II.12
Bait Acceptance by Different Grasshopper Species and Instars
Jerome A. Onsager, R. Nelson Foster, and Larry Jech
Data Collection
Relative
Susceptibility of Different Species
Relative
Susceptibility of Different Developmental Stages
Relative
Susceptibility of Different-Aged Populations
Summary
and Recommendations
References
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The Grasshopper Integrated Pest Management (GHIPM) Project provided
unique resources and opportunities that allowed investigators to
gather a large amount of data on the responses of rangeland grasshoppers
to carbaryl bait. A total of 39 different species were recorded
in 24 different control experiments at 14 different sites in the
western parts of North Dakota and South Dakota. All species were
not present in sufficient numbers to provide useful information,
but the data base allowed GHIPM-funded investigators to study many
questions that could not have been examined without it.
Data
Collection
The monitoring procedure was to establish from 4 to 10 monitoring
sites, each consisting of 40 0.1m2 rings spaced about 5
m apart in circles, both in plots that were scheduled for treatment
and in adjacent plots that remained untreated. Density counts and
sweep-net collections were made as close as possible (usually 24
hours) before scheduled treatments, and again as close as possible
to 48 hours after treatment. The information from all sample sites
per plot for each sampling date was then combined for further study.
Each sweep sample was examined to determine the species and stage
of development for every grasshopper in the sample. Each total density
count was then converted to density per instar per species by multiplying
observed total density times the appropriate proportions of composition
within the sweep samples. The procedure is identical to that described
in chapter II.2, Evaluation of
Rangeland Grasshopper Controls, except that density was estimated
for each instar of a species as well as for all individuals of a
species.
Computer tabulations of different species recorded in different
experiments revealed a potential for 253 independent determinations
of species-specific response to carbaryl bait. Pretreatment and
posttreatment data for each species in each experiment were then
examined to assess which of the possible determinations would be
meaningful. A total of 101 potential data sets were declared useless,
leaving 152 legitimate determinations.
Reasons for rejecting some data sets included initial presence
in such low density that subsequent reduction would not be measurable
(in most cases, at least five specimens in pretreatment samples
were required), absence of specimens at untreated sample sites (which
prohibited estimation of mortality in the absence of treatment),
and higher estimated mortality in untreated plots than in treated
plots (a common artifact of sampling error among low-density samples).
The 152 data sets accepted as legitimate provided opportunities
to study a variety of questions about response to carbaryl bait.
The simplest assessment concerned the average percent control among
all individuals of a species. This average percent control was calculated
with a variation of the formula by Connin and Kuitert (1952):
Percent control = 100(1-(Ta xUb / Tb / Ua)), where
Tb is density in treated plots before treatment,
Ta is density in treated plots after treatment,
Ub is density in untreated plots before treatment, and
Ua is density in untreated plots after treatment.
The formula does not yield simple or raw control data-that is,
the percentage of the total infestation that disappeared in treated
plots. Rather, it yields adjusted control data: the percentage of
the total infestation that most likely was killed by carbaryl bait.
The formula is useful for two major reasons. First, grasshopper
infestations suffer some mortality each day due to natural causes,
so the formula removes that natural mortality from consideration.
The formula essentially uses data from untreated sites to estimate
what the post-treatment counts at treated sites would have been
in the absence of treatment. Percent control then represents the
difference (if any) between expected and observed post-treatment
density in treated plots. Second, without the formula, the percent
control that is estimated will be grossly different, depending on
how much time elapses between pretreatment and posttreatment counts.
These problems can be illustrated with an example.
Let us assume that an infestation of 30 grasshoppers/yd2 comprises
6 Aeropedellus clavatus, 15 Melanoplus sanguinipes, and
9 Amphitornus coloradus. We decide to treat half and leave
half, and we sample both halves on the day before treatment (day-1),
and on days 2, 3, 4, and 5 after treatment. Table II.12-1 shows
typical density data.
Table II.12-1-A representative example of typical grasshopper
density data in untreated plots versus plots that were treated (on
day zero) with carbaryl bait
|
Time
(days after treatment)
|
A. clavatus
|
M. sanguinipes
|
A. coloradus
|
All species
|
|
Untreated plot
|
Treated plot
|
Untreated plot
|
Treated plot
|
Untreated plot
|
Treated plot
|
Untreated plot
|
Treated plot
|
|
|
|
-1
|
6
|
6
|
15
|
15
|
9
|
9
|
30
|
30
|
|
+2
|
3.68
|
2.95
|
13.69
|
3.42
|
7.71
|
7.56
|
25.08
|
13.93
|
|
+3
|
3.13
|
2.51
|
13.28
|
3.32
|
7.33
|
7.18
|
23.74
|
13.01
|
|
+4
|
2.66
|
2.13
|
12.88
|
3.22
|
6.96
|
6.82
|
22.5
|
12.17
|
|
+5
|
2.26
|
1.81
|
12.49
|
3.12
|
6.61
|
6.48
|
21.36
|
11.41
|
Looking only at the raw density for All species in only
the treated plot, a reader might believe that this bait treatment
achieved about 54- to 62-percent average control of the infestation.
