Wildlife of the Prairies Keith E-Evans and George E

USDA FOREST SERVICE GENERAL TECHNICAL REPORT NC-2 9

. WILDLIFE OF _THE PRAIRIES KEITH E-EVANS AND GEORGE E. PROBASCO

NORTH CENTRAL FOREST EXPERIMENT STATION FOREST SERVICE U.S. DEPARTMENT OF AGRICULTURE _ CONTENTS

The Resource ...... _ • . . , ...... 1 "\ • Habitat Management ...... 12 \, • Discussion ...... 14 Llterature Cited ...... 15

North Central Forest Experiment Statlon John H. Ohman, Dlrector Forest Service - U.S. Department of Agriculture 1992 Folwell Avenue St. Paul, Minnesota 55108

Manuscript approved for publication January 5, 1976

1977

° THE AUTHORS

Kelth E. Evans Is a Prlnclpal Wlldllfe o Biologist and Project Leader at Columbia, Missouri, specializing in research on the l_apacts of land use alternatlves on nongame • species on the Ozark Plateau. Kelth holds B.S. and M.S. degrees from Colorado State University, where he majored In w11d1Ife management, and a Ph.D. from Cornell Unl- verslty, where he majored In wildlife scl- ence and minored In animal nutrition and !i!iiiiii_!iii...... _i_!ii_,,...... i_::. forest conservation. Kelth was employed by

• :_iiii'_....i!i::: iment Station for 8 years before moving to • ii_ili__iiiii:i:_,__,_,....!:: ii_ :_'_.... :_ :.i'_: tColhe umRbockyia, Miossuntainouri Fiorestn 1972.and Range Exper- i

George E. Probasco is an Associate Research Wildlife Biologist at Columbia, Missouri. He received his B.S. in Zoology In 1966 from Ft. Hays, Kansas State College, and hls M.S. In Range Management In 1968 from the University of Wyoming. He Is currently studying the influence of eastern redcedar invasion on b_rd and small mammal populations of the Ozark gladelands.

• • .

I WILDLIFEOF THE PRAIRIESAND PLAINS . Keith E. Evans and George E. Probasco

The extensive and diverse grasslands plant and animal composition drastically. of North Amerlca were malntalned and per- Extensive areas are managed for livestock petuated under the land use policies of the production by perpetuating the native aborigines. Grasses dominated the low rain- flora. These areas are inhabited by many fall Great Plains for many thousands of wildlife species. yearsand still do except where cultivation or destructive grazing have occurred. Fire was required in the higher rainfall prairie THE RESOURCE and Savannah areas to control woody plant encroachment. Some of the wildlife species Grasslands encompass the most extensive adapted tO these grasslands included bison, and varied of all plant communities in antelope, elk, bear, rabbits, prairie North Amerlca (fig. I). The mldcontinent dogs, wolves, coyotes, and grouse. The ' grassland extends 2,500 miles north to river bottoms and badlands in the area south and averages 400 miles in width from provided habitat for deer, Audubon's big- the Rocky Mountalns east as far as Illinois. horn sheep, bobcat, mountain lion, turkey, It includes four of the elghtmaJor grass- quails and waterfowl, land types recognized in North America: (I) oak-bluestem savannah, (2) tall grass, In the late 1800's the American fron- (3) mixed grass, and (4) short grass. The tier expanded westward--the day of the true remaining four are: (5) bunch grass, "wild west". Settlement under the Home- such as the palouse prairie in Oregon, stead Act of 1862 encouraged many people Idaho, and Washington, (6) annual grass to settle on the prairies and plains. In in the Callfornla's Central Valley, (7) a shOrt span of time the area changed from desert grassland in New Mexico and Arizona, wild grasslands teeming with wildlife to and (8) the coastal prairies in Texas, civilized areas of farms, ranches, fences, Louisiana, Maryland, New Jersey, New York, towns, roads, and railroads. The 19th Rhode Island, and Massachusetts. century witnessed the beginning and end of the Oregon Trail, gold rushes, open range_ and cattle drives. The oak-bluestem savannah stretches from southeast North Dakota and middle- The wet years and high prices asso- west Minnesota south along the eastern edge ciated with World War I made the Great of the tall grass prairie to mldeastern Plalnslook llke the promised land to many Texas. It is actually the transition zone farmers. However, the years of productlv- or ecotone between the eastern deciduous ity and profits gained by cultivating low forest and the tall grass prairie. ralnfaliareas were short. First came low Scattered segments of savannah occur postwar prices, then high taxes, low rain_: throughout the oak-hlckory forest and the fall, and dust. Thusbegan the "dust eastern part of the bluestem prairie. bowl" days of the 1930's. The bluestem-oak-hlckory forest mosaic, cedar glades, and cross timber vegetation types of Kuchler's (1964) Potentlal For the most part, the economy and Natural VegetationMap are included as land use patterns on the North American savannah. The greater part of the Ozark grasslands have again become fairly stable. Dome is often included as part of the A large proportion of the original grass- savannah (Marbut 1911, Davis 1964). lands are now being cultivated. The tall grass prairieonce covered 42 million The tall grass prairie formerly acres, but nowhas been reduced to 18 occupied the ecotone between the savannah million acres. Over much of the remaining and the mixed grass prairie. Fire was an uncultivated tall grass prairie region, important factor in preventing the forest livestoc_ overgrazlng has altered the type from invading the prairie (Curtis ,

,I¢. ° KEY TO GRASSLANDTYPES:

",', \,, _,\,,,, \...... ,_ \\,,\_,\ OAK-BLUESTEMSAVANNAHANDTALL GRASS

-...... _. MIXED GRASS b_._._, BUNCHGRASS ..:..

_i! SHORTGRASS ,":.:.:'"'"".:. ANNUALGRASS _:o-_;;,..:. DESERTGRASS COASTAL

Figure 1.--P_nc_pal g_ssL_d _pes of the United States. I

1959). Currently most of the tall grass Terrain,5o11,and Climate prairie has been converted to forest, to mixed grass, or plowed. The conversion to The topography of the mldcontlnent mixed grass res t!ted from prolonged exten- grasslands is variable, but generally it is sire livestock grazing, moderately ro11Ing. Some areas, such as the Red River Valley, are virtually flat; other areas (river breaks) along the major The mixed grass or needlegrass- drainages are steep. Major soil groups wheatgrass-grama grass (Stipa-Agropyron- include Alflsols, Mollisols, Vertisols, Boutelouu) community occupies the area west Inceptisols, and Aridlsols. There is a • Of the tall. grass region, nearly to the gradient from the cool, moist soils in the foothills of the Rocky Mountains. There northeast to the warm, dry soils in the is no definite line between the mixed southwest. grass and short grass (Boutelo_-Buchloe) vegetation types. The thin soil and low Climate of the midcontinent grass- precipitation areas are dominated by the lands is temperate and subject to extreme short grass species whereas the northern fluctuations. Summers in the north are and eastern Great Plains areas contain a hot and winters are cold while in the south vegetation type dominated by the mixed winters are less severe. Temperatures grass species, range from-40 C to +40 C in the north and from-I0 C to +40 C in the south (Kincer 1941). The frost-free season General aspects of the other grass- ranges from less than 4 months in the land types are discussed in subsequent north to nearly the entire year in southern sections of this paper. Texas.

. When periodic droughts occur, the NatlveAmerlcans and early settlers preclpltatlonfailure usually occurs in utilized fire to attract and hold game and July and August. During these droughts to prepare the range for early spring ralnfa11 w_ll vary from 50 to 90 percent grazing. Fire suppression efforts early of normal (Borcher_ 1950). Average annual in this century effectlvely reduced the preclpltatlonvarles from 115 cm in the use of fire for these purposes. However, savannah to 65-80 cm in the tall grass research in the Kansas Flint Hills prairie, to 50 cmln the mixed grass (Anderson et al. 1970), Missouri (Kucera prairie, and to less than 35 cm in parts and Koelllng 1964), and Texas (Wright of theshort grass prairie. Evaporation 1974a, 1974b) reveal that obllgate is a s_gnlflcant factor in mldcontlnent relations may exist between fire and some grassland climate. Except in the savannah grassland plants. and ta!igrass prairie, the evaporation potentlalexceeds precipitation. Strong Drought, fire, cultivation, and grazing winds prevai ! over the grasslands inducing all influence vegetation composition. The high evaporation rates and exerting con- Influence of the vast herds of bison is siderablelnfluence on the thermoregulatory probably the least understood influence. behavior of homeothermlc wildlife. Winds For example, there is disagreement as to often become more intense during drought the successional status of the short grass

