the bobwhite in illinois:

The Bobwhite in Illinois:

Its Past, Present and Future

State of IllinoisDepartment of Natural Resources

The northern bobwhite is one of America’s mostpopular and widely distributed game birds. Its rangeextends from theAtlantic ocean west to the foothills ofthe Rockies, and from the Gulf of Mexico north to cen-tral Wisconsin and Minnesota. Highest populationshave historically occurred within the 23 states bound-ed by a line from Delaware west to Nebraska thencesouth to Texas. There is currently widespread concernamong wildlife professionals regarding the future ofthis species, and for good reason. From 1966 to 1999,quail populations declined nearly 4% per year in theSoutheast (12) and 3% per year in theMidwest (19). Ifthis rate of decline continues over the next 30-40years, quail will be virtually extinct in both regions.

Despite its current problems, the bobwhiteremains an integral part of our natural Illinois her-

itage, and its welfare should concern us all. Much hasbeen written about the species,both popular and tech-nical. Unfortunately, popular accounts are often moreentertaining than informative, while technical articlesare generally written by scientists for other scientists.Consequently, the interested laymen often finds itdifficult to obtain in-depth or current biologicalinformation – we hope this booklet provides suchinformation. In it, we discuss important aspects ofquail biology, life history, and population dynamics inthe context of basic ecological principles and currentmanagement concepts. In that way, we hope to sharewith the reader – biologist and nonbiologist, hunterand nonhunter – a better appreciation and under-standing of this remarkable bird and the challenges itfaces.

Illinois Department of Natural Resources

Introduction

1

2

As the War of 1812 ended, settlers from the eastand south began to arrive in the territory that wassoon to become the state of Illinois. It is difficult toimagine what this area actually looked like back then.The major geologic and geographic features – therivers, streams, hills, valleys, and plains – were prettymuch the same as today, but what was in and on thesefeatures was vastly different. Except for a few smallsettlements and Indian villages, the landscape wasvirtually devoid of human influence. The southern1/4 of the State and the counties bordering theWabash, Mississippi, and lower Illinois rivers werecovered with what must have been some spectacularmature hardwood forest. In contrast, the flat, fertilelands of east-central Illinois – now the great “corndesert” – was a vast sea of tall-grass prairie with fin-gers of forests extending along the rivers and streams.At the forest-prairie interface, and to a lesser extentimbedded in each, were areas of open canopy forestwith an understory of prairie grasses and forbs. Earlywriters referred to these as barrens,now they are morecommonly called savannas.

Early settlers not only encountered a differentflora than exists today, they encountered a differentfauna as well. The territory of Illinois was still hometo elk, bison, cougars, black bear, timber wolves, andpassenger pigeons. Also present were most of thefamiliar species that we know today including deer,turkey, and of course bobwhite. No scientific surveyshad yet been conducted so we can only speculate onthe bobwhite’s distribution and abundance based onaccounts of early travelers and our current knowledgeof the species’ habitat needs and preferences. Whilepossibly locally common, quail would almost certain-ly not have been widely distributed or abundantthroughout the entire territory. They would have beenrare or absent in the wooded wetlands along theIllinois and Mississippi rivers and in the large, unbro-ken tracks of mature forest and tall-grass prairie thatdominated the landscape. Not surprisingly, they were

reportedly restricted to areas where grasslands andforest converged. According to Captain AdamBogardus, a noted 19th century hunter from Elkhart,bobwhite were most abundant in savannas, sandyareas, and other less fertile, higher, drier locations thatfeatured a mix of scattered hardwoods, shrubs, andshort, open stands of little bluestem and variousprairie forbs (6). To this day, bobwhite seek out plantcommunities with similar structure and diversitythough species composition differs widely.

Almost immediately, settlers began to affect thelandscape and the resident wildlife. As the humanpopulation of Illinois increased from about 12,000 in1810 to almost 1million by 1850, forests, and especial-ly prairies, were gradually replaced by farmland.Some indigenous species simply could not withstandthese changes or the increased hunting pressure. In arelatively short period of about 40-50 years, elk, bison,cougar, bear, and timber wolves were essentially elim-inated from the State. Other species, however, actual-ly benefitted from some of the early human-inducedlandscape changes, one in particular was the bob-white. The bobwhite does not thrive in primitive,undisturbed areas. Instead, quail are almost alwaysfound in some kind of disturbed or manipulated set-ting, which explains their traditionally close associa-tion with agriculture. In pre-settlement days, theyprobably responded positively, but temporarily, to for-est or prairie fires started accidentally by lightning orpurposely by native Americans. But it was the activi-ties of the early settlers that really began to make thelandscape more suitable for quail. Initially, settlersbegan clearing forests to establish farm fields becausethey believed that forest land was more fertile thangrassland, and their primitive plows could not turnthe tough prairie sod (26). The small clearings creat-ed adjacent to woodland edges and the waste grainand seeds from annual weeds and grasses associatedwith soil disturbance provided bobwhite with abun-dant and accessible food supplies. Pioneer farmingmethods quickly exhausted the fertility of forest soils

History and Distribution in Illinois

Pre-settlement - 1860

3

forcing farmers to abandon their fields after a fewyears and clear new ones. These abandoned fieldswere quickly occupied by plants favored by bobwhitefor nesting and brood rearing. By all indications,quailthrived in this primitive agricultural setting and prob-ably reached peak distribution and abundance inIllinois about the time of the Civil War.

1860 - 1960The next 100 years produced tremendous cultur-

al, technological, and economic changes includingagricultural changes that altered the very face of theland. In 1860, 34% of Illinois was farmland; by 1900,the figure had risen to 90%. Much of this increase wasdue to the advent of the moldboard plow whichallowed virtual complete conversion of the prairie tocropland. As in the forested southern portions of thestate, early agricultural advances into the prairie wereundoubtedly beneficial to bobwhite. Cropfields andassociated annual weeds and grasses provided abun-dant food supplies while the establishment of osageorange hedgerows and native woody plants made pos-sible by fire suppression provided much needed cover.As the land became settled and citizens had moreleisure time, the abundant bobwhite became thefavorite quarry of sport hunters with fine shotgunsand beautiful, well-trained setters and pointers. Bythe beginning of the 20th century, bobwhite werehighly valued as a gamebird and the State Legislatureenacted hunting seasons and bag limits to protect thisimportant natural resource.

During the first 1/3 of the 20th century, agricul-ture continued to evolve from settlement to commer-cial farming while still remaining rather diverse andprimitive – but not for long. Rapid advances in mech-anization in the 1930s allowed farming to becomemore efficient,while development of synthetic fertiliz-ers and chemical pesticides around the time of WorldWar II led to the intensive,monocultural rowcrop agri-culture that currently dominates the Illinois land-scape.

We cannot say with certainty exactly how quailfared throughout this period. Two prominent Illinoisornithologists, comparing their results with earlierscientific surveys, concluded that bobwhite popula-tions in 1956-58 were roughly similar to what had

been observed by their earlier colleagues in 1906-09(34). What accounts for this apparent stabilitythrough the first half of the 20th century? Thoughagricultural technology was advancing, farms werestill small and fields, often bordered by shrubbyfencerows, were usually no larger than 20 to 40 acres.And while tractors had largely replaced horses as apower source, the production of livestock created acontinuing need for permanent pastures and hay-fields. Furthermore, crop rotation including smallgrains, legumes, and periodic fallowing was still wide-ly practiced.

1960 - PresentQuail were probably scarce, if not absent, over

much of the “Grand Prairie” region of Illinois by 1900(32) owing to the intensity of agriculture on this flat,incredibly fertile land. In much of westcentral andsouthern Illinois, however, quail continued to prosperas late as the 1960s simply as a byproduct of normalfarming operations. Many farms in these slightly hilli-er, less productive portions of the state still featured adiverse mix of rowcrops, small grains, pasture, hay-fields, fallow land, and woodlots all occurring in rela-tively small patches and tied together by a network ofbrushy fencerows and woody hedgerows. The prac-tice of regular fallowing of cropland and the mainte-nance of woody hedgerows was especially beneficial toquail. Fallowed fields provided the early successionalweeds and grasses critical for successful reproductionand brood rearing while the brushy hedgerows pro-vided protective cover and travel lanes that allowedthe birds to safely access food supplies in the cultivat-ed fields.

Agriculture in the Illinois quail range began tointensify and simplify in the 1960s and especially the1970s as we entered the era of “fencerow to fencerow”farming. Cropping patterns became less diversified asfarmers concentrated on 1 or 2 cash grains. Smallwoodlots and idle corners were cleared and planted torow crops. Fencerows and hedgerows were bulldozedout to enlarge fields in order to accommodate modernmachinery and increase tillable acreage. Heavy use offertilizers replaced crop rotation and fallowing, com-bines replaced old-fashion corn pickers, and fallplowing became an almost universal practice. Pasture


4

and hayfield acreage was greatly reduced as the cattleindustry moved from grassland to feedlot, and muchof the grassland that did remain was converted frommixed stands to monocultures, especially fescue. Ontop of all this, there was a large increase in the use ofherbicides and insecticides intended to eliminate thetwo primary sources of native bobwhite foods – weedsand insects. All of these things made farmland thatonce supported an abundance of quail, now basicallyinhospitable. The response of quail was dramatic, rel-atively sudden, and predictable – statewide abun-dance declined at least 60% from themid-1960s to themid-1980s (Fig. 1).

