FOOD PLANTS OF FEMURRUERU?' ^'? URRUBRUM (DEGEER) THE BLUESTEM GRASS REGION OF KANSAS

by

ORLO KENNETH JANTZ I

B. S., Kansas State University, 1957

A MASTER 1 S THESIS

submitted in partial fulfillment of the

requirements for the degree

MASTER OF SCIENCE

Department of Entomology

KANSAS STATE UNIVERSITY Manhattan, Kansas

1962 LD 2(ofc8 T^ \%a ii cm TABLE c £ OF CONTENTS

INTRODUCTION 1

REVIEW OF LITERATURE 2

MATERIALS AND METHODS 5

RESULTS AND DISCUSSION 10

SUMMARY 22

ACKNOWLEDGMENTS 77

LITERATURE 78 INTRODUCTION

The species of in a given area and their populations are determined to a considerable extent by food plants. Food prefer- ences and relationships of species and plant species in pastures of the Great Plains are not well understood.

Cage studies on plant food preferences are one of the current phases of the grasshopper project which has been in progress since

1957 and is part of Regional Project NC-52 on factors influencing the distribution and abundance of grasshoppers. The Kansas portion of the project involves a study of a series of habitats within a bluestem grass range, the Donaldson Pastures, near Manhattan, Kansas. Nine pastures, each under different experimental treatments, are studied.

Three of these pastures are included in an intensity-of-grazing treat- ment, three in a deferred grazing treatment and three in a time-of- spring-burning treatment. Range sites *ithin each of the nine treat- ments are: limestone breaks, ordinary upland, and clay upland. Axnett

(i960) was the initial investigator. Largest grasshopper populations were found on the early spring burned and the heaviest grazed pastures.

Lightly grazed pastures and those grazed under a def erred-rotation practice had the smallest populations. Clay upland sites, and to a slightly lesser extent ordinary upland range sites, supported larger populations than the limestone breaks. His study was intended to be exploratory in nature. A better understanding of grasshopper behavior, particularly in regard to feeding habits and preferences, and associa- tion with plants, was considered the next step.

This report describes results of cage studies on plant preferences of Melanoplus femurrubrum femurrubrum (DeGeer), conducted during the growing season of 196l. Two other species were studied: Melanoplu s differentials (Thomas), and Phoetaliotes nebrascensis (Thomas).

Further data for M. _f. femurrubrum (De(5eer), not presented here, along with that for the other two species, will be presented as a minor portion of the Ph.D. dissertation when more data are available.

REVIEW CF LITERATURE

Literature accounts of grasshoppers are probably more numerous

than for any family except possibly mosquitoes.

Although much of the literature includes general references to adult lists and associated habitats, relatively little exists on

explanation of relations of habitat to species distribution. Specific

studies on association between grasshopper species and range plant

species are limited in Kansas and occur only to a limited extent else-

where, oodruff (1937) did a survey of the grasshoppers inhabiting

the native grasses in Kansas. Shotwell (1939) summarized the species

according to habitats. A'ilbur (1936) reported on the injury to the

heads of native grasses and Wilbur and Fritz (19^0) studied the popula-

tions present in typical pastures of the bluestem region of Kansas.

Smith (195*0 assembled and analyzed information on annual fluctuations

of grasshopper populations in Kansas over a 100 year period from 185*4-

to 1954. One conclusion of his study of the relation of climate to

grasshopper populations was:

While the numbers of grasshoppers each year must be dependent in some way on the kind, amount, and quality of natural food available to them and their parents, modified directly by the weather as from dashing rains and by the extremes of temperature, and indirectly as weather affects plants, parasites, predators and diseases, the data avail- able on food do not permit a correlation to be made with grasshopper populations. It is generally accepted that plants, directly or indirectly,

influence the size of grasshopper populations. Isley (1957) stated

that plant distribution and the extent and vigor of plant growth are definitely soil-related, and soil make-up is a determining factor in

the choice of egg laying sites by many species. He therefore consid-

ered soil as the primary controlling environmental factor in local distribution and plants and vegetative soil cover, as they are related

to food, protection, temperature, and humidity in the microhabitats, as indirectly significant. The plants chosen as food vary between different species of grasshoppers, and all the plants present in any chosen location are not necessarily used as food plants. The old belief that grasshoppers will devour everything that is green has been accepted by many authors but was originally applied only to the

"Rocky Mountain locust", Kelanoplus spretus V.alsh, (Riley, et al.,

1877). work by Criddle (1933). Isley (1937. 1938, 19^, 19*f6), Hodge

(1933) » and Sanderson (1939) . showed that a large percentage of grasshoppers were associated with a restricted number of host plants.

Other investigations on relationships of habitat to grasshopper dis- tribution and host plant associations to various degrees include those of Hebard (1925. 1929. 193L 193^a, 193'+b, 1936, 1938), Uvarov (19?3),

Cantrall (19^3). Pfadt (19^9), Friauf (1953). Shotwell (1930, 1938),

Ball et al. (19^2), Anderson and bright (1952), Barnes (1955). and

Wakeland (1958). Some recent work, part of which is still in progress, includes population studies on Arizona desert, range, and cultivated land by Barnes (1959. 19&1), and effects of grasshopper and management practices on short-grass rangeland (Nerney 1959. 19^0, 196l). Using

the technique of crop analysis, (Mulkern and Anderson, 1959. Brusven and Mulkern, i960) Mulkern (i960, 196l) in North Dakota and Pruess

(i960, 196l) in Nebraska are currently involved in food habit and

preference studies with various grasshopper species.

Factors that affect a grasshopper's selection of host plants,

from the many available, are too numerous to mention. Smith (1959)

found that certain plants which are eaten are nutritionally inadequate

to certain grasshopper species. Painter (1953) states that plants

and different parts of the same plant may differ nutritionally and

implies that this may be a possible explanation for some resistance

in plants. In grasshoppers, however, the resistance seemed to be a preference phenomenon. A majority of the work on breeding for re-

sistance has been and is being done with field crops (Painter 1951*

1953* I960, 1961). Diver and Diver (1933) noted that degree of wet-

ness, vegetation height, density and type of plant community are among

the factors determining the distribution of species. Correlation between mandibular morphology and food plants (Isley, 19^) is important as an indication of general food habits. In certain cases

a grasshopper species will feed on one plant, and during later de-

velopment due to availability of plants or other factors, change food plants. This was the case with Melanoplus bruneri as reported by

Kreasky (i960). Nymphs and adults fed heavily on lupine and timothy.