periodsj (Borchert 1950). plains. Many ' including Shelford (1963) , distinguish the short grass type as a The other grassland areas have dif- , distinctive unit (climax vegetation type). ferent climates. The coastal prairies are Weaver and Albertson (1956) refer to the wet, humld, and generally mild, and not short grass as a dlscllmax reflecting past , subject-to the temperature extremes of the grazing use. Larson (1940) offered the intercontinental areas. Annual precipitation following evidence to support the view that for the coastal prairies ranges from 100 the short grass plains represent the true to 140 cm." Average annual temperatures in climax of the pristine biome: the northern areas range from a low of 0 C to a high of 25 C, while in the southern I. Historical records indicate the Theareasbuntcemph eragratssures(paranlousege)frpromair1i3e C retcoei30vesC. large herbivores stocked the plains most of _its moisture in the form of winter to carrying capacity and the in- troduction of livestock was merely snow. The California Central Valley and a substitution for grazing by desert grasslands usually receive winter wildlife. ralnwlth hot, dry summers. Annual precipitation inthese grasslands is generally 2. Explorers and pioneers referred 50 cm or less. Average annual temperatures in the bunch grass prairie range from to the short grass plains long -5 C .to 25 C. In the latter two grassland before livestock was introduced. areas the temperature range is 5 C to 27 C (Kincer 1941). 3. The marked ability of the short grass dominants to withstand grazing, indicating they evolved with grazing as an environmental factor. ..-. ' Vegetation Regardless, bison stocking rates .. Weaver (1954) summarized the Impacts probably varied from extremely heavy use of drough t and Daubenmlre (1968) of flre to no use because fences were not present ongrasslands. Under extended drought con- to restrict bison grazing. Moreover, the dltions, the grass cover ls reduced and grasslands in presettlement times probably 'the plant Species composition altered were subjected to frequent and erratic (Albertson et aZ. 1957, Coupland 1958). periods free of grazing during which changes in vegetation occurred because of Fire, whldh frequently accompanies climatic or moisture conditions. the drought, makes additional demands upon thevegetatlon. Some woody plants such Overgrazlng by bison, however, prob- as black oak (Quer_z8 velutina), eastern ably did encourage the growth of short redcedar (Juniper_is virginia), and ashe's grass species and produced conditions juniper (Juniperu8 ashei) are particularly favoring the "invasion" of some annual or vulnerable to fire. biennial species.

3 [

The.former tall grass, oak-bluestem duc_loides) occur on the dry and abused savannah, and mixed grass types today make sites throughout the mixed grass area and up one of the most productive crop-growing dominate the short grass plains. regions in the world: the first two com- bined are.known as the "Corn Belt"; the The mixed grass prairie contains corn- third, the "Wheatland". The vegetation of ponents of the tall grass prairie on the those portions of these areas still re- better sites and species from the short maining in grass has often been altered grass plains on the thin soll or dry sites because cool season species have been or where abusive grazing has been practiced. planted to replace native warm season There also are a number of grass species grasses, that reach heights of 60 to 90 cm and fill the niche between the tall grasses (often A vegetation gradient, corresponding over 2 m), and the short grass (less than to the soils gradient, occurs across the 20 cm). These include western wheatgrass midcontinent grasslands from east to west (Agropyron smithii), slde-oats grama and from north to south. Ecologists divide (Bouteloua curtipendula), several species the area into several vegetation types, of needlegrass (Stipa spp.), Junegrass however, the ecotones or transition zones (Koeleria cristata), and several species of between types are wide, variable, and depend dropseeds (Sporobolus spp.). Cool season -on past land use and climatic conditions, grasses play an important role throughout Much of the region is actually a mosaic the mixed grass prairie and are dominant of several types. A wide diversity of 0 on the northern Great Plains. plants occur on the grasslands. The Herbaceous broadleafed plants (forbs) following discussion is limited to a few increase the range forage value and provide of the dominant species, esthetical values to photographers and other rangeland recreatlonists. The com- Big bluestem (Andropogon gerardi) * posite family (Compositae) and the legume is the dominant component in the oak family (Leguminosae) are well represented. savannah and tall grass prairie. Big Common composites include sunflowers • 2 bluestem also occurs on better sites (Helianthus spp.), rosinweeds (Silphium • throughout the mixed grass prairie. Sand spp.), and coneflowers (Ratibida spp., bluestem (A. ball//), a close relative of Rudbeckia spp., and Echinacea spp.). big bluestem, occurs on sandy soils in Common legumes include leadplant (Amorpha the tall, mixed, and short grass prairie, spp.), false indigos (Baptisia spp.) Little bluestem (A. 8ooparius) dominates prairie clovers (Petalostemum spp.), and the drier, upland portions of the savannah Psoralea spp. and extends its range westward to become Common shrubs and trees occurring on an important dominant throughout the mixed the savannah and tall grass prairie include: grass prairie. Little bluestem is a warm smooth sumac (Rhus glabra), snowberry season bunch grass that commonly provides (Symphoricaz_o8 spp. ) dogwood (Cornu8 excellent cover for many wildlife species. spp.), hawthorn (CPataegu8 spp. ), plum There are many more species of Andropogons, (PPunus spp.), hazelnut (Cor_u8 ameT4eana), but the only other one noted here is oak (QuePcu8 spp.), elm (Ulmu8 spp.), and broomsedge (A. virginicus), which commonly eastern redcedar (Juniperus virginiana) occurs on old fields and overgrazed (Aikman 1929, Bruner 1931, Curtis 1959, pastures in the east. Kucera 1960, Weaver 1954).

Other warmseason grasses associated Periodic droughts restrict woody plant with the tall grass prairie include switch- invasion on the mixed and short grass grass (Panicum virgatum) and Indian grass plains to the moist areas along drainages. (Sorghastrum nutans). Prairie cordgrass These woodlands provide cover, food, and (Spartina pectinata) occurs around the landscape diversity for many wildlife prairie marshes and along the drainage species. Dominant trees include: ash systems throughout the tall and mixed grass (Framinu8 spp.), cottonwood (Populus spp.), prairie. Warm season grasses also dominate willow (Saliz spp.), elm (U_ spp.), and the short grass areas. Blue grama (Bouteloua boxelder (Aoer negundo). Many shrub species gracilis) and buffalo grass (Buchloe occupy the understory.

•Nomenclature for grasses follows 2Nomenclature for forbs, shrubs, and Hitchcock (1850). trees from Harrington (1854). The palouse region and Callfornia shovelers (Spatula clypeata) and gadwalls Valley were originally dominated by bunch (Anus strepera), which seek the grasslands; grass species, such as bluebunch wheat- green-winged teal (Anus cmeoline_sis), grass (Agropyron spica_) and Idaho fescue lesser scaup (Aythya affinis), bufflehead _estu_a idahoensis). The palouse prairie (Buoep_zla albeola), ring-necked ducks is now mostly under cultivation and the (Aythya collar_s), common goldeneyes CaliforniaValley has been converted to an (Bucephala cLrngula), and white-winged annual grassland containing many introduced scoters (Melanit_z deglmcdi), which are species including chess (Bromus spp.), more abundant in the semlforested park The desert grasslands are dominated by lands; and redheads (Aythya amez_ouna), curly mesquite (Hilaria belangeri), galleta canvasbacks (Aythya valisineria), and ruddy (Hi!ur_a jumesii), and grama (Bouteloun ducks (O_a jumaice_is), which seem con- spp.). The coastal prairies contain mostly tent with either a park land or a prairie tall grass species such as bluestem habitat (Llnduska 1964). (Ar_opogon spp.)_ and cordgrass (SpcD_tinu spp.). Over most of the northern unglaciated mixed grass plalns, natural ponds were Wi]dlife Species scarce under pristine conditions. Since the early 1930's more than 300,000 stock Waterfowl _ ponds have been constructed in North and • South Dakota, Minnesota, and Montana and

lag_eFeo-behdesindtructhetiodesn torucf watioterfn oowlf mohabstitoaft ' pr10o,v000inceshaveofbeenCanada.construcTheseted ponondsthepropducerairle the prairie by the plow because it was about 1,000,000 waterfowl each year easier to plow the dry uplands than to (Linduska 1964). Duck breeding populatlon drain the wetlands. Only a few years later, estimates vary between 3.3 pairs per with the deyelopment of large equipment square mile (1.8 pairs per surface acre (early 1900's) and govermental assistance of water), according to Lokemoen (1973) programs (1930's),1arge-scale drainage and 7.0 pairs per square mile (2.3 pairs programs Were initiated. By the 1950's, per surface acre of water), according to thesehad reduced the original wetlands in Bue et al. (1952). Broods averaged from the United States from 127 mi11ion acres 0.79 per acre of stock-watering pond to 82 miilion acres (Linduska 1964). (Lokemoen 1973) to 1.5 broods per acre of pond (Bue st al. 1952). Prairie potholes are the backbone of duckprodUction in North America. The During recent years, populations of prairie potholes region, 300,000 square the Canada goose (Br_tu ounu_is), such miles, makes up only 10 percent of the as those wintering on the short grass total waterfowl breeding area of this plains, have steadily increased because conti:nent, yet produces 50 percent of the prairie lands have been cultivated and duck crop in an average year--more during bodies of water impounded. Irrigation good Years (Linduska 1964). using water from these impoundments and from wells has provided large acreages of Wlth a few wet years the prairie wheat, milo, and corn that offer more and potholesprovlde habitat for a rapidly better foods than do native grasses expanding waterfowl population. Several (Grieb 1970). successive drought years bring an inevitable

