield than plants in-
fested at all three-leaf stages, and plants infested at the
three- and Þve-leaf stages produced greater biomass
yield than those infested at the seven-leaf stage. Bio-
mass yield in each variety was signiÞcantly affected by
leaf age in both study years (Tables 2 and 3). Biomass
April 2011 SANDHU ET AL.: SUGARCANE RESPONSE TO E. lignosellus 479

Table 2. Sugarcane tillers and yield measurements (least mean square ⴞ SEM) in 2008 study comparing varieties, leaf stages, and
their interaction

Tillersa Millable stalksb Biomass yieldc Sugar yieldd

Main effect
Variety
CP 78-1628 4.3 ⫾ 0.14A 3.6 ⫾ 0.10A 2.6 ⫾ 0.08A 0.26 ⫾ 0.009A
CP 88-1762 3.8 ⫾ 0.14B 3.2 ⫾ 0.10B 2.2 ⫾ 0.08B 0.23 ⫾ 0.009B
CP 89-2143 3.1 ⫾ 0.14C 2.2 ⫾ 0.10C 1.9 ⫾ 0.08C 0.21 ⫾ 0.009B
F; P; df ⫽ 2, 66 18.54; ⬍0.0001 46.71; ⬍0.0001 22.24; ⬍0.0001 7.98; 0.0008
Leaf stage
Untreated (un) 3.8 ⫾ 0.16ab 3.1 ⫾ 0.12 2.7 ⫾ 0.09a 0.28 ⫾ 0.011a
3-leaf (3L) 4.1 ⫾ 0.16a 3.2 ⫾ 0.12 2.3 ⫾ 0.09b 0.24 ⫾ 0.011b
5-leaf (5L) 3.8 ⫾ 0.16a 3.0 ⫾ 0.12 2.1 ⫾ 0.09b 0.23 ⫾ 0.011b
7-leaf (7L) 3.4 ⫾ 0.16b 2.8 ⫾ 0.12 1.8 ⫾ 0.09c 0.18 ⫾ 0.011c
F; P; df ⫽ 3, 66 3.88; 0.0129 1.64; 0.1895 19.39; ⬍0.0001 16.25; ⬍0.0001
Interaction effects
Variety ⫻ leaf stage
CP 78-1628 ⫻ un 4.0 ⫾ 0.27b 3.3 ⫾ 0.17b 2.9 ⫾ 0.17a 0.30 ⫾ 0.021a
CP 78-1628 ⫻ 3L 4.9 ⫾ 0.27a 3.9 ⫾ 0.17a 2.9 ⫾ 0.17a 0.28 ⫾ 0.021a
CP 78-1628 ⫻ 5L 4.5 ⫾ 0.27ab 3.8 ⫾ 0.17ab 2.6 ⫾ 0.17b 0.27 ⫾ 0.021a
CP 78-1628 ⫻ 7L 4.0 ⫾ 0.27b 3.3 ⫾ 0.17b 2.1 ⫾ 0.17c 0.19 ⫾ 0.021b
F; P; df ⫽ 3, 44 2.59; 0.0644 3.20; 0.0323 5.23; 0.0036 4.86; 0.0053
CP 88-1762 ⫻ un 3.7 ⫾ 0.31bc 3.1 ⫾ 0.18 2.7 ⫾ 0.11a 0.28 ⫾ 0.014a
CP 88-1762 ⫻ 3L 4.5 ⫾ 0.31a 3.6 ⫾ 0.18 2.2 ⫾ 0.11b 0.24 ⫾ 0.014a
CP 88-1762 ⫻ 5L 4.0 ⫾ 0. 31ab 3.2 ⫾ 0.18 2.1 ⫾ 0.11b 0.23 ⫾ 0.014a
CP 88-1762 ⫻ 7L 3.2 ⫾ 0. 31c 3.0 ⫾ 0.18 1.7 ⫾ 0.11b 0.17 ⫾ 0.014b
F; P; df ⫽ 3, 44 3.37; 0.0268 2.03; 0.1242 12.76; ⬍0.0001 10.12; ⬍0.0001
CP 89-2143 ⫻ un 3.7 ⫾ 0.24 2.8 ⫾ 0.25 2.6 ⫾ 0.19a 0.27 ⫾ 0.022a
CP 89-2143 ⫻ 3L 3.0 ⫾ 0.24 2.0 ⫾ 0.25 1.8 ⫾ 0.19b 0.20 ⫾ 0.022b
CP 89-2143 ⫻ 5L 3.0 ⫾ 0.24 2.1 ⫾ 0.25 1.7 ⫾ 0.19b 0.19 ⫾ 0.022b
CP 89-2143 ⫻ 7L 2.9 ⫾ 0.24 2.1 ⫾ 0.25 1.5 ⫾ 0.19b 0.17 ⫾ 0.022b
F; P; df ⫽ 3, 44 2.18; 0.1039 2.01; 0.1265 5.94; 0.0017 3.62; 0.0203

Means separation tests (orthogonal contrasts, SAS Institute 2008) were performed where ANOVA F values (Proc GLM, SAS Institute 2008)
were signiÞcant at P ⱕ 0.10. Means followed by different letters in a column are signiÞcantly different. Uppercase letters indicate differences
among varieties (main effects). Lowercase letters indicate differences among leaf stages (main effects) and among leaf stages of the same variety
(interaction effects).