Damage to timothy became especially noticeable as lupine became depleted.

Species differ in the amounts of food required. When considering

economic importance of a particular species, amounts of food eaten by

each individual must be considered. Gangwere (1959) noted that food

consumption increases in direct proportion to size during the nymphal

stages. Mulkern (l96l) working in this area has developed a Plant

Value Index based on palatibility of plants to cattle. 5

Many grasshopper species confine their feeding to a group of re- lated plants and in a few known cases to a single plant species. As an example* Hypochlora alba (Dodge) is often cited as feeding only on

Arteme3ia spp. Isley (1938) * however, states that in cage tests

H. alba will live for ten days supplied with broomweed and sunflower.

It has been stated before, but bears repeating, that an understanding of the host plant relationships of all major species would contribute valuable data toward an understanding of terrestrial communities.

The work reported on here is considered a first step in obtain- ing clues as to preferred plant species which might be used in a correlation that exists in the association between grasshopper and plant species, particularly in the bluestem regions of Kansas.

MATERIALS AND METHODS

The initial evaluation of the associations between plant species and grasshopper species was approached in the following manner. The population density of each species of plant from each pasture treat- ment and range site within each treatment was obtained from the De- partment of Agronomy and corresponding data on population of each grasshopper species from Arnett (i960). Evaluation was made of each grasshopper species present in relation to numbers of each plant species present on (l) various soil types and pasture treatments;

(2) the same soil type irrespective of pasture treatments; and (3) the same pasture treatment irrespective of soil type. Using a statis- tical formula, "Kendall's Tau" (Figure l), (Siegel, 1956), rank corre- lation coefficients were determined. The and plants are ranked according to relative abundance of each species after combining collections of each of the three soil types for each of the nine pas- tures. For example, hy looking at Figure 1, plant rank 8th is opposite ranked 1st. In the plant rank row only one plant r?nks higher than 8th (9)» so "1" is placed in the concordance row. Next, plant ranked *tth corresponds to insect ranked 2nd, and k is exceeded k times

(hy 9» 6, 7, 5), so "4" is recorded in the concordance row, and so on.

Thus concordance is the number of ranks higher than the one being ranked. Discordances, on the other hand, are the number of plants in the row ranking lower than the one being ranked. There are 7 numbers below or less than 8 (4, 6, 2, 1, 7» 5» 3) so "7" is recorded as the discordance of the plant ranking 8th. Next, (for the plant ranked ^th) there are 3 numbers smaller than k (2, 1, 3)f so "3" is recorded in the discordance row, and so on. As a check for errors, the following procedure is useds total concordance plus total dis- cordance should equal l/2N(N-l) where N is the number of pastures in which you have observations. Thus in the example in Figure 1, 12 +

2k a l/2 x 9 x 8 or 36. Plates I - VI are submitted as examples of the rank correlation coefficients for the various treatments and soil types as they are plotted for each plant and grasshopper species.

Coefficient range is from zero to a plus 1.0 or a minus 1.0. The plates are limited to plus or minus 0.9i however, since this was the highest correlation observed for any species, retailed report on the correla- tion studies will be given at a later date in the Ph.D. dissertation in connection with a proposed crop analysis study. It ia submitted here only becruse it was one of the methods used as a guide in determining which plants to use in food preference work. 7

Cage studies were conducted in an outdoor insectary which could be opened from three sides, thereby coming relatively close to outdoor temperatures (Plate VII). A water cooler was installed to maintain temperatures when necessary below 100° F. Each cage was composed of six compartments, Ik* x 11 lA" x 9 lA", with removable glass fronts

(Plate VIII).

The following plants were used (common names are those recommended by Anderson, 196l)»

Perennial Grasses!

Asm

Age

Asc

Bcu Bouteloua curtipendula (Michx.) Torr. - sideoats grama

Kcr Koeleria cristata (L.) Pers. ----- prairie junegrass

| sc

Pvi

Snu

Scr Sporobolus cryptandrus (Torr.) A. Gray- sand dropseed

Perennial Forbs:

fcla Achillea millefolium L. subsp. lanulosa (Nutt.) Fiper ----- western yarrow

Aps

Artet

Ave Asclepias verticillata L. ------whorled milkweed

Astei

Keu

Lpu 8

Ost Oxalis stricta L. ------common yellow oxalis

Ppu Petalostemum purpureum (Vent.) Rydb. - purple prairieclover

Pfl Psoralea tenuiflora Pursh var. floribunda (* ! utt.) Rydb. ----- manyflower scurfpea

Rci Ruellia humilis Nutt. ( "?. cilio3a of manuals, in part; R. carolinlensls of manuals, in part) ------fringeleaf ruellia

Sun Schrankia nuttallii (D.O.) Standi. (S. uncianata of manuals, not

villd. ) _----_--__-_ catclaw sensitivebriar

Sin Silphium integrifolium 7'ichx. (including 3. 3pecio3um Nutt.) - - wholeleaf rosinweed

Vba Vernonia baldwini Torr. ------baldwin ironweed

Annual Forbs:

Ael Ambrosia elatior L. ------common ragweed

Eca Erigeron canadensis L. ------horseweed fleabane i.oody Plants:

Aca Amorpha canescens Pursh ------leadplant amorpha

Cov Ceanothus ovatus Desf . ------inland ceanothus

The cage floors were lined with tinfoil and sand was placed in each, forming a slope approximately 1 inch deep in front and k inches deep toward the rear.

The plants were placed in water-filled plastic vials through a hole in the cap. The water kept the plants from wilting during the feeding period. Each vial was inserted into the sand equidistant from the screen, until the lip was level v.ith the sand surface. This gave each grasshopper an equal chance to walk directly from the sand onto the chosen plant. Water for the grasshoppers was supplied by soaked cellulose cotton placed in petri dishes. Twenty grasshoppers were placed in each compartment. An attempt

was made to keep instars of the same age togetner when nymphs were

studied. A spread of two consecutive instars was allowed in each

cage} i. e., 2nd and 3rd instar nymphs in one cage, 3rd and 4th in

another, etc. Since females of M. f. femurrubrum (DeQeer) may be

easily confused with other Melanoplus some of the specimens were sent

to the II. S. National Museum for confirmation.