Emergence at about this time of the ConservationReserve Program (CRP) carried with it widespreadanticipation of positive benefits to wildlife, includingbobwhite. This program was a provision of the 1985Food Securities Act (Farm Bill) that encouragedlandowners to retire highly erodible cropland for aminimum of 10 years and replace it with semi-perma-nent vegetation, usually grass. From the late 1980s tothe present, approximately 2 to 3% of the Illinois quailrange has been enrolled but despite initial optimism,CRP apparently has not benefitted regional orstatewide quail populations as much as was anticipat-ed (95). We’ll discuss possible reasons and remediesfor this in a later section (Bobwhite and Agriculture).

Current DistributionThe bobwhite’s geographic range includes all of

Illinois, although the northern 1/4 of the State, where

annual snowfall averages 25-40 inches, is considered“fringe” or marginal because of unfavorable winterclimate (102). While quail probably occur in everycounty in Illinois, they are most abundant and welldistributed in the southern and westcentral portionsof the State (Fig. 2). They are scarce north

of this region because winters are too harsh, and eastbecause rowcrop agriculture is too intense.Distribution within their primary range is stronglyinfluenced by human land use which in turn is largelygoverned by soil productivity and terrain. Quail habi-tat in Illinois is mainly limited to moderately rollingterrain with low to medium soil productivity, i.e.,areas with relatively diverse, less-intensive agriculture.Flat landscapes with highly productive soils are usual-ly devoid of quail and quail habitat because they areintensively rowcropped,whereas the steepest terrain isusually too heavily forested.

Fig. 1. Bobwhite population change from 1960 to pres-ent, short-term fluctuations are mainly due to weather;long term trends reflect habitat loss.

Fig. 2. Current distribution of potential bobwhite habi-tat (green) as determined from satellite imagery.

5

TaxonomyBecause common names can vary locally, animals

are given scientific or Latin names for universal iden-tification. The bobwhite’s scientific name is Colinusvirginianus. Biologists recognize several distinct sub-species or races within the United States; C. v. mexi-canus, or the interior bobwhite, occurs in Illinois andneighboring states. The namemexicanus has nothingto do with“Mexican quail”which some peoplemistak-enly believe have contaminated our present stock.While true that releases of a small southern sub-species, C. v. texanus were made in the 1920s and1930s, they failed because the birds were whollyunsuited to the midwestern climate. Despite popularopinion to the contrary, we can assure readers thatthe present-day Illinois quail is the same old “bob-white” that has always been here.

Taxonomically, bobwhites belong to the orderGalliformes or “fowl-like” birds. They are related tochickens, pheasants, grouse, turkeys, and the “westernquails” (e.g., scaled, Gambels, California, mountain).Most of the bobwhite’s closest relatives (other mem-bers of the genus Colinus) are relatively unfamiliarforms living in Mexico and central America. In fact, itis from this area that the species is thought to haveevolved (54). Fossil records tell us that quail-like birdsexisted at least a million years ago, but their appear-ance in what is now Illinois probably did not occuruntil sometime after retreat of the last glacier some10,000 years ago.

Size and GrowthAs a general rule, animals that occupy extensive

geographic ranges tend to be larger in the northernthan southern parts of their range, and quail are noexception. Bobwhite in Georgia and Florida averageabout 5.8 oz compared to nearly 7 oz inWisconsin. Asmight be expected, Illinois birds are intermediate insize. Fall and winter weights of 850 birds from south-ern Illinois averaged just under 6 oz (178.2 g) (97).Weights of males and females are virtually identical

this time of year but deviate during the breeding sea-son as will be described in a later section(Bioenergetics). Bobwhite chicks weigh only about6 g at hatching but gain rapidly and attain 80% ofadult weight by 11 weeks of age and 97% by 21 weeks.

One of the most persistent bits of quail folklore isthat present-day birds are smaller than they used tobe. We are at a loss to explain the persistence of thismisconception. The above mentioned southernIllinois weight study, which included samples collect-ed from the late 1940s to 1970, showed no differencein average weight over that time. Likewise, present-day, fully-grown wild quail in southern Illinois stillaverage a little over 6 oz, just as they always have.

Plumage and MoltsThe bobwhite is a small, plump, brownish bird

with stubby wings and short, dark legs. Plumage onthe back, wings, and tail is various shades of chestnutbrown while the underside consists of small dark barson a whitish background. The head has a dark cap, adark collar at the base of the neck, and a dark widestrip running from cheek to eye. Males sometimesshow just a hint of a crest or topknot. Males also havea distinctive white throat patch and white stripe run-ning from the neck over the eye to the beak. Infemales, these areas are buff colored. There also areslight differences in the wing markings of males andfemales (115). At a distance, the bobwhite appearsrather drab, but in hand their subtle shadings andmarkings are quite attractive.

Abnormal color variation is rare, but 3 distinctphases are sometimes seen in the wild. There is par-tial albinism in which some or most of the feathers arewhite (true albinism with pink eyes and white bill isextremely rare). A second variation is the “red” phasein which some or all of the feathers are reddish orauburn in color. Finally, there is a “blond” phase inwhich the birds have a light camel coloration over por-tions or all of their body.

The bobwhite goes through a series of molts pro-


Description

6

gressing from natal down to juvenile to adult plumage.Some flight is possible at 2 weeks of age. By 7 weeks,the birds are more than half grown but the sexes arestill indistinguishable. At 14-15 weeks, the young are

essentially fully grown and can be distinguished fromadults only by certain wing feathers. During their firstautumn of life, juveniles molt 8 of the 10 primary wingfeathers, retaining the outer 2 until the following year.Juveniles also retain their primary coverts which arebuff tipped and ragged in appearance as contrastedwith adult’s which are more rounded and have a uni-form charcoal color. Biologists use these criteria todistinguish juveniles from adults. Age of juveniles tothe day is then estimated from progression of molt ofthe primaries (Appendix A).

VocalizationOne of the most familiar and pleasing of all birds

sounds is the call for which the bobwhite is named.The clear, ringing bob-white or ah-bob-white is given

by males, usually unmated ones, throughout thespring and early summer. Another series of calls arevariously described as group movement, scatter, orcovey calls. These include a soft hoy, a louder hoy-poo,

and a clear loud musical hoi-lee or hoyee (110). Thelatter is heard just after daybreak on most fall andwinter mornings. It is given by one or more coveymembers prior to leaving their night roost. Variationsof the hoy or hoy-poo call are given by females duringthe breeding season and by coveymates attempting toregroup after being flushed. Less familiar are thesquee and“caterwaul” sound associated with agonisticbehavior before and during the breeding season.Finally, there are a number of very soft, gentle vocal-izations uttered among coveymates as they forage forfood. To enjoy these pleasant sounds, one must bevery close to an undisturbed covey.

Male Bobwhite Female Bobwhite

7

Most people remember from their high school orcollege biology that evolution involves such things as“genetic variation,” “struggle for existence,” and “sur-vival of the fittest.” Many assume the latter refers sole-ly to contests of strength and speed between membersof different species. This is not altogether true. Whilesuch competition does play an important role in evo-lution, the primary arena for natural selection isintraspecific relationships. The term fitness refers toan animal’s relative ability to contribute to the futuregenetic composition (gene pool) of its species. Thetype of individual that leaves the greatest number ofreproducing offspring will eventually predominate ifsuch success is due to inheritable characteristics.Conversely, those traits that render the bearer less

likely to do this will become rare or disappear entirely.It should be apparent then that the real “players” arenot individual animals, but their genes. In fact, some-one once said that a chicken is merely an egg’s way ofproducing another egg. In any event, we can safelyassume that natural selection produces types of indi-viduals that are most efficient at passing on theirgenes. Recognizing that this efficiency is manifestednot only in physical attributes, but in behavioral traitsand life history patterns as well, we might well ask –what is the best way for individuals to enhance theperpetuation of their genes? Should they search forfood in groups and share it, or forage alone? Shouldthey produce many fragile young, or a few hardy ones?Should they lavish care on these offspring at their ownexpense, or be mainly concerned with their own wel-fare so as to survive and breed again (49)? Obviously,animals do not consciously scheme about such things;in fact, they have little choice in the matter as most ofthese characteristics are inherited and thus “hard-wired.” Further,what is best for one species might notwork for another. That is why there are so many dif-ferent life styles in nature. Evolution, operating with-in an ecological framework,has produced a wide spec-trum of living patterns and adaptations, the effect ofwhich is to permit the broadest and most completepossible utilization of available resources. Each

organism occupies its own functional place or ecolog-ical niche in the overall system. This chapter, and por-tions of those following, examine the bobwhite’sniche, i.e., where it lives and how it makes a living. Todo this, we will describe important events that makeup the bobwhite’s annual cycle (Fig. 3). starting withearly spring.

Spring and SummerBobwhites spend the fall and winter months in

tight social groups called coveys. As days lengthenand warm in late March, certain behavioral changesbegin to occur among coveymates. Night roostsbecome less compact and individuals and smallgroups spend more time away from the main bodyduring the day. Males are noticeably more antagonis-tic toward one another, and their familiar “bobwhite”call is heard for the first time since the previous sum-mer. These are but the outwardmanifestations of pro-found physiological and hormonal changes takingplace in response to the increasing daylight hours(photoperiod). The male’s internal testes increase insize and spermatogenesis begins. The female’s onefunctional ovary, which has been quiescent through-out the winter, also begins to enlarge as does heroviduct in preparation for the development and pas-sage of eggs. Although these and related activities areprimarily stimulated by changing daylength, tempera-ture also plays a role. The timing of covey breakup and“bobwhite” calling seem at least partially dependenton late March and early April temperatures.