a
Number of secondary and tertiary shoots.
b
Number of stalks ⱖ1.5 m in height.
c
Total weight (kilograms) of millable stalks in each bucket.
d
Raw sugar weight (kilograms per bucket) calculated from brix and pol values by using the theoretically recoverable sugar method (Glaz
et al. 2002).

yield in CP 78-1628 was signiÞcantly reduced com-
pared with the untreated control when infested at the
Þve- and seven-leaf stages in 2008 (Table 2) and at the
seven-leaf stage in 2009 (Table 3). In both CP 88-1762
and CP 89-2143, the biomass yield was signiÞcantly
lower when infested at all leaf stages compared with
the untreated control in both years (Tables 2 and 3).
Sugar yield was signiÞcantly affected by variety and
leaf stage in both years (Tables 2 and 3). Sugar yields
were signiÞcantly greater in CP 88-1762 than in the
other two varieties in both years (Tables 2 and 3), and
greater in CP 88-1762 than in CP 89-2143 in 2008
(Table 2). In both years, plants had lower sugar yield
compared with the untreated control when infested at
all leaf stages, and those infested at the seven-leaf stage
had less sugar yield than those infested at the three-
and Þve-leaf stages (Tables 2 and 3). Sugar yield in
each variety was signiÞcantly affected by leaf age in
both study years (Tables 2 and 3). In CP 78-1628,
damage by lesser cornstalk borer resulted in sugar
yield reductions compared with the untreated control
only when infested at the seven-leaf stage in both
years. Sugar yield in CP 88-1762 was signiÞcantly re-
duced compared with the untreated control when
infested at the seven-leaf stage in 2008 (Table 2) and
at the Þve- and seven-leaf stages in 2009 (Table 3). E.
lignosellus damage to CP 89-2143 caused signiÞcant
sugar yield losses at all the infested leaf stages com-
pared with untreated controls in both years (Tables 2
and 3).
Discussion
Damage. Feeding damage to the plants occurred
over the 2Ð3 wk required for the third instar larvae to
complete development to the pupal stage. Dead hearts
and dead plants can be recognized soon after larval
feeding cuts off the supply of water to the shoots and
tillers from the roots. Expression of the symmetrical
rows of holes in leaves is delayed for at least 1 wk as
the leaves expand from within the stems to reveal the
damage. Damage symptoms were expressed as per-
centages of damaged shoots and tillers to standardize
the results across replicates and treatments, because
plants in each bucket grew and produced new tillers
at different rates in response to the treatments. In-
creases in the observed percentage damage symptoms
between sample dates resulted from continued dam-
age by larvae and delayed expression of symptoms