After some preliminary experimentation and consideration of the

various amounts of food required by different species, it was decided

to use only two plant species in each cage. The question arose con-

cerning the validity of this technique since in some instances, grass- hoppers might be forced to make a choice between two undesirable plants. It is believed, however, that the results are probably valid since an attempt was made to pick out the most obvious desirable or undesirable plant species based upon the before-mentioned association studies. Changes will be made in subsequent studies, allowing more choices.

The grasshoppers were allowed to feed for 48 hours on each set of two plants. After 48 hours the plants were removed and replaced by another series of two plant species. Three replicates of each series were included. Species of grasses vs. grasses, and grasses vs. forbs, were evaluated. Photographs were taken of the plants immediately after removal from the cages. (Plates IX - XV). These plates are only representative to show the type of damage done; however, a complete photographic record of all three replicates was taken. The photographs were ranked according to the intensity-of-f eeding as compared with before-feeding pictures. A grade of A, B, C and D was used: A = no 10

feeding! B = trace to l/2 of plant eaten; C = l/2 plus eaten; D • eaten entirely. This intensity-of-f eeding (IF) was graded for each series of plants. Vhen replacement of a plant was necessary before the end of

the ^8 hour period, that replicate was automatically rated a "D", how-

ever, the plant was replaced to keep from forcing the grasshoppers onto

the second, less preferred plant.

Counts of total numbers of grasshoppers resting (R) on the plants,

and whether or not they were feeding (F) were made at 7 a.m., 12 noon,

^8-hour feeding period. 5 p.m., and in some cases 10 p.m. during the

Feeding was recorded when movement of the mouthparts was observed

when in contact with the plant, and included in certain instances

case3 when the grasshopper was merely nibbling and did not leave any

visible damage. A high resting count without feeding on a particular

plant may indicate an association other than food preference. Tem-

peratures were not recorded in the first few experiments. In subsequent

studies room temperature was recorded during each observation. Results

of counts are presented in tables 1 through 7. The abbreviations

across the top represent the first letter of the genus and the first

two letters of the species of plant, i. e., "Age" = Andropogon gerardi.

Plants and grasshoppers were obtained directly from the Donaldson

Pasture area.

RESULTS AND DISCUSSION

Feeding data, which include degree of injury based upon (l)

photographs of feeding on each plant and (2) numbers of grasshoppers

resting and feeding on each plant, are presented in tables 1 through

7. Considerable difference is apparent between counts made for a

given species on a given plant at various times during the day. When 11

both plants read zero at a given time, all grasshoppers were perched either on the sand floor or screen sides of the cage.

Intensity-of-feeding (IF), as graded at the bottom of each series, generally match the resting and feeding counts. In certain cases a plant was graded "A", indicating from the photograph that it was appar- ently untouched, yet the count showed one or more cases of feeding. In other cases, little or no feeding was observed, yet the photographs indicated damage, Indicating that feeding occurred at other times than when the counts were made.

Table 1 shows the results of big bluestem (Age), compared to 23 other plants, 11 of which were preferred over big bluestem as indicated by the higher resting and feeding counts during most individual obser- vations. Each replicate was completely consumed in four instances

(grade "D"). The average numbers of grasshoppers resting and feeding respectively on these four plants as compared to big bluestem were J aster (Aster) (9.0 vs. 1.0; 5.2 vs. 0.?); horseweed (Eca) (8.9 vs. 1.4;

4.2 vs. 0.4); common ragweed (Ael) (5«7 vs. 2.2; 2.6 vs. 0.1); and scurfpea (Pfl) (3*6 vs. 0.2; 1.6 vs. 0.0). The average numbers resting and feeding respectively on each of the remaining seven preferred plant species as compared to big bluestem, when based upon grading of injury, werei wholeleaf rosinvteed (Sin) (7*4 vs. 0.4; 4.8 vs. 0.2); western yarrow (Ala) (6.4 vs. 0.4; 2,2 vs. 0.0); sagewort (Artem) (5«4 vs. 2.4;

2.7 vs. 0.7); purple prairieclover (Fpu) (4.7 vs. 0.2; 2.2 vs. 0.2); western ragweed (Aps) (4.5 vs. 0.7? 2.8 vs. 0.3); leadplent amorpha

(Aca) (3.4 vs. 0.2; 0.8 vs. 0.0); and sand dropseed (Scr) (2.8 vs. 3»2;

1.8 vs. 1.0). Equal damage based upon grading compared to big bluestem was observed on fringeleaf ruellia (Rci), catclaw sensitivebriar (Sun), 12 baldwin ironweed (Vba), indiangrass (Snu), 3cribner panicum (Psc), switchgrass (Pvi), and dotted gayfeather (Lpu). Big bluestem was pre-

ferred over falseboneset kuhnia (Keu) , inland ceanothus (Cov), prairie junegrass (Kcr), little bluestem (Asc), and sideoats grama (Ecu). In no case was a "D" assigned to big bluestem and only two were rated

"C+".

Vith reference to individual feeding observations, there was no definite time or sequence during which feeding occurred. However, it was obvious that fresh plant material was preferred. This accounts for the higher number present during the 5 p»m. count at the beginning of each new series. New plants were usually placed in the cages between

3 and 5 p»m. They were vigorously attacked as soon as the cage was left undisturbed. This was especially apparent in the 3 J>* m * counts on the highly preferred plants such as common ragweed (Ael), horseweed

(Sea), aster (Aster), and scurfpea (Pfl) (Table 1). Undesirable plants were also attacked shortly after being placed in the cage; however, they sustained little or no injury as indicated by the intensity-of- feeding (IF) as graded from the photographs. In most cases grasshoppers went directly to the desired plant; however, in a few instances it was observed that an individual grasshopper would jump onto a plant and shortly leave the plant without feeding and move to the other plant where vigorous feeding occurred, resulting in severe damage.

High counts which occurred at other times during the day, for example, 5 p»m« the second day for horseweed (Eca) and 7 a.m. for aster

(Aster), in most cases were the result of the plant being consumed and replaced in one or more of the three replicates at that time.

Temperatures in Table 1 do not apparently show any trend or association with either a high or a low incidence of resting or feeding. 13

Subsequent tables do show, however, that at 60° F. or below, feeding was severely reduced and in most instances terminated.