Life History

Fig. 3. Important events in the Bobwhite’s annual cycle

8

Pairing sometimes, but not always, takes placebetween coveymates, which brings up the question ofinbreeding. Those who contend that hunting is neces-sary to mix the birds and prevent inbreeding overlookthe obvious fact that the species did quite well withoutour help for thousands of years. Actually, there is suf-ficient natural reshuffling of groups from late summerthrough early spring to ensure a healthy mix of genet-ic material. Bobwhite courtship activities and displaysare perhaps less familiar to most people than those ofcertain other birds. When displaying to a hen, thecock puffs out his feathers and bends his fully extend-ed wings so that the tips just touch the ground. Hishead is lowered and turned to show off the whitethroat and cheek markings. All the while, the henkeeps her tailfeathers fully spread and when ready tocopulate, squats and utters a barely audible call (109).Several weeks may elapse between pairing and nestbuilding during which time several more bouts of cop-ulation may take place. The first nests are usuallystarted in late April or early May and take 1-2 days tocomplete. Both sexes participate,butmales domost ofthe work. The typical nest is a rounded, domed struc-ture with an oval entrance on one side. It is built ofgrass stems from the previous year’s growth and sitsin a shallow, saucer-like depression in the groundscratched out by the birds.

The female begins laying within a day or so ofnest completion and proceeds at the rate of approxi-mately 1 egg per day until the entire clutch is deposit-ed. During this time, she remains at the nest only long

enough to deposit an egg. The eggs themselves arewhite, pointed at one end, and about 1” x 5/6” in size.Most clutches contain from 12 to 16 eggs; 13.7 is theaverage in Illinois (64). Incubation takes about 23days to complete and usually begins within a day or soafter the last egg is laid. About 1/4 of the nests areincubated by males, the remainder by females. Thesexes apparently do not alternate these dutiesalthough the incubating bird often leaves the nest forseveral hours each afternoon to feed and relax with itsmate. For the first week or so, incubating birds areeasily disturbed, but as hatching time nears, they arevery reluctant to flush and will tolerate closeapproach.About 48 hours prior to hatching, the near-ly-developed chick begins to“pip” the egg with its“eggtooth.” Once out of the egg, the precocial chicks areusually dry and able to move about within a fewhours. If it is late in the day or raining, they may bebrooded in the nest overnight. Otherwise, the incu-bating bird is joined by its mate and the young arequickly led away from the nest. On average, >9 ofevery 10 eggs hatch. Of those that don’t, about 40% areinfertile and 60% contain dead embryos.

Bobwhite nests,which are built on the ground, arevulnerable to a wide variety of destructive agents. Infact, only about 1/3 of them hatch successfully. Duringa 15-year study in southern Illinois (64), about 11% ofthe 860 nests examined were abandoned before hatch-ing, and another 12% were destroyed by farmingactivities – primarily mowing. About 2% failedbecause of excessive heat and drought while an evensmaller percentage were flooded out. Miscellaneouscauses and research activity led to the failure of anadditional 4%. The primary cause of nest losses wasother animals; predators destroyed 37% of the nests,accounting for over half of all nest failures. Commonpredators on this particular rural study area were thehouse cat, striped skunk, and various snakes such asthe prairie kingsnake, black rat snake, and racer.Numerous other animals including dogs, foxes, coy-otes, weasels, raccoons, opossums, and crows alsodestroy nests on occasion andmay be a problem local-ly. Perhaps surprisingly, the attending adult is rarelykilled during nest attacks (about 1 time in 10), andusually only late in incubation when they are reluctantto flush from the nest.

Bobwhite nests are usually doomed-shaped structures, located atground level, and constructed of dead grass stems.

9

Because 60-85% of the autumn population repre-sents birds hatched the summer just ended, reproduc-tive success plays a major role in determining fallabundance. The number of young recruited into thefall population in a given year depends on several fac-tors including the number of breeding hens, the num-ber of chicks hatched per hen, and the survival ofthese chicks from hatch to autumn. The number ofchicks per hen is determined by: a) the number ofnesting attempts made, b) the success rate of thesenests, c) the number of eggs per nest, and d) thehatching rate of these eggs. Research has shown thatyear-to-year variation in clutch size and hatchabilityof eggs is of little importance. What is important is thetotal nesting effort (number built per hen) and theirrate of success (98).

The bobwhite’s nesting season is one of thelongest of any temperate avian species. In Illinois, egglaying has been recorded as early as mid-April andhatching as late as October. Most nests though (80%)are started sometime between the 2nd week in Mayand the 3rd week in July with peak hatching fromabout mid-June through mid-August (Fig. 4).

Scientists have long known that bobwhite henswill renest 1 or more times following initial nest fail-ure and that males sometimes incubate nests.

However, based on indirect, circumstantial evidence,most biologists assumed that quail weremonogamousand rarely if ever produced more than one brood perseason. More recent telemetry studies have shown,however, that bobwhite mating systems are much

more complicated and flexible than originallythought, combining elements of monogamy,polygamy,multiple clutches, renesting, and incubationby both sexes (14, 111). For example, reproductiveefforts can include monogamous pairing and produc-tion of single broods, production of consecutiveclutches with male incubation of the first and femaleincubation of the second, or renesting after success orfailure of the first nest with 1 or more mates involved.Although early biologists may have underestimatedthe extent of multiple broods, they did not necessarilyunderestimate actual reproductive output which isnormally measured by comparing spring and subse-quent fall population levels.

Another factor affecting overall productivity issurvival of chicks from hatch to autumn.Unfortunately, this is one of the most poorly docu-mented aspects of bobwhite life history. One reason isthe difficulty in capturing and studying very youngchicks in the wild to determine rates and causes ofdeath. Based on limited telemetry studies and obser-vations of brood shrinkage over time, it is estimatedthat from 50 to 70% of all chicks hatched usually sur-vive to autumn. However, a more recent study inOklahoma recorded <40% survival from hatch to 39days of age (22). The heavy use of insecticides in agri-cultural habitats has raised concern about their effectson quail chicks, both directly and indirectly.Researchers in the southeastern United States con-cluded that foliar insecticides did not directly poisonchicks (76). However, Illinois biologists suspect thatinsecticides may increase pheasant brood mortalityby reducing their invertebrate food supplies (118).Whether or not the same holds true for bobwhite isnot known, but the possibility certainly exists.A current belief popular among hunters in Illinoisand elsewhere is that wild turkeys destroy large num-bers of eggs and chicks and are therefore responsiblefor the widespread decline in quail abundance.Turkeys undoubtedly eat quail eggs and chicks occa-sionally, but food habits studies in the Southeast andMidwest clearly show that this is a rare event.Furthermore, the decline in quail abundance inIllinois and elsewhere started long before turkeysbecame abundant, andit is evident in areas where turkeys are rare or non-existent.


Fig. 4. Egg-laying and hatching dates for the bob-white in southern Illinois. (Roseberry and Klimstra,1974).

10

Therefore, we can state with absolute certainty thatturkeys are not the cause of the current decline inIllinois quail populations.

Some people believe that wet weather during thenesting season is detrimental to quail productionbecause it drowns chicks and floods eggs. In realitythough, nesting quail generally avoid poorly drainedareas or sites prone to flooding, and losses from heavyrainfall are rare. Actually, above-average precipitationduring spring and summer is more often associatedwith good reproduction. On the other hand, severedrought is potentially harmful because it can result inincreased nest abandonment, reduced hatchability ofeggs, and possibly reduced insect food supplies forchicks. Extremely high temperatures in late summeralso can result in early termination of nesting efforts(64), thereby reducing overall productivity.

Conditions prior to the breeding season also canaffect subsequent reproductive efforts. Research insouthern Illinois has shown that severe winters, espe-cially prolonged snow cover, tend to be followed bysummers of below-average productivity (98). Themost likely explanation is that hens with inadequatewinter food supplies enter the breeding season in lessthan prime physical condition. This theory is sup-ported by the fact that snow cover occurring in earlywinter seems less detrimental than when it occurs inlate winter.

Fall andWinterAs the breeding season winds down in early

autumn, individuals begin to organize into cohesivesocial units (coveys). This behavior may help avoidand escape predators and conserve body heat duringcold nights. Coveys are not strictly family groups andmay contain remnants of several summer broods aswell as their surviving parents and an occasionalunmated adult. Not much is known about covey socialstructure, but there appears to be remarkably littleantagonism amongmembers. Although fall covey sizeis commonly overestimated from casual observation,such groups generally contain from 8-20 birds andaverage slightly over 13 (98). Occasionally, as many as25-30 quail may flush from one location, but this usu-

ally represents 2 coveys temporarily feeding in closeproximity, andmore often than not, these large groupssplit in flight and go their separate ways.

Bobwhite coveys occupy more or less discretehome ranges as opposed to wandering randomlyabout the countryside. Because quail are often foundin the same place year after year, some people assumethat it is the same covey living there. Actually though,only about 15-30% of the birds alive in one autumnsurvive to the next, and there is no guarantee they willoccupy the same covey range although this sometimeshappens. Certain home ranges contain quail year afteryear not because they are the same birds, but becausethe area itself consistently attracts quail.

As autumn crop harvesting and frosts reduceavailable cover, there is considerable movement ofquail as they settle into their more permanent winterhabitat. Individual movements of several miles ormore have occasionally been recorded during this“fallshuffle.” Once established though, coveys are reluc-tant to abandon their territories unless forced out byhabitat disturbance or extreme harassment fromhunting or field trials. Prolonged snow coverage alsomay force some poorly situated coveys to abandontheir ranges in search of better conditions (86).

