FOOD PLAOT PREFERENCES OF (ORTOOPTERAt ) OF SELECTED PLANTED PASTURES IN EASTERN KANSAS

by

JAMES DALE LAMBLEY

B. S., Kansas State University, ftonhattan, 1965

A THESIS

submitted in partial fulfillment of the

requirements for the degree

MASTER OF SCIENCE

Department of Entomology

Kansas State University Manhattan, Kansas

1967

Approved byt

Major Professor LP alW ii

IP C-'-5- TABLE OF CONTENTS

^ INTRODUCTION •

^ REVIEV OF LITERATURE

MTHRIALS AND METHODS ^

^ Study Area

'° Field and Laboratory Studies

RESULTS AND DISCUSSION 21

Acridinae

1 22 Oedipodinae '

1 9S

SUWJ'ARY 128

LITERAPJRS CITED. 131

ACKKO'.VLEDGKENTS '•^®

APPENDIX 1^° INTOODUCnON

The purpose of this study, near Manhattan, Kansas, during 1965 and 1966, was to increase knowledge of the feeding and behavior of grasshoppers in

of the planted (tame) pastures. Emphasis was placed on the feeding habits more common . Great Grasshoppers have long been considered serious plant pests in the

of rangelands Plains area of the United States. Loss in production potential

(including pasture grass and other forage) has been estimated to be not include $80,000,000 per year for 1959 and 1960 (Anon., 1965). This does

funds spent for control. methods of Consequently, m-jch of the research has been directed towards biology immediate direct control. Little basic research dealing with the

less on and ecology of grasshoppers of rangeland has been done and even from such planted pasture species. Neglect in basic research has resulted

from cropland; factors as (l) lower economic return from grassland than

than in cropland: and, (2) damage is often less apparent in grassland intensive (3) recent recognition of grasslands as resources deserving

scientific investigation.

Publications dealing with and direct control of grasshoppers

often include some references on the feeding habits and behavior of the

various species. Past research on the biology of grasshoppers generally those deals with a single or small number of species. These are usually Hanson and species of greatest economic importance. The author agrees with Grasshoppers Vorhies (1938) in concluding that basic research is essential.

there is an as grassland pests cannot be dealt with scientifically unless

exact knowledge of their basic biology and ecology. Isely (1944) stated: "It should be evident that food specificity research will contribute to further progress in working out the control of pest grasshoppers."

This study was initiated to provide greater understanding of the biology and ecology of grasshoppers inhabiting tame pasture habitats through increased knowledge of their feeding and behavorial habits. REViaV OF LITERATURE

Of the vast amount of literature on the Acrididae, much is concerned with taxonomy and direct control of cropland species. Little information

is available on feeding habits and behavior, type and extent of damage to

plants, and host plant relationships of grasshoppers of planted grasslands.

Taxonomic works pertaining specifically to adult grasshoppers of Kansas Claassen include those by Brunner (1885, 1887), Isely (1905), Beamer (1915),

(1915), and Hebard (1931, 1934). Many taxonomic papers concerning grass-

hoppers of other areas also include species common to Kansas. Some are:

Coppock (1962), Alexander (1941), Ball et al . (1942), Brooks (1958),

Froeschner (1954), Gurney and Brooks (1959), Kebard (1925, 1929, 1936, 1938),

Knutson (1940), and Pfadt (1965).

Few taxonomic works have been published on grasshopper nymphal stages.

Criddle (1931) and Hanford (1946) described and published keys to nymphs,

some species of which are found in Kansas. Brusven (1966) published keys,

descriptions, pictures, and ecological notes for the nymphal stages of the

subfamily Acridinae of Kansas.

Many of the above taxonomic works include notations on favored

habitat, food plants, and seasonal occurrence of various species of the

Acrididae.

Lists of grassland species collected during non-taxonomic studies

have been made by Arnett (1959), Wilbur and Fritz (1940), Woodruff (1937),

and Hebard (1933).

Keys to the flora of Kansas have been prepared by Barkley (1963). furnished Illustrations as v;ell as keys to the grasses of Kansas have been

by Gates (1936). Launchbaugh (1964) illustrated and described many of the native grasses and broadleaf plants found in Kansas rangelands and, to some extent, in tame pasture areas. Many Kansas grassland plants are described by Hitchcock (1950), Stevens (1950), Phillips Petroleum (1963), and the

North Central Regional Publication No. 36 (i960).

Opinions of entomologists regarding food preferences of grasshoppers have changed greatly during the last 100 years. The old idea that grass- hoppers devour everything green has been accepted by many. Hovjever, it was originally applied only in describing Mel anoolus spretUS Walsh, (Riley,

Packard, and Thomas, 1877). Later investigations by Griddle (1933), Isely

(1937, 1938, 1944, 1946), Langford (1930), Henkle (1938), Shotwell (1941),

Brett (1947), Pfadt (1949), Anderson and V;right (1952), and Scharff (1954) showed that many species of grasshoppers were associated with a restricted number of host plants. Coppock (1962) and Hebard (1931, 1938) listed general

food preferences of most species of grasshoppers common to Kansas.

Isely and Alexander (1949) suggested determination of acridid food

preferences by examination and identification of plant fragment contents in

a the grasshopper crop (ventriculus) . Mulkern and Anderson (1959) developed

technique necessary for such a study. This technique utilized plant epidermal

characteristics for plant fragment identification. Mulkem and Anderson

(1959), Pruess (i960), Mulkern, Anderson, and Brusven (1962), Mulkern,

ToC2ek, and Brusven (1964), and Campbell (1966) have conducted crop content

analysis studies on most grasshopper species encountered in their studies.

Scharff (1961) used crop content analysis to a limited extent in his study of

the ecology of Melanoolus sanouinioes on Montana grasslands. Brusven and

Mulkern (1960) and Pruess (i960) have published keys for the identification

of plant fragments recovered from grasshopper crops. .

plant epidermal characters Plant taxonomic workers have also utilized varieties. Davis (1959) for the identification of plant species and using various plant epidermal differentiated some 40 species of grasses by several varieties of alfalfa, characteristics. Stoddard (1965) identified by leaf epidermal cell patterns. marigold, and some grasses (four Panicum spp.) Kentucky bluegrass (Poa oratensis Nittler (1966) differentiated varieties of

furnishes keys for the identification of L.) by this method. Metcalf (I960) plant anatomical characters. many European plant. species through use of these characters are included. Descriptions and illustrations of many of characters are illustrated and Several of the more common plant epidermal

discussed by Esau (1960). of food plants appear to be Factors affecting a grasshopper's selection that differential feeding on quite numerous. Uvarov (1928) pointed out seeking water. He concluded that water is plants is due primarily to the Acrididae nearly always show the only chemical compound toward which observed that Melanoolus bivittatus positive chemotropism. Griddle (1933) during dry Sonchus Srvensis L . (Say) was greatly attracted to sowthistle, ,

be due to the large amount of weather and that this attraction seemed to Wilbur (1936) reported that grasshopper moisture which the plant contained. grasses was largely caused by a search injury to the inflorescence of pasture of (1939) working with grass seedlings for moisture. Hermann and Hslick

bromegrass (Bromus inermis ) smithii.) . smooth western wheatgrass ( Acroovron species of Poa noted arundinacea , and several reed canarygrass (Phalaris ) authors species but within species. The differential feeding not only a^ng were perhaps caused both by the offered the explanation that differences grass- by the food preferences of the amount of green material available and

hoppers involved. Brunson and Painter (1938) observed that grasshoppers, when forced to

open- consume a single variety of corn, preferred some plants to others in pollinated varieties. These authors suggested that the genetic composition of the plant may influence its susceptibility to attack by grasshoppers.