relative to the number of undamaged shoots. De-
480 JOURNAL OF ECONOMIC ENTOMOLOGY Vol. 104, no. 2

Table 3. Sugarcane tillers and yield measurements (least mean square ⴞ SEM) in 2009 study comparing varieties, leaf stages, and
their interaction

Tillersa Millable stalksb Biomass yieldc Sugar yieldd

Main effect
Variety
CP 78-1628 5.4 ⫾ 0.18A 4.7 ⫾ 0.10A 3.1 ⫾ 0.06A 0.30 ⫾ 0.001A
CP 88-1762 5.1 ⫾ 0.18A 4.4 ⫾ 0.10B 3.0 ⫾ 0.06A 0.28 ⫾ 0.001B
CP 89Ð2143 3.6 ⫾ 0.18B 3.2 ⫾ 0.10C 2.5 ⫾ 0.06B 0.22 ⫾ 0.001C
F; P; df ⫽ 2, 84 30.45; ⬍0.0001 64.39; ⬍0.0001 29.97; ⬍0.0001 33.64; ⬍0.0001
Leaf stage
Untreated (un) 4.8 ⫾ 0.21 4.2 ⫾ 0.11a 3.6 ⫾ 0.07a 0.33 ⫾ 0.008a
3-leaf (3L) 4.9 ⫾ 0.21 4.3 ⫾ 0.11a 3.0 ⫾ 0.07b 0.28 ⫾ 0.008b
5-leaf (5L) 4.6 ⫾ 0.21 4.1 ⫾ 0.11a 2.8 ⫾ 0.07b 0.26 ⫾ 0.008b
7-leaf (7L) 4.5 ⫾ 0.21 3.6 ⫾ 0.11b 2.1 ⫾ 0.07c 0.19 ⫾ 0.008c
F; P; df ⫽ 3, 84 0.80; 0.4968 7.36; 0.0002 76.63; ⬍0.0001 49.11; ⬍0.0001
Interaction effects
Variety ⫻ leaf stage
CP 78-1628 ⫻ un 5.0 ⫾ 0.33 4.4 ⫾ 0.19b 3.7 ⫾ 0.14a 0.35 ⫾ 0.016a
CP 78-1628 ⫻ 3L 6.0 ⫾ 0.33 5.3 ⫾ 0.20a 3.6 ⫾ 0.13a 0.33 ⫾ 0.014a
CP 78-1628 ⫻ 5L 5.5 ⫾ 0.33 5.1 ⫾ 0.20a 3.3 ⫾ 0.13a 0.31 ⫾ 0.014a
CP 78-1628 ⫻ 7L 5.1 ⫾ 0.33 4.0 ⫾ 0.20c 2.0 ⫾ 0.13b 0.21 ⫾ 0.014b
F; P; df ⫽ 3, 56 1.82; 0.1534 9.80; ⬍0.0001 28.43; ⬍0.0001 14.38; ⬍0.0001
CP 88-1762 ⫻ un 5.0 ⫾ 0.41 4.3 ⫾ 0.19ab 3.6 ⫾ 0.15a 0.34 ⫾ 0.014a
CP 88-1762 ⫻3L 5.6 ⫾ 0.36 4.9 ⫾ 0.20a 3.1 ⫾ 0.13b 0.30 ⫾ 0.014ab
CP 88-1762 ⫻ 5L 5.1 ⫾ 0.36 4.3 ⫾ 0.20ab 3.0 ⫾ 0.13b 0.28 ⫾ 0.014b
CP 88-1762 ⫻ 7L 4.9 ⫾ 0.36 3.9 ⫾ 0.20b 2.1 ⫾ 0.13c 0.19 ⫾ 0.014c
F; P; df ⫽ 3, 56 0.62; 0.6043 4.20; 0.0095 17.00; ⬍0.0001 20.64; ⬍0.0001
CP 89-2143 ⫻ un 4.4 ⫾ 0.30a 4.0 ⫾ 0.24a 3.6 ⫾ 0.18a 0.31 ⫾ 0.018a
CP 89-2143 ⫻ 3L 3.1 ⫾ 0.36b 2.8 ⫾ 0.20b 2.2 ⫾ 0.13b 0.21 ⫾ 0.014b
CP 89-2143 ⫻ 5L 3.4 ⫾ 0.36b 3.0 ⫾ 0.20b 2.1 ⫾ 0.13b 0.19 ⫾ 0.014b
CP 89-2143 ⫻ 7L 3.4 ⫾ 0.36b 3.0 ⫾ 0.20b 2.0 ⫾ 0.13b 0.18 ⫾ 0.014b
F; P; df ⫽ 3, 56 3.80; 0.0150 5.09; 0.0035 17.91; ⬍0.0001 11.03; ⬍0.0001

Means separation tests (orthogonal contrasts, SAS Institute 2008) were performed where ANOVA F values (Proc GLM, SAS Institute 2008)
were signiÞcant at P ⱕ 0.10. Means followed by different letters in a column are signiÞcantly different. Uppercase letters indicate differences
among varieties (main effects). Lowercase letters indicate differences among leaf stages (main effects) and among leaf stages of the same variety
(interaction effects).
a
Number of secondary and tertiary shoots.
b
Number of stalks ⱖ1.5 m in height.
c
Total weight (kilograms) of millable stalks in each bucket.
d
Raw sugar weight (kilograms per bucket) calculated from brix and pol values by using the theoretically recoverable sugar method (Glaz
et al. 2002).

