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Relating feeding rates to sex and size in six species of grasshoppers (Orthoptera: Acrididae)

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Abstract

Nymphs and adults of both sexes of six species of grasshoppers, Camnula pellucida, Dissosteira carolina, Melanoplus infantilis, M. sanguinipes, M. packardii and M. bivittatus, were fed wheat seedlings to determine daily feeding rates. The relationships between the feeding rates and five indices of body size were checked with five types of equations — power, parabolic, exponential, linear, and logarithmic. For combined data for all species, the power equation (y = axb) produced the best correlations with the feeding rate being proportional to the 1.56 power of pronotum length, 1.87 of femur length, 1.94 of body length, 0.72 of live weight, and 0.66 of dry weight. The power equation underestimated nymphal feeding and overestimated adult feeding. Better correlations were obtained when separate power equations were used for nymphs and adults. While the b parameters for the nymph equations did not differ significantly from those for the adult equations, the a parameters for nymphs were much larger, reflecting that nymphs ate more relative to their size than adults. For individual species, the parabolic equation (y = a + bx + cx2) was as good or sometimes better than the power equation. Generally, ranking the indices of body size by their coefficients of determination (r2 values) produced the following hierarchy: pronotum length (best), femur length, live weight, dry weight, and body length (worst). However, as the differences between the best and worst indices were minor (usually less than 10%) and as live weight and body length are easy to determine, we recommend the use of either of these two indices for estimating feeding rates.
RELATING FEEDING RATES TO SEX AND SIZE IN SIX SPECIES OF
GRASSHOPPERS (ORTHOPTERA: ACRIDIDAE)I
ROBERT G. HOLMBERG
Athabasca University, Edmonton. Alberta T5L 2W4
JOHN MICHAEL HARDMAN2
Research Station, Agriculture Canada. Lethbridge, Alberta TIJ 4B I
Abstract Can. Em. 116:597-606 (1984)
Nymphs and adults of both sexes of six species of grasshoppers. Camnula pellucida,
Dissosleira carolina, Melanoplus infantilis. M. sanguinipes, M. packardii and M. biv-
illalus, were fed wheat seedlings to determine daily feeding rates. The relationships
between the feeding rates and five indices of body size were checked with five types
of equations -power, parabolic, exponential. linear, and logarithmic. For combined
data for all species. the power equation (y =axb) produced the best correlations with
the feeding rate being proportional to the 1.56 power of pronotum length, 1.87 of
femur length, 1.94 of body length, 0.72 of live weight, and 0.66 of dry weight. The
power equation underestimated nymphal feeding and overestimated adult feeding. Bet-
ter correlations were obtained when separate power equations were used for nymphs
and adults. While the bparameters for the nymph equations did not differ significantly
from those for the adult equations, the aparameters for nymphs were much larger,
reflecting that nymphs ate more relative to their size than adults. For individual species,
the parabolic equation (y =a+bx +cx2)was as good or sometimes better than the
power equation. Generally, ranking the indices of body size by their coefficients of
determination (r values) produced the following hierarchy: pronotum length (best),
femur length, live weight, dry weight, and body length (worst). However, as the dif-
ferences between the best and worst indices were minor (usually less than 10%) and
as live weight and body length are easy to determine, we recommend the use of either
of these two indices for estimating feeding rates.
Resume
On a determin~ Ie taux d'alimentation des nymphes et des imagos des deux sexes de
six es~ces de sauterelles (Camnula pellucida, Dissosleira carolina. Melanoplus in-
fantilis, M. sanguinipes. M. packardii et M, bivitlalus) nourris de pousses de bl~. On
a ensuite v~rifie la relation entre Ie taux d'alimentation et cinq indices de la taille
corporelle au moyen de cinq fonctions: puissance, parabolique, exponentielle, lineaire
et logarithmique. La fonction puissance (v =axb) donne la meilleur correlation pour
les donnres de toutes les es~es, Ie taux d'alimentation etant proportionnel it la lon-
gueur du pronotum it la puissance 1.56, it la longueur du femur it la puissance 1.87,
it la longueur du corps it la puissance 1.94, au poids vif it la puissance 0.72 et au poids
sec it la puissance 0.66, Cette equation sousestime Ie taux d'alimentation des nymphes
et surestime celui de I'inseete adulte. Ainsi, on obtient de meilleures correlations en
utilisarit des ~uations distinctes pour ces deux stades. Si Ie parametre best pratique-
ment identique dans les deux ~uations, Ie parametre aest beaucoup plus grand dans
celie des nymphes, ces demieres mangeant plus en fonction de leur taille que I'adulte.
Chaque espece prise individuellement, la fonction parabolique (y =a+bx +cx2)
donne d'aussi bons, sinon meilleurs resultats que la fonction puissance. En regIe ge-
n~raIe, Ie classement des indices de taille selon Ie coefficient de determination (valeur
r) est Ie suivant: longueurdu pronotum (meilleur),longueurdu femur, poids vif, poids
see et longueur du corps (pire). Toutefois, puisque la difference entre les deux extremes
est relativement faible (habituellement moins de 10%), et puisque Ie poids vif et la
taille du corps sont les plus facHes it d~terminer, on recommande I'emploi de ces deux
indices pour ~valuer Ie taux d'alimentation.
IConlribution No. 1793 of the KenlVilIe Research Station.
2Presenl ad~: Research Station, Agriculture Canada, Kenlville, Nova Scotia 84N US. '1
597
598 THE CANADIAN ENTOMOLOGIST April 1984
Introduction
Grasshoppers are notorious for reducing productivity of rangeland and planted crops
(Hewitt 1977; Smith and Holmes 1977). However, before managers can decide whether
grasshopper control is needed, they must be able to estimate how much damage would
occur without human intervention. Though the amount of grasshopper damage is affected
by variables such as weather; kinds and densities of vegetation, grasshoppers, predators
and competitors; and types of control measures used (Hewitt 1977), the feeding rates of
the grasshoppers themselves is of primary importance. Unfortunately, determining the
feeding rate of each "kind" of grasshopper would seem to be a major task because feeding
rates vary with species, stage of development, age and sex (Parker 1930). Also, many
species of grasshoppers attack economically important plants (Beirne 1972; Brooks 1958)
and many species may feed at one site (Hardman and Smoliak 1980). An easier approach
is using the relationship between feeding rate and body size as has been demonstrated in
other arthropods (Bertalanffy 1957; Reichle 1968).
We determined the weight of wheat eaten by different developmental stages and both
sexes of six species of grasshoppers. We then examined the relationship between feeding
rates and five indices of grasshopper size, namely pronotum, femur and whole body lengths
and live and oven-dried body weights.
Materials and Methods
Grasshoppers. The species of grasshoppers used (ranked from smallest to largest) Were:
Melanoplus infantilis Scudder, Camnula pelludda (Scudder), M. sanguinipes (Fabr.), M.
packardU Scudder, M. bivillatus (Say), and Dissosteira carolina (L.). Eggs were collected
from the field or obtained from field-collected specimens. Collections were made within
60 km of Lethbridge, Alberta, and with the exception of M. infantilis, all came from the
edges of cereal fields. The parents of M. infantilis came from Stipa-Bouteloua prairie.
Eggs were incubated in moist sand at 22°C for about 1month and then, to break diapause,
stored at 3°C for at least 2 months. When required, eggs were rinsed for I min in a 1%
bleach solution (to destroy most pathogens), placed in a dish of moist sand and allowed
to develop in wooden cages (Hunter-Jones 1961).
Except for M. infantiUs, all cages had a single 40 W incandescent light bulb that
raised the air temperature from 25 to 37°C (measured below the bulb). For M. infantilis,
25 W bulbs were used. These bulbs raised the air temperature tQ 32°C. The incandescent
lights of the cages and the fluorescent ceiling lights of the rearing room were turned on
at 6 a.m. and off at 10 p.m. each day.
Young nymphs were fed leaf lettuce; older nymphs and adults, head lettuce. All
grasshoppers were also supplied a dried powdered food mixture that consisted, on a weight
basis, of: 68% alfalfa, 23% skim milk, 4.5% brewer's yeast, and 4.5% egg yolk. Dental
rolls inserted in small plastic containers provided water.
Wheat. The food of the grasshoppers in th,; feeding trials consisted ofIeaves and a 1-2 cm
portion of the stem of two-or three-leaf stage wheat seedlings, Triticum aestivum L. var.
Neepawa. The plants were grown in 15 cm plastic pots of Cornell mix [comprised of 2
parts sphagnum moss: 2 parts vermiculite: 1 part sand plus CaCOJo fast release fertilizer
(0-20-0) and slow release fertilizer (18-6-12 plus trace elements)]. The pots, planted with
80 seeds each, were placed at 25°C in a cabinet provided with 15 h of fluorescent and
incandescent light and watered each day. The seedlings were ready for use within 9-18
days.
Feeding trials. Feeding rates were based on individual grasshoppers that were tested
within 3 days of their most recent moult. For the feeding trials, each grasshopper was
weighed and placed in a container along with a weighed amount of freshly cut wheat. For
nymphs smaller than 1 cm, the containers consisted of 90 mL paper cups that were closed
Volume 116 THE CANADIAN ENTOMOLOGIST 599
with small clear Petri dishes. Cylindrical I L cardboard containers, closed with muslin,
were used for larger grasshoppers. The wheat cuttings were trimmed of all dry growth and
their stems placed in water. The containers of wheat cuttings were stoppered with muslin
(small containers) or corks (large containers). Fresh weights of five samples of wheat were
taken from each pot used in the feeding trials to determine the equivalent dry weight of
the wheat available to the grasshoppers. The mean ratio of the dry:wet weight of the wheat
varied with age, from 0.096 for 9 day old plants (n = 40, range = 0.082 - 0.108) to
0.115 for 18 day old plants (n = 44, range = 0.096 - 0.125).
The containers along with their grasshoppers and wheat cuttings were placed in an
environmental chamber at 30.6:t0.5 (S.D.tC. The ambient relative humidity was
4O:t 1.9% R.H. Lighting conditions were the same as above. After 24 h, the containers
were checked to determine which, if any, grasshoppers had died. Then the containers and
their contents were chilled for 20 min at -5°C and frozen for several hours at - 40°C.
The frozen grasshoppers were reweighed and measured, dried at 65°C for 72 h and
weighed again (Riegert and Varley 1973). The wheat remains and the associated control
cuttings were dried at 65°C for 24 h before weighing.
Data from grasshoppers that died (0.8%), moulted (3.2%), ate all ofthe available
wheat (2.4%), or ate nothing (3.6%) during the trials were not included in the analysis.
The weight eaten by the grasshoppers was calculated as the difference between the esti-
mated initial dry weight and the actual dry weight of the remains.
Indices of grasshopper size. Five indices of grasshopper size were used: (I) length of
the dorsal surface of the pronotum, (2) length of femur of the third leg, (3) total body
length, (4) initial live weight, and (5) estimated initial dry weight. The initial dry weight
of each grasshopper was calculated as the product of its initial live weight and the mean
ratio of the dried:frozen weight of grasshoppers of the same class. With the exception of
body lengths over 2 cm, the length measurements were made with the aid of a stereo-
microscope fitted with an ocular micrometer. Body lengths of grasshoppers over 2 cm
were measured with a ruler. Microscope measurements were rounded to th~ nearest
0.1 mm, ruler measurements to 0.5 mm. Body lengths were dorsal measurements from
the head to the end of the abdomen, but excluded antennae. Pronotum and femur indices
were maximum lengths.
Data analysis. Means of the dry weights of wheat eaten were compared with pronotum,
femur, and body lengths and the live and dry weights for each class of grasshopper. Classes
were distinguished by species, stage, and sex. Adults were identified with the aid of Brooks
(1958). Developmental stages were distinguished by comparing the number of antennal
segments and examining wing development (Shotwell 1941). Sexes were determined by
examination of the external genitalia (Brusven 1967; Shotwell 1941). We did not distin-
guish the sexes of instars I and II of C. pellucida, nor of instars I to III of M. sanguinipes.
Thus the data for these stadia are averages of unknown mixes of males and females. There
were insufficient eggs of M. bivittatus to obtain feeding rates for instars I and III. These
factors reduced the possible number of grasshopper classes from 72 (6 species x 6 stages
x 2 sexes) to 63.
The method of least squares was used to estimate the parameters of five equations
relating feeding rate to the five indices of body size for each class of grasshopper. The
equations tested were: power (y =axh), exponential (y =aeh"), parabolic
(y =a+bx +cx2), linear (y = a+bx), and logarithmic (y = a+bIn(x)).