,crash. Although droughts cause short- Upland GameBirds term declines in duck numbers, they help to.malntain fertility and increase pond Before settlement, greater pralrle life These long-termadvantages probably chickens (_mpunuchus c_pido) were confined far oUtwelght the short-term disadvantages, primarily to the tall grass prairie. Their range gradually extended westward as the Fifteen species of ducks nest commonly native sod gave way to farms and wheat in the prairlepothole region. Most abundant are the mallard (Anus pla_rhy_hos), s fields throughout the mixed grass prairie types (Cooke 1909). Prairie chicken pop- plntall (Anus uuUf_z), and blue-wlnged teal ulations thrived with white-man's first (Anus diseors). Other species include "i i i| attempt at farming. This was attributed ,- . to the abundance of food and undisturbed 3 Aviun nomenclature follows American nestlng areas ("prairie-type" land inter- F O_nitho_ogists Union (1957). mingled with patch farming). With patch

5 farming operations, prairie chicken populations peaked in Iowa about 1880 when 69 percent of the State was cultivated. By 1900, 90 percent of Iowa was cultivated and prairie Chicken numbers were decreasing (Stempel and Rodgers 1961).

Prairie chickens were first recorded in North Dakota in the early 1880's (Johnson 1964) and in Colorado in 1897 (SCiater 1912). Their populations increased ii_! and flourished during the "good pinnate years,' between 1900 and 1930. With the dust bowl days of the 1930's small ranches _i_ and farms were abandoned or incorporated into larger holdings. This led to large areas of intensive cultivation on the better soils and grasslands only where the soils were too sandy or the land too hilly to f_rm. _With very little winter food on the grasslands and virtually no nesting cover on the farmlands, prairie chicken ° numbers decreased drastically. Prairie chicken numbers followed this same pattern over most of the Great Plains (Beck 1957, Evans and Gilbert 1963, Johnson 1964).

The lesser prairie chicken (_armo_s pal lidi_tus) inhabits a small area of Figure 2.--_-_iled grouse have been grassland and brushland located in and still are the pro_r_v_t upland g_e northern Texas, western Oklahoma, south- bird of the northern bz_sh_, park • western Kansas, southeastern Colorado, l_n_s, and plains. and eastern New Mexico. Their populations :are fairly stable. Habitat of the Attwater's 1. The climax sagebrush of the prairie chicken (Tymp_nuchus c_pido northern desert shrub area at_ateri) is disappearing along the Texas (Columbian sharp-tailed grouse). Cull Coast; consequently, this bird is currently listed as an endangered species. 2. The subclimax brush in the grass- The heath hen (Tympanuchus _pido cupido), lands east of the Rocky Mountains which formerly occupied the coastal and in the park lands of the prairies of the New England States, is Rockies (plains sharptail). now extinct. 3. The oak-savannah and logged or Since pioneering days the sharp- burned areas in the east (prairie tailed g_ouse (Pedioecetes phasianellus) sharptail) . has been a part of the animal llfe on the • brushlands, park lands, savannahs, and 4. Openings in the boreal forest plains of much of the northern United (northern sharptail, Alaskan States and Southern Canada (fig. 2). sharptail, other boreal forest JOhnson (1964) stated that sharptails races). Probably have been in North Dakota for . hundreds of years. As with the prairie The bobwhite quail (Colinus virginian, s) chicken, sharptail numbers decreased during is a savannah species, with its populations the drought of the 1930's, and never re- most stable in the southern two-thirds of gained their previous high. Intensive the savannah area (Rosene 1969, Johnsgard cultivation and certain grazing practices 1973). In the northern third of the have decreased sharptail range and pop- savannah, populations decrease during years ulations numbers throughout most Great of adverse climatic conditions. The Plains States. bobwhite has adapted to the habitats associated with agriculture; consequently, Habitats of the sharp-tailed grouse it is abundant even in extensively cultl- include (Aldrich 1963) : vated areas. The wild turkey (MeleaF_s gaIlopavo) Birds of prey occurring in grasslands is considered a forest bird; however, include the red-tailed hawk (Bu#eo archeological evidence indicates that _umuioensis), Swainson's hawk (B. Indians in the savannah utilized the turkey swai_oni), rough-legged hawk (B. Z.agopus), for both foodand ornamentation. Shortly sharp-shlnned hawk (Aoo_piter stz_zt_s), after 1900, exploitation and habitat and Amerlcan kestrel (Fa_oo sparver_,us). destruction resulted in the extirpation or The large hawks feed on rabbits, small near extirpation of the turkey in the mammals, and snakes, the small hawks and savannah (Hewltt 1967). Today, as a falcons on small mammals and birds, snakes, result of State restocking programs, and insects. In addition to their Impor- turkeys are found throughout all their tance in food chains, these birds contri- former range as well as in areas that bute to man's pest control efforts. The orlglnally did not support wild turkeys nocturnal raptors include the great horned (Sanderson and Schultz 1973). Good turkey owl (Bubo viPginianus), short-eared owl range contains trees for roosting and trees (Asio _u_rneus), long-eared owl (Asio that produce mast and fruit for food. otus), and burrowing owl (Speo_jto The grasses and forbs of the understory cunicularia). Many other avian species produce seeds and habitat for insects occur on the grasslands. Their role and upon which.young turkeys feed. Occasion- function in grassland ecosystems is ally turkeys will feed on cultivated discussed in Wiens and Dyer (1975). crops. Turkeys are found in the savannah as wellas along the wooded stream and , Bison river drainages throughout the mixed grass The American bison (Bison bison) _ was prairie, the one feature of the North American central grasslands that all historians, The mourning dove (ZenaidamacrouPa) naturalists, trappers, or travelers men- (American Ornithologists' Union 1973) is tioned in the diaries concerning the prlmarilymlgratory and depends on the pristine conditions of the area (fig. 3). northern prairies and savannahs for llfe Based on a series of assumptions about requirements only during the breeding carrying capacity, range area, habits, and season. Populatlons do winter in the population trends, Seton (1929) estimated southern grassland types, depending on a populatlon of from 40 to 60 million biaon food availabillCy. Preferred habitat for in North America. Bison were once spread the mourning dove consists of tree cover over one-third of the North American for nesting and fields with open cover for Continent w-Ith the largest herds dlstri- feeding. Hedgerows, shelterbelts, orchards, buted along the Mississippi River Valley. or woodlots provide acceptable nesting During the years previous to disturbance habitat (Hanson and Kossack 1963). When by white man, the total area inhabited by conifers are available, doves prefer them bison was about 3 million square miles for nesting (Hanson and Kossack 1963, (Scion 1929). Caldwe11 1964). The weak feet and bill of the mourning dove limits its feeding Overgrazing by bison, in association activities to areas of sparse cover. Hat- with trampling, rubbing, and wallowing, vested fields_ fleldmargins, and over- contributed to the creation and maintenance grazed pastures contain the waste grains of environmental conditions favorable to a (wheat and corn) and weeds (foxtail) pre- variety of other wildlife (England and ferredbythe dove (Korschgen 1958, Hanson Devos 1969). Bison are primarily grazers, and Kossack 1963, Ward 1964). so they depended on the herbaceous vegetation for food. Few herbivores possessed Nongame Birds the capability for altering the environment , as did the bison. These animals require Predatory birds have suffered because approximately 30 pounds of forage per day, of man's ignorance and apathy. Overzealous so even large areas could easily be control programs and pesticide use have overgrazed. serlously reduced some species populations and causedothers to be placed on the en- A dramatic change occurred on the dangered and threatened species llst. As Great Plains during the 1800's. Prior to one becomes knowledgeable of the feeding habits of these birds, it is evident that they, too, are important components in the _Ma_nalian nomenclature from Hall biotic communities to which they belong. (1965).