Data in Table 1 were concerned with 2nd and 3rd instar nymphs except those on: inland ceanothus (Cov), indiangrass (Snu), prairie junegrass (Kcr), little bluestem (Asc), and sideoats grama (Bcu), where 3rd and 4th instar nymphs were used} and sand dropseed (Scr), scribner panicum (Fsc), switchgrass (Pvi), and dotted gayfeather (Lpu), where adults were used.

Sand dropseed (Scr) as compared to 20 other plants is shown in

Table 2. Grading indicates that eight plants were preferred over sand dropseed. Only one plant in this series, horseweed (Eca), was complete- ly consumed in all three replicates (grade "D"). The average numbers of grasshoppers resting and feeding respectively on these eight pre- ferred plants as compared to sand dropseed were J aster (Aster)

(10.4 vs. 1.8; 5.6 vs. 1.4)} dotted gayfeather (Lpu) (9.0 vs. 1.4;

3.6 vs. 0.6); wholeleaf rosinweed (Sin) (6.8 vs. 0.5; 3.5 vs. 0.5); horseweed (Uca) (5«8 vs. 1.7? 2.3 vs. 0.5); western ragweed (Aps)

(4.4 vs. 1.8; 1.4 vs. 1.0); indiangrass (Snu) (2.8 vs. 1.6; 1.0 vs.

0.8); scurfpea (Pfl) (2.4 vs. 1.0; 0.8 vs. 0.8); and common ragweed

(Ael) (1.4 vs. 3*4; 1.0 vs. 1.4). Equal damage based upon grading compared to sand dropseed was observed on sagewort (Artem), and baldwin ironweed (Vba). Sand dropseed was preferred over inland ceanothus (Cov), falseboneset kuhnia (Keu), purple prairieclover (Ppu), catclaw sensi- tivebriar (Sun), leadplant amorpha (Aca), little bluestem (Asc), common yellow oxalis (Ost), scribner panicum (Psc), prairie junegrass (Kcr), and sideoats grama (Bcu). In no case was a "D" grade assigned to sand dropseed and only one was rated "C" and two a "C+". 14

High average resting counts are noted for aster (Aster) (10. 4), dotted gayfeather (Lpu) (9»0), wholeleaf rosinweed (Sin) (6.8), and inland ceanothus (Gov) (6.5) (Table 2). These four plants (three forbs and one woody plant) apparently were preferred resting sites.

Although the numbers feeding on each increased correspondingly, the degree of injury remained rel tively low.

Temperatures in Table 2 ranged from 48° F. to 99° F« No reduc- tion in resting or feeding is apparent at high temperatures up to

99° • There was some reduction in numbers feeding at 6l° F. although a high count remained for dotted gayfeather (Lpu). Below 54° F. all feeding terminated. Resting counts remained near normal at low temperatures compared to other temperatures, as indicated at 48° F.

Data in Table 2 were concerned with 3rd and 4th instar nymphs except those on: common ragweed (Ael), and leadplant amorpha (Aca) where 4th and 5th instar nymphs were used; and on dotted gayfeather

(Lpu), scribner panlcum (Psc), prairie junegrass (Kcr) , sideoats grama

(Ecu) j indiangrass (Snu), little bluestem (Asc), and common yellow oxalis (Ost) where adults were used.

Switchgrass (Fvi) compared to 22 other plants is shown in Table 3»

Grading indicates that 10 plants were preferred over switchgrass.

Again only one plant in this series, western yarrow (Ala), was com- pletely consumed in all three replicates (grade "D"). A grade is not available for sagewort (Artem) and horseweed (3ca). Accidentally the negatives were exposed to direct light and were not developed. The higher average resting and feeding counts as compared to switchgrass,

(sagewort, k.k vs. 1.2, 2.k vs. 0.4; horseweed, 7.0 vs. 0.4, 0.8 vs.

0.0) indicate that both were preferred. The average numbers of 15 grasshoppers resting and feeding respectively on the 10 preferred plants based on grading as compared to switchgrass were: dotted gay- feather (Lpu) (12.0 vs. 2.8} 4.0 vs. l.O); wholeleaf rosinweed (Sin)

(7.0 vs. 1.7; 2.8 vs. 0.5); western ragweed (Aps) (5.0 vs. 1.5; 1.5 vs.

0.7)5 common ragweed (Ael) (4.8 vs. 1.6; 2.6 vs. 0.6); aster (Aster)

(4.5 vs. 1.8; 1.0 vs. 0.5); western yarrow (Ala) (4.2 vs. 1.2; 0.8 vs.

0.2); scurfpea (Pfl) (4.0 vs. 1.2; 2.0 vs. 0.3); little bluestem (Asc)

(2*8 vs. 3.8; 2.2 vs. 3.0); sand dropseed (Scr) (1.8 vs. 4,2} 1,0 vs.

3.2); and leadplant amorpha (Aca) (1.2 vs. 0.8; 0.7 vs. 0.3) • Equal damage based on grading compared to switchgrass was observed on baldwin

ironweed (Vba), inland ceanothus (Cov) , and scribner panicum (Psc).

Switchgrass was preferred over purple prairieclover (Fpu), falseboneset kuhnia (Keu), catclaw sensitivebriar (Sun), common yellow oxalis (Ost), prairie junegrass (Kcr) » sideoats grama (Bcu), and indiangrass (Snu).

In no case was a MD" grade assigned to switchgrass and only one was rated "C". No feeding was observed on three plants (Table 3): false- boneset kuhnia (Keu), switchgrass (Pvi vs. Eca) and catclaw sensitive- briar (Sun); however, in two cases where a grade is available, injury had resulted. Apparently this was not the effect of temperature since

76° F. was the low and 99° F. the high in this case. The highest rest- ing count was observed on dotted gayfeather (Lpu) followed by horseweed

(Eca) and wholeleaf rosinweed (Sin).

Data in Table 3 were concerned with 3rd and 4th instar nymphs except those on: common ragweed (Ael), and inland ceanothus (Cov) where 4th and 5th instar nymphs were used; and on little bluestem (Asc), sand dropseed (Scr), indiangrass (Snu), scribner panicum (Psc), dotted gayfeather (Lpu), common yellow oxalis (Ost), prairie junegrass (Kcr) and sideoats grama (Bcu) where adults were used. 16

Indiangrass (Snu) as compared to 19 other plants is shown in Table

4. A roll of film was accidentally dropped on a concrete floor causing it to unwind part of the film. Therefore, the degree of injury could not be graded for eight of the plants. However, based on average num- bers of grasshoppers resting and feeding respectively as compared to indiangrass the following seem to be preferred: wholeleaf rosinweed

(Sin) (11.2 vs. 0.2; 9.4 vs. 0.2); horseweed (Eca) (8.6 vs. 0.2; 1.2 vs. 0.2); baldwin ironweed (Vba) (3.6 vs. 0.2; 2.2 vs. 0.2); western yarrow (Ala) (3.4 vs. 0.0; 1.2 vs. 0.0); sagewort (Artem) (2.8 vs. 0.8;

2.0 vs. 0.4); and leadplant amorpha (Aca) (2.2 vs. 0.6; 1.0 vs. 0.4).