In typical Illinois quail country, covey rangesaverage about 35-40 acres, but may be larger or small-er depending on habitat conditions, population densi-ty, and weather. The birds normally do not traversetheir entire range each day; instead, they will occupy aportion for several days, then shift activities to anoth-er part for a time. The ranges of neighboring groups

Bobwhites are almost always found near an “edge.”

11

may overlap to some extent, especially in feedingareas, but coveys tend to“keep their distance”when inheavy cover or roosting. In fact, their habit of callingjust before leaving the roost in early morning probablyserves to keep neighboring coveys apprised of eachother’s location and thus properly spaced (28). Coveyswill, however, readily accept stragglers into the group,and it is not uncommon as winter progresses for sur-vivors of 2 ormore shrinking coveys to recombine into1 larger group.

Bobwhite are among the most sedentary of allavian species. Under normal circumstances, dailymovements total no more than 1/4 mile – often lessduring inclement weather. Furthermore, virtually allmovement is via the ground. Except for possibly ashort flight to their night roost, quail seldom fly unlessdisturbed. When flushed though by man or predator,they explode into the air in a burst of noise andblurred shapes. This creates a rather startling effectand makes it difficult to focus attention on a singlebird – an obvious adaptation to escape predators.Upon landing, scattered individuals begin callingalmost immediately in an effort to regroup. Researchin Texas on the flight characteristics of bobwhitesfound that when flushed, coveys flew an average ofabout 50 yards at around 20 mph (57), althoughspeeds up to 38 mph have been recorded (109).

At night, coveymembers form a circular, compactroost for warmth and protection. The birds sit withtheir tails in and their heads pointing out, somewhatlike the spokes of a wheel. The small piles of brownand white droppings that mark these sites are a suresign of quail in an area. Bobwhites generally feedtwice daily in fall and early winter – once shortly afterleaving the roost in early morning, and again in mid-to late afternoon. Midday is often spent “loafing” inmoderately heavy cover. As winter progresses andfood supplies diminish, the birds spend an increasingproportion of their time foraging for food. Duringstormy weather, they may remain on their roost laterin the morning, feed only once at midday, and returnto the roost earlier in the afternoon. Formation oftemporary day roosts is also a common response tocold, windy weather.

Fall and winter are normally critical times forwild bobwhites. From 1/2 to 3/4 of all birds entering

the fall usually perish before spring. In addition topredators and the gun, quail sometimes succumb toaccidents, disease, starvation, and exposure, but theselatter mortality agents are of only minor importance.

For example, quail are sometimes killed by flying intoobstacles, and bird dogs occasionally capture and killotherwise healthy birds. There was even a recordedinstance of an entire covey drowning while attemptingto fly across a large reservoir (72). By and largethough, accidents are a minor mortality factor in thewild. The same can be said for disease. Penned bob-whites are notoriously susceptible to certain avian dis-eases such as avian pox, quail bronchitis, ulcerativeenteritis, and histomoniasis (blackhead). Cestodes(tapeworms) and nematodes (roundworms) aresometimes a problem too. Among wild birds though,parasites and diseases are generally of little conse-quence (21). There have been recorded instances ofextensive mortality following heavy applications ofthe pesticides dieldrin and heptachlor, especially in


Sure signs that quail are in the vicinity.

12

the southeastern United States (101, 102)Consumption of DDT also may have harmful meta-bolic effects but thankfully, these environmental con-taminants are much less prevalent now than 30 or 40years ago. Newer insecticides such as parathion andthe herbicide paraquat may cause indirect problems,however. Studies in the Southeast (11) have suggestedthat the former may increase bobwhite susceptibilityto avian predation while other research detected apossible negative effect of paraquat on bobwhitereproduction (5).

Predation is undoubtedly the major cause of bob-white mortality on a year-round basis althoughhunters may actually take more birds from fall tospring. Mammals most capable of capturing maturebirds during fall and winter include coyotes, red andgray foxes, and house cats. Other mammals such asbobcats, raccoons, opossums, skunks, mink, andweasels probably capture healthy bobwhites onlyoccasionally. Cooper’s and sharp-shinned hawks, aregenerally considered to be the most effective avianquail predators. Northern harriers (“marsh hawks”),great-horned owls, and barred owls also take quail onoccasion, but the broad-winged soaring hawks (red-shouldered, red-tailed, broad-winged) rarely do.Despite the fact that quail and their eggs are taken bya wide variety of mammals, birds, and reptiles,research has clearly shown that bobwhites are not astaple, or even contribute more than a minor part, tothe diet of any single predator. In Illinois, most quailpredators feed primarily on rabbits, small rodents,and to some extent other birds, and their actions maynot always be entirely harmful. A study in Texas foundthat removal of coyotes actually resulted in an increasein more effective quail nest predators such as redfoxes, raccoons, and skunks (47). It should be notedtoo that the proximate (immediate) cause of death isnot always the ultimate (fundamental) cause. If bull-dozing out a hedgerow displaces a secure residentcovey into marginal range where individuals are easytargets for predators, what factor is really to blame fortheir loss? Similarly, prolonged snow cover mayincrease quail vulnerability to predation by weakeningthem physically, reducing effectiveness of escapecover, and forcing them to feed in exposed areas forlonger periods of time. The point is that survival is

often determined by the interaction of several factors,and the final agent of death may be only incidental tothe primary cause.

Another example of the above is starvation andexposure which are difficult to separate as mortalityagents. Bobwhites are much more vulnerable to expo-sure when they are undernourished; likewise, they aremore susceptible to starvation at very low tempera-tures. Severe winters, especially the combination of

deep snow cover, low temperatures, and strong windscan kill bobwhites directly. Fortunately, such blizzardconditions rarely occur in the better Illinois quailrange (which partly explains why it is good range).Unquestionably, the environmental condition mostdetrimental to Illinois bobwhites is prolonged snowcover. Deep snow, especially when crusted, limitsaccess to food and reduces the effectiveness of certaintypes of cover. Ice cover also can be harmful, but suchconditions seldom last for more than a few days. Howbobwhites respond physically and physiologically toharsh environmental conditions is discussed in moredetail in the following section.

Winters with prolonged snow coverage are usually fol-lowed by 1 or more years of below-average quailpopulation

13

All living things require some form of energy, andthe ultimate source of all energy on earth is the sun.Through the process of photosynthesis, green plantsconvert this light energy into chemical energy or food.Herbivorous animals exist by feeding on plants and inturn provide energy for the carnivores that feed onthem. This exchange of energy between and amonganimals and their environment is called bioenergetics,and it serves as a useful framework not only forunderstanding ecosystems but for practical wildlifemanagement as well.

Bioenergetic relationships are governed by thelaws of physics. For example, the 2nd law of thermo-dynamics states that conversion of energy from oneform to another (always from more concentrated tomore diffuse) is never 100% efficient. At each transfer,some energy is lost as unavailable heat. Plants are ableto convert only about 1% of all incoming solar radia-tion into living tissue or biomass. Primary consumers(herbivores) are about 10% efficient (e.g., produce 100kcal of tissue for every 1,000 kcal consumed),whereassecondary consumers (carnivores) are perhaps 20%efficient. This stepdown effect has important implica-tions as it determines the length of food chains andthe relative abundance of animals at each trophiclevel. Consequently, the total biomass of herbivores inan area will always exceed that of carnivores becausethe former have more potential energy available tothem.

Animals are unable to utilize all of the gross ener-gy they ingest – some is lost through bodily wastesand some is needed for the work of digestion itself.That which remains is called net energy and is usedfor growth, maintenance (routine bodily functionsand upkeep), thermoregulation (maintenance of sta-ble body temperature), daily activities (food gather-ing, predator avoidance, etc.), reproduction, and stor-age of reserves (deposition of fat). When energyintake is insufficient to accommodate all of theseneeds, there is a definite priority of use.Thermoregulation and other life-sustaining bodily

processes are maintained at all cost, growth andreproduction are a secondary priority, and fat isdeposited only after all other needs are met.

Depending on their diet and needs, animals haveevolved a variety of feeding patterns and digestive sys-tems for acquiring and utilizing food. Bobwhite areprimarily seed eaters although they do consume somefruits, buds, and leafy material as well as animal mat-ter. Like other gallinaceous birds, quail have a sac-likeenlargement of the esophagus called the diverticulumor crop. This structure, which may have 4 times thecapacity of the stomach, allows the birds to temporar-ily store rapidly-gathered seeds thereby reducing feed-ing time and consequent exposure to predators andthe elements. After ingestion, food material passesfrom the crop into the anterior portion of the stomach(proventriculus) where digestion begins. It thenmoves into the muscular gizzard where hard seeds arepulverized with the aid of small particles of sand orgravel called grit. Some relatives of the bobwhite(grouse for example) have two pouches or caeca at thejunction of the small and large intestine which func-tion in the microbial breakdown of low quality, high-bulk foods. Game birds like turkeys, pheasants, andquail, that consume highly nutritious foods, have littleneed for, and therefore, lack well-developed caeca.

Bobwhites eat a variety of foods. A large numberof crops collected from southern Illinois huntersrevealed no fewer than 187 different items (67). Corn,soybeans, wheat, lespedezas, acorns, common rag-weed, sassafras, tick clover, and slugs were the mostcommon items found. Other foods taken with someregularity include jewelweed, beggar-ticks, wild bean,foxtail, crabgrass, lance-leaved ragweed, smoothsumac, ash, rush-foil, partridge pea, vetch, horse net-tle, grasshoppers, leafhoppers, ground beetles, andvarious insect larvae.