In order to judge the relative usefulness of the five types of equations and five size
indices we used mean coefficients of determination (fl). These were calculated as follows:
JK f2J..
f2=II
J~I '~I J'K [1]
Table I. Parameters for the power equations (y =axb) relating dry weighl of wheat eaten 10 5 indices of body
size for all 6 species of grasshoppers. Probability levels of all r2 values are < 0.001 excepl Ihose marked by an
asterisk which are < 0.005. The numbers of paired means used to determine the paramelers are 29. 29. 51. 12.
and 63 for M. F. N. Adl. and All. respectively. a=first parameter of the power equation. h=exponenlof
the power equation. S.E. h=standard error of b. M=male nymphs and adults. F =female nymphs and
adults. N =nymphs of both sexes. Adt =adults of both sexes. All =all stages and sexes
Sex or
Size index stage a h S.E. hr2
PronolUm length M2.022 1.428 1.93 x 10-1 0.88
F2.060 1.564 1.38 x10-1 0.93
N1.845 1.612 1.91x1O-1 0.92
Adt 1.568 1.598 3.96x 10-1 0.62*
All 1.871 1.560 1.69 x10-1 0.91
Femur length M 0.357 1.708 1.15 x 10-1 0.84
F0.311 1.850 8.00 x 10-2 0.91
N0.189 2.141 1.16 x10-1 0.92
Adt 0.060 2.331 4.95 x 10-1 0.69
All 0.274 1.869 9.98 x 10-2 0.89
Body length M0.102 1.769 6.74x 10-2' 0.81
F0.088 1.890 4.94 x 10-2 0.88
N0.036 2.258 7.37 x 10-2 0.89
Adt 0.029 2.100 4.90 x 10-1 0.65*
All 0.070 1.935 5.95 x10-2 0.86
Uve weight M0.514 0.661 2.64 x 10-3 0.84
F0.478 0.706 l.40x 10-3 0.92
N0.325 0.799 3.57 x 10-3 0.90
Adt 0.064 0.975 1.91 x 10-1 0.72
All 0.415 0.715 2.04 x 10-3 0.88
Dry weight M1.542 0.597 9.52 x 10-3 0.82
f1.521 0.654 5.32X 10-3 0.91
N1.180 0.748 1.39 x 10-2 0.90
Adt 0.259 0.956 1.94 x 10-1 0.71
All 1.336 0.661 7.19 x 10-3 0.87
600 THECANADIANENTOMOLOGIST April 1984
where J=number of species (up to 6) and K=number of size indices (up to 5). Jwas
I (all species combined), 5 (five individual species Le., some classes of M. sanguinipes
were excluded due to insufficient data) or 6 (six individual species). Kwas I (one of five
size indices), 2 (weight indices), 3 (length indices), or 5 (all size indices). Statistical tests
could not be used because the size indices and fl values of the equations were based on
common feeding rates.
Results
The total number of classes of grasshoppers tested was 63. The mean number of
grasshoppers per class was 23.3; range 10 to 36. The ratio of the standard error to the
mean feeding rate of each class was usually less than II %. The major exceptions occurred
in M. sanguinipes where the average ratio was 18%. The ratio of the standard error to the
mean lengths of the pronotum, femur and body were, with one exception out of 189 cases,
always less than 5%; and for the live and dry weights, always less than 10%.
When data for all species were combined (Le., 63 paired means (n) were used to
calculate the coefficients of determination (fl va~ues», the power equation best described
the correlations between the five indices of body size and the mean feeding rates. The r2
values for the power equations ranged between 0.86 and 0.91 and were about 22% higher
than the next best equations. For the power equations, there were only minor differences
(about 6%) between the best and the worst fl values for the five indices of body size.
Ranking the indices for all classes produced the following hierarchy (fl values in paren-
Volume 116 THE CANADIAN ENTOMOLOGIST 60
theses): pronotum length (0.91), femur length (0.89), live weight (0.88), dry weighl
(0.87), and body length (0.86) (Table I). Though body length was the worst size index,
its r2 values were still higher than the other four indices when they were used with the
other equations. The equation and size index hierarchies for male (n =29), female (11
=29), and nymphal (n =51) classes were similar to those found for all the classe§
combined.
When the same data were separated by species (n reduced to between 8 and 12), the
power equation was, generally, still the best of the five equations. However, the differ-
ences between the r2 values of the power equations and the next best equations were, 011
average, only 8% (cf. 22% for all species). Indeed, in M. packardii the parabolic equatiol1
was slightly better than the power equation (fl values of 0.98 and 0.96, respectively). For
the power equations, the average difference between the best and worst of the five indices
of body size for each species was not much different from all classes combined (Le., 8%
rather than 6%). The hierarchy of size indices was: pronotum length (0.95), live weight
(0.92), femur length (0.91), dry weight (0.91), and body length (0.89).
When only the data for nymphs of each species were examined (n reduced to between
6 and 10), the fl values of the power equation were only slightly higher than those of the
parabolic (Le., 0.96 vs. 0.95). The fl values of the length indices were nearly the same
(Le., femur 0.93, body 0.92, pronotum 0.92), but were higher than the weight indices
(Le., live 0.88, dry 0.87).
When the data for males (nymphs and adults) were separated by species (n reduced
to between 4 and 6; M. sanguinipes excluded as n=3), the fl ranking of the equations
was: parabolic (0.93), power (0.79), logarithmic (0.74), linear (0.64), and exponential
(0.58). The major change from previous rankings was the predominance of the parabolic
equation over the power equation. There was no clear hierarchy pattern for size indices.
The results for females were similar to the males.
The values of the terms of the power equations for all species combined, in different
combinations according to stage and sex, are also given in Table 'I. The avalues varied
between about 0.03 for body length to more than 2.0 for pronotum length. They were
lowest for the adults and usually highest for the males. The bvalues tended to be lowest,
about 0.7, for dry weight'and highest, near 2.0, for body length. The lowest bvalues were
associated with the males and the highest with the adults. High bvalues suggest that
feeding rates are strongly influenced by body size.
When graphs were made of feeding rate vs. the five size indices, it was apparent that
the power equations for all stages and sexes often underestimated the feeding rates of
nymphs and nearly always overestimated those of the adult~specially the males. For
this reason, we developed separate equations for nymphs and adults. Differences between
mean sizes and feeding rates of first to third instar males and females were negligible but
older females were significantly larger and ate more than males (Figs. I, 2). However
there were no. significant differences between the bvalues of the equations for male and
female nymphs or between male and female adults.
(NOTE: Graphs for each grasshopper species as well as individual r2 and equation values
are available, at cost of duplication, from the Depository of Unpublished Data, CISTI,
National Research Council of Canada, Ottawa, Ontario K IA OR6. This unpublished ap-
pendix also includes the mean sizes and feeding rates for each class of grasshopper.)
Discussion
It is well known that rates of metabolic processes, such as respiration, are related in
a predictable manner to various indices of body size, such as body weight (Bertanlanffy
1957; Brody 1945; Keister and Buck 1974; Zeuthen 1953). As expected, our results in-
100
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. M. infanti/is
. C.pellucida
. M.sanguinipes
*M. packardii
. M. bivittatus
. O.carolina
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Dry weight (mg) >
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2:
FIG. I. Relationship between dry weight of wheat eaten by and dry body weight of nymphs of six species of grasshoppers. Points are mean
observations as described in the text. Constants for the power curve are given in Table I. Curves above and below the line denote 95%
confidence limits for the regression equation. Open symbols are females: shaded. males.
.0
00
~
100 Adults <
4M. infantilis 0
c
70 .C. pellucida
~M.
sanguinipes -
.;:
-*M. packardii
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'0 D.carolina
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300
FIG. 2. Relationship between dry weight of wheat eaten by and dry body weight of adults of six species of grasshoppers. For further explanation
see Fig. I.
g
....
604 THECANADIANENTOMOLOGIST April 1984
dicate that such relationships also exist in grasshoppers between feeding rates and five
indices of body size.
Our results also support the common practice of relating such variables with the
power equation, at least when data for several species are combined.
As reported by Davey (1954), Misra and Putnam (1966) and others, we found that
adult grasshoppers fed less than immatures. The feeding rates not only leveled off when
the insects matured, but in some cases were actually lower than those of the fifth instars.
For this reason, we recommend the use of two power equations, one for nymphs and one
for adults.
There was one finding that differed from what has been published previously. Nagy
(1952) and Gangwere (1959) have reported that, on a gram for gram basis, adult male
grasshoppers may eat up to twice as much as adult females. Our results indicated that in
five of the six species, the females proportionately ate more than the males (range from
2-72% on a dry weight basis). In the sixth species, D. carolina, the males ate 20% more
than the females. The most likely explanation for this difference is variation in feeding
rates during the egg-laying cycle of the females and the mating condition of the males.
Our results were based on recently moulted adults. Nagy (1952) and Gangwere (1959)
used adults of unknown ages.
Size indices and the power equation. When Reichle (1968) examined feeding rate as
a function of dry weight in II species of forest floor arthropods, his power equation had
abtenn of 0.68 :t 0.129 i.e., close to the "surface law" of body weight raised to the two-
thirds power. Similar relationships also have been found for various millipedes and isopods
by Dunger (1958), Gere (1956), and Van der Drift (1951). In our work, the bvalue for
relating feeding rates and dry weights of all classes of grasshoppers was 0.661 :to.OO749.
This was not significantly different from Reichle's value of 0.68 or the surface law of 0.67
(t-test, P > 0.1). With grasshopper nymphs, the btenn 0.748:t 0.0139 was significantly
higher (P < 0.001) than the surface law value, though it did not differ significantly from
Reichle's value. The bvalue for adults, 0.956, had a much higher standard error, 0.194,
and so did not differ significantly from either Reichle's or the surface law values.
As live weight and dry weight values are directly dependent upon one another, it was
of no surprise that the correlations between feeding rates and live weights were close to
those found for the correlations with dry weights. However, we were surprised to find that
live weights usually produced higher r2 vall!es than dry weights (cf. Reichle 1968).
The feeding rates of sardines on plankton increase with the square of the body length
of the fish (Yoshida 1956). This was also the case for adult grasshoppers and all classes
taken together (P > 0.1) when bvalues were compared with the theoretical value of 2.0.
Nymphs, however, had a bvalue (2.258 :to.0737) significantly higher than 2.0
(P < 0.001).
The shape and hence the relative size of a pronotum or femur is liable to vary con-
siderably between even closely related species of insects. Thus their usefulness in pre-
dicting feeding rates should not be as generalizable as indices of weight or body length.
However in our work these two measurements proved to be excellent indicators of grass-
hopper feeding rates. The average bvalue of the power equation for the pronotum meas-
urements of nymphs and adults was about 1.60; for femur measurements, 2.24.
Practical applications. The main purpose of our work was to detennine which and how
well various indices of body size could eliminate the need to consider the influences of .
species, stage, an<;lsex on feeding rates of grasshoppers. The usefulness of our findings
will depend upon the precision required in detennining grasshopper feeding rates as well
as the time and facilities available to collect the necessary infonnation.
If it is possible to identify grasshopper samples to species, stage, and sex one could
use equations appropriate for each of these grasshopper classes (i.e., as given in our un-
Volume 116 THE CANADIAN ENTOMOLOGIST 605
published appendix). For example, one would get a more precise estimate of the basic
feeding rate of male M. infantilis by using an equation for male M. infantilisrather than
an equation produced for both sexes but the latter equation would be more accurate for
that species than a general equation for all species.
Often pest managers may not have the necessary resources to determine the species,
stage, and sex of most grasshopper populations under study. Thus it is likely that a mixture
of species, stages, and sexes would have to be treated. For this situation, we recommend
the use of two power equations, one for nymphs and one for adults. Adults of most grass-
hopper species have fully developed wings and may therefore be easily distinguished from
nymphs.
It is also unlikely that pest managers could afford to determine body size by dry
weights or microscopic measurements of pronotum or femur lengths. Very little precision
would be lost and a lot of convenience gained by the use of live weights or body lengths.
Both of these are also readily automated.
Though we are confident in the feeding rates and equations that we have determined,
we realize that some adjustments will have to be made for field conditions. For example,
Misra and Putnam (1966) showed that C. pellucid a reared in cages in the field required
33% more Poa pratensis to pass from hatching to adulthood than those reared indoors.
Thus, before our feeding rates can be used, they need to be adjusted for the effects of
temperature, selective feeding, available food supplies, and clipping rates. We believe
this is possible because:
J. Preliminary data (Hardman unpub.) suggest that feeding rates of M. sanguinipes are
related to temperature in the same way as developmental rates. If this is generally so,
then feeding rates may be computed on a day-degree scale.