Barnes (1955), Pfadt (1949b), Pickford (1963), Riegert, Pickford, and

Putnam (1965), and Smith (1959) found that certain plants are nutritionally

inadequate for som.e grasshopper species. Painter (1963) suggested that

plants and even different parts of the same plant may differ nutritionally.

Fraenkel (1959) suggested that insects were attracted by such secondary

plant compounds as alkaloids, gluoosides, organic acids, saponins, and tannins

rather than by the plants themselves.

The relationship of grasshoppers to their environment has long been

Swenk a point of study and discussion. Parker (1930), Shotwell (1941), and

and Bratt (1941) noted the effects of temperature and moisture on development

of of certain grasshopper species. Brett (1947) noted interrelated effects

temperature, humidity, and food on the development of Melanoolus sanquinipes..

Cantrall (1943), Friauf (1953), Shotwell (1930, 1948), Uvarov (1928),

Isely (1937, 1938), Pfadt (1949a), Vestal (1913), and others have investigated

the importance of habitat and host plant associations to grasshopper

distribution. Vestal (1913) and Cantrall (1943) concluded that grasshoppers

tend to seek habitats best suited for their optimum living conditions.

Isely (1937) considered soils and their related vegetation used as food or

shelter to be the primary factors in determining local grasshopper communities.

Vestal (1913) suggested that grasshoppers of a given habitat tend to increase,

decrease, or maintain themselves according to their tolerance to changes in

temperature, moisture, soil, plant succession, and in interspecies conflicts. Grasshoppers with a wide range of tolerance will be most capable of maintaining themselves.

Isely (1938) noted that the adaptation of certain acridids to introduced species of plants had complicated the problem of local distribution. He stated:

"The preference for Bermuda grass on the part of our acrididae

was acquired. It is obvious.. . .that vegetation absent in typical

virgin, optimum habitats of a species can not , belong to the original basic food requirements of the species."

Shotwell (1938) partially attributed the increase in numbers of certain grasshopper species, not previously prevalent, to introduced food plants.

The importance of land use on the abundance of grasshoppers has been shown by Amett (i960), Barnes (1959), Bei-Bienko and Mishchenko (1951),

Branson (1942), Buckell (1921), Coyner (1939), Nearney (1958, 1960), Odum

(1953), Shotwell (1953), Treherne and Buckell (1924), and Uvarov (1928).

Buckell (1921) and Nearney (1958) pointed out that grass sparseness as well as invasion of forbs is conducive to increased grasshopper numbers and injury.

Vegetation height, density, and degree of wetness have been listed as

factors affecting grasshopper distribution by such authors as Diver and

Diver (1933) and Kaufmann (1965).

Isely (1944, 1946) suggested that grasshoppers have evolved morphological

characters v/hich facilitate feeding in their preferred habitat. He found

that mandibular morphology of grasshoppers could be correlated with the type

of food plant utilized. Using this method, he classified 34 species as

graminivorous, 27 as forbivorous, and 18 as herbivorous. Kaufmann (1965)

found that Bavarian grasshopper species with differing food preferences

showed differences in maxillary lacinae. Also, geophilous grasshoppers, which

fed mainly on the ground, had either small arolia or none at all, while

acridids found in tall vegetation had large and well developed arolia. Criddle (1917), Putnam (1954), Numerous authors, including Brett (1947),

have linked grasshopper population and Parker, Newton, and Shotwell (1955), Nerney (i960) found that on increases with drought conditions. However, grasshopper populations were often poor or overgrazed Arizona rangeland, rainfall than after those with greater after years of normal or above-normal

said that a distinction should unusually light precipitation. Scharff (1961) different geographical areas. In be made among drought conditions in Midwest, drought and lowered explanation, he stated that in such areas as the from disease and parasites humidities, while probably furnishing release increase in numbers, seldom are thus allovdng ffelanoplus sanouinioes to

to a serai-starvation condition. severe enough to reduce nutrition levels present during more critical drought Such semi-starvation levels are often United States. periods in the more arid areas of the western fluctuations on grasshopper population Smith (1954) summarized information climatic to 1954 and correlated this with in Kansas for the period 1854 numbers must be dependent upon kind, factors. He concluded that grasshopper though modified by other environmental amount, and quality of food available, to be however, did not permit a correlation factors. Available data on food,

made with grasshopper population trends. indicated much study of the In summary, the literature cited above Food selection by grasshoppers seemed ecology of various grasshopper species. ecological studies. Most authors also to be of prime importance to such This study is intended to extend indicated the need for further research. relationship to the habitat, through the knowledge of the grasshopper and his of grasshoppers inhabiting planted further examination of the food preferences

oastures. MATERIALS AND METHODS

Study Area

planted pastures, This study was conducted during 1965 and 1966 in 6 Pottawatomie County, located in east-central Riley County and south-west elevation of 1,000 to Kansas. The terrain is rolling, to hilly, with an were located within 15 miles 1,100 feet above sea level. All study pastures of Manhattan, Kansas. forage growth, The climate of this area is normally favorable for

(81.28 cm.) and an having an average annual precipitation of 32 inches weather conditions were average frost-free season of 172 days. During 1965

spring was exceptionally cool some-A'hat near the average, but during 1966 the approximately half of and the summer hot and dry. Precipitation in 1966 was

normal (Appendix, Table 3).

of or more planted All pastures studied had vegetation consisting 1 pastures were forage species not native to eastern Kansas. Individual further selection was selected primarily with regard to vegetation type: but present, and proximity based upon grasshopper abundance, grasshopper species

to Kansas State University. southwestern Grasshopper collection site I (Fig. l) was located in miles east of Pottawatomie County, approximately 10 miles north and 3

site was predominately Manhattan, Kansas. The vegetation of the collection

and Korean lespedeza ( Lesnedeza western wheatgrass ( Aaroovron smithii) site was used stiDulacea), (Table l). The soil type was silty clay. This

for 1965 and 1966 studies. period. Mowing was This pasture was overgrazed throughout the study practices plus accomplished twice during each growing season. Management throughout 1966. severe drought led to a very poor pssture condition Fig. 1. Collection site I; an overgrazed western wheatgrass-Korean lespedeza pasture. Site collected during 1965 and 1966.

Fig. 2. Collection site II; non-grazed smooth brome-sweetcl over- alfalfa pasture. Site collected during 1965 and 1966.

Fig. 3. Collection site III; vegetation largely Kentucky bluegrass, Japanese brome, doiATiy brome, sideoats grama and big and little bluestem. Pasture grazed, then rested, at approximately one month intervals. Site collected during 1965. Fig. 4. Collection site IV: an overgrazed smooth brome pasture. Site collected during 1966.

Fig. 5. Collection site V: a moderately grazed smooth brome pasture. Site collected during 1966. Fig. 6. Collection site VI; a non-grazed reed canarygrass-smooth brome area. Site bordered by plantings of soybean, corn, sorghum, sunflower and by smooth brome pasture. Site collected during July, August, and September of 1965.