creases in the observed percentage of dead hearts and
symmetrical rows of holes between sample dates re-
sulted from the emergence of new tillers that in-
creased the number of undamaged shoots relative to
the number of damaged shoots. Larvae occasionally
cut through the unfurled leaves above the growth
point, resulting in initial dead heart symptoms. But
continued expansion of the undamaged leaf tissue
pushed out the dead leaf material nearby reverting
the shoot or tiller into the undamaged category at the
subsequent sample date. Decreases in the percentage
of dead plants between sample dates resulted from the
eruption of new tillers above the soil surface from
plants that showed no visible signs of life at the pre-
vious sample date.
Plant deaths caused by lesser cornstalk borer feed-
ing varied with sugarcane variety and leaf stage at
infestation. CP 89-2143 displayed greater susceptibil-
ity to lesser cornstalk borer damage than the other two
varieties. Damage differences among varieties may be
due to morphological, physiological, or biochemical
resistance in plants. Agarwal (1969) reported that the
physical and chemical make-up of sugarcane plants
greatly inßuences their protection from insect dam-
age. He reported that cell wall ligniÞcation and the
number of sclerenchymatous cell layers play an im-
portant role in imparting resistance against stem bor-
ers. Varietal differences in percentage lesser cornstalk
borer damage also were reported by Chang and Ota
(1989), with ⬍5% dead hearts in sugarcane varieties
H83-6818 and H83-7498 compared with 23% dead
hearts in H77-6359. Similar varietal differences have
been reported for damage caused by other pyralid
stem borers. Bessin et al. (1990) reported seasonal
differences in injury by sugarcane borer, Diatraea
saccharalis (F.) among tested sugarcane clones that
may have been due to resistance to stalk entry or
antibiosis (larval survival). Pfannenstiel and Meagher
(1991) reported signiÞcant differences among sugar-
cane varieties to injury by the Mexican rice borer,
Eoreuma loftini (Dyar).
Infestation at the three- and Þve-leaf stages resulted
in greater percentages of dead hearts and dead plants
than when infested at the seven-leaf stage. Infestation
at the seven-leaf stage resulted in a greater percentage
of plants with symmetrical rows of holes in the leaves.
This may be due to increasing stem thickness with
plant age that allowed fewer larvae to penetrate to the
stem core to cause dead hearts. In maize, Rojanaridi-
piched et al. (1984) found that the silica and lignin
April 2011 SANDHU ET AL.: SUGARCANE RESPONSE TO E. lignosellus
content of some varieties increased with age and were
major factors of resistance to stem borers at later plant
development stages. Greater Þber content has been
shown to generally increase D. saccharalis mortality in
sugarcane (Mathes and Charpentier 1962).
Tiller Production. The increased number of tillers
produced in 2009 compared with 2008 may have re-
sulted from different planting times. Sugarcane was
planted in January for the 2008 trial and in November
for the 2009 trials. Higher temperatures normally pres-
ent in southern Florida during November and De-
cember relative to January may have resulted in en-
hanced initial growth of the sugarcane plants.
The three varieties tested in this study produced
signiÞcantly different numbers of tillers in response to
feeding by lesser cornstalk borer at different leaf
stages. Although signiÞcantly more primary and sec-
ondary shoots of CP 78-1628 and CP 88-1762 were
killed when infested at the three- and Þve-leaf stages
compared with no shoot loss in the untreated control,
there was no difference in the number of tillers among
these treatments when sampled after 4 mo. This result
is probably due to increased tiller production in the
infested plants that compensated for the shoots killed
by lesser cornstalk borer during the Þrst 2 mo of
growth. Increased tiller production in infested plants
may be the result of changes in “growth-regulating
substances” in damaged primary shoots (Dillewijn
1952). Changes in sourceÐsink relationship due to
damaged primary shoots may also be responsible for
increased tiller production (Trumble et al. 1993).
They reported that the injury to rice stem tissue can
reduce the ability of a plant or tiller to capture light
and nutrients, resulting in increased availability of
these resources to neighboring plants or tillers. Lower
tiller production in plants infested late rather than
early may be due to lesser cornstalk borer larval dam-
age to both tillers and primary shoots of the seven-leaf
stage plants. In early-infested plants, only primary
shoots were damaged because tillers had not yet de-
veloped. Larvae completed their development by the
time of tiller emergence in early-infested leaf stages
allowing these tillers to escape injury. Tillers had al-
ready emerged by the time of the seven-leaf stage
infestation and both primary shoots and tillers were
available for borer damage. The greater percentage of
dead plants in CP 89-2143 probably reduced the po-
tential compensation from production of secondary
and tertiary shoots compared with the other two va-
rieties.