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i _ _ _ ..... _ %___%_ii_

!!_...... _i___ _!_i_i_ _iiiiiiii!!_'_:'_:_''_: ...... Figure 4.--The present day land-use patterns provide habitat for extensive herds of

• _ '__i_i_iii_:_I_ ::i!_!!_i:'_::i_iiii_!!_:_...... _...... __ iilJ_:...... iii!'i!i_:_iii_!I pro___ antelope, ,°_i_,i_,_ ._ .... _i__; _ _._i_'___._i%_,,...._,_...... _ _,_...._...... _,___,., _._i_,_,_i_i_......

F_gure 3.--T_e bison or American buffalo don't refer to excessive slaughter of w_s _e the dominant animal on the antelope. Therefore, _f antelope were North American central grasslands, as numerous as estimated, and _f they were not: slaughtered to the same extent as bison, this t_me the Indians were completely de- why d_d population numbers decrease so pendent on bison for their llvellhood, greatly? Their hunting methods did not cause major changes _n b_son numbers, and usually re- Perhaps "Itwas due to the reduction sulted in a harvest of surplus animals, in suitable habitat brought about by the It is est_nated that approximately 300,000 extermlnat_on of b_son and the extensive plains Indians existed primarily by hunting construction of fences. Perhaps their i withennipegb_son andherdsfrombetwetheen RockyMexicMounto andainsLaketo susceptidecline bilitycan be toattribdutedisease. to Travea h_lighng i the M/ssissippi River (Allen 1954). The from Fort Abraham Lincoln west to the part-Arab horses, escaping from the Little M/ssouri in 1873, J. A. Allen re- [ Spaniards, gave the Indian additional ported that he saw antelope almost con- t hunting ability. Even with the horse and stantly (Bailey 1926). In the interim bow the Indian could not seriously deplete before his return trip a few months later, his resource, as the gun and fence was to a fatal epizootic raged among the prong- do later _ Liquidation of the bison took horns over most of the area between the . about 50 years--by 1883 only a few animals Yellowstone and Missouri Rivers and 75 to remained. 90 percent of the pronghorns perished.

Pronghorn antelope populations de- Pronghorn Antelope creased from an estimated 40 million (Seton 1929) to 13,000 around 1915 (Hoover ' Before intensive settlement of the et al. 1959). The species was then afforded Unlted States, pronghorn antelope rlg_d protection by landowners and law (Anti_ouupr_ _mer__) (fig. 4) were the enforcement agencies, and their current codom_nant grazing species with b_son on population _s estimated at 365,000 the Great Pla_ns and with elk on the (Yoakum 1968). The increase _s attributed Paiouse and Callfornla Prairies. They to: (I) controlled hunting, (2) favorable were as numerous, if not more so, than the habitat, and (3) improved wildllfe man- bison (Rand 1945). The present range of agement practices such as transplantlng pronghorns is more restricted (England programs, water developments, and fences and Devos 1969). Most historical writings constructed to permit pronghorn travel.

, Pronghorns fill a forage utilization Tule elk. First, the Spanish settlements niche that isbetween the grazers (bison of the late 1700's and the invasion of the and elk) and the browsers (deer). Prong- annual grasses which resulted in the dis- horn antelope have adapted well to changes placement of the dominant bunch grasses, in prairie vegetation resulting from live- which no doubt had adverse effects upon stock grazing, the Tule elk population (McCullough 1969). Then the increase in competition by wild When populations of pronghorns and cattle and horses, the gold rush (1848), livestock are in adjustment with the forage and the resulting slaughter of wild animals capacity of a good condition range, corn- during the booming market-huntlng years, petition between cattle and pronghorns is further decimated the population. By negligible. The compatability of cattle 1855, the market-huntlng era brought the and pronghor n antelope is similar to that Tule elk to the end of its time as an which must have existed, in the past, important element in the fauna of the between bison and pronghorn. Competition region. Today there are about 400 to 500 between domestic sheep and pronghorns can Tule elk in California's Central Valley be so severe that the latter will be (McCullough 1969). The wapiti or Nelson's ellmlnated. Sheep, Which can be kept alive elk have been completely eliminated over with supplemental feeding, can overgraze the rest of the grassland habitats they the-vegetation to a point where it is formerly occupied. Their populations are unable to support pronghorn antelope now limited to high mountain ranges. (Buechner 1960),

Elk

The pristine range of the elk, or Deer wapiti, was as widespread in North America as that of the bison. At least two species The white-tailed deer (D_u v_rg_n_) Of elk inhabited the grasslands and wooded was a frequent part of the hunter's bag river bottoms: Nelson's elk or wapiti during the Lewis and Clark expedition of (Cer_s cunuder_sis), which inhabited the 1804 to 1806. Mule deer (Duma hen_or_s) midcontinent grasslands, and the northern shown in figure 5 were less abundant but Pacific coast grasslands, and the Tule elk occurred as far east as Chamberlain, South (C,:r_v_nodes), which occupied the Southern Dakota (Lewis 1961, Cutright 1969). The . Pacific area in the Central Valley of Astoria party, which ascended the Missouri • in 1811, made no mention of deer while on California (McCullough 1969). an overland Journey from the mouth of the Cannonball River until they reached the The carrying capacity of the Tule elk Black Hills of Wyoming (Irving 1868) was estimated at 500,000 animals in although they made frequent references to pristine time. Several factors contributed white-tailed deer while traveling along to the decline and near extinction of the rlverways.

:.__' Figure 5.--Mule deer populations ,:_ .....ii_,_ _:,_ _.._:_:,,_!_ increased on the midcontinent

"_:i_i_i_..... : grasslandsof hunting withand ththee increcontrolase ...... _:::_...... " _ in the able of woody

_.:.

9 Bailey(1926) reported mule deer to Small Game be plentiful throughout North Dakota untll about 1880 when their numbers began to Increased population size of the decrease as hunting pressure mounted. By eastern cottontail rabbit (SylviY.ugus 1888 deer Were fairly well exterminated flo_s) on the savannah and tall grass from NorthDakota because intensive live- prairie coincided with the advent and stock grazing eliminated the shrubby species expansion of agriculture (Lord 1963). that provided excellent browse and cover Its smaller home range makes it more tom- for deer. In the Great Basin and south- patible with intensive agriculture west, however, livestock grazing encouraged (Beck_fth 1954). Many miles of brushy the invasion of shrubs (Longhurst 1960); fencerows and grassy road ditches adjacent thus, deer habitat was often improved, to cultivated fields have furnished good Mackie (1970), in the northern Great Plains, habitats capable of supporting high cotton- reported that the food and range-use habits tail populations. The unspecialized of mule deer and cattle differed substan- habits of the cottontail make it likely tially. Deer ingested primarily shrubs to remain an abundant game animal. and forbs, cattle mostly grass. Deer inhabited relatively steep slopes, cattle The tree squirrel most commonly the more open ridgetops and coulee occurring in the savannah and along river battoms, bottoms is the fox squirrel (Sc__s niger). It depends on fruits and nuts but also With the disappearance of the bison' feeds on crops when these are available. and elk, the white-tailed deer became the One to three squirrels per acre is consid- only large herbivore left on the tall grass ered optimum for this type of habitat prairie and savannah. The forest clearing (Schwartz and Schwartz 1959). and patch farming practiced by the _mmigrating settlers increased the edge Two lagomorphs inhabit the extensive habitat preferred by the white-tail and upland grassland types. The white-tailed this contributed to a substantial increase Jackrabbit (Lepus t_onser_ii) on the in the deer population (Severinghaus and northern Great Plains and the black-tailed • Cheatum 1956). On the northern savannah, Jackrabbit (Lepus califor, nic, us) on the however, their numbers declined to less southern Great Plains. Both depend on than 2,000 in the face of increasing human good eyesight and speed to avoid enemies populations pressures and year-round because they are not burrowing animals hunting. The 1930's depression caused like many other small mammals. Rather, farm abandonments resulting in widespread they rest and hide in shallow depressions, secondary succession, and the restoration or "forms", at the base of a shrub or of the habitat (Beckwith 1954). By 1968, clump of grass deer numbers on the northern savannah had PrairieDogs increased to an estimated 845,000 (Nixon 1970). The most conspicuous of the grassland rodents were the prairie dogs (Cy_s Btghorn Sheep spp.). Prairie dogs were once incredibly numerous. Because prairie dogs eat grass Before disturbance by the white man, and compete with the livestock industry, • bighorn sheep (Oois oaru_n_sis) were their numbers have been reduced by exten- numerous in mostmountain ranges of the sive poisoning progrms. Dobie (1949) west, and in the "badlands" along portions reported information £rom writers during of the Missouri, Little M/ssourl, the late 1800's of prairie dog towns cover- Yellowstone, North Platte, Arkansas, ing 16 million acres with an estimated Colorado, Green, and Gila Rivers, and their 400 million animals. These prairie dog principal tributaries. They were never towns provide food and/or cover for prairie characteristic prairie dwellers as some- rattlesnakes (Cro_Zus vir_dis), burrowing times rePorted, but they did occupy some owls, (Speo_jto cwzi__), and the of the buttes and badlands of the Great black-footed ferret (_uste_znig_pes). Plains region. They were exterminated The black-footed ferret is completely early in the history of white man settle- dependent upon the prairie dog town for ment. The last Audubon or badlands bighorn both shelter and food. Man's wholesale sheep (0. c. audubcmi) was killed in 1916, destruction of the prairie dog has so re- near the White River of South Dakota duced ferret populations that this hand- (Hipschman 1959). some weasel is now one of America's rarest