2. Though feeding rates are affected by food selection mechanisms peculiar to each grass-
hopper species and by availability of alternative hosts (Gangwere 1972; Hardman and
Smoliak 1980, 1982; Hewitt 1977), these factors can be taken into account by the use
of selection coefficients (Holmberg and Turnbull 1982; Rodell 1977).
3. Though grasshoppers may sever plant leaves without eating them, and so waste more
than they eat, the ratio of the amount destroyed to the amount eaten can be calculated
because it is related to factors such as the size and shape of the plant, the width of the
grasshopper's head, and whether the plant is a preferred or non-preferred host (Gang-
were 1972; Mitchell and Pfadt 1974).
With the use of modified power equations as described above and the application of
automation to determine grasshopper weights or lengths, we think that it should be possible
to estimate grasshopper damage for a wide variety of field conditions.
Acknowledgments
We thank Agriculture Canada for granting permission to R. G. H. to work at the
Lethbridge Research Station while he was on sabbatical leave from Athabasca University,
R. C. Andrews and C. Cody for technical assistance, N. Hall for advice and supplies of
grasshopper eggs, and Paula Wintink-Smith for help with the figures.
References
Beirne, B. P. 1972. Pest insects of annual crops in Canada. IV. Hemiptera-Homoptera. V. Onhoptera. VI.
Other groups. Mem. ent. Soc. Can. 85. 75 pp.
Bertalanffy, L. von. 1957. Quantitative laws in metabolism and growth. Quart. Rev. Bioi. 31: 217-231.
Brody, S. 1945 (reprinted 1964). Bioenergetics and Growth. Hafner, N. Y.
Brooks, A. R. 1958. Acridoidea of southern Alberta, Saskatchewan, and Manitoba (Onhoptera). Can. Em.
Suppl. 9. 92 pp.
Brusven, M. A. 1967. Differentiation, ecology and distribution of immature slant-faced grasshoppers (Acri-
dinae) in Kansas. Tech. Bull. Kans. St. Univ. 149. 59 pp.
606 TIlE CANADIAN ENTOMOLOOIST April 1984
Davey. P. M. 1954. Quantities of food eaten by the desert locust, Schistocerca gregaria (Forsk.), in relation
togrowth.Bull.ent.Res.45:539-551. .
Dunger. W. 1958. Ober die Zersetzung der Laubstreu durch die Bodenmakrofauna im Auenwald. Zoo/. Jb.
(Syst.) 86: 139-180.
Gangwere, S. K. 1959. Experiments upon the feeding consumption of the grasshopper Me/anop/us s. scudderi
(Uhler). Mich. Acad. Sci., Arts, Len. 44: 93-96. .
- 1972. Host finding and feeding behaviour in the Ot:thopteroidea, especially as modified by food avail-
ability: a review. Rev. Univ. Madrid 11: 106-158.
Gere, G. 1956. The examination of the feeding biology and me humificative function of Diplopoda and lso-
poda. Acta Bioi. Acad. Sci., Hung. 6: 257-271.
Hardman. J. M. and S. Smoliak. 1980. Potential economic impact of rangeland grasshoppers (Acrididae) in
southeastern Alberta. Can. Ent. 111: 277-284.
- 1982. The relative impact of various grasshopper species Dn Stipa-Agropyron mixed prairie and fescue
prairie in southern Alberta. J. Range Mgmt 35: 171-176.
Hewiu, G. B. 1977. Review of forage losses caused by rangeland grasshoppers. U.S. Dep. Agric. Misc. Pub/.
1348. 24 pp.
Holmberg. R. G. and A. L. Turnbull. 1982. Selective predation in a euryphagous invertebrate predator, Par-
dosa vancouveri (Arachnida: Araneae). Can. Ent. 114: 243-257.
-Hunter-Jones. P. 1961. Rearing and breeding locusts in the laboratory. Anti-Locust Research Centre,
London.
Keister, M. and J. Buck. 1974. Respiration: some exogenous and endogenous effects on rate of respiration.
pp. 469-509 in M. Rockstein (Ed.), Physiology of Insecta. Academic Press, N.Y.
Misra. S. D. and L. G. Putnam. 1966. The damage potential of the grasshopper. Camnula pellucida (Scudd.)
(Orthoptera:Acrididae) on pastures and ranges in Canada. Indian J. Ent. 18: 224-233.
Mitchell. J. E. and R. E. Pfadl. 1974. A role of grasshoppers in a short grass prairie ecosystem. Environ. En!.
3: 358-360.
Nagy, B. 1952. Food consumption of Dociostaurus crucigerus brevicollis Eversm. and Oedipoda coeru/esC"ens
L. (Orth. Acrididae). Acta Bioi. Acad. Sci.. Hung. 3: 41-52.
.Parker , J. R. 1930. Some effects of moisture and temperature upon Melanoplus mexicanus mexicanus Saussure
and Camnula pellucida Scudder (Orthoptera). Mont. agric. Exp. Stn Bull. 233. 132 pp.
Reichle; D. E. 1968. Relation of body size to food intake. oxygen consumption. and trace element metabolism
in forest floor arthropods. Erology 49: 538-542.
Riegert, P. W. and J. L. Varley. 1973. Above-ground invertebrates: II. Population dynamics and biomass
production of grasshoppers. Can. Comm. into Bioi. Prog., Matador Project, Tech. Rep. 16. 134 pp.
Rodell. C. F. 1977. A grasshopper model for a grassland ecosystem. Ec%gy 58: 227-245.
Shotwell. R. L. 1941. Life histories and habits of some grasshoppers of economic importance on the Great
Plains. U.S. Dep. Agric. Tech. Bull. 774. 47 pp.
Smith. D. S. and N. D. Holmes. 1977. The distribution and abundance of adult grasshoppers (Acrididae) in
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Van der Drift. J. 1951. Analysis of the animal community in a beech forest floor. Tijdschr. Ent. 94: 1-168.
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-Reference not seen in the original.
(Received 29 April 1983; accepted 27 October 1983)
Unpublished Appendix for:
Holmberg, R.G. and J.M. Hardman..
1984.
Relating feeding rates to sex and size in six species of
grasshoppers (Orthoptera: Acrididae).
Canadian Entomologist 116(4): 597-606.
Original on deposit in the:
Depository of Unpublished Data, CISTI,
National Research Council of Canada,
Ottawa, Ontario
KIA OSA.
Copies are available at cost of duplication.
This appendix provides details of the information summarized in our
published paper that should be of use to those who are deeply
interested in the feeding rates of grasshoppers.
The information is presented in 6 figures and 41 tables as follows:
Figures 1~6, dry weight of wheat eaten t'er8U8 the five
body size indices for each of the six species of
grasshoppers. Lines indicate the power equation for an
stages.
Tables Al~A6, values for the power equations for the six
individual species.
Tables A7-AI3, values for the parabolic equations for
combined and individual species.
Tables AI4-A20, values for the exponential equations for
combined and individual species.
Tables A21-A27, values for the linear equations for
combined and individual species.
Tables A28-A34, values for the logarithmic equations for
combined and individual species.
Tables A35-A40, mean values used to generate Figs. 1-6
along with standard errors.
Table A41, numbers of antennal segments of the sex
species of grasshoppers studied. This should be of help in
separating instars.
25
20
15
10
5
-..
>
ca 00
"'CS
"
'"
E
-25
C
CI)
fa 20
CI)
....
ca 15
CI)
z: ~10
II-
0
.... 5
z:
'"
.- 00
CI)
~>
..
Q25
20
15
0
10
00
.20
12 3 4
Pronotum length (mm)
0
0
.
5 10 15 20
Body length (mm) 25
00
.
25 50 75 100
Dryweight (mg) 125
0
.
5015
5 10
Femur length (mm)
0
0
.
0 50 150 250 350
Live weight (mg) 450
FIG. 2. C. pellucida. relationship between
weight of wheat eaten and five indices of body
size. Points are mean observations, replication
given in top left-hand graph. Constants for the
power equations are given in Table A2.
Male=., females = 0 , both males and
females=CI .
30
25
20
15
10
5
0
>. 0
ca
"1:'
.......
~
E30
-
C 25
eI)
....
Z20
-; 15
eI)
~10
'0 5
0
i, 0
.-
eI)
~
~30
Q
25
20
15
10
5
00
300
2 3 4
Pronotumlength (mm)
0
5 10 15 20
Body length (mm)
08
50 1 0
Dry weight (mg)
.19
0
.
1 0
55 10
Femur length (mm)
015
0
8
25 0 50 150 250 350
Live weight (mg) 450
0
FIG. 3. M. sangllinipes. relationship between
weight of wheat eaten and five indices of body
size. Points are mean observations, replication
given in top left-hand graph. Constants for the
power equations are given in Table A3.
Males =8 , females=O, both males and
females=() .
45
0
35
.22 .
25
15
25
524 22
20
00 1 2 3 4 .5 6
Pronotum length (mm)
-
>
as
"'C
.......
'"
E45
-C
!35
as
G)
.. 25
ca
G)
.J:.
~15
.....
0
... 5
.J:.
.~ 00
G)
~
>-
.. 45
Q
35
25
15
5
00
10 20
Body length (mm)
50 100
Dry weight (mg)
0
.
.
1575 10
Femur length (mm)
0
0
.
30 0 50 150 250 350 450
Live weight (mg) 550
0
FIG. 4.M. packardii, relationship between
weight. of wheat eaten and five indices of body
size. Points are mean observations. replication
given in top left.hand graph. Constants for the
power equations are given in Table A4.
Males=. .females=o
.
150
80
60
40
20
-
~00
'C
-....
~
E
-80
c
Q)
....
CO 60
CU
....
CO
CU 40
,r.
~
'S 20
....
.c
~
8; 00
~
>
..
C 80
60
40
20
1 23 4 5 6
Pronotum length (mm)
10 20
Body length (mm)
.
50 150
Dry weight (mg)
.
250
.
7246810 12 14 16 18
Femur length (mm)
0
.
30 0 50 150 250 350 450 550 650 750
Live weight (mg)
FIG. 5. M. bivittatus. relationship between
weight of wheat eaten and five indices of body
size. Points are mean observations. replication
given in top left-hand graph. Constants for the
power equations are given in Table A5.
Males=. . females=Q .
45 230
35
25
15
-
>
ca
"'D
.......
0)
,g 45
C
.s 35
ca
CD
1a 25
CD
.c
~ 15
...
05
.c 0
Ie' a
CD
~
~45
C
0
0
..
0
5 15 25
Body length (mm)
0
35
25 .
15
50 100 150 200
Dry weight (mg)
0
0
.
10 05 10 15
Femur length (mm) 20
0
.
35 150 350 550 750
Live weight (mg)
0950
0
FIG. 6. D. carolina, relationship between
weight of wheat eaten and five indices of body
size. Points are mean observations, replication
given in top left-hand graph. Constants for the
power equations are given in Table A6.
Males=. ,females=O.