The fallacy is that if a similar strategy is applied to data from
untreated plots, a reader could estimate 16- to 29-percent control
where nothing was done. Use of the formula yields more conservative
and more realistic estimates of about 44- to 46-percent adjusted
control of all species.
Raw estimates for individual species can also be very misleading.
For example, A. clavatus usually is the first species that
hatches in the spring. By the time of typical bait treatments to
control later-hatching major pest species, A. clavatus often
is present as very old adults that suffer very high daily mortalities
likely associated with the process of aging. Raw estimates indicate
51- to 70-percent population reduction, but adjusted estimates reveal
only 20-percent control due to the bait, meaning the raw estimates
placed control at 2.5 to 3.5 times higher than it actually was.
Notice in the example that discrepancies between raw and adjusted
mortalities for A. coloradus are even greater than they were
for A. clavatus. This is because adjusted response to treatment
(2-percent control) was less than the daily loss due to natural
mortality (5 percent per day). In such a case, raw estimates yield
greatly distorted results. As one might then expect, raw estimates
are closest to adjusted estimates in cases like the M. sanguinipes
example, where natural mortality was relatively low (3 percent
per day) and adjusted control was relatively high (75 percent).
Nevertheless, it should be noted that all raw estimates for M.
sanguinipes still were too high, and the degree of error increased
as the amount of time between pretreatment and posttreatment samples
was increased. Similar errors are guaranteed to occur in real life
(in field experiments or commercial control projects) if natural
mortality is ignored.
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Relative
Susceptibility of Different Species
The results of GHIPM experiments were combined with a number of
previous studies by the authors and others (see Swain [1986] and
Quinn et al. [1989]) to produce table II.12-2. It divides grasshoppers
into three broad classes of susceptibility. The sensitive class
contains species that readily seek out and eat wheat bran bait and
therefore usually suffer a high degree (average = 56-87 percent)
of adjusted (true) mortality. The vulnerable class contains species
that usually either suffer only a moderate degree (30-55 percent)
of adjusted mortality or else exhibit such great variation among
different tests that one cannot safely depend on more than moderate
results. The nonsusceptible class (less than 30-percent adjusted
mortality) contains species that eat little or no bait and therefore
usually are not markedly affected by bait.
Most of the experiments that contributed to table II.12-2 were
applied when the majority of target pest grasshopper species were
in third, fourth, or fifth instars. A few very early species like
A. clavatus and M. confusus typically were treated
as adults or fifth instars, while some relatively late species like
P. nebrascensis and P. quadrimaculatum were occasionally
treated as first or second instars where they were incidental rather
than primary target species.
Table II.12-2-Classification of grasshopper
species according to susceptibility to carbaryl wheat bran bait
|
Class and expected levels of control
|
Species
|
|
|
|
Sensitive (>55-% control)
|
Ageneotettix deorum
|
|
Anabrus simplex
|
|
Control is expected to average about 70%. Worst-case and
best-case scenarios will be about 55% and 85%, respectively.
|
Aulocara elliotti
|
|
Camnula pellucida
|
|
Hadrotettix trifasciatus
|
|
*Melanoplus bivittatus
|
|
Melanoplus confuses
|
|
Melanoplus dawsoni
|
|
Melanoplus foedus
|
|
*Melanoplus infantilis
|
|
*Melanoplus occidentalis
|
|
*Melanoplus packardii
|
|
Melanoplus sanguinipes
|
|
Spharagemon equale
|
|
Stenobothrus brunneus
|
|
*Mermiria bivittata
|
|
|
|
Vulnerable (30- to 55-% control)
|
*Aulocara femoratum
|
|
Eritettix simplex
|
|
Control is expected to average about 42%. Worst-case
and best-case scenarios will be about 12% and 72%, respectively.
|
Melanoplus femurrubrum
|
|
Oedaloenotus enigma
|
|
Opeia obscura
|
|
Phoetaliotes nebrascensis
|
|
Psoloessa delicatula
|
|
|
|
Nonsusceptible (<30-% control)
|
Aeropedellus clavatus
|
|
Amphitornus coloradus
|
|
Control is expected to average about 15%. Worst-case
and best-case scenarios will be about 0% and 30%, respectively.
|
Cordillacris crenulata
|
|
Cordallacris occipitalis
|
|
Hesperotettix viridis
|
|
Metator pardalinus
|
|
*Phlibostroma quadrimaculatum
|
|
Trachyrhachys kiowa
|
|
|
|
*These species are not likely to suffer best-case
scenario levels of control.