Fig. 7. Welanoolus differentialis damage to reed canarygrass within collection site VI. 16

The study site was bordered by a roadside dominated by alfalfa ( .'fedicaqo

sativa ) . bindweed ( Convolvulus arvensls ) and smooth brome ( Bromus inermis ).

Grasshopper collection site II (Fig. 2) was located in Pottawatomie

County, approximately 4 miles north-east of Manhattan, Kansas, and at the

south edge of the Tuttle Creek Reservoir spillway. This collection site was utilized for both 1965 and 1966 studies.

During reservoir construction, the topsoil had been stripped from the

area, leaving a silty clay subsoil at the surface. The area was then sown

to smooth brome, alfalfa, and yellow and white sweetclover (Kelilotus spp.),

(Table 2). The area was neither mowed nor grazed prior to or during the

study period.

Grasshopper collection site III (Fig. 3) was established as a Kansas

State University sheep pasture, approximately l/2 mile north of the city of

Manhattan in Riley County, Kansas. The vegetation of this site was pre-

dominately Kentucky bluegrass, Japanese brome ( Bromus iaoonicus ) . downy brome

( Bromus tectorum ) . and big bluestem ( Androgooon gerardi ). (Table 3). The

soil type v;as silty clay loam. The pasture was periodically heavily grazed,

then rested at approximately one month intervals throughout the summer.

Because of lov; grasshopper numbers, this site was utilized only for 1965

studies.

Grasshopper collection site IV (Fig. 4) v;as located in the Kansas State

University horse pasture at the north edge of the university campus. The

vegetation of the collection site was principally smooth brome with bindweed

being the most common weed species (Table 4). The soil type was silty clay.

This collection site vras used for both 1965 and 1966 studies.

This pasture was heavily grazed throughout the 1965 and 1966 study period.

The area was treated for weed control with 2,4-D herbicide on July 21, 1965, 17

Pasture and on July 16 and September 9, 1966. Mowing was done periodically. condition was extremely poor during the drought months of July, August, and

September of 1966. By August 1, 1966, little green vegetation was apparent.

Grasshopper collection site V (Fig. 5) was located in Riley County, approximately 3/4 mile north of Kansas State University proper. This pasture is managed by the Kansas State University Department of Husbandry.

Smooth brome constituted the major portion of the vegetation. Baldwin ironweed (Vernonia baldwini) and western ragweed ( Ambrosia osilostachva) were two commonly occurring weeds (Table 5). The soil type was silty clay.

This pasture was continuously grazed by cattle at a moderate rate throughout the study period. Mowing or herbicidal treatm.ents were not practiced.

Grasshopper collection site VI (Fig. 6 and 7) was located approximately one mile north of Kansas State University campus. It was collected only during 1966. This collection site was situated within a small lowland area at the northeast corner of the university agronomy farm. This area of silt loam soils has an underlying watertable approximately 9 feet from the ground

surface.

The collection site was located within and adjacent to a drainage gully v/hich traverses the area. An almost pure stand of reed canarygrass (PhalarJS

a mixed stand of arundlnacea ) was to be found within the gully basin with

smooth brome graduating to a pure stand of smooth brome at a distance from

the gully edge (Table 6). This area was not grazed, however, the gully banks

v;ere mowed in the fall of 1956.

The collection site was bordered by study plots of corn, soybeans, and

sorghum on the west and by a smooth brome pasture on the east. 18

Field and Laboratory Studies

each site was made three A critical evaluation of the vegetation of

order to gauge the influence of times during the 1965 growing season. In were made four times during drought upon plant availability, evaluations period, plant species making the 1966 gro-A-ing season. At each evaluation

and given a numerical relative up the vegetation of each site were identified and -1 abundance rating (Tables 1-6). Plants were ranked 5, 4, 3, 2, 1 (adapted from Cain and Castro, (trace) in decreasing order of abundance

and relative availability of a 1959). Such ratings denoted both numbers Identification of plant species was plant species to the grasshopper. descriptions, and pictures cited partially accomplished through use of keys,

in the literature review. were made, including slides of A complete series of histological slides

sites. Leaf and stem samples each plant species from individual collection different plants of the sam.e were taken from the top, center, and base of plant. These samples were species and from different parts of the same preserved material was ground preserved in 70 percent alcohol. Later, the that found in the grasshopper in a Waring blender to a size approximating procedure utilized was similar to crop (r-lulkern and Anderson, 1959). The and TAjlkern (i960). that of Mulkem and Anderson (1959) and Brusven were compiled by Descriptions for the identification of plant fragments Literature were also used. the author. Keys cited in the Review of intervals at each collection Grasshopper collections were made at weekly were not taken on rainy or site throughout the growing season. Collections

was belo^i 80° F. Five excessively windy days or when the temperature

15" diameter and 28" deep, hundred standard sweeps with a sweep net, in 19

constituted one site collection. Collected specimens were preserved in 75 percent alcohol until the crop contents could be analyzed.

Adult grasshopper identifications were verified by taxonomic keys and by comparisons with specimens in the collection of the Kansas State University

a large number of Department of Entomology. This collection contains , Fritz specimens from many previous studies, including those by Wilbur and

in the (1940), Brusven (1966), and Campbell (1966). Many of the specim.ens collection were identified by A. B. Gurney of the U. S. National lAiseum and by Hebard (1931, 1934).

Identification of nymphal Kansas Acridinae was covered by Brusven (1966).

Identification of nymphal Oedipodinae and Cyrtacanthacridinae of Kansas was

and difficult because of a scarcity of published keys. Criddle (1931) differentiated Handford (1946) observed that most grasshopper genera could be author found in nymphal stages on the basis of adult characteristics. This adult characters to be of most value in differentiating nymphs of later

in compiling stages. The author aided Merlyn A. Brusven and John B. Campbell grasshoppers found in a reference collection of nymphs of fifty species of complete, Kansas pasture and rangeland. This reference collection, though not

proved a valuable aid to nymphal identification. to Crop content analysis data were recorded on I. B. M. data sheets as instar, collection date, location, collection site number, grasshopper species,

of pollen, sex of adult, and plant species ingested. Empty crops and incidence

fungus, and parts were also recorded. The I. B. M. data sheets processing were then sent to North Dakota State University where I. B. M. data

converted to tabular v/as done by Dr. Gregory B. Mulkern. These data were

of a species form, giving: the total number of specimens of each life stage 20

collected per site; percentage of the total numbers of each specific life

stage of a grasshopper species having ingested a particular plant species

in each site; percentage of each life stage with empty crops or v.dth

pollen, fungus or arthropod parts ingested (Tables 7-30). In addition,

each species was given a grass-forb index number for each site (Tables 33

and 34). The grass-forb index was obtained by subtracting the total percent

ingesting of individuals of a species ingesting grasses from the total percent

forbs. Grasshopper species whose ratings were 100 to 76 were classified as

forb feeders; 75 to 26, mixed feeders preferring forbs; 25 to -25, mixed

feeders; -26 to -75, mixed feeders preferring grasses, and; -76 to -100,

grass feeders. 21

RESULTS AND DISCUSSION

A total of 2754 grasshopper crops were analyzed in 1965: 5781 in 1966.