250
Table AI. M. infantiUI, parameters far the pawer equations relating dry weight of
wheat eaten to 5 indices af body size. The number af paired means(n) used to
det.ermine the parameters are 6, 6, 10 and 12 for M, F, N and A - respectively. a = first
parameter of the power equation, b = exponent of the power equation, SE b =
standard error of b, M = male nymphs and adults, F =female nymphs and adults, N
=nymphs of both sexes, A = all stages and sexes, Prob. = prabability level af the r2
values
Size Sex/ a b SE b r2 Prob.
index stage
Pronotum M2.096 1.556 2.93 X to-I 0.92 <0.005
lengt,h F2.209 1.592 1.52 X 10-1 0.98 <0.001
N2.128 1.744 1.47 X to-I 0.98 <0.001
A2.149 1.577 ~.16X 10-1 0.95 < 0.001
Fem ur M 0.314 1.784 1.42 X 10-1 0.90 <0.005
length F0.307 1.833 6.97 X 10-2 0.97 <0.001
N0.227 2.078 5.69 X 10-2 0.99 <0.001
A0.309 1.812 1.02 X 10-1 0.94 <0.001
Body M0.061 2.040 1.08 X 10-1 0.85 <0.01
length F0.074 2.014 6.44 X 10-2 0.93 .<0.005
N0.025 2.570 4.25 X 10-2 0.98 <0.001
A0.067 2.025 8.20 X 10-2 0.89 <0.001
Live M 0.462 0.709 8.00 X 10-4 0.87 <0.01
weight F0.519 0.704 3.79 X 10-3 0.94 <0.005
N0.314 0.888 2.61 X 10-3 1.00 <0.001
A0.489 0.707 5.15 X 10-3 0.90 <0.001
Dry M1.414 0.663 2.99 X 10-2 0.88 <0.01
weight F1.606 0.641 1.32 X 10-2 0.93 <0.005
N1.283 0.824 8.51 X 10-3 1.00 <0.001
A1.507 0.652 1.76 X 10-2 0.90 <0.001
Table A3. M. unguinipe3, parameters for the power equations. n =7 and 9; male and
female equations not given as sexes not separated in first three instars
Size Sex/ a b SE b r2 Prob.
index sta~e
Pronotum N1.510 1.480 1.69 X 10-1 0.95 <0.001
length A1.444 1.619 1.86 X 10-1 0.94 <0.001
Femur N 0.284 1.694 7.59 X 10-2 0.96 <0.001
length A0.256 1.771 6.01 X 10-2 0.97 <0.001
Body N0.088 1.681 6.02 X 10-2 0.88 <0.05
length A0.066 1.817 5.09 X 10-2 0.91 <0.001
Live N 0.471 0.623 2.81 X 10-3 0.91 <0.001
weight A0.375 0.658 1.79 X 10-3 0.94 <0.001
Dry N1.118 0.566 9.04 X 10-3 0.91 <0.001
weiJ!:ht A 1.075 0.589 5.09 X 10-3 0.93 <0.001
11
Table A4. M. packardii, parameters for the power equations. n=6, 6,10 and 12
Size Sex! abSE b r2 Prob.
index staJ1;e
Pronot-um M1.411 1.810 1.47 X 10-1 0.96 <0.001
length F1.512 1.848 1.19 X 10-1 0.98 <0.001
N1.401 1.932 1.50 X 10-1 0.97 <0.001
A1.459 1.831 1.26 X 10-1 0.97 <0.001
Femur M 0.145 2.194 8.62 X 10-2 0.95 <0.001
length F0.163 2.154 7.44 X 10-2 0.95 <0.001
N0.105 2.443 7.82 X 10-2 0.97 <0.001
A0.154 2.174 7.48 X 10-2 0.95 <0.001
Body M0.028 2.323 6.23 X 10-2 0.91 <0.05
length F0.034 2.266 5.59 X lO-2 0.92 <0.05
N0.009 2.890 4.51 X 10-2 0.98 <0.001
A0.031 2.293 5.55 X lO-2 0.91 <0.001
Live M0.284 0.833 2.40 X lO-3 0.93 <0.05
weight F 0.262 0.851 1.50 X 10-3 0.96 <0.001
N0.183 0.973 2.38 X 10-3 0.98 <0.001
A0.272 0.842 1.79 X lO-3 0.95 . <0.001
Dry M1.008 0.788 6.87 X 10-3 0.96 <0.001
weight F1.013 0.805 5.65 X lO-3 0.96 <0.001
N0.849 0.909 6.51 X lO-3 0.99 <0.001
A1.008 0.798 5.80 X 10-3 0.96 <0.001
12
Table AS. M. bivittatus, parameters for the power equations. n = 4, 4, 6 and 8 as no
data for instal'S I and III
Size Sex/ abSE b r2 Prob.
ind{'x stage
Pronotum M 1.974 1.728 1.68 X 10-1 0.94 <0.05
length F1.898 1.909 1.01 X 10-1 0.98 <0.02
N1.651 2.036 1.16 X 10-1 0.98 <0.001
A1.895 1.838 1.31 X 10-1 0.95 <0.001
Femur M0.382 1.818 1.05 X 10-1 0.88 >0.1
lengt.h F0.268 2.060 6.74 X 10-2 0.95 <0.02
N0.149 2.394 7.07 X 10-2 0.97 < 0.001
A0.306 1.962 7.84 X 10-2 0.92 <0.001
Body M0.128 1.798 4.92 X 10-2 0.91 <0.05
length F0.073 2.081 2.92 X 10-2 0.97 <0.02
N0.045 2.282 3.10 X 10-2 0.98 <0.001
A0.092 1.958 3.91 X 10-2 0.93 <0.001
Live M 0.508 0.722 1.59 X 10-3 0.94 .<0.05
weight F0.439 0.784 6.35 X 10-4 0.98 <0.02
N0.327 0.854 9.97 X 10-4 0.99 <0.001
A0.455 0.761 9.75 X 10-4 0.96 <0.001
Dry M1.626 0.678 5.88 X 10-3 0.94 <0.05
weight F1.573 0.727 2.43 X 10-3 0.98 <0.02
N1.260 0.811 3.56 X 10-3 0.99 <0.001
A1.564 0.710 3.31 X 10-3 0.96 <0.001
13
Table A6. D. carolina, parameters for the power equations. n =6, 6, 10 and 12
Size Sexl a b SE b r2 Prob.
index staSte
Pronotum M2.419 1.154 8.19 X 10-2 0.96 <0.001
length F 2.260 1.298 5.34 X 10-2 0.98 <0.001
N2.317 1.231 8.09 X 10-2 0.96 <0.001
A2.320 1.235 6.85 X 10-2 0.97 <0.001
Femur M 0.609 1.506 7.99 X 10-2 0.90 <0.005
length F0.574 1.585 8.16 X 10-2 0.88 <0.01
N0.362 1.988 6.50 X 10-2 0.92 <0.001
A0.581 1.555 7.63 X 10-2 0.89 <0.001
Body M0.148 1.647 5.04 X 10-2 0.87 <0.01
length F 0.135 1.705 4.88 X 10-2 0.85 <0.01
N0.039 2.255 4.33 X 10-2 0.96 <0.001
A0.138 1.685 4.62 X 10-2 0.86 <0.001
Live M0.684 0.604 1.72 X 10-3 0.90 <0.005
weight F0.602 0.648 1.08 X 10-3 0.92 <0.005
N0.431 0.745 1.88 X 10-3 0.95 <0.001
A0.631 0.630 1.18 X 10-3 0.91 <0.001
Dry M 1.901 0.562 6.59 X 10-3 0.90 <0.005
weight F1.770 0.617 4.42 X 10-3 0.92 <0.005
N1.482 0.710 8.10 X 10-3 0.95 <0.001
A1.815 0.594 4.84 X 10-3 0.91 <0.001
14
Table A7. All species combined, parameters for the parabolic
equations.
n= 29, 29, 51 and 63. All probabilities <0.001
Size Sexf a b cr2
index stage
Pronotum M -4.271 5.711 -1.38 X 10-1 0.75
length F -9.134 9.254 -3.66 X 10-1 0.74
N -4.541 6.029 -6.90 X 10-2 0.79
A -5.713 6.737 -1.46 X 10-1 0.74
Femur M -5.619 2.628 -1.49 X 10-2 0.71
length F -8.976 3.540 -1.99 X 10-2 0.75
N0.941 -0.478 3.05 X 10-1 0.86
A-5.930 2.576 1.82 X 10-2 0.74
Body M-7.933 1.779 -1.72 X 10-2 0.68
length F -13.419 2.616 -2.77 X 10-2 0.72
N-5.671 1.125 2.82 X 10-2 0.77
A-9.169 1.914 -1.23 X 10-2 0.70
Live M 2.656 0.098 -9.90 X 10~ 0.70
weight F2.605 0.110 -7.38 X 10~ 0.76
N 1.644 0.124 -9.20 X 10~ 0.79
A 2.744 0.093 -5.69 X 10~ 0.72
Dry M2.657 0.412 -1.85 X 10-3 0.64
weight F4.926 0.317 -6.12 X 10-4 0.70
N0.754 0.675 -4.08 X 10-8 0.77
A4.261 0.283 -5.11 X 10-4 0.66
15
Table A8. M. in/antil'lJ, parameters for the parabolic equations. n =
6, 6, 10 and 12
Size Sex/ ab c r2 Pro b.
index sta!1;e
Pronotum M-6.197 10.154 -1.48 X 10° 0.76 <0.05
length F-3.219 5.605 -8.28 X 10-2 0.92 <0.005
N2.182 -2.532 2.35 X 10° 0.97 <0.001
A-3.313 5.906 -2.50 X 10-1 0.84 <0.001
Femur M -10.276 5.404 -3043 X 10-1 0.77 <0.05
length F-5.282 2.729 -4.53 X 10-2 0.92 <0.005
N2.268 -1.298 4.13 X 10-1 0.98 <0.001
A-5.898 3.136 -1.00 X 10-1 0.82 <0.001
Body M-17.870 4.626 -1.85 X 10-1 0.78 <0.02
length F -11.739 3.0n -8.13 X 10-2 0.93 <0.005
N2.450 -1.007 1.68 X 10-1 0.99 <0.001
A-11.510 2.984 -8.78 X 10-2 0.81 <0.001
Live M -1.184 0.355 -2.11 X 10-s 0.95 <0.001
weight F-0.199 0.269 -9.50 X 10--4 0.99 <0.001
N0.435 0.199 -1040 X 10--4 1.00 <0.001
A0.960 0.196 -6.70 X 10--4 0.78 <0.001
Dry M-0.800 1.405 -3.33 X 10-2 0.97 <0.001
weight F0.372 0.984 -1.22 X 10-2 1.00 <0.001
N0.437 0.926 -9.92 X 10~ 1.00 < 0.001
A1.626 0.662 -7.58 X 10-3 0.76 <0.001
18
Table A9. C. peUucidfJ, parameters for the parabolic equations. n =
4, 4, 8 and 10
Size Sex! ab c r2 Prob.
index stage
Pronotum M -16.654 17.507 -2.87 X 10° 0.67 >0.1
length F -4.668 5.087 1.34 X 10-1 0.98 <0.02
N 0.271 0.868 8.70 X 10-1 0.06 <0.001
A0.557 0.976 6.80 X 10-1 0.82 <0.001
Femur M -22.908 8.389 -5.20 X 10-1 0.95 <0.05
length F -22.168 6.991 -2.83 X 10-1 0.97 <0.02
N -1.396 0.643 1.51 X 10-1 0.96 <0.001
A -5.261 2.604 -6.12 X 10-2 0.71 <0.005
Bod)" M -28.261 5.313 -1.79 X 10-1 0.96 <0.05
length F-26.861 4.295 -9.64 X 10-2 1.00 <0.005
N -2.399 0.547 3.28 X 10-2 0.95 <0.001
A-4.519 1.200 -9.21 X 10-3 0.74 <0.005
Live M 1.025 0.150 -5.24 X 10-4 1.00 <0.005
weight F1.847 0.121 -1.83 X 10-4 0.99 <0.01
N1.450 0.105 -1.40 X 10-4 0.91 <0.001
A2.821 0.65 -5.77 X 10-6 0.72 <0.005
Dry M3.043 0.422 -5.17 X 10-3 0.98 <0.02
weight F2.879 0.471 -2.70 X 10-3 0.98 <0.02
N1:672 0.468 -3.02 X 10-3 0.88 <0.001
A3.566 0.240 -9.53 X 10-4 0.63 <0.01
17
Table AID. M. unguinipel, parameters for the parabolic equations.
n=7 and 9
Size Sex/ a b c r2 Prob.
index sta,;e
Pronotum N -1.910 3.316 2.21 X 10-2 0.87 <0.005
length A3.804 -3.456 1.51 X 10° 0.85 <0.001
Femur N -3.036 1.662 -4.24 X 10-3 0.88 <0.005
length A 2.118 -0.637 1.97 X 10-1 0.92 <0.001
Body N -5.794 1.426 -2.66 X 10-2 0.81 <0.01
length A1.416 -0.133 4.25 X 10-2 0.80 <0.005
Live N 0.188 0.119 -2.78 X 10-4 0.87 <0.005
weight A2.828 0.031 3.88 X 10-5 0.88 <0.001
Dry N0.378 0.441 -3.51 X 10-3 0.86 <0.005
wei,;bt. A3.345 0.099 1.97 X 10-4 0.87 <0.001
18
Table All. M. pa~kardil~ parameters for the parabolic equations. (n
=6, 6, 10 and 12. All probabilities <0.001
Size Sex! b"
acr*
index staSte
Pronotum M -4.267 5.152 1.28 X 10-1 0.97
lengt.h F-5.641 6.566 5.16 X 10-2 1.00
N -2.186 3.031 6.14 X 10-1 0.99
A-4.422 5.362 1.66 X 10-1 0.98
Femur M-10.256 3.856 -5.37 X10-2 0.96
lengt.h F-12.511 4.673 -8.40 X 10-2 0.98
N-4.037 1.353 1.67 X 10-1 0.98
A-10.666 4.010 -5.22 X 10-2 0.97
Body M-19.467 3.663 -6.72 X 10-2 0.95
length F -19.378 3.691 -5.83 X 10-2 0.98
N-2.720 0.133 9.70 X 10-2 1.00
A-18.037 3.427 -5.39 X 10-2 0.95
Live M0.437 0.199 -3.04 X 10-4 0.98
weight F0.462 0.180 -2.05 X 10-4 0.99
N-0.850 0.242 -4.73 X 10-4 0.99
A1.331 0.164 -1.94 X 10-4 0.95
Dry M0.146 0.738 -4.20 X 10-3 1.00
weight F0.932 0.701 -3.08 X 10-3 0.99
N0.196 0.774 -4.50 X 10-3 0.99
A1.419 0.624 -2.76 X 10-3 0.95
19
Table A12. M. bivittatu" parameters for the parabolic equations. n
=4, 4, 6 and 8
Size Sex/ abcr2 Prob.