|
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Relative
Susceptibility of Different Developmental Stages
Some of the GHIPM experiments provided data that allowed the comparison
of the relative susceptibility of different instars of a species
to bait. In general, the requirements for a meaningful test were
the presence of at least four or more different stages in reasonable
numbers (usually at least five individuals per instar in pretreatment
sweep samples) in two or more different experiments. In those cases,
the authors calculated adjusted percent control for each instar
and used analyses of covariance, with instar as the covariant, to
test susceptibility by instar. When covariance was significant (when
percent control was affected by instar), the slope of the relationship
indicated whether larger or smaller instars were most susceptible.
A total of eight species were tested, six of which were considered
in table II.12-2 to be sensitive. Younger instars of three species,
A. deorum, M. packardii, and M. sanguinipes, were
found to be significantly more susceptible to bait than older instars.
Susceptibility was not affected by instar in the cases of A.
elliotti, C. pellucida, M. infantilis, P. nebrascensis, or T.
kiowa.
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Relative
Susceptibility of Different-Aged Populations
Some of the GHIPM experiments provided data that allowed the researchers
to examine the effect of age on susceptibility of populations to
bait. Age was expressed as average instar, which is calculated as
the sum of each instar number multiplied by the number of grasshoppers
in the instar (adults are considered instar 6 for this procedure)
divided by the total number of grasshoppers present. The requirements
for a meaningful test were significant adjusted control observed
in three or more experiments (incidences of zero control were excluded
from these calculations). The relationship between average instar
and percent adjusted mortality was examined by linear regression
techniques.
A total of 17 species was tested, 10 of which were considered in
table II.12-2 to be sensitive or vulnerable. For three of those
species, A. elliotti, A. deorum, and M. sanguinipes, percent
adjusted control increased significantly with average instar.
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Summary
and Recommendations
Grasshopper species vary considerably in their inclination to feed
on wheat bran and in their susceptibility to carbaryl-treated bait.
In addition, levels of control that follow bait treatments are considerably
lower and much less predictable than control achieved with liquid
sprays. The GHIPM Project greatly increased the knowledge base for
both acknowledged pest grasshopper species (the primary target species)
and for incidental (nontarget) species. Project researchers now
feel that they can offer some general guidelines, based on species
susceptibility (table II.12-2),
for the appropriate use of carbaryl bait.
Individuals should not attempt to control nonsusceptible pest species
with bait. If such species comprise a significant proportion of
an infestation, a conservative manager should simply assume that
bait will give no control of that proportion. Vulnerable species
may or may not be markedly controlled by baits, but what regulates
that degree of success remains unknown, and at this time those results
cannot be predicted. Past situations have documented dramatic reductions
in vulnerable species from the use of bait, as well as cases of
almost total failure. In the future, managers should not use bait
against vulnerable species without seriously weighing the consequences
of failure. Control of the sensitive species with bait is generally
reliable.
Questions about optimum timing for bait treatments remain somewhat
perplexing, but it fortunately appears that timing is not of extreme
importance, perhaps because of compensatory factors. Some tests
support early treatments in that, at least for some species, younger
instars were more susceptible than older instars. This is logical
because smaller grasshoppers are killed by smaller doses of toxicant.
Another advantage of early bait treatment is that natural control
agents have more time to act upon surviving grasshoppers.
Other tests, however, support late treatments in that total percent
control was greater for older populations than for younger populations.
While these results may seem contrary, they also can be considered
strong evidence that something like changes in behavioral traits
(perhaps searching capabilities) or habitat characteristics (perhaps
cover, litter, or bare ground) make baits more accessible as the
season progresses. If such compensating factors exist, the mechanisms
cannot be accurately described at the present time. Fortunately,
however, for most species (14 of 17 tested), adjusted percent control
was not markedly affected by population age. It therefore appears
that timing of bait treatments is not of extreme importance as long
as it occurs when most of the primary target grasshoppers are in
third, fourth, or fifth instars.
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References
Cited
Connin, R. V.; Kuitert, L. C. 1952. Control of
the American grasshopper with organic insecticides in Florida. Journal
of Economic Entomology 45: 684-687.
Quinn, M. A.; Kepner, R. L.; Walgenbach, D. D.;
Bohls, R. A.; Pooler, P. D. 1989. Immediate and 2nd-year effects
of insecticide spray and bait treatments on populations of rangeland
grasshoppers. Canadian Entomologist 121: 589-602.
Swain, J. L. 1986. Effect of chemical grasshopper
controls on non-target arthropods of rangeland in Chaves County,
New Mexico. Master's thesis. Las Cruces, NM: New Mexico State University.
102 p.
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