Thirty four species were represented. Crop content analyses data are presented in Tables 7 to 30, inclusive. Summaries of the grass-forb indices for each species taken from each collection site are presented in Tables 33 and 34. In some instances, insufficient numbers of individuals of a grass- hopper species were collected and studied to show significant results. Crop content analysis tables for those were not included. Results are discussed, however, as they may be important indicators of food preferences. Summaries of numbers of individual species analyzed from each of the six collection sites are presented in Tables 31 and 32.

There is possible bias when comparing numbers of a grasshopper species

among individual collection sites. Four species ( Arohia simolex . Pardaloohora

haldemanii . Chortonhaga viridfaciata and simplex ) overwinter in

Kansas in the nymphal stage. Hence, few early spring, third and fourth instar, specimens were collected. In -addition, some species are more difficult to capture with a sweep net such as the subfamily Oedipodinae, in general, and Dissosteria Carolina and Chortoohaqa viridfaciata in particular.

Aqeneotettix deorum often drops to the ground when disturbed rather than jumping, thus being more difficult to collect. On the other hand, it prefers short vegetation which favors collecting.

Pollen, fungus and arthropod parts were commonly found in the grass- hopper crop; the ingestion rate varying with grasshopper species (Tables 7 to 30). Of these materials, fungus mycelium and spores were most commonly found. In many cases, the fungus could be seen growing on the plant fragments.

In other instances the crops were almost totally filled with fungus exclusive of much plant material. This apparently indicated direct grasshopper feeding 22

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In suggested by Mulkern, et. al. (1962). upon the fungus. Such is also highest during early instars (Tables 7 ™st instances, uptake of fungus was preference are suggested: (a) grasshopper to 30). Several possibilities moisture within dan,p plant residues; and, toward fungus; (b) a search for nearby fungus- instars thus forcing them to eat (c) lack of mobility of early

covered plant residues. from overgrazed western wheat- Ingestion of fungus by adults collected

(Table l) and overgrazed smooth grass-Korean lespedeza, collection site I increased greatly during the drought brome, collection site IV (Table 4) September, 1966. In these areas at this period of late July, August, and with little living vegetation present, time pasture condition was very poor upon dead plant materials. thus forcing grasshopper feeding with grasshopper species Ingestion of arthropod parts also varied

Oedipodinae and Cyrtacanthacridinae, in (Tables 7 to 30). However, the

of arthropod parts than did the general, showed higher ingestion rates most often legs, antennae, wings, Acridinae. Such arthropod fragments were whole and . In a few instances and scales of Diptera, Lepidoptera, many and Pfadt (1962) concluded that mites or spiders were found. Lavigne insects for food or moisture. grasshoppers actively seek injured or dead however. the grasshopper diet is unknown, The actual role of such materials in the widespread ingestion of such Mulkern, et. ai. (1962) suggest that play seems to indicate that they may materials by certain grasshopper species

nutrition of those species. a significant part in the Orphulella soeciosa and Syrbula Females of some species, notably did moles. greater volume of plant material than .dmi.abilis . ingested a grasshoppers, also recorded greater Kaufmann (1965), in studying Bavarian His studies showed, however. food ingestion by females of many species. 119

ingested that males of these same species assimilated a larger percent of

by the food than did the females. Thus, less volume of food was required

of most males. This author suggests that the smaller size of the males

species must also be taken into consideration.

The severe drought period of 1966 (Table 3, Appendix) afforded an

stresses. opportunity to study grasshopper feeding reactions to drought preferences were. During this period, no significant changes in food plant areas noted. Little variance in pasture condition was noted on those

under good grazing management. Sites I and IV were, however, overgrazed

throughout the study period, accompanied by greatly decreased pasture

conditions. As the vegetation dried, the grasshopper population largely

migrated into lower areas of green vegetation. By late July, 1966, the

vegetation within these areas was largely dried. At this time, there was a

significant increase in ingestion by grasshoppers of fungus-covered plant increased residue and arthropod parts. The proportion of empty crops also

overall greatly, often to 50 percent of those examined. By mid-August, 1966,

grasshopper numbers had greatly declined.

Acridinae

A total of 3288 grasshopper crops representing 11 species were analyzed

the during 1965 and 1966. Grass-forb indices ranged from -63 to -100 during

two year period, indicating a high preference for grasses. Acridinae

species represented 40 percent of the individuals collected throughout 1965;

38 percent during 1966.

on 55 specimens during Aoeneotettix deorum . Crop analysis was conducted

-100 over the 1965; 307 during 1966. Grass-forb indices ranged from -71 to ingestion tv;o years, indicating a grass feeder (Tables 17 and 18). Grass 120

relative abundance of species of percentages generally ranked closely with wheatgrass-Korean lespedeza collection site I grasses present. In western indicating ingested s^oth brome at a level (Table 1), however, speciniens

selection for that plant. well during the 1966 drought The species n,aintained itself relatively

slightly as the drought continued. period, with numbers declining only greatly increased during this period.- Ingestion of fungus and arthropod parts unpastured smooth brome collection This species was collected largely from smooth brome collection site V site II (Table 2) and moderately grazed site III in Kentucky bluegrass collection (Table 5). No specimens were found collection site IV (Table 4). (Table 3) or reed canarygrass analyses were completed on 14 specimens Boooedon oracile . Crop content The grass- 1966. The data are not tabled. during 1965 and 5 specimens during was -100 indicating a grass feeder. forb index for the specimens analyzed wheatgrass, smooth brome, and yellow The ingested grasses were western only at western wheat- . gracile was found bristlegrass (Setaria lutescens ) B. (Table l). grass-Korean lespedeza collection site I western of 25 specimens collected at Boooedon nubilim . Crop contents

(Table l) were analyzed but the wheatgrass-Korean lespedeza collection site I -100, index for the specimens analyzed was data are not tabled. The grass-forb wheat- The ingested grasses were western placing it in the grass feeder group.

grass and smooth brome. Of of 38 specimens were analyzed. nichromornha viridis . Crop contents canarygrass-smooth brome collection site VI these, 37 were collected at reed index for are included. The grass-forb (Table 6). No food ingestion tables reed canarygrass and tall dropseed the specimens studied was -100 with

often ingested. (Soorobolus asoer ) being most X21

Eritettix siinplex . Forty specimens were analyzed as to crop contents.

Of these, 31 were collected during 1965 from Kentucky bluegrass collection

site III (Table 3). Crop analyses data are not tabled. All grass-forb

indices were -100 with Kentucky bluegrass and sideoats grama most often

ingested.

Mermiria maculioennis . Crop analysis was conducted upon 8 specimens,.

all found at smooth brome-sweetclover site II (Table 2). Data are not tabled.

The grass-forb index placed it in the grass feeder group with smooth brorae

most often ingested.

Mermiria neomexicana . Eleven specimens were analyzed for crop contents with smooth brome being most often ingested. Specimens were found only at

smooth brome-sweetclover collection site II (Table 2). Data are not tabled.

Opeia obscura . Twelve specimens were found during 1966 at smooth brome-

sweetclover collection site II (Table 2). The grass-forb index for the

specimens v

Orohulella soeciosa . A total of 799 crops were analyzed during 1965;

1688 during 1966 (Tables 7 and 8). The grass-forb indices ranged from -71

to -100 over the tvjo years, thus designating it as a grass feeder. Ingestion

percentages of cultivated grasses ranked closely with relative abundance of

the grass species present. Thus, this species showed definitely higher ingestion

percentages of western wheatgrass in site I; of smooth brome in collection

sites II, IV, and V: and of Kentucky bluegrass and sideoats grama in site III.