index stage
Pronotum M 18.918 17.187 -lAD X 10° 1.00 <0.005
length F-10.285 9.668 1.88 X 10-1 1.00 <0.005
N12.885 11.755 -2.84 X 10-1 0.97 <0.001
A-6.192 6.927 3.41 X 10-1 0.89 <0.001
Femur M -27.485 9.543 -3.68 X 10-1 1.00 <0.005
length F19.430 6.442 -7.47 X 10-2 1.00 <0.005
N-20.099 6.721 -1.20 X 10-1 0.98 <0.001
A-13.416 4.989 -4.15 X 10-2 0.87 <0.001
Bod:y M -24.228 4.651 -9.27 X 10-2 1.00 <0.005
length F-14.592 2.708 2.84 X 10-3 1.00 <0.005
N-15.947 2.970 -1.50 X 10-2 0.98 <0.001
A -13.248 2.637 -1.00 X 10-2 0.84 <0.005
Live M 0.902 0.191 -2.53 X 10-4 0.99 <0.005
weight F2.039 0.168 -1.09 X 10-4 0.99 < 0.005
N0.634 0.198 -4.69 X 10-5 0.98 <0.001
A5.646 0.110 -2.27 X 10-4 0.87 <0.001
Dry M1.546 0.729 -3.66 X 10-3 0.99 <0.005
weight F2.532 0.646 -1.60 X 10-3 0.99 <0.005
N1.074 0.778 -3.56 X 10-3 0.98 < 0.001
A6.904 0.397 -6.62 X 10-4 0.86 <0.001
20
Table A13. D. carolina, parameters for the parabolic equations. n =
6, 6, 10 and 12
Size Sexf abcr2 Prob.
index stage
Pronotum M 1.150 1.393 2.68 X 10-1 0.97 <0.001
length F-3.556 5.217 -9.06 X 10-2 0.92 <0.005
N2.181 0.607 4.14 X 10-1 0.93 <0.001
A-0.560 2.806 1.40 X 10-1 0.92 <0.001
Femur M -12.895 5.557 -2.05 X 10-1 0.91 <0.005
length F -20.274 7.933 -2.83 X 10-1 0.92 <0.005
N 0.704 -0.095 2.99 X 10-1 0.98 <0.001
A-14.813 6.135 -2.07 X 10-1 0.85 <0.001
Body M -17.007 3.295 -6.48 X10-2 0.89 <0.005
length F -26.328 4.648 -8.64 X10-2 0.94 <0.005
N-2.575 0.396 6.55 X 10-2 0.98 <0.001
A-19.063 3.548 -6.26 X 10-2 0.86 <0.001
Live M0.353 0.149 -1.95 X 10"" 0.95 <0.001
weight F1.221 0.144 -1.23 X 10"" 1.00 <0.001
N1.596 0.119 -6.12 X 10-0 0.98 < 0.001
A2.659 0.108 -8.77 X 10-0 0.86 <0.001
Dry M0.774 0.614 -3.13 X 10-3 0.96 <0.001
weight F1.954 0.597 -2.10 X 10~ 1.00 <0.001
N1.759 0.538 -1.49 X 10-3 0.98 <0.001
A3.273 0.446 -1.52 X 10-3 0.84 <0.001
n
Table A14. All species combined, parameters for the exponential
equations. n = 29, 29, 51 and 63. All probabilities <0.001
Size Sex! a b r2
index sta,;e
Pronotum M1.831 0.468 0.76
length F2.175 0.452 0.76
N 1.396 0.574 0.77
A1.675 0.495 0.76
Femur M 1.479 0.240 0.73
length F1.707 0.233 0.77
N0.723 0.375 0.86
A1.311 0.255 0.76
Body M1.541 0.124 0.68
length F1.807 0.122 0.73
N0.739 0.197 0.80
A1.351 0.134 0.72
Live M 4.270 0.005 0.54
weigh t F5.472 0.003 0.52
N2.998 0.009 0.62
A 4.248 0.004 0.52
Dry M 4.646 0.015 0.46
weight F 5.694 0.012 0.50
N3.208 0.032 0.57
A4.457 0.015 0.49
22
Table A15. M. in/antili" parameters for the exponential equations.
n=6, 6, 10 and 12
Size Sex/ a b r2 Prob.
index staJte
Pronotum M0.858 0.879 0.84 <0.02
length F0.940 0.854 0.92 < 0.005
N0.669 1.090 0.96 <0.001
A 0.898 0.866 0.88 <0.001
Femur M0.750 0.361 0.80 <0.02
length F 0.779 0.359 0.91 < 0.005
N0.491 0.479 0.97 <0.001
A 0.763 0.361 0.86 <0.001
Body M0.653 0.261 0.74 <0.05
length F 0.823 0.202 0.82 <0.02
N0.327 0.325 0.96 <0.001
A 0.739 0.208 0.78 <0.001
Live M2.393 0.014 0.52 <0.01
weight F 2.996 0.010 0.56 <0.05
N 1.665 0.031 0.86 <0.001
A2.767 0.011 0.53 <0.001
Dry M 2.467 0.055 0.52 <0.1
weight F 3.243 0.032 0.51 <0.1
N 1.826 0.114 0.82 <0.001
A 3.010 0.037 0.48 <0.02
23
Table A16. C. pellucida, parameters for the exponential equations.
n=4,4,8 and 10
Size Sex/ a b r2 Prob.
index stage
PronotuID M4.253 0.207 0.37 <0.1
length F2.395 0.492 0.98 <0.02
N0.709 0.876 0.84 <0.005
A0.902 0.725 0.78 <0.001
Femur M5.585 0.045 0.14 <0.1
length F3.326 0.159 0.84 <0.05
N0.477 0.424 0.83 < 0.005
A1.044 0.262 0.64 <0.01
Body M5.282 0.028 0.17 <0.1
length F3.237 0.086 0.84 <0.05
N0.463 0.226 0.83 <0.005
A0.958 0.146 0.67 <0.005
Live M6.645 0.001 0.15 <0.1
weight F7.338 0.003 0.72 <0.1
N2.357 0.011 0.57 <0.05
A3.145 0.006 0.47 <0.05
Dry M7.126 0.003 0.06 <0.1
weight F8.135 0.009 0.63 <0.1
N2.449 0.045 0.56 <0.05
A3.571 0.018 0.39 <0.05
24
Table A17. M. aangu;n;pea, parameters for the exponential
equations. n =7 and 9
Size Sex/ a b r2 Prob.
index stage
Pronotum N0.845 0.676 0.90 < 0.005
length A0.804 0.703 0.93 <0.001
Femur N0.717 0.316 0.90 < 0.005
length A0.802 0.293 0.93 <0.001
Body N0.863 0.141 0.79 <0.01
length A0.821 0.146 0.86 < 0.001
Live N 2.341 0.007 0.64 <0.05
weight A2.587 0.006 0.75 <0.005
Dry N2.624 0.020 0.57 <0.05
weifz:ht A 2.945 0.015 0.69 <0.01
25
Table A18. M. p(Jck(J,di.~ parameters for the exponential equations.
n= 6, 6, 10 and 12
Size Sex/ a b r2 Prob.
index sta~e
Pronotum M 1.081 0.625 0.86 <0.01
length F1.205 0.612 0.85 <0.01
N0.893 0.742 0.87 <0.001
A1.141 0.619 0.86 <0.001
Femur M 0.847 0.313 0.84 <0.02
length F1.082 0.278 0.81 <0.02
N0.565 0.399 0.88 <0.001
A0.973 0.293 0.82 <0.001
Body M0.965 0.162 0.75 <0.05
length F1.204 0.148 0.73 <0.05
N0.369 0.267 0.90 <0.001
A1.087 0.154 0.74 <0.001
Live M 3.756 0.006 0.53 <0.01
weight F3.935 0.005 0.56 <0.05
N2.562 0.012 0.66 <0.005
A3.862 0.005 0.54 <0.01
Dry M3.604 0.023 0.56 <0.05
weight F4.168 0.019 0.53 <0.1
N2.409 0.050 0.71 <0.005
A2.920 0.020 0.54 <0.01
ze
Table A19. M. bivitttJtU8, parameters for the exponential equations.
n=4, 4, 6 and 8
Size Sex/ abr2 Prob.
index stage
Pronotum M2.277 0.510 0.81 =0.1
length F2.370 0.528 0.87 >0.05
N10405 0.711 0.92 < 0.005
A2.283 0.524 0.85 <0.005
Femur M2.560 0.210 0.72 >0.1
length F2.499 0.223 0.82 >0.05
N1.067 0.353 0.91 <0.005
A20477 0.219 0.78 <0.005
Body
M
2.424 0.114 0.76 >0.1
length F 2.142 0.131 0.86 >0.05
N1.123 0.184 0.92 <0.005
A2.230 0.124 0.81 <0.005
Live M6.684 0.004 0.58 >0.1
weight F8.077 0.003 0.64 >0.1
N4.044 0.009 0.83 <0.02
A7.511 0.004 0.62 <0.05
Dry M
7.114 0.015 0.55 >0.1
weight F9.325 0.010 0.57 >0.1
N4.152 0.034 0.82 <0.02
A8.579 0.011 0.56 <0.05
17
Table A20. D. carolina, parameters for the exponential equations. n
=6, 6, 10 and 12
Size Sex/ abr2 Prob.
index stage
Pronotum M 2.149 0.355 0.92 <0.005
length F 2.363 0.337 0.86 <0.01
N 2.084 0.375 0.88 <0.001
A2.261 0.345 0.88 <0.001
Femur M 2.258 0.197 0.75 <0.05
length F 2.782 0.175 0.66 <0.05
N 1.178 0.332 0.90 <0.001
A 2.533 0.183 0.70 <0.001
Body M 2.096 0.105 0.72 <0.05
length F 2.652 0.091 0.64 >0.05
N0.983 0.181 0.88 <0.001
A 2.402 0.096 0.67 <0.005
Live M 4.982 0.004 0.55 >0.05
weight F 6.182 0.002 0.47 >0.1
N 3.842 0.007 0.73 <0.005
A 5.788 0.003 0.49 <0.02
Dry M5.374 0.013 0.50 >0.1
weight F6.419 0.009 0.45 >0.1
N 4.020 0.029 0.71 <0.005
A5.999 0.010 0.47 <0.02
28
Table A21. All species combined, parameters for the linear
equations. n = 29, 29, 51 and 63. All probabilities <0.001
Size Sex/ abr2
index sta~e
Pronotum M -2.835 4.683 0.75
length F -4.196 6.097 0.73
N-3.924 5.540 0.79
A -4.068 5.573 0.74
Femur M -4.844 2.388 0.71
length F-7.668 3.173 0.75
N -9.446 3.473 0.82
A -6.951 2.886 0.74
Body M -4.561 1.245 0.68
length F -6.861 1.652 0.71
N-9.361 1.835 0.77
A -6.611 1.512 0.70
Live M 4.688 0.056 0.68
weight F 6.844 0.054 0.66
N2.599 0.095 0.79
A 5.177 0.055 0.67
Dry M6.048 0.162 0.53
weight F7.250 0.193 0.66
N3.451 0.341 0.70
A 5.732 0.191 0.64
29
Table A22. M. infantilis, parameters for the linear equations. n =
6,6, 10 and 12
Size Sexl a b r2 Prob.