All foods, regardless of type, consist of the samebasic components: carbohydrates, proteins, lipids(fats), vitamins, and various inorganic minerals. Mostdietary energy is derived from carbohydrates and fats;


Bioenergetics

14

proteins are used mainly for tissue building andrepair, while vitamins and minerals are essential for avariety of life processes. It is sometimes said that if awild animal is able to consume sufficient energy, thenits other nutritional needs will also be met.Nevertheless, seasonal shifts in diet may reflect, inaddition to availability, nutritional demands otherthan energy. For example, bobwhites need only about12% crude protein in their diet for normal wintermaintenance, but laying hens require up to 23% formaximum egg production, and growing chicks needat least 28%. Proteins are available from both plantand animal sources, 2 of the best in fall and winter aresoybeans and Korean lespedeza. Insects and otheranimal matter provide even more protein and fortu-nately, their availability coincides with increased bob-white demands during spring and summer. Animalmatter may constitute only about 10% of the adult’swinter diet compared to 25-30% in summer. Growingchicks, on the other hand, consume up to 80% insectsduring their first 2 weeks of life with small grass seedsmaking up much of the remainder of their diet.

The most vital nutrient for all life, including bob-whites, is water. Quail obtain water in 3 ways: as abyproduct of metabolizing foods (metabolic water),frommoisture in foods (preformed water), and as sur-face (free) water. Most researchers agree that the 1st 2sources are sufficient to meet the bobwhite’s needs.

Thermoregulation is a fundamental process of lifethat involves a constant balancing of heat productionand loss through both physiological (chemical) andphysical (mechanical) means. Bobwhites and otherwarm-blooded animals may be thought of as consist-ing of an inner core and an outer shell. The core,which contains the central nervous system, visceralorgans, and much of the skeletal muscles, must bekept at a constant temperature (a remarkably high106.7o F for bobwhite) (38). The outer shell of feath-ers, scales, skin, and some subcutaneous fat and mus-cle acts as insulation and is often considerably coolernear its surface than is the core.

Animals exchange heat with their environment ina number of ways. Heat is lost through radiation(directly into the air), conduction (contact with coolerobjects), convection (wind effect), and evaporation.Heat is gained through direct solar radiation, radia-

tion from surrounding objects, and from the conver-sion of chemical food energy to heat. The rate of heatexchange depends primarily on the differencebetween ambient (air) temperature and the animal’ssurface temperature, but is also related to body size.Large animals have proportionately less surface areain relation to volume, and therefore lose relatively lessheat by radiation and convection. Bobwhites partiallycompensate for their small size by spending the nightin tightly formed, circular roosts – a behavior thatreduces the total amount of exposed body surface andthus heat loss for all members of the covey.

Of course an animal is not always trying to retainheat, sometimes the goal is to dissipate as much heatas possible. Both situations require expenditures ofenergy. For all warm-blooded animals, however, thereexists a range of ambient temperatures that do notinfluencemetabolism. Maintenance of a constant coretemperature within this thermal neutral zone requireslittle or no energy and is accomplished by minoradjustments in the insulation properties of the outershell (e.g., fluffing feathers, standing or squatting, etc).One study set this zone for bobwhites at 86-104o F(70). If true, this would be quite high in relation toreported ranges for other gallinaceous birds and mayreflect the species’ subtropical evolutionary heritage.

As air temperatures fall below the neutral zone,thermoregulation is achieved mainly throughincreased metabolic activity and changes in behaviordesigned to minimize heat loss (e.g., covey roosting).The energy cost of thermoregulation can be appreciat-ed by comparing it to the basal or standard metabolicrate, which is the amount of energy required to exist(not grow or reproduce) under thermal neutral condi-tions. This rate can be expressed in a number of ways,perhaps the most common being kcal of energy need-ed per kg of body weight per day. One study deter-mined that bobwhites weighing about 190 g needed 24kcal per day just to survive in cages at 86o F under a10-hour photoperiod (15).About twice as much energy was required at 32o F.The important question to wildlife managers is howdifficult is it for bobwhites to acquire this necessaryenergy day after day. Several things must be consid-ered. First of all, life in the wild takes considerablymore energy than does life in a cage – perhaps 50%

15

more. Secondly, Illinois bobwhite often encountertemperatures below freezing which impose evengreater energy demands. For example, a temperaturedrop from 50 to 320 F increases energy demands by28%. Under normal late-winter conditions in theMidwest, bobwhites may need to forage almost con-stantly during daylight hours in order to consume suf-ficient energy (83). When intake fails to meetdemands, the birds may temporarily subsist bymetabolizing fat reserves, or as a last resort, catabolizeprotein from their own tissues. Such depletion of fatreserves may, however, jeopardize reproductive per-formance the following summer. It is possible, ofcourse, for temperatures to drop so low that quail can-not balance heat loss no matter how much they con-sume (especially if the cold is accompanied by strongwinds). When this occurs, body temperature declines(hypothermy) and the birds generally cease all activi-ty in a desperate attempt to conserve body heat. Deathis usually imminent at this point. Fortunately, this sit-uation is not common in Illinois and bobwhites aresurprisingly tolerant of cold temperatures per se.Anecdotal evidence from field studies in the Midwestsuggest that temperatures as low as -30o F do not nec-essarily result in heavy quail loss (98).

Nevertheless, the importance of adequate andaccessible food supplies to wintering birds cannot beoveremphasized. Not all foods are of equal quality.Yellow corn was once considered the most completequail food, but feeding trials during one study rankedit below millet, sorghum, and sassafras in sustainingvalue (73). Soybeans and common ragweed were nottested, but are known to be highly nutritious for quail.On the other hand, such items as smooth sumac,mul-tiflora rose, and Japanese honeysuckle fall into the cat-egory of emergency foods that may be taken occasion-ally, but have little sustaining value.

Bobwhites are considered generalists rather thanspecialists in their feeding habits – in other words,their diet depends primarily on what is available.Nevertheless, some items such as corn, soybeans,wheat, acorns, sassafras, beggar-ticks, and commonragweed may be selected for, i.e., taken in amountsdisproportionate to their availability (7). In contrast,the popular quail food Korean lespedeza may be con-sumed more in relation to its abundance. This is per-

haps not surprising since studies have shown thateven though a good source of protein, this legume isnot especially high in metabolizable energy, and hasonly moderate sustaining value (73, 82). Of course,other factors, such as palatability and durability, alsoinfluence food selection in the wild.

Although winter is potentially the most stressfulseason for bobwhite, significant energy“costs”also areassociated with reproduction. Demanding activitiescan be divided into behavior (courtship, territoryestablishment and defense, nest building), gonadaldevelopment, egg production, incubation, and brood-ing. The gross body weight of both males and females

remain generally stable throughout autumn and earlywinter, then declines in late winter as food suppliesdiminish (Fig. 5). Males continue to lose weight untilearly summer whereas females gain during April andMay, then decline throughout summer to a low in earlyautumn. Spring weight losses among males probablyreflects physiological and possibly psychologicalstresses associated with courtship, territorial estab-lishment, and competition for mates. In contrast,females gain weight during the pre-breeding perioddue to growth of reproductive organs and accumula-tion of fat. The caloric“cost”of developing a function-al oviduct and ovary is probably not great,but egg pro-duction requires a considerable expenditure of energy.Because developing young of oviparous (egg-laying)animals are not attached to the mother via a placenta,sufficient nutrients must be stored in each egg to per-mit development of the embryo to term. Incubation


Fig. 5. Annual weight fluctuations for male and femalebobwhite (Roseberry and Klimstra, 1971)

16

time is longer in birds like quail that hatch precocial(more fully developed) chicks, consequently the ener-gy content of their eggs must be higher. A bobwhiteegg contains about 16-19 kcal of energy, which mayapproximate 25% of the hen’s total daily intake (61).In fact, the energy content of a complete clutch ofabout 14 is roughly equivalent to the caloric value ofthe hen’s entire body.According to one study, however, the energy neces-sary for egg production and existence at normal sum-mer temperatures does not exceed that required forexistence alone in winter, and therefore should notimpose undue hardships (15). However, this study didnot consider the energetic costs of incubation, brood-ing, and the post-nuptial molt, nor the fact that hensmay renest 1 or more times. The major function ofincubation is to provide a suitable thermal environ-ment for eggs to hatch. To accomplish this,most birds

develop a brood patch where the transfer of heat isuninhibited by feathers. Early in incubation, the adultbird must supply most of the heat necessary to main-tain proper egg temperatures.As incubation progresses, an increasing proportion ofthis heat is supplied by the embryo’s own metabolicactivity. Avian physiologists disagree as to whetherincubation poses a significant energy demand on theadult. Some pheasant researchers, however, feel thatthe entire process of egg production, incubation, andbrooding constitute a serious physical drain on thefemale (8, 55). Additionally, the post-nuptial molt(also an energy requiring process) begins immediate-ly after reproduction has ceased. Based on weight lossand suspected mortality rates during this period, itdoes appear that late summer is a stressful time forbobwhite hens. And, as noted earlier, extremely hightemperatures may cause early cessation of nesting

17

activities.