Compensation through increased tiller production
in response to damage also was observed by Carbonell
(1978), who reported the recovery of lesser cornstalk
borer-damaged sugarcane plants through production
of additional tillers. Similarly, Hall (1990) and Cherry
and Stansly (2009) reported that early stand loss due
to wireworm (Elateridae) damage was compensated
by increased tiller production during the sugarcane
growing season. Compensatory plant growth in re-
sponse to insect damage caused at early growth stages
also has been reported in many other Þeld crops. For
example, rice, Oryza sativa L., compensates for injury
481
at the vegetative stage by the stem borer Scirpophaga
incertulas (Walker) (Lepidoptera: Pyralidae) (Rubia
et al. 1990, 1996) by producing new vegetative (Soe-
jitno 1979, Tian 1981) and reproductive tillers (Luo
1987). They also indicated that 33% dead hearts in
30-d-old plants did not have a signiÞcant effect on
productive tillers or grain yield. Jiang and Cheng
(2003) reported that rice plants infested with Asiatic
rice borer, Chilo suppressalis (Walker) (Lepidoptera:
Crambidae), produced approximately one more tiller
than untreated control plants 2 wk after infestation.
Studies also have shown that low insect infestation
levels at early growth stages may increase plant yield
in Þeld beans, Phaseolus vulgaris L. (Banks and
Macaulay 1967); wheat (Triticum spp. L.) (Gouch
1947, Bardner 1968); potato, Solanum tuberosum L.
(Skuhravy 1968); and cotton (Gossypium spp. L.)
(Kincade et al. 1970).
Sugarcane Yield Traits. Biomass and sugar yields
were greater in 2009 than in 2008. This resulted from
increased tiller and millable stalk production in the
second year over the Þrst year of the study. Although
the mean number of millable stalks produced by the
untreated controls was less than or equivalent to the
number produced by CP 78-1628 and CP 88-1762 in-
fested at the three- and Þve-leaf stages, biomass yields
of untreated plants were greater than or equivalent to
infested plants. The greater biomass yield in the un-
treated controls than in the infested leaf stages was
due to greater millable stalk weight in the untreated
controls, because biomass yield was a product of the
number of millable stalks and stalk weight in each
bucket. This disparity in millable stalk weight among
treatments was probably due to proportionately fewer
primary shoots remaining and more tillers produced in
early-infested than in untreated control plants at har-
vest. Early research by Rodrigúes (1928) observed a
gradient in stalk weight and sugar content among
shoots with primary shoots having the greatest weight
and “richest juice” (i.e., greater brix values) followed
by secondary and then tertiary tillers.
Although all the varieties were infested with the
same number of lesser cornstalk borer larvae at each
infested leaf stage, there were signiÞcant differences
in the responses to plant feeding damage that varied
by leaf stage, variety and their interaction. Similar
levels of damage resulted in signiÞcant differences in
responses by variety. At the last sample date for plants
infested at the three-leaf stage over the two study
years, 60 Ð 64% dead hearts resulted in no signiÞcant
yield loss in CP 78-1628, whereas 66 Ð 68% dead hearts
resulted in a 11Ð18% yield loss in CP 88-1762, and
74 Ð 88% dead hearts resulted in a 25Ð38% yield loss in
CP 89-2143. Across the three varieties tested, the re-
sults of this trial indicated a 14 Ð16% loss in yield when
attacked at the three-leaf stage with a 35% increase in
loss between the three- and Þve-leaf stages and an-
other 100% increase in loss between the Þve- and
seven-leaf stages. Based on these Þgures, plant cane
producers would be warranted to apply an effective
control strategy at the Þrst sign of dead hearts or dead
plants to prevent increasingly signiÞcant yield losses
482 JOURNAL OF ECONOMIC ENTOMOLOGY
up to at least 8 wk postemergence. Likewise, scouting
efforts should increase as the plants age, because yield
losses increase as the age at Þrst attack increases, at
least to the seven-leaf stage. The varieties CP 78-1628
and CP 88-1762 compensated for plant damage by
increasing tiller production above that of the un-
treated control when infested at the three- and Þve-
leaf stages. This trend was carried into the number of
millable stalks for these two varieties with a net in-
crease compared with the untreated control when
attacked at the three- and Þve-leaf stages. It is not clear
whether CP 89-2143 compensated for early season
damage, because tiller production was reduced by
18 Ð29% and millable stalk production was reduced by