index stage
Pronotum M -1.822 4.357 0.73 <0.05
length F-2.g25 5.247 0.g2 <0.005
N-3.g3g 6.012 0.g2 <0.001
A-2.488 4.870 0.84 <0.001
Femur M -2.482 1.7g0 0.70 <0.05
length F-4.084 2.210 O.gl <0.005
N-5.666 2.64g 0.g4 <0.001
A -3.413 2.030 0.82 <0.001
Body M-2.g65 1.04g 0.62 >0.05
length F -4.116 1.282 0.88 <0.01
N-8.0g4 1.818 0.g5 <0.001
A -3.726 1.186 0.76 <0.001
Live M3.254 0.071 0.46 >0.1
weight F3.82g O.06g 0.68 <0.05
N0.564 0.188 1.00 <0.001
A3.543 0.070 0.5g <0.005
Dry M3.354 0.277 0.47 >0.1
weight F4.324 0.218 0.63 <0.05
N1.021 0.704 O.gg <0.001
A3.g84 0.233 0.56 <0.01
30
Table A23. C. pellucidtJ, parameters for the linear equations. n=
4,4,8 and 10
Size Sexf a b r2 Prob.
index sf-alte
Pronotum M3.566 1.549 0.32 >0.1
length F -5.942 5.958 0.98 <0.02
N -3.519 4.984 0.94 <0.001
A -3.014 4.532 0.80 <0.001
Femur M 5.758 0.314 0.10 >0.1
lengt.h F -2.353 1.968 0.88 >0.05
N -5.942 2.439 0.94 <0.001
A-2.498 1.693 0.70 < 0.005
Body M5.278 0.204 0.13 >0.1
length F-2.841 1.075 0.89 >0.05
N -6.045 1.294 0.94 <0.001
A -3.053 0.944 0.74 <0.005
Live M 6.914 0.010 0.12 >0.1
weight F7.266 0.037 0.80 >0.1
N 2.411 0.072 0.89 <0.001
A3.844 0.042 0.70 <0.005
Dry M7.485 0.019 0.05 >0.1
weight F 8.551 0.119 0.71 >0.1
N 2.669 0.306 0.87 <0.001
A4.778 0.136 0.59 <0.01
31
Table A24. M. 3anguipipe3, parameters for the linear equations. n
=7 and 9
Size Sex! a br2 Prob.
index stage
Pronotum N -2.019 3.430 0.87 <0.005
length A-4.864 4.952 0.78 <0.005
Femur N -2.911 1.610 0.88 <0.005
length A-5.765 2.180 0.87 <0.001
Body N-2.079 0.727 0.79 <0.01
length A-5.294 1.068 0.78 <0.005
Live N 2.835 0.038 0.71 <0.02
weight A 2.073 0.047 0.87 <0.001
Dry N3.401 0.110 0.66 <0.05
weiJ!:ht A2.920 0.128 0.86 <0.001
32
Table A25. M. packardii, parameters for the linear equations. n =
6, 6, 10 and 12
Size Sexl a b r2 Prob.
index sta!?;e
Pronotum M-5.302 6.019 0.97 <0.001
length F-6.096 6.936 1.00 <0.001
N -6.259 6.756 0.98 <0.001
A -5.837 6.527 0.97 <0.001
Femur M-7.720 3.018 0.95 <0.001
length F-7.748 3.209 0.98 <0.001
N-10.188 3.597 0.97 <0.001
A -7.898 3.138 0.97 < 0.001
Body M-6.936 1.599 0.89 <0.005
length F-7.055 1.747 0.93 <0.005
N-13.784 2.383 0.98 < 0.001
A-7.178 1.688 0.91 <0.001
Live M5.670 0.063 0.77 <0.05
weight F5.990 0.067 0.84 <0.02
N2.246 0.123 0.94 <0.001
A5.772 0.066 0.81 <0.001
Dry M5.403 0.246 0.78 <0.02
weight F6.638 0.245 0.82 <0.02
N1.836 0.504 0.97 <0.001
A5.987 0.246 0.80 <0.001
33
Table A26. M. bivittatulJ} parameters for the linear equations. n =
4, 4, 6 and 8
Size Sexf a b r2 Prob.
index stag;e
Pronotum M-3.748 6.802 0.92 <0.05
length F -12.756 11.232 1.00 <0.005
N -10.627 9.952 0.97 <0.001
A -10.442 9.657 0.88 <0.001
Femur M -2.635 2.844 0.84 >0.05
length F -13.296 4.919 1.00 <0.005
N-14.795 4.981 0.98 <0.001
A -10.226 4.176 0.86 <0.001
Body M-3.149 1.540 0.87 >0.05
length F -15.303 2.809 1.00 <0.005
N -13.711 2.567 0.98 <0.001
A-10.823 2.288 0.84 <0.005
Live M 10.002 0.057 0.72 >0.1
weight F 10.433 0.080 0.92 <0.05
N3.408 0.126 0.95 <0.001
A 8.832 0.076 0.85 <0.005
Dry M10.817 0.204 0.69 >0.1
weight F13.219 0.250 0.88 >0.05
N 3.760 0.493 0.95 <0.001
A10.930 0.247 0.83 <0.005
34
Table A27. D. carolina, parameters for the linear equations. n=6,
6, 10 and 12
Size Sexf a b r2 Prob.
index sta~e
Pronotum M -1.890 3.632 0.96 <0.001
length F-2.207 4.327 0.91 <0.005
N-2.687 4.124 0.90 <0.001
A-2.482 4.102 0.91 <0.001
Femur M -1.676 2.054 0.81 <0.02
length F-0.055 2.232 0.70 <0.05
N-10.275 3.956 0.95 <0.001
A-1.251 2.199 0.73 <0.001
Body M -2A90 1.101 0.78 <0.02
length F-0.871 1.174 0.69 <0.05
N -12.890 2.194 0.96 <0.001
A -2.083 1.167 0.72 <0.001
Live M6.336 0.040 0.65 >0.05
weight F9.744 0.032 0.55 >0.05
N2.611 0.095 0.98 <0.001
A8.150 0.034 0.59 <0.005
Dry M 7.135 0.141 0.59 >0.05
weight F10.214 0.125 0.53 >0.1
N3.102 0.398 0.97 <0.001
A8.619 0.133 0.56 <0.01
35
Table A28. All species combined, parameters for the logarithmic
equations. n = 29, 29, 51 and 63. All probabilities <0.001
Size Sex/ abr2
index staJ1:e
Pronotum M -0.177 12.621 0.68
length F -1.206 18.269 0.67
N 0.445 13.428 0.70
A-0.531 15.160 0.66
Femur M-15.880 15.292 0.67
length F -24.986 22.133 0.69
N -18.582 17.871 0.71
A-20.043 18.621 0.67
Body M -27.681 16.074 0.66
length F -41.002 22.973 0.69
N -32.944 19.094 0.70
A-34.193 19.517 0.66
Live M-12.995 6.056 0.66
weight F -19.571 8.376 0.68
N -14.118 6.685 0.70
A-15.705 7.077 0.66
Dry M -2.357 5.204 0.62
weight F -5.883 7.782 0.68
N -3.206 6.206 0.68
A-4.054 6.510 0.64
3e
Table A29. M. infantili~l parameters for the logarithmic equations.
n= 6,6, 10 and 12
Size Sex/ ab,2 Prob.
index stage
Pronotum M2.781 7.389 0.74 <0.05
length F2.659 9.202 0.87 <0.01
N2.697 9.006 0.83 <0.001
A2.700 8.361 0.80 <0.001
Femur M-6.322 8.525 0.73 <0.05
length F -8.831 10.647 0.87 <0.01
N -8.974 10.811 0.85 <0.001
A-7.692 9.674 0.80 <0.001
Body M-14.041 9.691 0.68 <0.05
length F-18.098 12.164 0.91 <0.005
N-21.119 13.669 0.88 <0.001
A-16.315 11.040 0.79 <0.001
Live M-4.403 3.366 0.70 <0.05
weight F -6.199 4.218 0.90 <0.005
N-7.468 4.681 0.88 <0.001
A -5.435 3.834 0.80 <0.001
Dry M0.893 3.158 0.71 <0.05
weight F0.517 3.863 0.90 <0.005
N -0.075 4.361 0.88 <0.001
A0.631 3.556 0.81 <0.001
37
Table A31. M. eanguipipe" parameters for the logaritmic: equations.
n=7 and 9
Size Sexf b"Prob.
arW
index stage
Pronotum N 1.280 7.081 0.81 <0.01
length A0.174 10.488 0.67 <0.01
Femur N w6.806 8.159 0.83 <0.005
length Awll.992 11.980 0.74 <0.005
Body Nw12.918 8.282 0.80 <0.01
length Aw21.269 12.347 0.71 <0.005
Live N w5.038 3.020 0.80 <0.001
weight Aw9.479 4.464 0.73 <0.005
Dry NwO.303 2.762 0.80 <0.01
wei~ht A w2.510 4.054 0.74 <0.005
30
Table A33. M. bivittatulf, parameters for the logarithmic equations.
n = 4, 4, 6 and 8
Size Sex! a b r2 Prob.
index sta~e
Pronotum M -4.581 22.166 0.98 <0.02
length F -13.106 37.222 0.94 <0.05
N -7.234 27.451 0.96 <0.001
A -10.233 31.007 0.83 <0.005
Femur M -26.541 23.746 0.95 <0.05
length F -54.261 41.537 0.97 <0.02
N--40.327 32.644 0.97 <0.001
A -43.343 34.211 0.85 <0.005
Body M -40.245 23.283 0.97 <0.02
length F -77.999 41.050 0.95 <0.05
N -56.133 30.835 0.96 <0.001
A -61.893 33.273 0.82 <0.005
Live M-21.863 9.232 0.98 <0.02
weigbt F-41.429 15.251 0.94 <0.05
N -28.638 11.414 0.96 <0.001
A -34.412 12.867 0.84 <0.005
Dry M -6.962 8.671 0.98 <0.02
weigb t F -17.137 14.286 0.95 <0.05
N -10.558 10.829 OJ)5 <0.001
A -13.947 12.115 0.85 <0.005
41
Table A34. D. carolina, parameters for the logarithmic equations. n
= 6, 6, 10 and 12.
Size Sex/ a b r2 Prob.