The term habitat refers to the physical setting thatan animal occupies. Good habitat must provideessential life requisites (food,shelter,breeding sites) insuch a way that they can be successfully utilized by theanimal. Different species have their own unique habi-tat requirements and preferences, and how well theseare met in a particular area determines whether or notthe species will be present and in what abundance.Some wildlife species have very specific requirements.For example, Kirtland’s warbler, a small songbird,nests only in groves of young jack pines in northernMichigan. Other species, such as the bobwhite, areable to satisfy their basic needs from a variety of veg-etative communities thus enabling them to occupy awider geographic range.

Bobwhite are found in a wide variety of ecologicalsettings throughout their range including openpinelands in the Southeast, grass/brush rangeland inTexas, native prairie in Kansas, and of course, agricul-tural settings in theMidwest. This wide range of habi-tats may lead some to conclude that quail are adapt-able with flexible demands, but in fact, they are not.They require a rather specific set of conditions at boththe site and landscape levels, and if these conditionsdo not exist in a particular area, then neither willquail. The types of plant communities that can meetthese needs appear varied to us humans, but from thebird’s perspective, they are probably much more simi-lar than we realize with respect to such things as bareground exposure, stem densities, and canopy cover.The fact that a number of different plant communitiescanmeet these fundamental needs does notmean thatthe requirements themselves are flexible or that thebirds can alter them in order to adapt to changing con-ditions.

Experienced hunters and biologists can usuallydifferentiate between excellent and poor quail habitatand perhaps 1 or 2 intermediate categories. Most ofus, however, know of mediocre-looking spots that

hold birds year after year while other seemingly moreattractive areas go unused. Obviously we do not per-ceive habitat exactly as animals do. How then do ani-mals select a place to live? First of all, it must bestressed that their selection process is instinctive andinherent, i.e., it represents “evolutionary wisdom” notconscious thought. For quail, habitat selection isprobably based not so much on the species of plantspresent, but on their structural properties. Primarycues might include such things as litter depth, lightpenetration, relative obstruction of visibility andmovement (including flight), and thermal properties.

Quail seek and utilize settings where they can for-age effectively while remaining relatively secure from


Habitat

The close interspersion of cropland, herbaceous vegeta-tion, and woody cover provides ideal fall/winter habitatfor bobwhite (top), whereas good summer habitatrequires a more open mix of grasses and forbs (bottom).

18

predators and the elements. The types of situationsthat meet these criteria are limited by the behavioraland physical limitations of the birds themselves. Quaileat mainly seeds and insects located on or just abovethe ground, but they are ill-equipped to scratch or digthrough thick litter to reach these food items.Consequently, foraging areas must be relatively openat ground level.Also, because raptors are their primary predators,quail show a strong preference for sites providingsome degree of overhead concealment. And finally,because their anatomy precludes sustained flight, allusable habitat must be relatively close to protectivecover. In the Midwest, this set of requirements is usu-ally met in diverse, patchy, predominately open land-scapes with abundant woody edge and early succes-sional vegetation. Biologists in both Missouri (18)and Illinois (100) found that highest quail populationsoccurred in settings that were approximately 10-25%wooded and 75-90% open. Despite the relatively smallamount required, the presence of woody cover isclearly the most important factor in determining thedistribution of quail in Midwestern landscapes.Bobwhites are not forest dwellers, but they seldomstray far from forest edges, wooded ravines, or shrub-by fencerows. To be useful, however, such sites musthave sufficiently dense understory vegetation to pro-vide the birds both cover and concealment. Frequentflooding,heavy grazing, and insufficient light penetra-tion all greatly diminish the value of woodlands forq u a i l .Whereas woody cover provides necessary protectionfor bobwhite, more open habitats are essential forreproductive activities and feeding.

Non-breeding SeasonPrimary requisites during the fall and winter

months are adequate protective cover and an accessi-ble, reliable food supply. These essentials are not nec-essarily mutually exclusive. Some of the vegetationthat provides cover and concealment also supplies orharbors food and vice versa.

Individual covey ranges generally contain 1 ormore dense, woody or brushy patches that provideescape from predators and protection from severe

weather. These “headquarters” are usually about 1/10to 1/2 acre in size and may consist of bramble patchesor even large brushpiles. In southern Illinois, most

covey headquarters are in the Japanese honeysuckleunderstory of woodlots, wooded ditches, orhedgerows. When growing prostrate, honeysuckle haslittle value for quail, but if supported by other vegeta-tion or structure, its protective value is unsurpassed.Honeysuckle is especially valuable during heavy snowcover because it retains leaves throughout winter thusforming a protective canopy for the birds.

Another essential ingredient of winter habitat isan adequate and accessible food supply. Plant foodsutilized by bobwhite are found in a variety of succes-sional stages; however, early stages associated with soildisturbance, particularly row crop agriculture providethe greatest abundance of plant foods for Illinois bob-white including both waste grains (e.g., corn, soy-beans, milo) and seeds from annual grasses and

Diversity and edge characterize both non-breeding (top)and breeding (bottom) habitat.

19

broad-leafed weeds (e.g., foxtail, ragweed). Certaintypes of woody plants also supply food items in theway of hard and soft mast (i.e., acorns, berries, fruit,etc.).

A 3rd requirement for wintering birds is roostingcover. Quail spend most nights in sparse, short vege-tation with little surface duff and little or no overheadcover (65). Favored sites are lightly to moderatelygrazed pastures, wheat stubble fields, and early old-fields. One of the few uses that quail make of fescuepastures is for roosting. During periods of severeweather or prolonged snow coverage, quail will shifttheir roosting sites to heavier cover, especiallyJapanese honeysuckle.

The ability of a particular area to support quaildepends not only on the presence of essential habitatcomponents, but on their distribution or interspersionas well. One of the more important concepts inwildlife management is edge effect.Wildlife is generally most diverse and abundant in the

transitional zone where 2 or more distinct vegetativecommunities come together. There are exceptions ofcourse. Some species prefer large unbroken forests orgrasslands. By and large though, game and otherwildlife are apt to be found at or near the borders ofadjacent cover types where there is a distinct changein vegetative structure. Such areas usually offer richer,more varied plant communities thus providing for theneeds of more species. For individual species, theseareas provide“simultaneous access” to multiple habitat compo-nents. This is especially important to relatively seden-tary forms like the bobwhite. Quail are very reluctantto venture more than a few dozen yards or sofrom protective cover in order to feed. Consequently,high populations can be achieved only where food andcover patches are small (preferably no larger than 10-20 acres) and within 100-200 feet of each other.


Nests are generally found in relatively dense ground cover (top), whereas brood foraging areas are somewhat moreopen (bottom).

20

Breeding SeasonHabitat utilization during the spring and summer

is dictated not only by specific needs associated withnesting and brood rearing, but possibly also by theneed to avoid extremely high temperatures.

Researchers in Texas (33) found that bobwhitesgenerally avoided sites with operative (basicallyunshaded) temperatures >102o F. Prime breedinghabitat is generally somewhat more open than winterrange and usually contains proportionately less woodsand cropland and more grassy and herbaceous cover.While a reasonable mix of breeding and nonbreedinghabitat is desirable, the interspersion of cover typesthat satisfy seasonal requirements need not be as tightas those which accommodate daily needs. Research insouthern Illinois suggested that nesting habitat up to1/2 mile distant from winter range was well utilized(98).

An earlier study in southern Illinois examinedover 750 nest sites to determine the type of settingpreferred by nesting bobwhites (64). Virtually allnests were constructed of dead grass stems. Typicalnesting sites were well-drained and in moderatelydense stands of herbaceous and grassy ground coverwith scattered small shrubs or brambles. Within suchareas, nests were usually located close to more open,often bare, ground and often near some type of edge.These conditions may be found in a variety of settingsincluding idle fields, weedy fencerows, unmowedroadsides, unimproved pastures, and some hayfieldsand CRP fields. Quail may also occasionally nest inweedy cropfields, and notill areas but tend to avoiddense monocultural stands of legumes or grasses(especially fescue).

Habitat needs of growing chicks are less well doc-umented than the summer and winter requirementsof adults. In general though, successful brood rearinghabitat must have an abundant supply of insects andbe open enough at ground level to permit the smallchicks to move freely about without continuallyencountering dew-soaked vegetation. The thinstands of annual grasses, broad-leafed forbs, andlegumes that provide a desirable substrate may befound in early oldfields, early CRP fields, and somehayfields.

The Critical Role of Plant SuccessionIt should be clear to the reader by now that quail

are completely dependent upon weedy/grassy (i.e.,herbaceous) vegetation for their very existence. It notonly provides much of their natural food items (bothseeds and insects), but it is absolutely essential forboth nesting and brood rearing. The problem,howev-er, is that under natural conditions, this type of vege-tation is transitional, not permanent – and that bringsus to the subject of plant succession. Plant successionis the natural, progressive change in vegetative com-position and structure that takes place over time.