10 _---,-is. Many other wildlife and plant to the virtual elimination of the mountain species are associated with the influence llon by 1900 (Schwartz and Schwartz 1959). of a pralrle dog town. The bobcat has survived, but its range is much more limited (Schwartz and Schwartz Manmal_ian Predators 1959). It. has also undergone persecution because of the "all-predators-are-bad" The grassland herbivores originally philosophy. This legend seems to rest served as food supplies for a number of more on fiction than fact because Korschgen mammalian predators. The best known species (1957a) determined that the bobcat diet are the gray wolf (C_is lupuS), red wolf is primarily rabbits and squirrels. (C..rufus), coyote (C. Lz_ans), red fox (Vulpes. vulpes), gray fox (Urocyon The striped skunk (Mephitis mephitis), cinereoargenteus), mountain lion (Felis spotted skunk (Spilogale putorius), and concolor), and bobcat (Lynx rufus). The raccoon (Procyon lotor) have survived in ' gray wolf depended primarily on the bison close proximity with settlement; and in and elk fur food. Originally they ranged some cases they llve around homeslte throughout the United States, but now the buildings. Their food supply consists of only remaining populations are in northern small mammals, small birds, reptiles, Minnesota, Wisconsin, and Michigan (Mech amphibians, insects, and eggs. 1966). Loss of preyspecies and heavy hunt'ing led to the demise of the gray Fish wolf. It had disappeared from the central, unlted;States by 1900 (Schwartz and The important fishes occurring in the Schwartz 1959). grassland streams and lakes are the minnows (Cyprinidae), the suckers The fate of the red wolf, which (Catostom_dae), the catfishes (Ictalumidae), originally ranged from Illinois and Indiana the sunfishes (Centrarchidae), and the south through the Ozarks and into Texas, perches (PePcic_e). Members of the sunfish was similar to that of the gray wolf. and perch families have been widely in- The current status of its population and its troduced into manmade reservoirs and diStribution is questionable; however, it stock-waterlng ponds. The best known are 'is generally conceded that the red wolf is the large-mouth bass (MicroptePu8 8almoides)s restricted in both categories. Thus, it and the walleye (Stizostedion vitreum), has been placed on the endangered species (Schultz 1936, Koster 1957, Bailey and list by the United States Department of Allum 1962, Cross 1967). interior, in the 1940's Leopold and Hall (1945) reported seeing red wolves in the western Ozarks but Plmlott and Joslln Amphibians (1968) have since reported that the distribution of this wolf currently is Arid climates pose special problems limited to the coastal prairie areas, for amphibians. Grassland amphibians have been able to adapt to the arid climates The coyote is the only canine pre- through the evolution of behavioral or dator of any significance remaining on physiological traits that tend to conserve the grassland and savannah. Coyotes are moisture. For example, the tiger salamander continuously subjected to harsh control (Ambystoma tigrinum),e plains spadefoot programs and heavy hunting pressure, yet (Scccphiopus bombifrons), Rocky Mountain Coyotes continue to expand their range, toad (Bufo woodhousei), and Great Plains The coyote prefers a grass-shrub or grass- toad (B. cognut-us) are able to escape tree cover combination and its apparent drying conditions by burrowing. Rapid adaptability enables it to rapidly occupy development of their young enables the all such areas created by man. spadefoot to use large temporary rain pools ° for breeding. The red fox and gray fox are found in the tall grass and mixed grass prairies. The most widely distributed amphibian These two species seem to be adapting to in the grassland is the leopard frog the habitat changes caused by current land use.

Depredations by mountain llon on SFish nomenclature follows Eddy (18,57) domestic livestock and the resultant con- SAmphibian nomenclature follows Conunt trol campaign plus habitat destruction led (18,58).

ii (R_pipier_), which occurs in some dry nearly 200 million acres of midcontinent habitat butgenerally Within traveling short grass-mixed grass type. This area distance Of permanent water sources. The supports a large livestock industry and bullfrog (Rune catesbeiar_) is widespread has a large potential for wildlife protec- in the central and lower Great Plains and tion. Livestock management systems are now has been widely introduced elsewhere. The in operation on private and public lands bullfrog is aquatic and must have permanent that strive for multiple use of the grass- water sources to survive, land with coordinated livestock and wildlife use.

Reptiles-. The large number and diversity of Many of the reptiles (turtles, llz- wildlife species plus the variability with- ards, and snakes) have adapted to arid in the grassland plant communities provides conditions. The more common turtles in- for a large number of wildlife habitat clude the common snapping turtle (ChelydPa management options. These options are yellow mud turtle se_ent_na), further complicated by year-to-year changes (KinoS_e_m flavesce_s ) , painted turtle in weather and precipitation. Rangeland (Ch_se_js pica%u), three-toed box turtle wildlife and livestock depend on forage (TePPuper_ e_Poli_n), ornate box turtle production for food and cover. Forage (T. oPRu_), and spiny soft-shelled turtle production can vary from nearly nothing (Tr_onyx spinifer). Except for the box on the high plains during droughts to turtle, all require permanent sources of ' 9,000 pounds per acre (Curtis 1959) on water, the tall grass prairie. The discussion Most lizards and snakes don't need that follows relates to the food and cover free water to llve. Species found through- requirements of many of the species out the grasslands include: earless liz- previously mentioned. The land manager ards (Holbrookia spp.), horned lizards must use imagination and foresight to (Ph_nosomG spp_), spiny lizards (Scelopor_s design specific systems to provide for an spp.), whiptails (Onem_dopho_s spp.), optlm_balance of different habitat skinks _E_meoes spp.), collared lizards components. " (6_o_hy_8 coILlri8), hognose snakes .(Hetero_:)n spp.), coachwhips (Masticophis spp.), racers (Coluber constrictor), king- Wetlands snakes (Laz_ropeltis spp.), black-headed snakes (Tantilla spp.), and rattlesnakes Many species of wildlife are associ- (C_o_lUs spp.). Garter snakes (Tha_nophis ated with the natural wetlands and manmade spp,) and water snakes (Natr_ spp.) are ponds of the grassland areas. Waterfowl f0undwherever there are streams or lakes and shorebirds are of primary importance. (Stebbins 1954, Conant 1958). Grassy shorelines support nearly three times as many breeding pairs of ducks as the bare mud type (Linduska 1964). The HABITAT MANAGEMENT composition, density, and height of the shoreline vegetation depends largely on Despite widespread vegetational the type of grazing system involved. changes, the potential value of much of Season-long grazing systems will deplete the original grassland areas as rangeland the shoreline vegetation even if livestock and wildlife habitat remains very good. stocking rates are low, unless there are Some of theassociated wildlife species several ponds or watering areas in each are now absent; populations of others have pasture. Shoreline vegetation'can be been reduced, but the majority of species maintained by fencing all or part of a haveadjusted to habitat alterations and stock dam or by using carefully designed remain important components in the modern rest-rotation grazing systems. Shorelines biotic communities. The oak-bluestem that are completely protected from livestock savannah, tall grass prairie, and the trampling and grazing often contain dense western bunchgrass types have been most emergent vegetation or shrubs that are altered by land use practices. The largest not sultable for dabbllng duck habitat. grassland typeremalning in America is the Management plans should be designed to produce some mudflats for shore bird feeding ' areas, grassy shoreline areas for duck 7Reptile nomenclature follows Stebbins nesting, and some emergent vegetation (195@_ and C_unt (_8). growth for duck brood protective cover.