index stalte
Pronotum M 0.487 10.807 0.84 <0.02
length F -0.683 14.987 0.84 <0.02
N0.203 11.870 0.76 <0.005
A -0.360 13.202 0.80 <0.001
Femur M -13.734 14.821 0.87 <0.01
length F -17.849 19.008 0.82 <0.02
N-22.546 21.568 0.84 <0.001
A -16.560 17.382 0.81 <0.001
Bod:r M27.920 16.304 0.85 <0.01
length F36.305 20.873 0.82 <0.02
N -46.350 25.029 0.87 <0.001
A-33.464 19.162 0.81 <0.001
Live M-11.637 5.723 0.81 <0.02
weight F16.974 7.700 0.84 <0.02
N -18.359 7.911 0.79 <0.001
A -15.117 6.924 0.80 <0.001
DI')' M -1.943 5.322 0.80 <0.02
weight F -4.216 7.358 0.84 <0.02
N-5.242 7.535 0.79 <0.001
A -3.493 6.523 0.79 <0.001
42
Table A35. M. in/antilis, mean values used for Fig. 1. Values in
parentheses are standard errors of means. m = male, f = female, a
= adult
Stage Sex Dry weight Pronotum Femur Body Live Dry
eaten . length length length weight weight
(mlt) (mm) (mm) (mm) (mlt) (mlt)
1m1.03 0.70 2.19 4.62 4.33 0.86
(0.15) (0.01) (0.02) (0.14) (0.225) (0.05)
1f 1.26 0.70 2.15 4.43 4.26 0.88
(0.14) (0.01) (0.02) (0.10) (0.216) (0.04)
2 m 2.40 1.03 2.98 5.87 9.87 2.19
(0.12) (0.01) (0.03) (0.06) (0.406) (0.09)
2 f 2.18 1.05 3.12 5.77 9.17 1.97
(0.15) (0.01) (0.03) (0.08) (0.320) (0.07)
3m4.51 1.53 4.18 7.66 20.37 4.70
(0.22) (0.02) (0.06) (0.14) (0.733) (0.17)
3f5.26 1.59 4.41 7.32 20.79 -4.80
(0.33) (0.02) (0.07) (0.16) (0.928) (0.21)
4m8.84 2.41 5.84 10.17 44.49 10.30
(0.45) (0.02) (0.04) (0.19) (2.15) (0.50)
4 f 7.81 2.41 6.06 9.62 40.05 9.19
(0.44) (0.02) (0.04) (0.11) (1.82) (0.43)
5m15.35 2.89 7.34 12.23 81.12 21.31
(0.71) (0.04) (0.07) (0.18) (3.63) (1.36 )
5f15.79 2.96 7.44 12.29 78.98 20.85
(0.89) (0.04) (0.07) (0.12) (2.01) (1.84 )
am8.26 3.22 8.36 14.90 132.6 33.80
(0.09) (0.03) (0.05) (0.30) (4.07) (1.04)
a f 15.16 3.68 9.38 16.86 203.0 60.87
(1.25) (0.11) (0.09) (0.33) (8.76) (2.56)
43
Table A36. C. pellurido, mean values used for Fig. 2. Values in
parentheses are standard errors of means. m,f = mixed males and
females. Other abbreviations as in Table A35
Stage Sex Dry weight Pronotum Live Dry
eaten length weight weight
mmm mm
1m,f 0.54 0.69 5.76 1.02
(0.16) (0.01) (0.113) (0.02)
2m,f 2.29 1.06 3.15 6.55 12.55 2.56
(0.17) (0.02) (0.03) (0.10) (0.637) (0.13)
3m5.40 1.74 4.72 9.03 32.79 6.92
(0.45) (0.03) (0.10) (0.19) (2.14) (0.46)
3f5.95 1.94 4.91 9.65 42.30 8.84
(0.48) (0.03) (0.21) (0.22) (2.93) (0.61)
4 m 8.50 2.68 6.27 11.56 64.93 14.35
(0.76) (0.04) (0.07) (0.20) (2.62) (0.58)
4 f 11.18 3.04 6.92 12.54 78.14 16.80
(0.85) (0.06) (0.09) (0.23) (5.16) .(1.11)
5 m 11.46 3.48 8.26 15.72 166.93 39.73
(1.33) (0.06) (0.10) (0.74) (12.02) (2.86)
5f19.92 4.11 9.54 18.33 234.51 53.0
(1.45) (0.07) (0.10) (0.57) (21.42) (4.84)
am7.09 3.84 10.71 19.40 237.11 70.66
(0.80) (0.04) (0.90) (0.17) (6.17) (1.84 )
a f 20.63 4.58 12.72 23.68 419.12 119.45
(1.82) (0.06) (0.15) (0.35) (20.42) (5.82)
44
Table A37. M. IIanquinipe3, mean values used for Fig. 3. See Tables
A35 and A36 for further explanation
Stage Sex Dry weight Pronotum Live Dry
eaten length weight weight
m mm mm
1m,f 0.97 0.78 5.23 1.15
(0.26) (0.01) (0.41) (0.09)
2m,f 2.50 1.24 3.32 6.12 11.87 2.73
(0.49) (0.02) (0.03) (0.16) (1.00) (0.23)
3m,f 3.41 1.98 4.89 10.41 46.91 11.82
(0.59) (0.07) (0.13) (0.24) (2.54) (0.64)
4 m 7.05 3.12 6.80 13.20 93.26 24.06
(0.85) (0.03) (0.05) (0.20) (4.11) (1.06)
4 f 12.06 3.23 7.42 13.42 102.2 25.86
(2.17) (0.08) (0.14) (0.39) (6.24) (1.58)
5 m 10.29 4.22 9.60 20.74 272.2 87.36
(2.45) (0.04) (0.07) (0.40) (7.23) (2.32)
5 f 14.03 4.22 9.73 20.32 268.1 87.14
(2.38) (0.05) (0.10) (0.32) (11.14) (3.62)
am18.37 4.49 11.28 21.35 384.7 139.63
(2.20) (0.05) (0.13) (0.24) (7.88) (2.86)
af26.14 4.71 12.12 22.85 431.1 153.91
(3.28) (0.07) (0.16) (0.45) (15.91) (5.68)
to
45
Table A38. M. packardii, mean values used for Fig. 4. See Table
A35 for further explanation
Stage Sex Dry weight Pronotum Femur Body Live Dry
eaten length length length weight weight
(mg) (mm) ~mm) fu!.g) (.!!!SL
1 m 0.83 0.91 2.66 5.39 5.75 1.15
(0.12) (0.01) (0.02) (0.06) (0.15) (0.03)
1f0.88 0.88 2.60 5.18 5.72 1.19
(0.10) (0.01) (0.04) (0.10) (0.24) (0.05)
2m4.00 1.41 3.80 7.74 20.09 4.44
(0.26) (0.04) (0.08) (0.13) (0.81) (0.18)
2f3.65 1.40 3.87 7.58 20.95 4.65
(0.26) (0.02) (0.03) (0.22) (1.35) (0.30)
3 m 6.67 2.29 5.60 9.65 42.2] 9.16
(0.27) (0.02) (0.04) (0.28) (2.95) (0.64) ,
3r8.04 2.31 5.65 9.59 44.64 10.00
(0.70) (0.04) (0.07) (0.17) (2.37) (0.53)
4 m 15.65 3.74 7.81 13.36 64.67 26.96
(1.10) (0.04) (0.06) (0.24) (4.54) (1.23)
4r20.53 3.72 7.97 14.28 113.31 28.33
(1.00) (0.06) (0.10) (0.26) (5.04) (1.16)
5m28.52 5.05 10.47 17.59 221.98 53.72
(1.40) (0.05) (0.09) (0.24) (7.15) (1.73)
5 f 31.06 5.29 11.20 17.82 250.20 61.80
(2.29) (0.06) (0.12) (0.30) (18.62) (4.60)
a m 28.99 5.95 13.06 25.24 445.80 116.8
(1.82) (0.12) (0.20) (0.45) (14.50) (3.80)
af37.63 6.35 14.92 28.06 543.70 146.8
(2.95) (QJ!) (0.23) (0.46) (20.74) (5.60)
46
Table A39. M. bivittatu6, mean values used for Fig. 5. See Table
A35 for further explanation
Stage Sex Dry weight Pronotum Femur Body Live Dry
eaten length length length weight weight
(m,;) (~ ~(m,;) (mJd
2 m 3.18 1.44 3.73 6.74 15.6 3.35
(0.27) (0.01) (0.02) (0.20) (1.05) (0.23)
2f3.31 1.42 3.75 6.70 14.8 3.27
(0.16) (0.01) (0.02) (0.11) (0.603) (0.14)
4m21.9 3.34 7.29 14.12 121.9 30.1
(2.2) (0.05) (0.09) (0.31) (6.48) (1.60)
4r28.7 3.58 8.02 15.22 158.7 39.84
(1.4) (0.04) (0.11) (0.28) (6.38) (1.60)
5 m 32.2 4.71 10.22 19.47 255.6 64.4
(2.4) (0.09) (0.17) (0.58) (13.1) (3.30)
5 f 42.8 5.19 11.33 21.27 320.5 81.4
(2.5) (0.11) (0.25) (0.43) (15.4) .(3.90)
a m 33.5 6.06 14.39 26.88 498.5 134.6
(2.9) (0.11) (0.18) (0.53) (18.5) (5.00)
af66.8 6.96 16.50 29.02 749.7 230.9
(6.0) (0.08) (0.21) (0.57) (38.0) (I 1.7)
4'1
Table A41. Number or antennal segments or the diiJerent stages or
sexes of six species of grasshoppers
Species Stage Sex Mb. Mode Max. Speries Stage Sex Min. Mode Max.
M. in! antili, C.pellucida
1 m 11 12 13 1m,f 10 10 11
1r11 12 13 2m,f 10 13 14
2m14 15 16 3 m 14 17 18
2 f 14 15 16 3 f 15 16 18
3m17 19 19 4 m 17 19 19
3r17 18 20 4f17 19 20
4 m 19 20 22 5 m19 20 21
4 f 19 20 21 5 f20 21 22
5 m 21 22 23 a m 20 23 23
5 f 20 21 22 af20 22 23
a m 22 23 24
a f 22 23 24
M.Banqu;nipe, M. packard;;
1m,f 11 12 13 1m12 13 13
2m,f 14 15 16 1f11 13 13
3m,f 17 18 20 2 m15 17 19
4 m 19 20 21 2 f 16 17 17
4 f 19 20 22 3m18 19 21
5 m 22 22 24 3f 18 19 21
5 f 21 22 23 4 m21 22 23
a m 21 23 25 4 f 21 22 22
ar21 23 26 5m20 23 24
5r21 23 24
am22 25 25
ar23 25 27
M. b;t,';ttatu8 D. carolina
2m15 16 17 1 m10 11 13
2r16 16 18 1 f11 11 13
4 m 21 21 26 2 m 13 14 15
4f20 21 23 2 f12 14 15
5m23 24 25 3 m16 18 19
5r'22 24 24 3r17 18 20
a m 23 24 26 4 m19 21 22
ar23 24 28 4 r19 21 22
5m22 23 24
5r20 23 24
am23 25 25
af23 25 25
"
49
... Holmbeq~, R. G. and J. M. Hardman. 1984. Relating feeding rates to s"x and size in 6 species of grasshoppers (Orthoptem: Acridicla••). Canadian Entomologist 116: 597-606," Depository of Unpuhlish ••d Data, CISTI, National Research Council of Canada, Montreal Hoad Campus, Ottawa, ON, KIA OH6, Canada. E-mail CISTl.INFO@NHC.CA models of pest phenology and crop production to simu ...
... Available feeding rates for grasshoppers are either singletemperature values or averages over varying temperatures (Barnes 1955, Smith 1959, Misra and Putnam 1966, Mitchell and pfadt 1974, Hewitt et al. 1976, Hewitt 1978, Holmberg and Hardman 1984, Johnson and Pavlikova 1986. These results are of limited value because insect feeding rates are temperature dependent (for example, Hammond et al. 1979, Ferro et al. 1985, Logan et al. 1985, Trichilo and Mack 1989, Lactin et a1. ...
... The trend in feeding rate optima might be an evolutionary response to this apparent constraint on thc ability to increase body temperature. Holmberg and Hardman (1984) measured fceding rates of M. sanguinipes at 30.6°e. Converted from 24-h totals, their mean feeding ratc measurements were 0.04, 0.10, 0.14, 0.40, and 0.51 mg/h for 1st to 5th instars, respectively (R. G. Holmberg and J. M. Hardman, unpublished data 2 ). ...
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Feeding rates of laboratory-reared Melanoplus sanguinipes (F.) nymphs were measured in 3-h trials at 15, 20, 26, 30, 35, 40, or 45°C. Feeding rates increased with temperature to a maximum, then decreased at higher temperatures. Maximum feeding rate occurred at 35°C in 1st and 2nd instars, and at 40°C in 3rd, 4th, and 5th instars. In a subsequent 24-h trial, mean hourly feeding rates at 30, 35, or 40°C agreed with those of the 3-h trials. A linear model was used to describe feeding rates at temperatures at or below that at which rate was maximum, and a nonlinear model was used to describe the feeding rate trends over the full range of experimental temperatures. Using the linear model, estimated low-temperature feeding thresholds ranged from 13.5°C for 2nd instars to 15.9°C for 1st instars. Using the nonlinear model, estimated low-temperature feeding thresholds were near 13°C for all but 3rd instars, for which it was near 8°C; modeled temperatures at which feeding was maximal agreed with measured values, and the modeled upper temperature threshold for feeding was between 45 and 46°C for all instars. The linear and nonlinear models are similar at temperatures below that at which feeding is maximal, but the linear model overestimates feeding at higher temperatures. Application of the feeding rate functions to field conditions requires improved understanding on the determinants of body temperature under field conditions, and on the relationship between feeding and wastage.
... The body weight of the females of the three species used in the present study was greater than that of the males (Table 1). Holmberg & Hardman (1984) report a strong correlation between food consumption and body weight in six species of grasshopper. Moreover virgin females of most species of Orthoptera produce a few eggs, which have a lower hatching success and the hatchlings suffer a higher mortality than those produced by mated females (Schmidt & Albutz, 1994;Chapman, 1998;Walker et al., 1999). ...
... How-ever, it is also important to have a measure of the population density (number of individuals/m 2 ) of the orthopteroid species (Hewitt & Onsager, 1983) and mean environmental temperature for an accurate estimate of crop damage. In addition to the laboratory data on consumption, for the estimation of the potential yield losses due to Orthoptera, other insect-caused damage, such as cutting off of plant parts without consuming them (Holmberg & Hardman, 1984) and the fact that Orthoptera can increase their body temperature above ambient by means of thermoregulation (Begon, 1983;Willott, 1997), need to be taken into consideration. ...