When an area is cleared to bare soil and leftunplanted or undeveloped, annual weeds and grassessuch as foxtail, common ragweed, pigweed, beggar-ticks, cocklebur, etc. will begin to appear during the1st growing season.After another year or so, perennials such as aster andgoldenrod start showing up. Then in a few more yearswoody shrubs and small trees (e.g., briars, poison ivy,sumac, sassafras, persimmon, winged elm, etc.) beginto invade. In time, a mature forest develops. The rateat which these changes take place and the duration ofeach succession or seral stage depends mainly on soilfertility and annual rainfall. In southern Illinois, theusual pattern is annual weed stage (1-3 years), peren-nial weed stage (3-10 years), shrub and bramble stage(7-15 years),pioneer tree stage (10-25 years), and latertree stage (after 30 years) (116). The 1st 3 stages areparticularly important to bobwhite; the annual andearly perennial weed stages for brood rearing, laterperennial and early shrub and bramble for nesting,and late shrub and bramble for protective cover. Theearliest stage provides optimum brood rearing habitatbecause it typically harbors good insect populationsand is still open enough at ground level to permityoung chicks to move about easily and find food. Thisstage,however,normally lacks sufficient accumulationof dead grass stems for nest building and sufficientstanding vegetation for nest concealment.Consequently, most nests are found in slightly laterseral stages characterized by scattered shrubs and bri-ars interspersed with a moderately dense groundcover of weeds and grasses.Unforunately,prime broodrearing and nesting habitat is ephemeral. Through thenatural process of succession, brood habitat will ulti-

21

mately become too rank and nesting habitat toowoody. In fact, it can be safely stated that left undis-turbed, good quail habitat will inevitably progresstoward something less desirable. To keep this fromhappening, there must be some type of regular distur-bance every few years to restart or retard successioneither by fire or some type of soil disturbance. We willdiscuss this in more detail in a later section (Habitat

Restoration).

Habitat From Various PerspectivesBobwhite habitat can be described and evaluated

at spatial scales ranging from site level (e.g., nest siteor field), to local (e.g., covey range or farm), to land-scape (e.g., several sections). Specific requirements


Left undisturbed, plant communities naturally progress from bare ground through annual, perennial, shrub/sapling,and forest stages.

22

exist at each scale. At the field level for example, arethere enough dead grass stems to build a nest, orenough litter-free ground for chicks to forage effec-tively, or sufficiently thick forest understory to serve ascovey headquarters? At the farm level, are all essentialhabitat components present and well interspersed?And finally, at the landscape level, is the habitat patchlarge enough to support a viable population? Becausehabitat formerly existed in larger blocks, and becauseindividual birds and coveys normally utilize rangesmeasured in acres rather than square miles, earlyquail biologists did not pay particular attention tolandscape conditions. As human developments andintensified agriculture began to dissect and fragment

existing quail range, some populations were forcedinto smaller, more isolated patches of habitat. Quailresearchers at Southern Illinois University (SIUC)voiced concern early on that these types of situationsmay put quail at risk for a variety of reasons (93, 98).Subsequent research has, in fact, suggested that habi-tat blocks or connected patches need to total at least5,000 acres or more to support viable, sustainablebobwhite populations (39). We repeat that individualbirds do not need this much space. However, as wewill talk about in the following section on demograph-ics, there may be a critical minimum population sizewhich will ensure viability, and that minimum popu-lation size may require as much as 5,000 or more acresof contiguous habitat.

A recent study examined satellite imagery todetermine if such remotely-sensed data could be usedto identify and differentiate between landscapes thatwere potentially suitable for bobwhite and those thatwere not. As it turned out, the researchers found thatin Illinois, bobwhites were usually associated withdiverse, patchy landscapes that contained moderateamounts of grassland and rowcrops and abundantwoody edge (100). More specifically, they seemed toprefer areas with about 15-30% grassland, 30-65%cropland, and at least 30 meters of woody edge perhectare (preferably more). In addition, they preferredlandscapes with a variety of cover types occurring insmall, well-interspersed patches. A later study (112)also found that bobwhite abundance was seeminglyhighest in landscapes featuring relatively largeamounts of small grains (i.e., winter wheat). Whetherthis reflects direct positive benefits of the wheat fieldsthemselves, or is merely indicative of general land-scape conditions favorable to quail is not yet clear.

The Question of WaterThe importance of permanent surface water to

bobwhite habitat is a question that comes up fromtime to time. Like all animals, quail need water insome form to survive. However, bobwhite in theMidwest are apparently able to satisfy most or all oftheir requirements from succulent vegetation, insects,and dew. Some experts believe that quail can live forseveral weeks without free water (102), while othersdoubt that they require surface water at all for drink-ing (38, 109). In any case, the absence of permanent

Successful bobwhite management must address habitatneeds at the landscape (top), farm (middle), and field(bottom) level.

23

surface water does not appear to limit the distributionof bobwhite in Illinois.

The primary interest of most sportsmen andother laymen often centers on the individual animal –its size, appearance, behavior, etc. Professional wildlif-ers, on the other hand, are generally more concernedwith the entire population. A population may bedefined as an assemblage of individuals in which eachhas a reasonable chance of mating with any memberof the opposite sex. In the preceding sections,we dealtwith characteristics, living habits, and needs of indi-viduals, pairs, and coveys.We now focus on demographic attributes that areunique to the population as a whole including birthrates, death rates, sex-age ratios, and abundance.Despite a shifting emphasis from single species to anecosystem approach in wildlife conservation, popula-tions generally remain the basic unit of management.

Mortality and Reproductive RatesWhether a population increases, declines, or

remains stable over time depends on the relationshipbetween mortality and reproductive rates. If birthsconsistently exceed deaths, the population willincrease; if more animals die than are produced,num-bers will decline – neither situation can be perma-nent. Under stable living conditions, mortality andreproductive rates tend to produce a relatively stablepopulation size consistent with the carrying capacityof the habitat occupied. Different species arrive at thisbalance in different ways. Some, like elephants andhumans, tend to be long-lived with relatively lowmor-tality and reproductive rates. Others, like bobwhite,are short-lived with high mortality rates balanced byhigh reproductive rates. Mature bobwhites normallysuffer a 65-85% loss annually, making them amongthe shortest-lived of all vertebrate species. Most ofthis loss occurs during late fall and winter when non-

hunted populations in the Midwest are normallyreduced by about 50% and hunted populations by65% or more.

Fortunately, the bobwhite’s high reproductivecapacity is generally adequate to cope with theseheavy losses. Quail biologists measure annual repro-duction as percent summer gain (net summer pro-duction expressed as a percentage of the breedingpopulation). A 26-year study of one hunted bobwhitepopulation in southern Illinois revealed that summergains most commonly ranged from 100 to 300%(mean = 205; range 17-383) (98). For any givenamount of fall to spring mortality, there is a specificamount of reproductive gain necessary to exactly bal-ance the losses. This is not, however, a simple 1:1 rela-

tionship, at least not in terms of rates or percentages(Fig. 6). The reason for this is that each bird dying inwinter increases the amount of summer recoveryneeded while decreasing the base (breeding stock)from which that recovery is made (98). Thus, itbecomes progressively harder for populations to com-pensate reproductively for previous fall and winterlosses as those losses increase. This has importantimplications for management. For example, eventhough an increase in fall-spring mortality from 50 to60% is relatively greater than going from 60 to 70%,the latter is considerably more serious in terms ofpotential recovery. In fact, researchers found that the


Demographics

Fig. 6. Summer gain/winter mortality relationship nec-essary to maintain population stability.

24

study population mentioned above lacked sufficientreproductive capacity to consistently make up for pre-breeding losses >70%.

The Phenomenon of Density DependenceAs with most wildlife species, bobwhite reproduc-

tion and mortality tends to be density dependent. Inother words, their rates vary depending on the relativesize of the population. Specifically, percent summergains tend to be lower when breeding populations arehigh in relation to carrying capacity. Conversely, fallpopulations at or near carrying capacity usually sufferhigher mortality rates during the subsequent winterthan populations below carrying capacity. Obviously,this phenomenon does not represent a consciouseffort on the part of the animals to balance births anddeaths. Instead, density dependence is likely an inci-dental consequence of many factors – particularlysocial interactions within the population and the com-petition for preferred space. We know, for example,that when numbers are high, some coveys are forced tooccupy marginal, less secure ranges. Conversely, agreater proportion of individuals from small popula-tions are usually able to find suitable territories.Therefore, it would not be unexpected for survivalrates to be higher for these smaller populations.Likewise, it is not hard to imagine how crowding dur-ing the breeding season could interfere with reproduc-tive activities either through harassment, attraction ofpredators, or inequities in the distribution of primenesting sites.

Density dependence is more than just an academ-ic curiosity – it is a key reason why bobwhites are ableto successfully absorb heavy losses from predationand hunting year after year. Density dependenceaffords populations a kind of built-in stabilizer suchthat winters of above-average losses are often followedby summers of above-average gains, and vice versa.However, it is important to remember that this phe-nomenon is just a tendency evident over time, not ahard, fast rule. In other words, reproductive and mor-tality rates do not automatically and completely adjustto density each and every year. If they did, populationsize would never vary, which of course it does.

Furthermore, certain conditions (e.g., severe winters)can cause heavy mortality or depress reproductionregardless of current population density. And finally,the compensatory nature of density dependence isonly partially effective. Even though the per capitarate of reproductive gain might be somewhat less forlarge breeding populations, generally the greater thenumber of breeders, the greater the absolute numberof young produced.

Sex and Age CompositionOwing to their rapid population turnover, autumn

quail populations in the Midwest typically consist ofabout 65-85% juveniles (birds hatched the previoussummer), and about 15-35% adults (birds that havegone through at least 1 breeding season). A 50-yearcollection of hunter-harvested birds from Illinoisrevealed an average fall age ratio (juveniles:adult) of4.9:1 (112). Traditionally, such ratios have been con-sidered an index of productivity the previous sum-mer. The rationale being that good nesting yearsshould result in proportionately more juveniles in thefall population. As with many biological parametersthough, correct interpretation of age ratios is not assimple as it might first appear (87). It turns out thatautumn age ratios are actually more indicative ofannual population change than population size. Thisis because high recruitment rates, and thus highyoung:adult ratios, are often associated with lowbreeding populations owing to density dependence.Consequently, they do not necessarily imply maxi-mum total production or large fall populations. Inaddition, the absolute ratio of young to adult in the falloverestimates actual production for 2 reasons. First, itis inflated by adult mortality following the breedingseason, i.e., it is not a ratio of young per breedingadult, but young per surviving adult. Secondly,research has shown that juveniles are almost 25%more vulnerable to the gun than are adults and thusare over-represented in harvested samples (99).