12 Uplands (Robinson 1957). Dense grassland cover can be opened by using fire, cultivation, Brushy cover is to sharptails what or grazing. Croplands provide crop residue grass is to prairie chickens, second growth and weed seeds which are valuable bobwhite is to ruffed grouse, and mature coniferous fall and winter foods. Corn (Zea muys), forest is tospruce grouse. On good Korean lespedeza (Lespedeza stiFalacea) , moisture and soil sites, lack of grazing and sorghum (Sorghum v_lgaPe) are cul- or rest will bring about an increase in tlvated crops which rank high in the bob- such desirable species as wild plum (Prunus white diet. Ragweeds (Ambrosia spp. ), ameP__), chokeberry (P. virginiana), wild beans (Strophostyles spp.), CPoton silver buffaloberry (Shephe_dzz _Pgenteu), spp., foxtall grasses (Setu_a spp. ), and and hawthorn (C_u_egu8 spp.), sunflowers (Helianthus spp. ) are native I annual plants and are prolific seed Food habits of the sharp-tailed producers commonly associated with disturbed grouse are nearly as varlable as the areas (Baumgartner et al. 1952; Korschgen habitats they occupy. Several plant species 1952, 1960; Robinson 1957). Insects ' do occurln most food habits studies, in- associated with cultivated crops and forbs diCating their importance for food and/or are used for food in spring and summer cover. Tkese species include dandelion (Rosene 1969). The brushland and wood- (Taraxac_m officinale), rose (Rosa spp.), land areas are used for escape, roosting, hawthorn (CPataegus spp.), snowberry and feeding. Casey (1965) questioned the (SymphO_ca_pos occidentalis), Russian ° need for tree cover if brushland was olive (EZ_e_s _gustifolia), silver available. Robinson (1957) provided buffaloberry, and buds from Populus species, support for this idea when he found the Cultivated crops are eaten when available number of coveys in an area to be dependent (Evans 1968). Several of these berry- on the number of "headquarter areas", which producing species provide grouse with a were brushy cover dense enough to provide winter diet sufficient to survive winter adequate protection during periods of high Conditions on the northern Great Plains light intensity. Cover type distribution (Hillman and Jackson 1973, Evans and must be planned and incorporated into a Moen1975). total management plan so that access to all cover types will be available within Evans and Dietz (1974) tested seven the home ranges of a covey; a home range diet materials and found nitrogen-corrected varies from 12 to 20 acres. metabolizable energy (Kcal/g. dry matter) to range from 3.91 for corn to 1.39 for The cottontail rabbit is a close rose hips. The fruit of silver buffaloberry associate of the bobwhite over much of was the best native winter food item their ranges. The most preferred foods tested. Fleshy hawthorn berries were in- for cottontail rabbits in Missouri, are gested in larger quantity than other air- bluegrass (Poa pPatenses), wheat (TPitic_#_ dried foods. For maintenance, winter aestivum), and white clover (Trifolium foods needed to be consumed in large enough repens) (Korschgen, unpublished data). A quantity to provide metabolizable energy in wide variety of food species appearing in excess of 1.5 times basal metabolic rate. smaller amounts serve to illustrate the A keyto intensive management of winter catholic taste of the cottontail. habitat for sharp-tailed grouse is the Cottontails may become a nuisance if in- propagation and/0rencouragement of high clement weather forces them to feed on the • energyaproviding plants. Several native bark of shrubs, shade trees, orchards, and shrubspecies provide good protective cover windbreaks, thus damaging or killing in- for sharptails and produce berries that dividual plants. Hunting or trapping can are palatable and high in energy, be used to effectively control cottontail populations which are exceeding the Edmlnster (1954) listed four types of carrying capacity of the habitat. cover required by the bobwhite: grassland, cropland, brushland, and woodland. Within the various rangeland habitats The grassland provides nesting and feeding from sea level to above timberline, many cover durlngsPrlng and s,,mmer. Parmalee management practices are available for in- (1.955) found 85 percent of the nests creasing grazing capacity, improving range located during hls study to be in grass- condition, and creating new range. All land. Grassland must be open enough to are designed to favor livestock. The non- permit birds to move about unhampered game bird section of most management plans

13 are shortand generally useless for required to remove range forage sufficient evaluatlngeffects of different practices to feed 1 cow (Stoddart and Smith 1955). on birds and their habitats. The interest and awareness of animals other than cattle The coyote often preys on livestock and sheep is rapldly growing as managers and blg game species; however, Korschgen recognlzethe value of diversified wild- (1957b) reported that the coyote diet in llfe populatlons. Buttery and Shlelds M_ssourl consisted mainly of rabbits and (1975) have summarized information per- rodents. Thls agrees wlth Murle (1940) talning to bird habitat values and have who determined that coyotes in Yellowstone pointe d out research needs, consumed primarily rodents and insects. Consumption of 11vestock or wild mammals Big Game other than those mentioned above was 10 percent or less for each. The evidence indicates that often the coyote could be Mule deer, white-tailed deer, and considered a valuable aide in controlling pronghorn antelope currently offer the rabbit and rodent populations. In local best potential for big game management, instances coyotes can cause economically Deer are browsers and require a supply of woody plants for food and cover. Good significant sheep mortality. sharp-tailed grouse habitat on the northern In spite of the fox-chicken legend Great Plains is usually compatible wlth good deer habitat. Some livestock grazlpg it appears that the foxes still depend on systems are very compatible wlth deer and wild animals for the major portion of their diet (Korschgen 1957a). Rabbits grouse populations. Grazing systems that and rodents comprised more than 50 percent reduce brushy cover in the small drainage areas probably also reduce deer habitat beenof theconsidereddiet for btoothhavespecboies.th deFtorixesmenthaveal values, and beneficial attributes. Detrimental

Whlte-tailed deer range can best be because they occasionally take domestic described as a mosaic of croplands or poultry but beneflclal because of the pasturelands and oak-hlckory woodlots, impact they make on rodent populatlons and brushy fencerows, and wooded stream bottoms the value of the pelt. (Crawford 1970). A survey of deer food habits in Missouri (Murphy 1970) revealed DISCUSSION the importance of cultivated crops In the diet, especially corn and soybeans. This Past experience indicates that future survey showed substantial dependence on habitat management objectives are going to shrubs and small trees as a food source, require conslderable thought and imaglna- • Important shrub species are buckbrush tlon. The bison, elk, and gray wolf are (Sgvr;phoz_o_o8 spp.), sumac (R_s spp.), gone from the grasslands and savannahs-- and hawthorn. Acorns are an important victims of changing land use and indlscrlmant contribution of the tree component in the shootings. Those game species remaining more heavily wooded areas. Favorable have been able to adapt to changing land habitat extends along the major drainage use but even these species are limited in across the Great Plains. the amount of change that they can tolerate. We have hardly considered the multitude of Pronghornantelope differ considerably nongame species present. We know that land from deer in their habitat preferences, use changes influence these species Deer prefer woody or brushy areas, and (Warbach 1958, Graber and Graber 1963, antelopeprefer upland and open expansions Buttery and Shlelds 1975), but insufficient of grasslands. The forb component of datamakes accurate evaluation of these , grassland vegetation communities Is changes difficult. Wildlife has continually important to pronghorns, been relegated to areas that are difficult to cultivate or to intensively manage, and these areas continue to dwindle as new Predator-PreRelay tions economic uses are discovered. There are two broad alternatives available to JaCkrabbits occaslonally become so society. The first is to continue man- numerous that control campaigns are aging with a dwindling habitat base usurped launched. Because of their size and by an expanding urban population and inten- differences in food habits, it is estimated siftcation of agriculture practices; or that approximately 200 Jackrabbits are secondly, stimulate incorporation of

14 habitat management practices in land use American Ornithologists' Union. 1973. planning through education and economic Thlrty-second supplement to the American