28 Ð30% when infested at the three-leaf stage com-
pared with the untreated control. Relative increases in
percent yield loss compared with the untreated con-
trol were dependent on both variety and leaf stage at
Þrst attack. When CP 78-1628 was attacked at the
three-leaf stage, it compensated for the attack by pro-
ducing roughly equivalent levels of biomass and sugar
yield compared with the untreated control. However,
later attacks resulted in subsequently greater increases
in percentage yield loss at each step compared with
the other two varieties. Yield loss increased by 50 Ð
100% between the three- and Þve-leaf stages and by
250 Ð280% between the Þve- and seven-leaf stages for
CP 78-1628. Overall yield loss was greater for CP
88-1762 than for CP78-1628, but yield loss increased by
only 25Ð50% between the three- and Þve-leaf stages
and by 100 Ð125% between the Þve- and seven-leaf
stages. Overall yield loss was the greatest for CP 89-
2143, but yield loss increased by only 17Ð18% between
the three- and Þve-leaf stages and by 10 Ð35% between
the Þve- and seven-leaf stages. Therefore, both the
variety and age of the plants should have a signiÞcant
affect on control recommendations. For a variety with
a response to damage similar to CP 78-1628, our Þgures
suggest that the producer could effectively wait until
the Þrst signs of infestation occurred before initiation
of effective treatments without a signiÞcant yield loss.
In contrast, producers with varieties similar to the
apparent sensitivity of CP 89-2143 would probably
need an early warning system based on a combination
of pheromone traps and degree-day models for sug-
arcane growth and insect population development to
forecast and prepare for treatments to prevent dead
hearts and dead plants caused by lesser cornstalk
borer. Arceneaux (1935) demonstrated that compen-
sation for mechanical damage also varied by sugarcane
variety with compensatory growth greatest for the
variety Co 290 and least for POJ 234. Variations in
compensatory biomass yield responses of sugarcane
with plant age at the time of mechanical shoot removal
also were reported by White and Richard (1985).
They manually removed 25, 50, and 75% of randomly
selected sugarcane shoots (at 2Ð5 cm below soil level)
in mid-March, mid-April, and mid-May to determine
the effect of this shoot removal on biomass yield. They
reported that the reductions in sugar yields tended to
increase with the delay of shoot removal date during
the crop season.
Vol. 104, no. 2
In summary, the lesser cornstalk borerÕs ability to
cause damage to sugarcane was dependent on variety
and time of infestation. Equal numbers of lesser corn-
stalk borer larvae produced more dead hearts and
dead plants in CP 89-2143 than in CP 78-1628 and CP
88-1762. Infestation of plants at the three- and Þve-leaf
stages produced more dead hearts and dead plants
than infestation at the seven-leaf stage. Tiller counts
were greater in early damaged plants than late dam-
aged plants, because E. lignosellus larvae damaged
more tillers in late-infested plants that were not pres-
ent in early-infested plants. Early-infested plants re-
sulted in more yield response than late-infested plants
in CP 78-1628 and in CP 88-1743 with CP 78-1628
demonstrating the greatest ability to compensate for
lesser cornstalk borer damage among the three tested
varieties. The above-mentioned conclusions were
made based on controlled experiments conducted in
a greenhouse. In Þeld conditions, many climatic (sun-
light, temperature, moisture) and biological factors
(insect population, other pests, natural enemies)
could affect the results. Field trials will be conducted
in to determine how season-long exposure to lesser
cornstalk borers and insect population limiting factors
(e.g., soil moisture and high summer soil surface tem-
peratures) may affect damage throughout the season
and resulting yields.
Acknowledgments
We thank K. Pernezny (EREC, Belle Glade, FL) for pro-
viding laboratory facilities for diet preparation. Moth collec-
tion for colony established was assisted by N. Larsen and B.
Thappa (EREC). Statistical assistance was provided by M.
Brennan and J. Colee (University of Florida, Gainesville,
FL), and Drs. M. Josan and T. Lang (EREC). Sandeep
Sandhu (Belle Glade, FL) was instrumental in experimental
setup and support. Funding for this study and the land for
collecting insects and sugarcane were provided by the Sugar
Cane Growers Cooperative of Florida and the United States
Sugar Corporation. This work was made possible by student
Þnancial support from the Wedgworth Graduate Fellowship
and the University of Florida College of Liberal Arts and
Sciences.