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In the Mediterranean area the orthopterans Calliptamus barbarus barbarus, Dociostaurus maroccanus and Tettigonia viridissima are considered to be pests of major economic importance crop plants. The aim of the present study was to evaluate the area of leaf of vine, cotton and potato plants consumed by these Orthoptera at two temperatures. Leaf area was determined for each plant species before the leaves were provided to the insects. Virgin male and female adults were allowed to feed on a leaf of each plant species and the area of leaf consumed was determined after 24h at 25 and 30°C. Results showed that temperature had a significant affect on consumption by both males and females of all the species examined, and it was higher at 30°C. The influence of plant species was not significant for C. barbarus barbarus. D. maroccanus and T. viridissima, however, consumed a greater area of potato leaves than of the other species of plants. Females consumed more food than males in all the species tested. The results of the present study provide a first approximation of the area of leaf consumed and the potential loss of foliage these plants might suffer if attacked by the three species of Orthoptera tested.
... sanguinipes), in most cases approximatedly linearly. The power relationship (b = 0.68) of consumption as a function of body weight described from previous studies (Holmberg & Hardman, 1984) is a manifestation of the common relationship between body size and age : log (size) = a + b log (age), O < b < 1 (e.g., see Johnson & Wellington, 1980). Within an age group, the relationship of consumption to body weight is generally linear, variable, and exhibits a small positive slope. ...
... Consumption of safflower averaged 64 mg dry weight per day, comparable to amounts measured on other food plants ( Table 1). Holmberg & Hardman (1984) estimated the rates of consumption of wheat by adult C . pellucidu, M . ...
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Laboratory experiments were conducted to measure the feeding rates, preferences, survival rates, maturation times, and weight gain of four common species of grasshoppers (Orthoptera: Acrididae) on four lines of safflower (Carthamus tinctorius). When safflower lines were presented individually, all were readily consumed although consumption differed significantly according to grasshopper species, age and sex, and to safflower line. Consumption was greatest on a safflower line devoid of spines. However, grasshoppers also fed readily on spiny lines, and results of experiments in which all safflower lines were presented together indicated preferences that did not necessarily correspond to the results of the single-line feeding trials. Camnula pellucida, a graminivorous species, did feed on safflower, but restricted feeding almost exclusively to stem cambium, whereas Melanoplus bivittatus, M. packardii and M. sanguinipes fed mainly on the heads, flowers and leaves.Highly significant differences in grasshopper maturation time and weight gain were apparent among grasshopper species, and among safflower lines. Seedtec-5, the line which was least digestible, least consumed and least preferred nevertheless yielded rapid development and the greatest body weights of grasshoppers. All species of Melanoplus developed more quickly and gained more weight on safflower than on wheat. One species, M. packardii, was more able to utilise safflower than its congeners, and may gain competitive advantage if safflower becomes widely grown.
... The amount of food consumed by grasshoppers varies with both plant-specific (e.g., quality, presence of antifeedant compounds, and water content of food plants) and individual insect specific attributes (e.g., length of food deprivation, sex, instar, and volumetric changes in the gut) (Holmberg and Hardman 1984;Johnson and Mündel 1987;Chapman and Sword 1994;O'Neill et al. 1994). In addition, infection with entomopathogens has been found to reduce feeding by grasshoppers and locusts (Johnson and Pavlikova 1986;Olfert and Erlandson 1991;Moore et al. 1992;Sieglaff et al. 1997;Thomas et al. 1997). ...
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Blaesoxipha atlanis (Aldrich) is a common parasite of Melanoplus sanguinipes (Fabr.) in western Canada. We tested the hypothesis that parasitism by B. atlanis reduces food consumption by adult M. sanguinipes. Unparasitized grasshoppers serving as controls and grasshoppers infected with a single parasite larva were fed known quantities of freshly cut wheat (Triticum aestivum L. ‘Katepwa’) (Poaceae) leaves in the laboratory. The median development time in hosts of larvae of both male and female B. atlanis was 5.5 days. Two thirds of parasitized grasshoppers died within 9 days of infection, but all control insects survived. The dry mass of leaves consumed each day did not differ between parasitized insects that died and insects that survived parasitism; both groups fed less than unparasitized controls. The influence of parasitism on food consumption differed between host sexes, with feeding being depressed earlier and more severely in female than in male grasshoppers. The reduction in food consumption was most pronounced on day 6 after infection, when parasitized males and females consumed only 10% and 7%, respectively, of the food consumed by unparasitized controls. Parasite sex did not influence food consumption. Grasshoppers that survived parasitism by B. atlanis resumed feeding, consuming as much as unparasitized counterparts. Reduced food consumption limited the ability of grasshoppers to compensate for the nutritional demands of developing parasite larvae. As a consequence, parasitized grasshoppers lost body mass during the interaction. We propose that the temporary reduction in feeding by grasshoppers parasitized by B. atlanis that survive parasitism is not evidence of host regulation, but is consistent with a stress-induced alteration in host behaviour.
... With regard to male beetles, the feeding responses to a great extent resemble those of their female congeners. In spite of the similarity appeared by the current results, Nagy (1952) and Gangwere (1959) reported that adult male grasshoppers may eat up to twice as much as adult females, while Holmberg and Hardman (1984) reported that the females proportionately eat more than males. The most likely explanation for this controversy is the variation in feeding rates during the egg-laying cycle of females and the mating condition of males. ...
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In the Mediterranean area the orthopterans Calliptamus barbarus barbarus, Dociostaurus maroccanus and Tettigonia viridissima are considered to be pests of major economic importance crop plants. The aim of the present study was to evaluate the area of leaf of vine, cotton and potato plants consumed by these Orthoptera at two temperatures. Leaf area was determined for each plant species before the leaves were provided to the insects. Virgin male and female adults were allowed to feed on a leaf of each plant species and the area of leaf consumed was determined after 24h at 25 and 30°C. Results showed that temperature had a significant affect on consumption by both males and females of all the species examined, and it was higher at 30°C. The influence of plant species was not significant for C. barbarus barbarus. D. maroccanus and T. viridissima, however, consumed a greater area of potato leaves than of the other species of plants. Females consumed more food than males in all the species tested. The results of the present study provide a first approximation of the area of leaf consumed and the potential loss of foliage these plants might suffer if attacked by the three species of Orthoptera tested.
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First-instar grasshoppers infected with Melanoplus sanguinipes entomopoxvirus consumed 25% less food within 5 d (75 degree-days (DD), base 10°C) after inoculation with 5 × 103 virus occlusion bodies. Grasshoppers ate 25 and 40% less food by the end of the second week (200 DD, base 100°C) after being infected as first instars with 5 × 102 and 5 × 103 occlusion bodies per individual, respectively. When infected as third instars with 1 × l04 and 1 × 105 occlusion bodies per individual, grasshoppers ate 20 and 35% less food by the second week of the study, respectively. The mean amount of food consumed by infected grasshoppers over the course of the study was 36, 31, 76, and 57% that of noninfected grasshoppers at dosages of 5 × 102 or 5 × 103 occlusion bodies per first instar, and 1 × 104 or 1 × 105 occlusion bodies per third instar. This study suggests that an application of entomopoxvirus to first-instar grasshoppers will reduce their damage potential by causing initial mortality, by reducing the amount of plant material consumed by the surviving grasshoppers, and by preventing grasshopper maturation and reproduction (at higher doses). The negative influence of entomopoxvirus on grasshoppers in the interim between infection and death is integral to future evaluations of its potential as a crop protection measure.
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To determine the effectiveness of montane barriers to gene flow of the two-striped grasshopper, Melanoplus bivittatus, collections were made at four sites at the corners of a square location, each bisected by one of five Colorado mountains (total: five locations, 18 sites). The premise of this design is that samples from opposite sides of an effective barrier should have more genotypic differences than samples from the same side. Genotypic differences were measured for each grasshopper as 18 morphological measurements and six enzyme loci. Dissimilarities in morphology or enzyme loci were expressed as significant estimates of Mahalanobis squared distance (respectively, D2 or G2) between two sample sites. The presence of foothills had no influence on the number of significant G2 (age and sex classes combined) or significant D2 (age and sex classes separated). For samples near the foothills, adult females had more significant D2 than adult males. For samples near the two higher mountains, adult males had more significant D2 for site samples from opposite sides. A combination of gene flow and selection is the most plausible explanation for most of these results.
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The effect of ingestion of Nosema locustae Canning spores on feeding by grasshoppers was measured in simultaneous laboratory and field experiments. After 21 days, fourth-instar Melanoplus sanguinipes (F.) nymphs, administered spores at the rates of 0, 2.0 × 104, 2.0 × 105, and 2.0 × 106 per grasshopper, showed dry matter consumption of 102, 87, 64, and 26 mg in 48 hr, respectively. Rate of inoculation was a significant factor in suppression of feeding after correction for the effects of developmental stage, sex, and body weight. The quantity of dry matter consumed decreased linearly with increasing rate of spore ingestion. Experiments on 50 caged 1-m2 plots on pasture grass yielded similar trends in per capita consumption independent of the effects of mortality. Field consumption per integrated grasshopper-day was 108, 77, 31, and 27 mg dry wt at the four inoculation rates, over 20 days.
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Dichroplus maculipennis (Blanchard) and D. elongatus Giglio-Tos are two of the most important melanoplines in Argentina, both ecologically and economically. The postembryonic development and forage loss (consumption of Bromus brevis Ness + fallen material) caused by older nymphs (instars IV, V, VI) and adults of both species were studied under controlled conditions (30ºC, 14L:10D, 40% RH). Five nymphal instars were recorded in D. elongatus, and six in D. maculipennis. Total nymphal development was similar in both species (D. elongatus: 32 ± 0.70 days; D. maculipennis: 34.5 ± 0.37 days). Daily consumption increased from nymphal instars to pre-reproductive adult stage. In both species, pre-reproductive females had higher consumption rates than other stages considered (D. elongatus: 30.6 ± 0.56 mg dry weight/day; D. maculipennis: 48.7 ± 0.74 mg dry weight/day). In the reproductive stage, consumption decreased significantly in both sexes. When feeding, D. maculipennis let some plant material to drop, increasing total loss. The percentage of fallen material was greater in reproductive adults, representing 3.9% and 2.9% of the total daily loss for males and females, respectively. Females and males of D. maculipennis were heavier than those of D. elongatus (P < 0.05), and daily consumption was significantly higher (P < 0.05). Regardless sex and reproductive status, adults of D. maculipennis consumed 29.1 ± 0.64 mg dry weight/day on average, while one of D. elongatus 20.0 ± 0.3 mg dry weight/ day.
Article
Sweep-net samples of grasshoppers were taken annually in late August at Stavely (1970-78) on Festuca scabrella prairie and at Coalhurst (1971-79) on Stipa-Agropyron prairie. Mean catches of grasshoppers were higher (170 vs 112 per 50 sweeps) and more species were sampled (27 vs 13) at Coalhurst. Melanoplus dawsoni (Scudder) was the dominant grasshopper at Stavely while Encoptolophys sordidus costalis (Scudder) and Melanoplus infantilis (Scudder) were codominants at Coalhurst. Grasshoppers were also sampled at two other sites in 1971 and one in 1971 and 1972 on Stipa-Agropyron prairie. Mean catches per 50 sweeps were 122, 164, and 234, respectively, at these sites with 14, 12, and 11 species of grasshoppers sampled. The dominant species were Ageneotettix deorum (Scudder), M. infantilis, and M. dawsoni. Of the 35 species collected at the study sites, 21, those forming at least 1% of the grasshoppers collected at one or more sites, were evaluated for their potential impact on rangeland. Population counts and published data on phenology, damage to rangeland, and feeding preferences were considered. The per capita feeding rate of adults-assumed to be proportional to the 0.68 power of body weight-was also assessed. Using these criteria, all but two species-Melanoplus femurrubrum femurrubrum (DeGeer) and M. dawsoni-were considered potentially damaging. Adult weights varied such that an adult M. infantilis, the smallest species, would feed at 28% the rate of an adult Metator pardalinus (Saussure), the largest species. Published data on habitat preferences of the 21 species show that most of the damaging species prefer sparsely vegetated habitats and thus would be favored where range is overgrazed by cattle.
Book
Brody, S. (1945) Bioenergetics and Growth. Reinhold, New York.
Article
1. The Sardine seems to strain plankton, by swimming with opening mouth and further controlling the swimming velocity and the diameter of the mouth, so as to make the velocity of current on the gill-rakers the optimum velocity (about 2-3cm./sec.). Therefore, the feeding velocity of the sardine seems to be proportional to the surface of the gill-rakers, when the plankton dencity in the water is constant. 2. By whether the time, which the gill-rakers require to be closed by plankton, is short or long, the sardine seems to recognize the plankton density in the water where it is feeding, in a short time (about one minute). 3. The secondary concentration for the plankton gathered by the gill-rakers in the oral cavity seems to be performed by “the pharyngeal pockets” and “the pharyngeal rakers”.