Of the 11 plants which were graded, six were ranked with a higher degree of injury than indiangrass. The average numbers of grasshoppers resting and feeding respectively on these six preferred plants based on grading as compared to indiangrass were: aster (Aster) (10.3 vs. 0.2;

3.5 vs. 0.0); 3curfpea (Ffl) (7.5 vs. 1.0; 2.0 vs. 0.2); scribner panicum (Fsc) (4.2 vs. 3«4; 2.8 vs. 2.8); common ragweed (Ael) (2.8 vs.

2.0; 1.8 vs. 1.4); inland ceanothus (Cov) (2.6 vs. 0.8; 1.2 vs. 0.4); and western ragweed (Aps) (1.5 vs. 1.5; 0.7 vs. 0.5). Indiangrass was preferred over purple prairieclover (Ppu), prairie junegrass (Kcr), sideoats grama (Bcu), dotted gayfeather (Lpu), and little bluestem

(Asc). In no case was a "D" grade assigned to any plant in Table 4; however, In the comparison of indiangrass vs. sideoats grama (Bcu),

H indiangrass is graded "C- , and footnoted indicating that one replicate was completely consumed. This replicate was graded "D" yet the average of three replicates remains a "C-". No feeding was observed at 60° F., however, four grasshoppers were feeding at 59° t* below which all feeding terminated. ,

17

Preferred resting sites were wholelesf rosinweed (Sin), aster

(Aster), and horseweed (Eca).

Data in Table 4 were concerned vith 3rd and 4th instar nymphs except those on: common ragweed (Ael), and inland ceanothus (Cov), where 4th and 5th instar nymphs were used; and on little bluestem

(Asc), prairie junegrass (Kcr), sideoats grama (Bcu), scribner panicum

(Psc), and dotted gayfeather (Lpu) where adults were used.

Sideoats grama (Bcu) as compared to l6 other plants is shown in

Table 5. Grading indicates that nine plants were preferred over sideoats grama. Two plants, aster (Aster) and common ragweed (Ael), were completely consumed in all three replicates (grade "D"). In four cases one of the three replicates was completely consumed as indicated

by footnote . These four cases were: purple prairieclover (Ppu) western ragweed (Aps), western yarrow (Ala), and scribner panicum

(Psc). The average numbers of grasshoppers resting and feeding respec- tively on the nine preferred plants based on grading as compared to sideoats grama were: scribner panicum (Psc) (8.2 vs. 2.8; 5.4 vs. 2.2); aster (Aster) (7.8 vs. 1.2; 6.'+ vs. 1.0); common ragweed (Ael) (7.4 V3.

1.6; 6.4 vs. 1.0); inland ceanothus (Cov) (6.2 vs. 1.4; 4.2 vs. 1.4); little bluestem (Asc) (4.2 vs. 4.4; 3.6 vs. 3. 10; purple prairieclover

(Ppu) (3.6 vs. 2.6; 1.4 vs. 0.8); baldwin ironweed (Vba) (2.8 vs. 1.2;

?.k vs. 0.6); western ragweed (Aps) (2.6 vs. 1.6; 1.8 vs. 0.8); and western yarrow (Ala) (1.4 vs. 2.0; 1.4 vs. 1.8). Equal damage based on grading compared to sideoats grama was observed on leadplant amorpha

(Aca), cntclaw sensitivebriar (Sun), common yellow oxalis (Ost), prairie junegrass (Kcr), and dotted gayfeather (Lpu). Sideoats grama was preferred over falseboneset kuhnia (Keu), and sagewort (Artem). 18

In no case was a M D" grade assigned to sideoats grama. Preferred rest- ing sites were scribner panicum (Fsc), aster (Aster), leadplant amorpha

(Aca), dotted gayfeather (Lpu) and common ragweed (Ael). No feeding was observed at 63 F«

Data in Table 5 were concerned with 4th and 5th instar nymphs except those on scribner panicum (Fsc), little bluestem (Asc), prairie junegrass O'er), and dotted gayfeather (Lpu) where adults were used.

Prairie junegrass (Kcr) compared to 17 other plants is shown in over prairie Table 6. Grading indicates that 11 plants were preferred ca^es: junegrass. All three replicates were completely consumed in two common ragweed (Ael) and scurfpea (Pfl). Two plants, aster (Aster) indicating that and western yarrow (Ala) are graded "D+" and footnoted amorpha two of the three replicates were completely consumed. Leadplant indi- (Aca) and baldwin ironweed (Vba) are graded "C-" and footnoted average numbers cating that one replicate was completely consumed. The the preferred of grasshoppers resting and feeding respectively on 11 aster plants based on grading as compared to prairie junegrass were! vs. (Aster) (10.2 vs. 0.4; 6.2 vs. 0.2); leadplant amorpha (Aca) (4.8

vs. 0.0); 0.2; 3.0 vs. 0.2){ western yarrow (Ala) (4.0 vs. 0.0; 2.0

dotted gayfeather (Lpu) (3.6 vs. 0.4; 0.4 vs. 0.0); scurfpea (Pfl)

vs. 0.2); (3.6 vs. 0.4; 1.4 vs. 0.0); horseweed (Sea) (3*0 vs. 0.6; 1.4

baldwin ironweed (Vba) (2.8 vs. 0.4; 0.8 vs. 0.2); common ragweed (Ael)

(2.2 vs. 0.4; 0.4 vs. 0.2); scribner panicum (Fsc) (1.6 vs. 0.8; and 0.0 vs. 0.0); western ragweed (Aps) (1.2 vs. 1.2; 0.8 vs. 0.4);

catclaw sensitivebriar (Sun) (0.2 vs. 1.0; 0.2 vs. 0.6). Equal damage false- based on grading compared to prairie junegrass was observed on junegrass boneset kuhnia (Keu) and purple pralrieclover (Ppu). Prairie ,

19 was preferred over sagewort (Artem), inland ceanothus (Cov), and whole- leaf rosinweed (Sin). In no case was a "E" grade assigned to prairie junegrass. Aster (Aster) and inland ceanothus (Cov) were the two most obvious preferred resting sites. Two grasshoppers were observed feed- ing at 56° F., however, only minor injury resulted. All other feeding in the 50° F. to 60° F. range terminated.