The species was found in all tame pasture sites with the exception of reed

canarygrass-smooth brome collection site VI (Table 6), being most abundant in

moderately grazed smooth brome collection site V (Table 5), overgrazed smooth

brome collection site IV (Table 4) and v;estern wheatgrass-Korean lespedeza

collection site I (Table l). It was first in overall order of abundance. This species seemed better able to sustain itself throughout the drought period of 1966 than most other species studied. Numbers declined only slightly as the drought continued. Ingestion of fungus and arthropod parts greatly increased during the drought period.

on 7 PseudODomala brachvotera . Crop content analyses were conducted specimens during 1965 and 4 during 1966. The data are not tabled. All specimens were collected from ungrazed smooth brome-sweetclover collection site II (Table 2), and had ingested smooth brome.

conducted on 171 specimens Svrbula admirabilis . Crop content analysis was during 1965: 102 during 1966. Grass-forb indices ranged from -85 to -100 over two years, indicating it was a grass feeder (Tables 19 and 20). This species was found in all collection sites with grass ingestion percentages ranking closely y/ith relative abundance of species of grasses present.

Oedipodinae

Crop contents of 481 specimens representing 9 species were analyzed during 1965 and 1966. Grass-forb indices ranged from 12 to -100 over the two years. Oedipodinae species were generally classed as grass feeders or mixed feeders preferring grasses. Species of this subfamily represented 9

percent of the individuals collected throughout 1965; 4 percent during 1966.

analyzed during 1965. Arohia conspersa . Crop contents of 6 specimens were

No specimens were found during 1966. The data are not tabled.

The grass forb index for the specimens analyzed was -60 indicating a

mixed feeder preferring grasses. Kentucky bluegrass was most commonly ingested.

specimens during 1965 Arohia simplex . Crop analysis was conducted on 44

and 16 specimens during 1966. Data are not tabled. Grass-forb indices

normally ranged from -85 to -100 during the two year period, thus establishing This species seemed better able to sustain itself throughout the drought period of 1966 than most other species studied. Numbers declined only slightly as the drought continued. Ingestion of fungus and arthropod parts greatly increased during the drought period.

on 7 PseudODomala brachvotera . Crop content analyses v;ere conducted specimens during 1965 and 4 during 1966. The data are not tabled. All specimens were collected from ungrazed smooth brome-sweetclover collection site II (Table 2), and had ingested smooth brome.

on 171 specimens Svrbula admirabilis . Crop content analysis was conducted during 1965; 102 during 1966. Grass-forb indices ranged from -85 to -100 over two years, indicating it was a grass feeder (Tables 19 and 20). This species was found in all collection sites with grass ingestion percentages ranking closely with relative abundance of species of grasses present.

Oedipodinae

Crop contents of 481 specimens representing 9 species were analyzed during 1965 and 1966. Grass-forb indices ranged from 12 to -100 over the two years. Oedipodinae species were generally classed as grass feeders or mixed feeders preferring grasses. Species of this subfamily represented 9

percent of the individuals collected throughout 1965; 4 percent during 1966.

during 1965. Arnhia consnersa . Crop contents of 6 specimens were analyzed

No specimens were found during 1966. The data are not tabled.

The grass forb index for the specimens analyzed was -60 indicating a

mixed feeder preferring grasses. Kentucky bluegrass was most commonly ingested.

during 1965 Arohia simplex . Crop analysis was conducted on 44 specimens

and 16 specimens during 1966. Data ore not tabled. Grass-forb indices

normally ranged from -85 to -100 during the two year period, thus establishing 123

it as a grass feeder. In western-wheatgrass-Korean lespedeza collection site I (Table 1) during 1965, however, the grass forb index v/as -48 because of a higher ingestion of Korean lespedeza. The grass-forb index for the same site during 1966 was -100.

Arohia xanthooera . Crop contents of 67 specimens were analyzed during the tvio year period. Data are not tabled. Grass-forb indices ranged from

-33 to -100, indicating it to be a mixed feeder preferring grasses.

Chortoohaqa viridifasciata . Crop analysis was conducted on 68 specimens during 1965: 39 during 1966. Grass-forb indices ranged from 12 tO -100 over two years (Tables 27 and 28). Highest rates of forb ingestion occurred at western wheatgrass-Korean lespedeza collection site I (Table l) where there vjas much feeding upon Korean lespedeza. Grasses were ingested at a high rate throughout all planted pasture collection sites.

Fungus and arthropod parts were commonly ingested, being at a higher rate during the drought period of 1966.

Dissosteria Carolina . Crop content analysis was conducted on 5 specimens collected during 1966. Data are not tabled. Four specimens found at western wheatgrass-Korean lespedeza collection site I (Table l) had ingested Korean lespedeza.

Eneootolophus sordidus . Crop contents of 34 specimens were analyzed.

Of these, 33 were collected at western wheatgrass-Korean lespedeza collection site I (Table l). No food ingestion tables are included.

The grass-forb indices indicated that the species preferred grasses; these being ingested at a rate approximating the relative abundance of the species of grasses present. Korean lespedeza was the only forb ingested.

Hadrotettix trifasciatus . Crop analysis was done upon 12 specimens, 11 of 'which v;ere found during 1966. Food ingestion tables are not included. 124

collection site II All specimens were collected at smooth brome-sweetclover

analyzed indicated a (Table 2). The grass-forb index for the specimens commonly ingested. mixed feeder preferring grasses. Smooth brome was most

conducted upon 44 Hippiscus ruaosus . Crop content analysis was

from -50 specimens during 1965; 78 during 1966. Grass-forb indices ranged grasses to -100 over two years, indicating a mixed feeder preferring wheatgrass- (Tables 23 and 24). Most specimens were taken from western was Korean lespedeza collection site I (Table 1) where western wheatgrass were preferred. Ingestion rates of smooth brome and Korean lespedeza comparable although the latter was more abundant within the collection site. from High ingestion rates of fungus and arthropod parts were recorded third instar through adults.

of 5 specimens were analyzed. Pardaloohora apiculata . Crop contents

and were found at Data are not tabled. All specimens had ingested grasses Kentucky bluegrass smooth brome-sweetclover collection site II (Table 2) and

collection site III (Table 3).

were analyzed during 1965; Pardaloohora heldemanli . A total of 35 crops ranged from -50 28 during 1966. Data are not tabled. Grass-forb indices feeder preferring to -100 during the two years, thus designating it a mixed relative grasses. Grasses were normally ingested according to their sweetclover were abundance within the collection sites. Korean lespedeza and lespedeza collec- two most commonly ingested forbs in western wheatgrass-Korean collection site II tion site I (Table l) and smooth brome-sweetclover

(Table 2). .

125

Cyrtaoanthacridinae

A total of 4766 grasshopper crops representing 13 species were analyzed

during 1965 and 1966. Grass-forb indices ranged from 100 to -100 indicating greatly diverse feeding habits. Most, however, v;ere classed as forb feeders,

mixed feeders, or mixed feeders preferring forbs. Cyrtacanthacridinae

species represented 51 percent of the individuals collected during 1965;

58 percent during 1966.