. incentives. Ornithologists' Union checklist of North American birds. Auk 90(2):411-419. The land manager must have some in- Anderson, Kllng L., Ed F. Smith, and centive forproduclng wildlife. It is Clenton E. Owensby. 1970. Burning unreasonable to expect a private land man- bluestem range. J. Range Manage. agerto voluntarily donate his most pro- 23(2):81-92. ductlve land to wildllfe habitat when the Anderson, Wallace L., and Lawrence V. demands of an urban population for farm Compton. 1958. More wildlife through andranch products are so great (Crawford soil and water conservation. U.S. Dep. 1970) The practice of charging hunters a Agric., Soil Conserv. Serv. Agric. Inf. fee to useprivate lands is relatively new Bull. 75, 15 p. in manyStates, but well established in Bailey, Reeve Maclaren, and Marvin O. A11um. more populated areas. This economic in- 1962. Fishes of South Dakota. Univ. centive is becoming increasingly popular Mich V Mus. Zool., Misc. Publ. ll9, 131 p. (Severson and Gartner 1972). Consumptive Bailey, Vernon. 1926. A biologlcal survey users havetradltionally shouldered the of North Dakota. U.S. Dep. Agrlc. burden of_game management costs through North Am. Fauna 49, 226 p. license fees and special taxes on hunting Baumgartner, Frederick M., Meredith J. equipment. Very llttle of this money has Morris, John L. Steele, and Jack E. gone directly to the landowner as an ' Williams. 1952. Oklahoma bobwhite food economic return for the wildlife produced relations. North Am. Wildl. Conf. on his land. Future wildllfe habitat 17:338-359. programs will depend on whether or not man Beck, John V. 1957. The greater prairie is willing topay a portion of the wild- chicken in history. Nebraska Bird Rev. llfe productions costs. Adequate procedures 25(1):8-12. for managing wildllfe and methods of application are readily available (Wing Beckwsucclthe,ssioStnephenon abandonedL. 1954.farmEcologllandscaland 1951, Anderson and Compton 1958, Giles its relationship to wildlife management. _1969, Allen 1972), but wld_spread Ecol. Monogr. 24(4):349-376. application awalts a basic philsophlcal Borchert_ John R. 1950. The climate o_ the change_in which wildllfe will be recognized Central North Amerlcan grassland. Assoc. as an esthetic necessity rather than an Am. Geogr. Ann. 40(1):1-39. esthetic luxury. The fate of wildllfe Bruner, W. E. 1931. The vegetation of is inexorably llnked with the fate of man. Oklahoma. Ecol. Monogr. 1(2):99-188. Bue, I. G., Lytle Blankenship, and • LITERATURECITED William H. Marshall. 1952. The relationship of grazing practices to Aikman, John M. 1929. Distribution and waterfowl breeding populations and structure of the forests of eastern production on stock ponds in western Nebraska. Univ. Nebraska Stud. South Dakota. North Am. Wildl. Conf. 26(1 & 2):1-75. 17:396-414. Albertson, F. W., G. W. Tomanek, and Buechner, H.K. 1960. Regulation of A, Riegel. 1957. Ecology of drought numbers of pronghorn antelope in cycles andgrazlng intensity on grass- relation to land use. l__n_nEcologyand lands of Central Great Plains. Ecol. management of wild grazing animals in Monogr 27_I):27-44. temperature zones, p. 226-285. F. Aldrlch, John W. 1963. Geographic Bourliere, ed. Eighth Tech. Meet. orientation of American tetraonldae. IUCN, Warszawa. J. WildlManage. 27(4):529-545. Buttery, Robert F., and Paul W. Shlelds. Allen, Durward L. 1954. Our wildllfe 1975. Range management practices and

legacy. 422 p. Funk and Nagnalls Co., bird habitat values. -.In=-. Management of New York. forest and range habitats fer nongame Allen, Durward L. 1972. The future of birds Symp. Proc. Dixie R. Smith, wildlife resources. J. Soil & Water Tech. Coerd. USDA For. Serv. Gen. Tech. Conserv. 27(6):244-249 Rep. NO-l, p. 183-189. American Ornlthol0gists' Union. 1957. Caldwe11, Larry D. 1964. Dove production Checklist of North American birds, and nest site selection in southern 5th ed. 691 p. Am. Ornlthol. Union, Michigan. J. Wild1. Manage. 28(4): Ithaca, New York. 732-738.

15 Casey, William H. 1965. Some speculations Evans, Keith E., and Aaron N. Moen. 1975. on theminimumhabitat requirements of Thermal exchange between sharp-tailed b6bwhite quail. In 19th Annu. Conf. grouse (Ped_oeoet_8 p_s__IZu8)and Southeast. Assoc. G--ame & Fish Comm. their winter environment. Condor 77 Proc., p. 30-39. (2):160-168. Conant, R. 1958. A field guide to reptiles Giles, Robert H., Jr., ed. 1969. Wild- and amphibians of the United States and life management techniques. 3rd ed. Canada east of the 100th Meridian. 623 p. The Wildl. Soc., Washington, DC. 366 p. Houghton Mifflin Co., Boston, Graber, Richard Rex, and Jean W. Graber. Massachusetts. 1963. A comparative study of bird CooMe, Wells W. 1909. The birds of populations in Illinois, 1906-1909 and Colorado. Auk 26(10):400-422. 1956-1958. Ill. Natl. Hist. Surv. Bull. Coupland, Robert T. 1958. The effects of 28(3):377-528. fluctuations in weather upon the grass- Grieb, Jack R. 1970. The shortgrass lands of the Great Plains. But. Rev. pra±rie Canada goose population. Wildl. 24(5):273-317. Monogr. 22, 49 p. Crawfo_d, Hewlette S., Jr. 1970. Mid- Hall, E. Raymond. 1965. Names of North western deer habitat. In White-tailed American m-mm_ls north of Mexico. Univ. deer in the Midwest. US-DA For. Serv. Kansas Mus. Nat. Hist. Misc. Publ. 43, Res_ Pap. NC-39, p. 19-22. North 16 p. Cent. For. Exp. Stn., St. Paul, Minn. Hanson, Harold Carster, and Charles W. Cross, FrankB. 1967. Handbook of fishes Kossack. 1963. The mourning dove in in Kansas. Univ. Kansas Mus. Nat. Illinois. Ill. Dep. Conserv. Tech. Hist. Misc. Publ. 45, 357 p. Bull. 2, 133 p. Curtis, John T. 1959. The vegetation of Harrington, H. D. 1954. Manual of the Wisconsin. 657 p. Univ. Wis. Press, plants in Colorado. 666 p. Sage Books, Madison, Wis. Denver, Colorado. Cutrlght, Paul Russell. 1969. Lewis and Hewitt, Oliver H., ed. 1967. The wild Clark: pioneering naturalists. 506 p. turkey and its management. 589 p. The Univ. Ill. Press, Urbana, Ill. Wildl. Soc., Washington, DC. Daubenmire, Rexford F. 1968. Ecology of Hillman, Conrad N., and Warren W. Jackson. fire in grasslands. Adv. Ecol. Res. 1973. The sharp-tailed grouse in South 5:209-226. Dakota. South Dakota Dep. of Game, Davis, A. M 1964. The Ozark savannah. Fish, and Parks. Tech. Bull. 3, 62 p. (Abstr.) 17th Annu. Meet. Am. Soc. Hipschman, Don. 1959. Department history, Range Manage. p. 7-8. 1909-1959. l_.n_nLooking back over the Dobie, James Frank. 1949. The voice of past 50 years. South Dakota Dep. Game, the coyote. 386 p. Univ. Nebraska Fish, and Parks 1959 Annu. Rep., p. Press, Lincoln, Nebraska. 13-73. Eddy, Samuel. 1957. How to know the Hitchcock, A. S. 1950. Manual of the freshwater fishes. 253 p. Wm. C. Brown grasses of the United States. U.S. Dep. Co., Dubuque, Iowa. Agric. Misc. Pub1. 200, 1051 p. Edm/nster, Frank Custer. 1954. American Hoover, Robert L., C. E. Till, and game birds. 490 p. Charles Scribner's Stanley Ogilvie. 1959. The antelope of Sons, New York. Colorado. 110 p. Colorado State Dep. England, Raymond E., and Antoon Devos. of Game and Fish. 1969. Influence of animals on pristine Irving, Washington. 1868. Astoria: Condltions on the Canadian grasslands, anecdotes of an enterprise beyond the J. Range Manage. 22(2):87-94. Rocky Mountains. Rev. ed., 649 p. G.P. Evans, Keith E. 1968. Characteristics Putnam and Son, New York. and habitat requirements of the greater Johnsgard, Paul A. 1973. Grouse and pralrlechlcken and sharp-tailed grouse-- quails of North America. 533 p. Univ. a review of the literature. U.S. Dep. Nebraska, Lincoln, Nebraska. Agric. Conserv. Res. Pap. 12, 32 p. Johnson, Morris D. 1964. Feathers from EvanS,°Keith E., and Donald R. Dietz. 1974. the prairie. Pittman-Robertson Proj. Nutritional energetics of sharp-tailed W -67-R-5, North Dakota Game and Fish grouseduring winter, J. Wildl. Manage. Dep., 240 p. 38(4):622-629. Kincer, J. B. 1941. Cllmate and weather Evans, Keith E., and Doug L. Gilbert. 1963. data for the United States. l_n_nCllmate Grouse of the grasslands. Colorado and man. Yearbook Agric. US. Dep. Outdoors 12(6):15-18. Agric., Washington, DC.