Data in Table 6 were concerned with 3rd and 4th instar nymphs except those on inland ceanothus (Cov), and wholeleaf rosinweed (Sin) where 4th and 5th instar nymphs were used; and on dotted gayfeather

(Lpu), scribner panicum (Fsc), and common yellow oxalis (Ost) where adults were used.

Little bluestem (Asc) as compared to 19 other plants is shown in

Table 7. Grading indicates that nine plants were preferred over little bluestem. The average numbers of grasshoppers resting and feeding respectively on the nine preferred plants based on grading as compared to little bluestem were: western yarrow (Ala) (4.7 vs. 1.0; 3*7 vs.

0.5); baldwin ironweed (Vba) (4.? vs. 1.2; 3.8 vs. 0.8); horseweed (Sea)

(4.0 vs. 1.2; 2.6 vs. 0.8); aster (Aster) (3.6 vs. 1.2; 3.0 vs. 0.8); wholeleaf rosinweed (Sin) (3. 5 vs. 0.5; 2.8 vs. 0.3); common ragweed

(Ael) (2.2 vs. 0.7; 1.5 vs. 0.5); ocribner panicum (Psc) (1.8 vs. 1.4;

0.4 vs. 0.2); leadplant amorpha (Aca) (1.2 vs. 0.6; 1.2 vs. 0.6); and

sideoats grama (Bcu) (0.6 vs. 1.0; 0.0 vs. 0.2). Equal damage baaed on

grading compared to little bluestem was observed on sagewort (Artem),

scurfpes (Pfl), and common yellow oxalis (Ost). Little bluestem was

(Ppu) preferred over falseboneset kuhnia (Keu) , purple prairieclover junegrass western ragweed (Aps) , catclaw sensitivebriar (Sun), prairie

(Kcr), dotted gayfeather (Lpu), and inland ceanothus (Cov). In no case 20

was a "D" grade assigned to any plant in Table 7; however, in the com- parison of little bluestero (Asc) vs. horseweed (Eca), horseweed is graded "C- M and footnoted indicating that ore replicate was completely consumed. This replicate was graded "I)" yet the average of three replicates remains a "C-". Western yarrow (A.la), baldwin ironweed

(Vba), and horseweed (Fca) were preferred resting sites. No feeding was observed in the 50° F. to 60° F. temperature range.

Data in Table 7 were concerned v.ith kth and 5th instar nymphs

except those on sideoats grama (Bcu), prairie junegrass (Kcr) , scribner panicum (P3c), dotted gayfeather (Lpu), and common yellow oxalis (Ost) where adults were used.

M. f. f emurrubrum (CeGeer) has been considered a severe pest and of great economic importance for many years. Ball (19^2) states that it is one of the most destructive grasshoppers of the United States and Canada. The importance of this species has been based on damage done to cultivated crops. Feair3 and Davidson (1956) consider it as one of the five species which cause about 90 percent of the grasshopper damage to cultivated crops in the United States. Claassen (1915) states that in the summer of 1913 Helanoplus f emurrubrum (DoGeer) did consider- able damage in Kansas especially to alfalfa. Ketcalf and Flint (1939) list it as very destructive in legume fields and common along roadsides.

The importance of this species as a rangeland pest is unknown.

Blatchley (1920) stated that it occurred everywhere in bluegrass pas-

tures and meadows , along roadsides and borders of cultivated fields, on city lawns and in open woodlands. In Kansas, r.ilbur (1936) listed it as one of several species present in limited numbers doing damage to pasture grass, especially brome grass, in 1932. Wilbur and Fritz 21

(19^+0) reported this species to be more evenly distributed over the three pastures studied than were any of the other species. Hebard

(1931) noted that M. f . femurrubrum (DeGeer) was a very abundant and generally weed-loving species which occurred over all of Kansas, being particularly injurious to alfalfa. In 1936 he listed this species as present in weedy cultivated area3 throughout North Dakota. Knutson

(1937) found it in a variety of habitats in northeastern Texas but stated that immatures and adults were in an alfalfa field in great numbers and had completely stripped off the leaves over a five acre area. Isley (19V+) listed it as having forbivorous mandibles.

During the summer of I96I this species was observed in large quantities in certain areas of the bluestem range, yet very little apparent damage to grasses resulted. It is believed that the cage results (Table 8) closely indicate actual food preferences in the field because (1) the grasshoppers appeared to behave normally in the cage while crawling, feeding and resting; and (2) the literature on this species has suggested a forb habitat. Observations and experiments under artificial holding and experimental conditions, such as in labora-

tories, insectaries or cages in the field, are generally accepted as the

best substitute for biological studies when direct field studies are

impossible. Such was the case during these studies because it was be-

lieved better data on preferences could be determined by limiting choice

to two species of plants rather than multiple choice which would have

been the case in field studies. Even then, a cage would have been

necessary. importance M. f . femurrubrum (DeQeer) probably is of more economic

as a beneficial insect rather than a harmful one on rangeland. This is 22

in contrast to the generally accepted assumption on field crops, viz., that it is one of the four major crop pests, particularly on alfalfa.

SUMMARY

The economic importance of grasshoppers is determined by food plants. Food preferences and relationships of grasshopper species and plant species in bluestem pastures are not well understood. Data in this study are the results of cage studies of Melanoplus femurrubrum femurrubrum (TeCJeer) conducted during the summer growing season of 19<6l.

Twenty grasshoppers of a given species were placed in each cage.

A spread of two consecutive instars was the maximum allowed in each cage. The grasshoppers were allowed the opportunity to feed for k& hours on either or both of two plant species for a given time period.

Each experiment was replicated three times. Counts were made at 7 a.m.,

12 noon, 5 p.m., and in some cases 10 p.m. during the ^8 hour feeding period, recording total number of grasshoppers (l) resting on the plants; and (2) feeding on plants. Photographs also were taken of the plants immediately after removal from the cages. Photographs were ranked according to intensity-of-f eeding as compared with before-feeding pictures. A grade of A, B, C, and D was used: A = no feeding; B = trace to l/2 of plant eaten; C = l/2 plus eaten; D eaten entirely.