Camovlacantha olivacea. Crop analysis was conducted on 22 specimens

during 1965 and 1966. Food ingestion tables are not included. The grass-

forb index for the specimens analyzed was 100 indicating a forb feeder.

host plant. Western ragweed ( Ambrosia osilostachva ) was the prim.ary

Twenty one specimens were collected at western vjheatgrass-Korean

lespedeza collection site I (Table l) and smooth brome-sweetclover collection

site II (Table 2) where western ragweed was readily available.

on 4 Hesoerotettix soeciosus . Crop content analysis was completed

specimens, all collected during 1965 from smooth brome-sweetclover collection

site II (Table 2) and smooth brome collection site IV (Table 4). Food

ingestion tables are not included. The grass-forb index for the specimens

was 100 indicating a forb feeder. Common ragweed ( Ambrosia artemisiifolia)

alfalfa, and sweetclover were ingested.

made on 12 Hesoerotettix viridis pratensis . Crop content analysis was

specimens during 1965 and 1966, and data are not tabled. The grass-forb were index for the specimens was 100 indicating a forb feeder. All specimens

and found at western heatgrass-Korean lespedeza collection site I (Table l)

smooth brome-sweetclover collection site II (Table 2). Missouri goldenrod

(Vernonia baldwini) were ( Solidaqo missouriensis ) and baldwin ironweed

ingested v;ith most regularity. 126

Melanoglus bivittatus . Crop content analyses was completed on 49 specimens during 1965 and 39 during 1966. Grass-forb indices ranged from

42 to -100 and varied with the collection site (Tables 29 and 30). In general, however, it was classified as a mixed feeder preferring forbs in those sites where forbs were abundant. In sites where forbs were not abundant, the species readily ingested grasses. Smooth brorae seemed most preferred of the grasses.

Melanoglus confusus . Two specimens were found on smooth brane- sweetclover collection site II (Table 2) during 1965. Both had eaten sweet- clover. No food ingestion tables are included.

Melanoplus differentialis . Crop content analysis was conducted on

137 specimens during 1965; 781 during 1966. This species was found in all planted pasture sites vdth the exception of Kentucky bluegrass collection site III (Table 3), but was most abundant in reed canarygrass-smooth brome collection site VI (Table 6) vAere 625 specimens were taken during 1966.

Grass-forb indices (Tables 13 and 14) ranged from 40 to -100 with a two year average of -64 indicating a mixed feeder preferring grasses.

The grass-forb index for specimens taken from reed canarygrass-smooth brome collection site VI (Table 6) was -84 with 73 percent of the specimens collected having ingested reed canarygrass (Fig. 7).

Melanoplus femurrubrum . Crop analysis was completed upon 2143 specimens during the tv;o year study period. Specimens were found at all collection sites v/ith the exception of Kentucky bluegrass collection site III (Table 3), but were most abundant in western v;heatgrass-Korean lespedeza collection site I (Table l). M. femurrubrum ranked second in overall abundance through- out the study. 127

The grass-forb indices ranged from 75 to -63 but generally indicated that the species was a mixed feeder preferring forbs (Tables 9 and 10).

The grass-forb indices changed appreciably among the different collection sites.

In reed canarygrass-smooth brome collection site VI (Table 6) the grass-forb index was -63 (Table 10). This resulted from the high ingestions of reed canarygrass and smooth brome which had a high abundance rating in this site. In western vjheatgrass-Korean lespedeza collection site I

(Table l) the grass-forb index v.'as 75 resulting from high ingestions of

Korean lespedeza, the most abundant forb.

MelanoDlus keeleri . Crop analysis vras completed on 115 specimens, largely collected during 1966 from western wheatgrass-Korean lespedeza collection site I (Table l) and smooth brome-sweetclover collection site II

(Table 2).

Grass-forb indices generally indicated this species to be a mixed feeder preferring forbs (Tables 25 and 26). Korean lespedeza and sweetclover were most often ingested.

f.'elanoolus packardii . Crop content analysis was conducted on 172

specimens during the two year study period. Of these, 162 were collected

during 1966 and largely from smooth brome-sweetclover collection site II

' (Table 2).

Grass-forb indices (Table 21 and 22) ranged from 15 to 94, the latter

being the index for collection site II (Table 2). Sweetclover was the most

commonly ingested plant.

to caused Melanoplus sanquinipes . This species has been reported have

heavy damage to grasslands of the United States during drought years. It 128

was found in moderate numbers in all collection sites with the exception of reed canarygrass-smooth brome collection site VI (Table 6).

The grass-forb indices ranged from 53 to -68, but where most abundant this species v;as rated as a mixed feeder (Tables 15 and 16). Alfalfa, sweetclover, and smooth brome showed the greatest ingestion percentages.

Third instar to adult specimens ingested moderate amounts of fungus and arthropod parts. As with many other species, ingestion of fungus and arthropod parts increased during the 1966 drought period.

Phoetaliotes nebrascensis . Crop contents of 1018 specimens were analyzed during the two years. Of these, 898 specimens were collected during 1966 from smooth brome-sweetclover collection site II (Table 2).

Grass-forb indices normally ranged from -95 to -100 indicating a grass feeder (Tables 11 and 12). The grass forb index for all specimens

collected from smooth brome-sweetclover collection site II (Table 2) was -95.

at western lineata . Seven specimens were found during 1965 wheatgrass-Korean lespedeza collection site I (Table l); 5 from smooth brome-

sweetclover collection site II (Table 2). No specimens were found during

1966. Food ingestion tables are not included.

Crop content analysis shov;ed ingestions of Korean lespedeza, baldwin

ironv;eed, and smooth brome at collection site I (Table 1). All specimens

analyzed from smooth brome-sweetclover collection 'site II (Table 2) had

ingested smooth brome.

SUM/fflRY

The investigation to determine the feeding habits and behavior of

grasshoppers in planted (tame) pastures of eastern Kansas was conducted

pastures studied had during the frost-free seasons of 1965 and 1966. All 129

not native to vegetation consisting of one or more planted forage species

preferences of the more the study area. Emphasis was placed on the food common grasshopper species. general, are Crop content analyses indicated that grasshoppers, in

of selectivity varying selective in their choice of food plants; the degree

in food plant with the grasshopper species. No significant changes

of the species studied preferences during various stages' of the life cycles

classified as grass feeders: grass were noted. Most Acridinae species were with relative abundance of grass ingestion percentages often ranking closely collection sites. Oedipodinae species species present within the grasshopper mixed feeders preferring grasses. were normally classed as grass feeders or habits. Crop analyses Cyrtacanthacridinae species had more diverse feeding ranged from 100 (forb feeder) grass-forb indices of species of this subfamily

classed as forb feeders, mixed to -100 (grass feeder). Most, however, were Phoetaliotes nebrascensis was feeders, or mixed feeders preferring forbs. consistently classed as a grass feeder. the only Cyrtacanthacridinae species numbers of individuals Some grasshopper species, as evidenced by particular planted pasture habitat. collected, had definite preferences for a proportions of forbs were normally Those species consistantly ingesting high pasture collection sites where such found in far greater numbers at planted olivacea was found only at collection forbs were most available. Camovlacantha plant, was present. Orohulell a sites where western ragweed, the preferred collection sites, but maintained highest .oeciosa was found at all grasshopper grazed smooth brome and western populations in overgrazed and moderately Aa^neotettix deorum was found most wheatgrass grasshopper collection sites. smooth brome grasshopper abundantly at ungrazed and moderately grazed

collection sites. 130

commonly found in grasshopper crops; Fungus and arthropod parts were materials, the grasshopper species. Of these the ingestion rates varying with seen some instances, fungus could be fungi was most often found. In fungus other times the crops contained inclusive with plant fragments. At indicating direct grasshopper feeding with little plant material present, instars, fungus was greatest with early upon the fungus. Ingestion of ingested drought period of 1966 also had although adults collected during the

large amounts. with grasshopper species. Ingestion of arthropod parts also varied