References Cited
Agarwal, R. A. 1969. Morphological characteristics of sug-
arcane and insect resistance. Entomol. Exp. Appl. 12:
767Ð776.
Arceneaux, G. 1935. Studies of gaps in sugarcane rows and
their effect on yield under Louisiana conditions. U.S.
Dep. Agric. Circ. 521: 1Ð20.
Banks, C. J., and E.D.M. Macaulay. 1967. Effects of Aphis
fabae Stop. and of its attendant ants and insect predators
on yields of Þeld beans (Vicia fabae L.). Ann. Appl. Biol.
60: 445Ð153.
Bardner, R. 1968. Wheat bulb ßy Leptohylemyia coarctata
Fall., and its effect on the growth and yield of wheat. Ann.
Appl. Biol. 61: 1Ð11.
Bardner, R., and K. E. Fletcher. 1974. Insect infestations
and their effects on the growth and yield of Þeld crops:
a review. Bull. Entomol. Res. 64: 141Ð160.
Bessin, R. T., T. E. Reagen, and F. A. Martin. 1990. A moth
production index for evaluating sugarcane cultivars for
April 2011 SANDHU ET AL.: SUGARCANE RESPONSE TO E. lignosellus
resistance to the sugarcane borer (Lepidoptera: Pyrali-
dae). J. Econ. Entomol. 83: 221Ð225.
Carbonell, E.E.T. 1978. Descripcion de danos causados por
Elasmopalpus lignosellus (Zeller) en cana de azucar y de
algunos de sus controladores biologicos. Saccharum 6:
118 Ð145.
Chang, V., and A. K. Ota. 1989. The lesser cornstalk borer
and its control, pp. 34 Ð37. In Annual Report of the Hawaii
Agricultural Experiment Station, University of Hawaii,
Honolulu, HI.
Cherry, R., and P. Stansly. 2009. Impact on yield of wire-
worm (Coleoptera: Elateridae) populations in Florida
sugarcane at planting. J. Am. Soc. Sugar Cane Technol. 29:
137Ð148.
Demandt, E. 1929. De resultaten van de uitsnijproeven; een
bijdrage tot het compensatie vraagstuk in den rietaan-
plant. Meded. Proefst. Java-Suikerind. 1929: 717Ð780.
Dillewijn, C. V. 1952. Botany of sugarcane. The Chronica
Botanica Co., Waltham, MA.
Falloon, T. 1974. Parasitoids and predators of the lesser
cornstalk borer, Elasmopalpus lignosellus (Zeller), on a
Jamaican sugar estate. M.S. Thesis, University of Florida,
Gainesville.
Gilbert, R. A., D. R. Morris, C. R. Rainbolt, J. M. McCray,
R. E. Perdomo, B. Eiland, G. Powell, and G. Montes.
2008. Sugarcane response to mill mud, fertilizer, and soy-
bean nutrient sources on a sandy soil. Agron. J. 100:
845Ð 854.
Glaz, B., S. J. Edme, J. D. Miller, S. B. Milligan, and D. G.
Holder. 2002. Sugarcane response to high summer water
table in the Everglades. Agron. J. 94: 624 Ð 629.
Gouch, H. C. 1947. Studies on wheat bulb ßy (Leptohylemia
coarctuta Fall.) numbers in relation to crop damage. Bull.
Entomol. Res. 37: 439.
Hall, D. 1990. Stand and yield losses in sugarcane caused by
the wireworm Melanotus communis (Coleoptera: Elateri-
dae) infesting plant cane in Florida. Fla. Entomol. 73:
298 Ð302.
Jiang, M. X., and J. A. Cheng. 2003. Interactions between the
striped stem borer Chilo suppressalis (Walk.) (Lepidop-
tera: Pyralidae) larvae and rice plants in response to
nitrogen fertilization. Anz. Schadlingskd. 76: 124 Ð128.
Kincade, R. T., M. L. Laster, and J. R. Brazzel. 1970. Effect
on cotton yield of various levels of simulated Heliothis
damage to squares and bolls. J. Econ. Entomol. 63: 613Ð
615.
Littell, R. C., P. R. Henry, and C. B. Ammerman. 1998.
Statistical analysis of repeated measures data using SAS
procedures. J. Anim. Sci. 76: 1216 Ð1231.
Loh, C. S., Y. G. Wen, and B. W. Shee. 1948. The effects of
topping on young canes. J. Sugar Cane Res. 2: 1Ð23.