Species of grasses vs. grasses, and grasses vs. forbs, were evaluated.

A total of nine perennial grasses, Ik perennial forbs, two annual forbs, and two woody plants were used in the cage studies. A total of 6k plant species was used in correlation studies.

Preferred most over big bluestem were* aster, horseweed, common ragweed and scurfpea. All four species were completely consumed in each of three replicates. 23

Preferred most over sand dropseed were: horseweed, aster, dotted gayfeather and wholeleaf rosinweed. Horseweed was completely consumed in each of three replicates.

Preferred most over switchgrass were: western yarrow, sagewort, horseweed and dotted gayfeather. Western yarrow was completely con- sumed in each of three replicates.

Preferred most over indiangrass were: wholeleaf rosinweed, horse- weed, baldwin ironweed and western yarrow.

Preferred most over sirieoats grama were: aster, common ragweed, purple prairieclover and western ragweed. Aster and common ragweed were completely consumed in each of three replicates.

Preferred so^t over prairie junegrass were: common ragweed, scurf- pea, aster and western yarrow.

Freferred most over little bluestem were: western yarrow, baldwin ironweed, horseweed and aster.

Horseweed appeared in the four most preferred plants in each cast

except sideoat3 grama, in which case it was not evaluated.

The work reported on here is considered a first step in obtaining

clues as to preferred plant species which might be used in interpreting

the reason for associations which exist between grasshopper and plant

species, particularly in the bluestem regions of Kansas.

M. f. femurrubrum (DeGeer) probably is of more economic importance

as a beneficial insect rather than a harmful one on rangeland. 2k

"KENDALL'S RANK CORRELATION COEFFICIENT" KENDALL'S TAU

Coefficient is; T t/?t;(N-1)

I = Number of pastures in which you have observations.

I/2N(N-1) = 36 for 9 pastures l/2N(N-l) = 21 for 7 pastures

1/2 x 9 x 8 = 36

1/2 x 7 x 6 = 21

S = (Number of concordances) - (Number of discordances)

Rank the observations (insects) from 1 to 9 and put opposite each insect rank the corresponding plant rank.

For example: Hypochlora alba vs. Andropogon gerardi

1957 - ordinary upland

Insect Rank 123^56789 Plant Rank 8^9621753 Concordances 1^013300 = 12 Discordances 73641021 = 2*f

36

(Total Concordances) plus (Total Discordances) l/2N(N-l)

S = 12 minus 2k m -12 3 -12 __ Tm = = -»33 ,1/2N(N-1);„»,;„ , \ rz

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Insectary in which cage studies of Melanoplus f etnurrubrum f emurrubrum (DeGeer) food prefer- ences were conducted* 60

PLATE VII

.' PLATE VIII

Cage used to study food preference and behavior of grasshopper species between plant species. 62

PLATS VIII u

« n

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h • • e en C5 3 «) -H O U %_/• (8 A s 3 u U & 3 G c O • u &a 60 • 3 o 3 faO I P •ri 3 I o •o •rl «M u « T> T i V 4> j 2 4) M JO >.Q £> O (4 « O 6C 3 (B 05 1. m o 76

• do •H

0^ <*.

I

bo

I* 77

ackw ts

The writer wishes to express appreciation to Dr. Herbert Knutson,

Head of Department of Entomology, for suggesting the problem, for ad- vice and assistance during the period of research and the preparation of this thesis.

Appreciation is due Dr. illiaa B« Arnett, now Assistant Professor,

University of Nevada, for use of grasshopper data that he recorded dur-

ing 1957 i 1958 and 1959.

Appreciation is due Dr. Kling L. Anderson, Professor of Agronomy, for use of plant counts of the study area as well as cooperation in making available the area for study and advice and encouragement.

Appreciation is due Professor Donald A. Wilbur for advice and cooperation. Furthermore the cages used were originally built under his direction. Thanks to Dr. Carl I* Rettenmeyer, Assistant Professor of Entomology, for help received in photography.

Thanks to Dr. Ashley B. Gurney, Insect Identification Branch,

Entomology Research Division, U. S. Department of Agriculture, for veri- fication of the representative species collected. Thanks to Dr. Leslie F.

Marcus, Professor of Statistics, for advice on statistical analysis.

Thanks are conveyed to Gerald Greene for help in familiarization of

the study area and to Dr. Ruff L. Gentry and Wilfred E. McMurphy for

identification of ^lant species. Thanks to Charles Frankenfeld for his

untiring help in the cage studies.

The writer wishes to express appreciation to his wife, June, for

constant support and material help throughout the period of research

and writing. 78

LITERATURE

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Barnes, 0. L. Effects of food plants on the lesser migratory grasshopper. Jour. Econ. Ent. 48(2) :119-24. 1955»

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E.R.D. , Special report z-136. 1961.

Blatchley, «. S. Orthoptera of northeastern America. Indianapolis: Nature Pub. Company. 784 pp. 1920.

Brusven, Merlyn A. and Gregory B. Mulkern. The use of epidermal characteristics for the identification of plants recovered in fragmentary condition from the crops of grasshoppers. N.D.Agr. Sxp. Sta. Research Rept. 3'1-H. i960.

Cantrall, I. J. The ecology of the Orthoptera and Dermaptera of the George Reserve, Michigan. Kus. Zool. Univ., Mich. Misc. Pub. 54:1-182. 1943.

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79

Diver, C. and P. Diver. Contributions towards a survey of the plants and of South Haven Peninsula, Studland Heath, Dorset. Jour. Ml. 2(l):36-69. 1933.

Friauf, J. J. An ecological study of the Dermaptera and Orthoptera of the Welaka area in northern Florida. Ecol. Monog. 23:79-126. 1953.

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Hebard , Morgan The Orthoptera of South Dakota. Proc. Acad. Nat. Sci. Phila. 77:33-155. 1925.

The Orthoptera of Colorado. Proc. Acad. Nat. Sci. Phila. 81:303-^25. 1929.

The Orthoptera of Kansas. Proc. Acad. Nat. Sci. Phila. 83*119-227. 1931.

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Orthoptera of North Dakota. N. Dak. Agr. Exp. Sta. Bui. 28^: 1-66. 1936.