16 Korschgen, Leroy J. 1952. Food habits of cattle in the l_ssouri River breaks. the bobwhite quail in Missouri. Montana Wildl. Monogr. 20, 79 p. Missouri Conserv. Comm. P-R Rep., 59 p. Ksrbut, Curtis F. 1911. Soil reconnaisance Korschgen, Leroy J. 1957a. Food habits-- of the Ozark region of Missouri and coyotes, foxes_ house cats, bobcats-- Arkansas. Field Operations of the in Missouri. Missouri Conserv. Comm. Bureau of Soils, Rep. 13, p. 1727-1873. P-E Eep. 15, 6_ p. Mech, L. David. 1966. The wolves of Isle Korschgen, LeroyJ. 1957b. Food habits Royale. Fauna of the Natl. Parks of of the coyote in Missouri. J. Wild1. the U.S. Fauna Set. 7, 210 p. Manage. 21(4):424-435. Murle, Adolph. 1940. Ecology of the Korschgen, Leroy J. 1958. Food habltsof coyote in the Yellowstone. Fauna of the mourning dove in Missouri. J. the Nat. Parks of the U.S. Fauna Set. Wildl. Manage. 22(1):9-16. 4, 206 p.

Korschgen, Leroy J. 1960. Production of Murphy, Dean A. 1970. Deer range appraisal game bird foods in Missouri. J. Wildl. in the midwest. In White-tailed deer Manage. 24(4):395-401. in the midwest. US---DAFor. Serv. Res. Koster, Wiiliam Jacob. 1957. Guide to the Pap. NC-39, p. 2-10. North Cent. For. fishes of New Mexico. 116 p. Univ. New Mexico, Alberquerque, New Mexico. NixonExp., ChStan.rl,es St.M. Pa1ul,970. Minn.Deer populations Kucera, Clair L. 1960. Forest encroachment in the midwest. In White-tailed deer in native prairie. Iowa State J. Sci. in the midwest. US---DAFor. Serv. Res. 34(4):635-640. Pap. NC-39, p. 11-18. North Cent. For. Kucera, Clair L., and Melvin Koeliing. Exp. Stn., St. _Paul, Minn. 1964. The influence of fire on com- Parmalee, Paul W. 1955. Some factors position of central Missouri prairie, affecting nesting success of the bobwhite Am. Midl. Nat. 72(1):142-147. quail in east-central Texas. Am. Mid1. Kuchler, August Wilhelm. 1964. Potential Nat. 53(1):45-55. natural vegetation of the conterminous Pimlott, Douglas H., and Paul W. Joslin. United States. 116 p. plus colored map. 1968. The status and distribution of Am Geogr. Sot., New York. the red wolf. North Am. Wildl. Conf. Lars0n, Floyd. 1940. The role of the bison 33:373-389. in maintaining the short grass plains. Rand, A. L. 1945. The 1945 status of the Ecology 21(2):113-121. pronghorn antelope (Antolocapra Leopold, A. Starker, and E. Raymond Hall. umericawn) (ORD), in Canada. Natl. 1945. Some mammals of Ozark County, Mus. Can. Bull. 106, 34 p. MissOuri. J. _--,,,1. 26(2):142-145. Robinson, Thane S. 1957. Ecology of Lewis, Meriwether. 1961. The Lewis and bobwhite in south-central Kansas. Univ. clark expedition. Vol. I. 1814 ed., Kansas Mus. Nat. Hist. Misc. Publ. 15, 286 p. J. P. Lippincott Co. 84 p. L!nduska, Joseph P., ed. 1964. Waterfowl Rosene, Walter. 1969. The bobwhite quail, tomorrow. 770 p. U.S. Gov. Print. its life and management. 418 p. Rutgers Off., Washington, DC. Univ. Press, New Brunswick, New Jersey. Longhurst, W.M. 1960. Big-game and Sanderson, Glen C., and Helen C. Schultz, rodent relationships to forest and grass- ed. 1973. Wild turkey management. lands in North America. I_nnEcology and 355 p. Univ. Missouri Press, Columbia, management of wild grazing animals in Missouri. temPerature zones, p. 305-326. F. Schultz, Leonard P. 1936. Keys to the Bourliere, ed. Eighth Tech. Meet., fishes of Washington, Oregon, and IUCN, Warszawa. closely adjoining regions. Univ. Lokemoen, John T. 1973. Waterfowl Washington Pub1. Biol. 2(4):123 p. production on stock-waterlng pond s in Schwartz, Charles W., and Elizabeth R. the northern plains. J. Range Manage. Schwartz. 1959. The wild animals of 26(3):179-184. Missouri. 341 p. Univ. Missouri Press, Lord, Rexford D., Jr. 1963. The cotton- Columbia, Missouri. tail rabbit in Illinois. 94 p. South. Sclater, W. L. 1912. A history of the I11. Unlv. Press, Carbondale, 111. birds of Colorado. 576 p. Witherby, McCu110ugh, Dale Richard. 1969. The London. rule elk. 209p. Univ. Calif. Press, Seton, Ernest Thompson. 1929. Lives of Berkeley, Calif. game anlmals. Vol. III. 780 p. Mackie, RichardJ. 1970. Range ecology Doubleday, Doran, and Co., Xnc., New and relations of mule deer, elk, and York.

17 Severtnghaus, C. W., and g. L. Cheatum. J. Wtldl. Manage. 28(1):152-157. 1956. Life and times of the white- Weaver, John Ernest. 1954. North American talled deer. In The deer of 'North prairie. 348 p. Johnsen Publ. Co., America. p. 57-186. Walter P. Taylor, Lincoln, Nebraska. ed. The Stackpole Co., Harrlsburg, Weaver, John Ernest, and F. W. Albertson. Pennsylvania. 1956. Grasslands of the Great Plains. Severson, Kieth E., and F. Robert Gartner. 395 p. Johnsen Pub1. Co., LSncoln, 1972. Problems in commerical hunting Nebraska. SYstems: South Dakota and Texas Wiens, John A., and Melvin I. Dyer. 1975. compared. J. Range Manage. 25 (5): Rangeland avlfaunas: their composition, 342-345. energetics, and role in the ecosystem. Shelford, Victor E. 1963. The ecology of l__nManagement of forest and range North America. 610 p. Univ. Ill. habitats for nongame birds Syrup. Proc. Press, Urbana, Ill. Dixie R. Smith, Tech. Coord. USDA For. Stebbins, Robert Cyril. 1954. Amphibians Serv. Gen. Tech. Rep. WO-1, p. 146-182. and reptiles of western North America. Wing, Leonard W. 1951. Practice of wild- 528p. McGraw-Hill, New York. life conservation. 412 p. John Wiley Stempel, M. E., and Sam Rodgers, Jr. 1961. and Sons, Inc., New York. Hist0ry of prairie chickens in Iowa. Wright, Henry A. 1974a. Effect of fire on Iowa Acad. Sci. Proc. 68:314-322. southern mixed prairie grasses. S. Stoddart, Laurence A., and Arthur D. Smith. Range Manage. 27(6):417-419. 1955. Range Management. 2nd ed. 433 p. Wright, Henry A. 1974b. Range burning. McGraw-Hill Book Co., New York. J. Range Manage. 27 (1):5-11. Warbach, Oscar. 1958. Bird populations Yoakum, Sire. 1968. A review of the in relation to changes in land use. S. distribution and abundance of American Wildl. Manage. 22(1):23-28. pronghorn antelope, p. 4-14. I__nnThird Ward, A_ Lorin. 1964. Foods of the Antelope States Workshop Proc., Casper, mourning dove in eastern Colorado. Wyoming.

•_,U.S. GOVERNMENT PRINTING OFFICE: 1978-768-3437130-6

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Evans, Keith E., and George E. Probasco. 1977. Wildlife of the prairies and plains. USDA For. Serv. Gen. Tech. Rep. NC-29, i8 p., illus. North Cent. For. Exp. Sin., St. Paul, Minn.

The North American grasslands are the most extensive and varied of all plant communities in North America. The grasslands are divided into eight subtypes and the wildlife resources, past and present, are discussed for each subtype. Objectives and practices of wildlife habitat management are discussed.

OXFORD: 182.3:175.2:149:148.2. KEY WORDS: bison, big game, waterfowl, upland birds, ram,reals,range mangement.

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Evans, Keith E., and George E. Probasco. 1977. Wildlife of the prairies and plains. USDA For. Serv. Gen. Tech. Rep. NC-29, 18 p., illus. North Cent. For. Exp. Sin., St. Paul, Minn. . . The North American grasslands are the most extensive and varied of all plant communities in North America. The grasslands are divided into eight subtypes and the wildlife resources, past and present, are discussed for each subtype. Objectives and practices of _rlldlife habitat management are discussed.

OXFORD: 182.3:175.2:149:148.2. KEY WORDS: bison, big game, waterfowl, upland birds, mammals, range management.

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" Leave parks and forests clean.., or cleaner.