Luo, S. 1987. Studies on the compensation of rice to the
larval damage caused by the Asian rice borer Chilo sup-
pressalis (Walker). Sci. Agric. Sin. 20: 67Ð72.
Mathes, R., and L. J. Charpentier. 1962. Some techniques
and observations in studying the resistance of sugarcane
varieties to the sugarcane borer in Louisiana. Proc. Int.
Soc Sugarcane Technol. 11: 594 Ð 604.
Painter, R. H. 1951. Insect resistance in crop plants. Uni-
versity of Kansas Press, Lawrence, KS.
483
Pedigo, L. P. 1991. Entomo1ogy and Pest Management.
Macmillan, New York.
Pfannenstiel, R. S., and R. L. Meagher, Jr. 1991. Sugarcane
resistance to stalkborers (Lepidoptera: Pyralidae) in
South Texas. Fla. Entomol. 74: 300 Ð305.
Rice, R., L. Baucum, and B. Glaz. 2009. Sugarcane variety
census: Florida 2008. Sugar J. 72: 6 Ð12.
Rodrigúes, H. J. 1928. Relation between high sucrose in
cane and suckering. Plant Sugar Manuf. 80: 101Ð102.
Rojanaridipiched, C., V. E. Gracen, H. L. Everett, J. G. Coors,
B. F. Pugh, and P. Bouthyette. 1984. Multiple factor re-
sistance in maize to European corn borer. Maydica 29:
305Ð315.
Rubia, E. G., B. M. Shepard, E. B. Yambao, K. T. Ingram, G. S.
Arida, and F.W.T. Penning de Vries. 1990. Stem borer
damage and grain yield of ßooded rice. J. Plant Prot.
Tropics 6: 205Ð211.
Rubia, E. G., K. L. Heong, M. Zalucki, B. Gonzales, and G. A.
Norton. 1996. Mechanism of compensation of rice plants
to yellow stem borer Scirpophaga incertulas (Walker)
injury. Crop Prot. 15: 335Ð340.
Sandhu, H. S., G. S. Nuessly, S. E. Webb, R. H. Cherry, and
R. A. Gilbert. 2010. Temperature-dependent develop-
ment of lesser cornstalk borer, Elasmopalpus lignosellus
(Lepidoptera: Pyralidae) on sugarcane under laboratory
conditions. Environ. Entomol. 39: 1012Ð1020.
SAS Institute. 2008. PROC userÕs manual, version 9th ed.
SAS Institute, Cary, NC.
Schaaf, A. C. 1974. A survey of the damage caused by
Elasmopalpus lignosellus (Zeller) (Lepidoptera: Phy-
citidae) to sugarcane in Jamaica, pp. 488 Ð 497. In Pro-
ceedings, 15th Congress of International Society of
Sugar Cane Technologists, 13Ð29 June 1974, Durban,
South Africa. Executive Committee of the ISSCT,
Quatre-Bornes, Mauritius.
Shapiro, S. S., and M. B. Wilk. 1965. An analysis of variance
test for normality (complete samples). Biometrika 52:
591Ð 611.
Skuhravy, V. 1968. Einßuss der entblatterung und des kart-
offel-kaferfrasses auf die kartoffelernte. Anz. Schadling-
skd. 41: 180 Ð 88.
Soejitno, J. 1979. Some notes on the larval behavior of
Tryporyza incertulas (Walker) (Lepidoptera: Pyrali-
dae). In Kongress Entomologi I, 9 Ð11 January 1979,
Jakarta, Indonesia. (http://agris.fao.org/agris-search/
search/display.do?f⫽1981/XB/XB81019.xml;XB8100078).
Tian, C. T. 1981. Reasons for the ßuctuation in populations
of Schoenobius incertulas (Walker). Yunnan Agric. Sci.
Technol. 3: 29 Ð34.
Trumble, J. T., D. M. Kolodny-Hirsch, and I. P. Ting. 1993.
Plant compensation for arthropod herbivory. Annu. Rev.
Entomol. 38: 93Ð119.
[USDA] U.S. Department of Agriculture. 2008. National
Agricultural Statistics Services. (www.nass.usda.gov).
White, W. H., and E. P. Richard, Jr. 1985. Sugarcane recov-
ery from spring stand losses associated with simulated
insect feeding as inßuenced by soil-applied herbicides.
Crop Prot. 14: 483Ð 489.
Received 10 May 2010; accepted 22 December 2010.