An ecological survey of the Orthoptera of Oklahoma. Okla. Agr. Exp. 3ta. Tech. Bui. 5*1-31. 1938.

Hodge, C, Iv. Growth and nutrition of Kelanoplus diff erentialis (Thos) (Orthoptera, Acrididae). I. Growth on a satisfactory mixed diet and on diets of single food plants. Physiol. Zool. 6:306-328. 1933.

Isley, F. B. Seasonal succession, soil relations, numbers, and regional dis- tribution of northeastern Texas Acridians. Bcol. Monog. 7:317-3^. 1937. •

80

The relations of Texas Acrididae to plants and soils Scol, Konog. 8:551-604. 1938.

Correlation between mandibular morphology and food specificity in grasshoppers. Ent. doc. Amer. Ann. 37*^7-67, illus. 19V+.

Differential feeding in relation to local distribution of grasshoppers. Ecology 27(2) :128-138. 19^6.

Knutson, Herbert. Notes on the Acrididae (Orthoptera) of Iowa. Field and Lab. 5(2):37-^7. 1937.

Kreasky, J. B. Extended diapause in eggs of high-altitude species of grass- hoppers* and a note on food plant preference of Kelanoplus bruneri. Ent. 3oc. Araer. Ann. 53(3) :V56-*f38. i960.

Ketcalf, C. L. and « P. Flint. Destructive and useful insects — their habits and control. McGraw-Hill Book Company, Inc. 98l pp. 1939

Mulkern f 0. B. and John F. Anderson. A technique for studying the food habits and preferences of grasshoppers. Jour. Econ. Ent. 52s3^2. 1959«

Mulkern, 3. B. , Merlyn A. Brusvon and Jerome A. Onsager. Biology and ecology of North Dakota grasshoppers. N. D. Annual Progress Kept., (Mimmeographed) i960.

Mulkern* 3. B., Jerome A. Onsager, Donald R. Toczek, and Roman J. Ertelt. Biology and ecology of North Dakota grasshoppers. It D. Annual Progress Rept., (Mimmeographed) 196l.

Nerney, N. J. Interrelated effects of grasshoppers and management practices on short-grass rangeland. U.S. Dept. Agr., A.R.S., S.R.D., Special report z-103. 1959*

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Interrelated effects of grasshoppers and management practices on short-grass rangeland. U.S. Dept. Agr., A.R.3., S.R.D., Special report z-137» 196l. 81

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Breeding plants for resistance to insect pests. Publ. Amer. Assoc. Advance. Sci. 61:245-266. i960.

Insect resistance in plants. Ann. Epiphyties, 12(4)597-403. 1961.

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Pruess, Kenneth P. Factors influencing the distribution and abundance of grass- hoppers in Nebraska. Neb. Agr. Expt. Sta. Annual Rept. (Mimmeographed) i960.

Factors influencing the distribution and abundance of grass- hoppers in Nebraska. Neb. Agr. Expt. Sta. Annual Rept. (Mimmeographed) 196l.

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The species and distribution of grasshoppers in the 1937 out- break. Insect Pest Survey 18, supplement to 6:385-^3. 1938.

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IVilbur, D. A. Grasshopper injury to the inflorescence of pasture grasses. Kan. Ent. Soc. Jour. 9*1-12. 1936.

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'.'.oodruff, Lawrence C. A grasshopper survey for Eastern Kansas, 1936. Jour. Kan. Ent. Soc. 10(3):75-83. 1937. FOOD PLANTS OF MELANOPLUS FEMURRUBRUM FEKUKRUFHUM (D3GSER) IN THE BLUESTEM GRASS REGION OF KANSAS

by

ORLO KENNETH JANTZ

B. S., Kansas State University, 1957

AN ABSTRACT OF A THESIS

submitted in partial fulfillment of the

requirements for the degree

MASTER OF SCIENCE

Department of Entomology

KANSAS STATE UNIVERSITY Manhattan, Kansas

1962 .

Food plants and relationships of grasshopper species and plant species in bluestem pastures are not well understood. This report is the result of cage studies on food plant preferences conducted during the summer growing season of 1961 on Helanoplus femurrubrum femurrubrum

(DeQeer)

Initial selection of plants to be evaluated, was based upon cal- culation of a correlation coefficient between each plant species population density, and grasshopper species population density.

Cage studies were conducted in an outdoor insectary. Twenty grasshoppers of a given species and plant species were placed in each cage. The grasshoppers were given the opportunity to feed for k8 hours on either or both of two plants for a given time period. Plants were replaced if completely consumed. After k8 hours the plants were re- moved and replaced by another series of two plant species. Species of grasses vs. grasses, and grasses vs. forbs, were evaluated. A total of nine perennial grasses, 1^+ perennial forbs, two annual forbs, and two woody plants were used in the cage studies. A total of Sh plant species were used in correlation studies. Counts were made three or four times per day during the ^8-hour period, recording total number of grasshoppers (1) resting on the plants; and (2) feeding on plants.

Photographs also were taken of the plants immediately after removal from the cage. The photographs were ranked according to the intensity- of-feeding as compared with before-feeding pictures. A grade of A, B,

C and D was used: A = no feeding; B trace to l/2 of plant eaten;

C = l/2 plus eaten; D = eaten entirely. i

Preferred most over big bluestem were: aster, horseweed, common ragweed and scurfpea. All four plants were completely consumed in each of three replicates.

Preferred most over sand dropseed were: horseweed, aster, dotted gayfeather and wholeleaf rosinweed. Horseweed was completely consumed in each of three replicates.

Preferred most over switchgrass were: western yarrow, sagewort, horseweed and dotted gayfeather. Western yarrow was completely con- sumed in each of three replicates.

Preferred most over indiangrass were: wholeleaf rosinweed, horse- weed, baldwin ironweed and western yarrow.

Preferred most over sideoats grama were: aster, common ragweed,

purple prairieclover and western ragweed. Aster and common ragweed

were completely consumed in each of three replicates.

Preferred most over prairie junegrass were: common ragweed

scurfpea, aster and western yarrow.

Preferred most over little bluestem were: western yarrow, bald-

win ironweed, horseweed and aster.

Horseweed appeared in the four most preferred plants in each case

except sideoats grama, in which case horseweed was not evaluated.

The work reported on here is considered a first step in obtaining

clues as to preferred plant species which might be used in interpreting

the reason for associations which exist between grasshopper and plant

species, particularly in the bluestem regions of Kansas.