Cyrtacanthacridinae normally showed higher However, the Oedipodinae and often Such arthropod parts were most ingestion rates than did the Acridinae. of arthropod and Orthoptera. The proportion fragments of Diptera, Lepidoptera, 1966. increased during the drought period of fragment ingestions by m.any species maturity of the plants within the Pollen ingestions coincided with the

grasshopper collection sites. second instar grasshoppers had empty A high percentage of first and with empty species. The number of specimens crops as did the males of several drought period of 1966. crops increased greatly during the Syrbula admirabilis ingested a greater Females of Orohulella sneciosa and period, the males. During the 1966 drought volume of plant material than did sites by specimens found in overgrazed the volume of plant material ingested

generally declined. 131

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ACKNOV/LEDOffiNTS

Sincere thanks and appreciation is extended to Dr. Herbert Knutson,

Professor and Head, Department of Entomology, for advice and guidance during the research and preparation of this thesis.

Sincere thanks are also given to .'.'r. Clenton Owensby, Instructor,

Department of Agronomy, who gave generously of his time and talents in plant identification and establishment of plant abundance ratings.

A special v.'0rd of thanks is reserved for Dr. John B. Campbell for his invaluable aid during the research problem.

Acknowledgment is extended to Dr. Merlyn A. Brusven, Assistant Professor,

University of Idaho, for aiding in the establishment of techniques used in the identification of crop contents and for helping with initial identification of the Acrididae of Kansas.

Aopreciation is given to Dr. Gregory B. Mulkern, Associate Professor,

North Dakota State University, for his talents and time expended in lEN'i processing of the research data.

I also wish to thank Dr. Kling L. Anderson, Professor of Pastures and

Range Management, Dr. Derrick H. Blocker, Assistant Professor of Entomology and Dr. Lloyd C. Hulbert, Associate Professor of Botany and Plant Pathology for their advice and information during preparation of this manuscript.

"in.-'nc' n' -"-"ort. '^rov' ^'"f-' W the Kansas Agricultural Experiment Station

00 Project 5-859 and :. :.. 0., U. S. D. A. Grant No. 12-1 "5-1 -8047 (33) is gratefully acknowledged.

My deepest appreciation is given to my wife, Nellie, for her help,

encouragement, understanding, and patience.

140

Table 1. Scientific naraes(3)of grasshopper species included in crop analysis studies.

Acridinae

Aoeneotettix deorum deorum (Scudder) Boopedon qracile Rehn Boopedon nubilim (Say)

Dichromorp'na viridis ( Scudder) Eritettix sit^plex tricarinatus (Thomas) Mermiria maculipennis macclunqi Rehn Mermiria neomexicana (Thomas) obscura (Scudder) Orphulella speciosa (Scudder) Pseudooomala bracyptera (Scudder) Syrbula adrrirabilis (Uhler)

Oedipodinae

Arohia conspersa Scudder simplex Tscudder) Arphia xanthootera (Burneister) Chortophacra viridifasciata (DeGeer) Dissosteira Carolina (Linneaus) Encoptolophus sordidus costalis (Scudder) trifasciatus (Say) Hippisous ruaosus Scudder apiculata (Harris) Pardalophora haldemanii (Thomas)

Cyrtacanthacridinae

Campvlacantha olivacea olivacea (Scudder)

Hesperotettix scecior/. -. ' Scudder) Hesoerotettix viridis pratensis Scudder Hesperotettix viridis viridis (Thomas) ffelanoplus bivittatus (Say) ffelanoplus confusus Scudder Melanoplus dif ferentialis (Thomas) Ifelanoplus f emu rru brum femurrubrum (DeGeer) Melanoplus keeleri luridus (Dodge) ffclanoolus oackardii Scudder Ifelanoplus sanquinipes (Fabricius) Phoetaliotes nebrascensis (Thomas) Schistocerca linonta Scudder

^^'pollovdng Hebard (1931, 1934) and Coppock (1962). 141

(a) Table 2. Scientific and common names of plants ingested by grasshoppers or found in grasshopper collection sites.

Scientific name Common name

Perennial Grasses

Aoropvron smithii Rydb. western wheatgrass Andror'^qon oerardi Vitnan big bluestem Andropocon scooarlus Kichx. little bluestem Bouteloua curtipendula (ffichx.) Torr. sideoats grama Bromus inermis Leyss. smooth brome Chi or is verticillata Nutt. windmillgrass Eraqrostis soectabilis (Pursh) Steud. purple lovegrass Festuea arunjinacea Schreb. tall fescue Panicur! scribnerianum Nash scribner panicum Panicum viroatum L. switchgrass Phalaris arundinacea L. reed canarygrass Poa pratensis L. Kentucky bluegrass Schedonnardus paniculatus (Nutt.) Trel. tumblegrass Sorohastrum nutans (L.) Nash indiangrass Spartina pectinata Link prairie cordgrass Soorobolus asoer (Michx.) Kunth tall dropseed Soorobolus crvptandrus (Torr.) A. Gray sand dropseed Tridens fla-/u3 (L.) Hitchc. purpletop

Annual Grasses

Aristida oliqontha Michx. prairie threeavm Bromu s iaponicus Thunb. Japanese brome Bromu s tectorum L. dovmy brome Diqitaria sanquinalis (L.) Scop. crabgrass Eraqrostis cilianensis (/Ul.) Lutati stinkgrass Hordeum pusillum Fv'utt. little barley Setaria lutescens (VVeigel) Hubb. yellow bristlegrass Setaria viridis (L.) Beauv. green bristlegrass

Grass-like Plants

Carex spp. L. sedge Typha latifolia L. common cattail

Perennial Forbs

Achillea millefolium var. lanulosa (Nutt.) Piper western yarrow Ambrosia psilostachya DC. ( A^ coronopi folia T. and G.) western ragweed ) ) ) ) )

142

Table 2. (cont.)

Scientific name Common name

Perennial Forbs

Amoelamus albidus (Nutt.) Britt. (Gonolobus laevis of manuals, not Michx._ climbingmilkweed Antennaria neglecta Green (incl. A;, campestis pussytoes Rydb. and A^ lonaifolia (Green)" field Artemesia ludoviciana Nutt. (incl. var. anaoholodes (Nutt.) T. and G. common sagev;ort milkweed Asclepjas sulivantii Engelm. sullivant milkweed Ascleoias tuberosa L. butterfly Asclepiodora viridis (Walt. ) Gray (Asclepias antelopehorn v^ Walt. green Aster ericoides L. heath aster Astragalus carvocarous Ker. (A^ crass- milkvetch icarpus Kutt. groundplum Callirhoe involucrata (T. and G.) Gray purple poppymallow Convolvulus arvensis L. bindweed bundleflower Desmanthus illinoensis (Kichx.) MacM. Illinois sunflower maximillani Schrad. maximilian Kuhnia eupatoroides L. (K^. suaveolens Fresin; falseboneset K. hitchcockii A. Nels.) lespedeza Lespedeza capitata Michx. roundhead alfalfa f.'.edicaoo sativa L. white sweetclovcr ifeli lotus alba Desr. sweetclover Melilotus officinalis (L.) Lam. yellow oxalis Oxalis stricta L. common yellovj prairie- Petal ostemum multiflorum Nutt. roundheaded cl over groundcherry Physal - s spp. L. Polyaonum 3 en svl van! cum L. (incl. P;. smartweed omissum Greene) Pennsylvania Ruellia humilis Nutt. (R^ ciliosa . in part: fringeleaf ruellia R. carolinicnsis . in part. Rumex crispus L. curled dock Saoittaria latifolia Willd. (including Sj. arrowhead esculenta Hov;ell) common Salvia azurea Lam. var. orandiflora Benth. pitcher sage ( S. pitcher! Torr. Schrankia nuttallj (DC.) Standi. sensitivebriar (S^ uncinata of manuals, not Willd.) catclaw Solidapo missouriensis Nutt. (incl. S^ goldenrod nlaberrima Martens and S^ moritura Steele) Missouri dandelion Taraxacum officinale VJeber (T^ vulqare Lam.) common Vernonia altissima Nutt. tall ironweed baldwin ironweed Vernonia bald-.vini var. baldwini Torr. . vervain Verbena stricta Vent. woolly . )

143

Table 2. (cont.)

Scientific name Common name

Biennial Forbs

Cirsium altissimum (L.) Spreng. tall thistle Cirsium undulatum (Nutt.) Spreng. wavyleaf thistle Gaura narvi flora Dougl small flovjer gaura Grindelia souarrosa (Pursh) Dunal curlycup gumv;eed Verbascum thaosus L. flannel mullein

Annual Forbs

Amaranthus retroflexus L. rough pigweed

Ambrosia artemisiifolia L. ( A. elatior L. common ragweed Chenopdium album L. lambsquarters Convza canadensis (L.) Cron. ( Erioeron c^. L., Leotilon c^ (L.) Britt) horseweed Croton spp. L. Croton Erioeron strigosus (JAjhl.) (E. ramosus (V.alt.) daisy fleabane Euphorbia maroinata Pursh snow-on-the-mountain Euphorbia spp. L. (incl. Tithvmalus (Tourn.) Adans. and Chamaesyce Small) matforming spurge Euphorbia serpens HBK serpent euphorbia Hibiscus trionum L. flowerofanhour Kochia scoparia (L.) Roth kochia

Lactuca scariola L. ( L. serriola L.; L. cirosa of manuals, not L.) prickly lettuce Leoidium densiflorum Schrad. peppergrass Lespedeza stioulacea Maxim. Korean lespedeza Solanum carol inense L. Solanum rostratum Dunal bu --i- nightshade Tribulus terrestris L. puncturevine

V/oody Plants

Amorpha fruticosa L. indigobush amorpha Populus spp. L. Cottonwood Rhus radicans L. (Rj. toxicodendron , of manuals, in part, not L.) poisonivy Rosa arkansana Porter var. suffulta (Grei-ne) Cockerell (R^ suffulta Greene) Arkansas rose Salix spp. L. willow Sambucus canadensis L. American elderberry Symohoricarpos orbiculatus Moench buck brush Vitis vulpina L. wild grape

(a) Folloy/ing Anderson, K. L. (1961), Fernald (1950), and Stevens (1950). 144

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FOOD PLANT PREFERENCES OF

GRASSHOPPERS ( ORTKOPTERA ACRIDIDAE) OF SELECTED PLANTED PASTURES IN EASTERN KANSAS

by

JAACS D.ALE LAMBLEY

B. S., Kansas State University, Manhattan, 1965

AN ABSTRACT OF A MASTER'S THESIS

submitted in partial fulfillment of the

requirements for the degree

MASTER OF SCIENCE

Department of Entomology

KANSAS STATE UNIVERSITY

ti'.anhattan , Kansas

1967 ABSTRACT

An investigation to determine the feeding and behavioral habits of grasshoppers in planted (tame) pastures of eastern Kansas was conducted during the frost-free seasons of 1965 and 1966. Emphasis was placed on the food preferences of the more common grasshopper species.

Grasshopper collection sites were selected primarily with regard to vegetation type: each consisting of one or more planted forage species not native to eastern Kansas. Collection sites selected were of (l) western

wheatgrass ( Aoroovron smithii ) -Korean lespedeza ( Lesoedeza stioulacea ).

(2) smooth brome ( Bromus inermis )-svjeetclover ( Wedicaoo spp.), (3) Kentucky

bluegrass ( Poa pratensis ) . (4) overgrazed smooth brome, (5) moderately

grazed smooth brome, and (6) reed canarygrass ( Phalaris arundinacea ) -smooth brome.

Plant species making up the vegetation of each site were identified and given a numerical relative abundance rating. Such ratings denoted both numbers and relative availability of a plant species to the grasshopper.

Grasshopper collections v;ere made at vjeekly intervals at each collection site throughout the growing season.

Grasshopper food ingestions were determined by analysis of the crop contents. Findings were then I. B. M. data-processed and data were con- verted to tabular form giving: the total number of specimens of each life

stage of a grasshopper species collected per site: percentage of the total numbers of each specific life stage of a grasshopper species having ingested a particular plant species in each site: percentage of each life stage vjith

empty crops or v;ith pollen, fungus, or arthropod parts ingested. Also,

each species was given a grass-iorb index number obtained by subtracting the total percent of individuals of a species ingesting grasses from the total percent ingesting forbs. Grasshopper species whose ratings were 100 to 76 were classified as forb feeders; 75 to 26, mixed feeders preferring forbs;

25 to -25, mixed feeders; -26 to -75, mixed feeders preferring grasses, and; -76 to -100, grass feeders.

Crop content analyses indicated that grasshoppers, in general, are selective in their choice of food plants; the degree of selectivity varying with the grasshopper species. No significant changes in food plant preferences during various stages of the life cycles of the species studied were noted. Wost Acridinae species were classified as grass feeders; grass ingestion percentages often ranking closely with relative abundance of grass species present. Oedipodinae species v;ere normally classed as grass feeders or mixed feeders preferring grasses. Cyrtacanthacridinae species had more diverse feeding habits, but were usually classed as forb feeders, mixed feeders, or mixed feeders preferring forbs.

Some grasshopper species showed definite preferences for a particular planted pasture habitat, being found only or most abundantly at collection

sites v/here the preferred food plant was available.

Fungus and arthropod parts were commonly found in the grasshopper

crops; the ingestion rates varying with the grasshopper species. Ingestions

of these materials greatly increased during the drought period of 1966.

A high percentage of first and second instar grasshoppers had empty

croos as did the males of several species. The numbers of specimens with

empty crops increased greatly during the drought period of 1966.