Bobwhites hatch at about a 50:50 ratio but femalessuffer slightly higher mortality thereafter, especiallyduring and after the breeding season. Therefore, fallpopulations normally have about equal numbers ofmales and females among young birds, but a prepon-


25

derance of males (60:40) among adults. Data fromhunter harvests suggest relatively little annual varia-tion in sex ratios, especially among juveniles, and nodiscernable long-term trends. The data also show thatfemales of both ages are slightly more vulnerable tothe gun than theirmale cohorts but the difference is sosmall as to be biologically unimportant (99).

MetapopulationsJust as a population is a group of interacting indi-

viduals, a metapopulation can be thought of as agroup of interacting populations. Metapopulationsoccur when habitat is moderately fragmented, i.e.,when occupied patches do not form a contiguousblock, but are not so greatly separated as to precludeoccasional interchange among neighboring groups.This exchange probably takes place mainly during the“fall shuffle” and spring covey breakup. Habitats can,however, become so fragmented that remnant popula-tions are truly isolated and at greater risk than nor-

mal. This risk is primarily determined by two factors:size of the population and degree of isolation fromneighboring groups. Small populations may simplygo extinct due to normal population fluctuations.This is especially true of species such as bobwhitewhich typically suffer 50-80% mortality from fall tospring. Isolation means that compensatory adjust-ments to severe winters, or excessive predation orhunting via ingress is limited or non-existent makingthese factors much more critical than they normallywould be. Finally, hunters and predators may be moreeffective when cover is limited and obvious.Additionally, isolated groups may lose genetic het-erozygosity through excessive inbreeding and geneticdrift. Ecologists speak of minimum viable popula-tions, i.e., the number of breeders necessary to ensurecontinued survival of the population. According toone leading quail biologist, the minimum viable fallpopulation may be at least 800 birds (44). Anotherunknown is what exactly constitutes an isolated pop-

26

ulation, i.e., what is the effective dispersal distance ofbobwhite? Several studies have suggested that popu-lations separated by much more than 1.25 to 1.5 milesmay be effectively isolated (112).

Relative numbers of quail present at any time orplace is of interest to sportsmen and biologists alike.This section discusses abundance across space(regional, local) and time (short-term, long-term).Distribution and abundance of quail nationwide isdetermined by basic vegetative patterns and by thefrequency and duration of potentially detrimentalenvironmental conditions. To a large extent, both aredetermined by climate. Within potentially suitablerange, local variation in abundance reflects the avail-ability and quality of essential resources such as food,cover, and reproductive opportunities, which, to alarge extent, are determined by human land use prac-tices.

A basic tenet of wildlife management is that agiven piece of land (unit of range) can support only somany animals of any one kind. The maximum num-ber of individuals of a particular species that an areacan support on a sustained basis is called carryingcapacity. This capacity depends primarily on 2things: the amount and quality of usable habitat, andthe species’ inherent tolerance to crowding. Biologistsoften express quail densities in terms of birds peracre, or some other land unit. Obviously, this is asomewhat artificial concept which does not reflectactual distribution patterns, but it is useful for com-parative purposes. It was once thought that their ownintolerance to crowding would limit bobwhites to nomore that about 1 bird per acre even in the best habi-tat. Studies in northern Florida have now shown thatthis limit can be exceeded under ideal conditions (59).In good Illinois quail range, populations may averageabout 25 birds per 100 acres, with local densitiesreaching >100/100 acres (98). By comparison, primequail range in the Southeast consistently supports 60-90 birds per 100 acres (24, 102), and managed localareas may have densities as high as 270/100 acres

(58).Both hunters and biologists are keenly aware that

bobwhite abundance can vary over time. There are 2distinct types of population change: short-term fluc-tuations and long-term trends. Short-term fluctua-tions are those periodic highs and lows that quail pop-ulations typically go through. These ups and downsnormally last only a few years and are primarily relat-ed to weather conditions although there may be acyclic component as well. Long-term trends arechanges in prevailing densities over several decadesand are generally habitat related (Fig. 1).

Short-term FluctuationsShort-lived species with high mortality and

reproductive rates typically show considerable fluctu-ation in year-to-year abundance, and quail are noexception. One local population occupying 2+ squaremiles near Carbondale fluctuated an average of about25% per year over a 26-year period. Maximum annu-al increase and decline was 80 and 60%, respectively(98). The various population phases (highs, lows,increases, declines) each tended to persist for severalyears suggesting at least a moderate “carryover” effectfrom year to year. The dominant factor influencingshort-term quail population fluctuations in Illinois iswinter weather – especially snow cover. Long-termrecords clearly show that annual population change isstrongly related to duration of snow cover the previouswinter. Furthermore, particularly hard winters canresult in major population declines. This was espe-cially evident during the late 1970s when 3 consecu-tive severe winters drove quail populations to all-timelows throughout the Midwest. Population responsesto summer conditions are much less noticeable or fre-quent although occasionally, extreme heat/droughtcan negatively impact production and fall abundance(91).

No discussion of bobwhite population fluctua-tions would be complete without mention of cycles.The fact that some wildlife species fluctuate betweenabundance and scarcity on amore or less regular basis


Abundance

has long fascinated and puzzled wildlife biologists.Two predominant types of cycles are recognized: the3-4 year microtine (e.g., voles, lemmings) cycle andthe 9-10 year cycle of snowshoe hare, Canadian lynx,and ruffed grouse in boreal habitats. The bobwhite isgenerally not considered a classic cyclic species,although earlier workers did not dismiss the ideaentirely (29, 60), and a long-term study of a local bob-white population in southern Illinois revealed a dis-tinct 8-10 year cyclic pattern in abundance (98).Recently, SIUC researchers statistically examined over70 long-term population records and found evidenceof cyclic behavior (4-17 years) in the northern andwestern portions of bobwhite’s United States range butnot the southeast (114). They speculated that thisbehavior resulted from a combination of externalinfluence (i.e., weather) and internal mechanismssuch as delayed density dependence and momentum.Even if present, cyclic behavior in bobwhite is at bestan underlying tendency,not a dominant force, and canbe easily masked by weather events and habitat loss.In fact, both simulated and real observations suggestthat as habitat declines, cycles become less pro-nounced primarily because the high phase isdepressed (98).

Long-term TrendsThe temporary ups and downs in quail numbers

described above are clearly evident from a half-centu-ry record of estimated quail abundance in Illinois(Fig. 1). Underlying these short-term fluctuations,however, is an obvious long-term, downward trend.Present day quail numbers in Illinois may be onlyabout 1/3 to 1/4 of what they were 40 years ago. Thissituation is evident in all parts of the Illinois quailrange (Fig. 7). There are multiple reasons for this,some of which may not as yet be fully understood.The primary reason, however, is that there are simplynot as many places for quail to live as there used to be.Some of this “quail space” has been lost to our everexpanding human population and insatiable demandsfor more living area. From 1964 to 2002, some 2.7mil-lion acres of Illinois farmland were converted to otheruses including housing, shopping malls, factories, air-ports, and roads. While we cannot assume that all ofthis agricultural land was quail habitat, some of it cer-

tainly was. Farmland that once supported several cov-eys of quail may now be the location of a sportinggoods store where Illinois hunters purchase suppliesfor their next bird hunting trip to Kansas. Severalspecies of Illinois wildlife (e.g., deer) have adapted tothese suburban/exurban developments and are doingquite well, but the same cannot be said for bobwhite.Human residents generally won’t tolerate the weedy,unmowed vegetation that quail need, and quail won’ttolerate the constant disturbance from humans, vehi-cles, and pets. Consequently, suburban or rural devel-opments, whether carved out of forested areas orexpanded into formerly agricultural lands, generallyoffer little or no quail habitat. Quail hunting opportu-nities also have been affected by the expansion ofdwellings and businesses into formerly rural areas.According to a recent SIUC study, 30% of rural Illinoisis within 275 m of a structure thus making it legallyclosed to hunting without permission (46).

Despite the above, nearly 75% of Illinois is stillfarmed so there can be little doubt that the major fac-tor in loss of quail habitat has been changes in the waywe farm the land. Forty to 50 years ago, most farm-land in westcentral and southern Illinois was usableby quail; today,much of it isn’t. The trend away fromdiversified, chemical-free, small-scale farming tointensive, chemically-dependent, row-crop agriculturehas negatively impacted quail habitat at all levels andspatial scales – field, farm, and landscape. Site (field)level conditions have been adversely affected mainly

27

Fig. 7. Population trends in the three quail managementregions of Illinois.

28

by technological advances. Valuable quail foods in theform of waste grains have been greatly reduced oreliminated by the combination of more efficient cropharvesting and fall plowing, while natural foods(annual weeds and insects) were being reduced byherbicides and insecticides. Advances in agronomyeliminated the need for periodic fallowing or idling ofcropland thus eliminating essential nesting and broodrearing cover while the widespread use of fescue cou-pled with earlier andmore frequent mowing have lim-ited the usefulness of agricultural grasslands for thispurpose. At the farm level, the shift from diversifiedcropping patterns to 1 or 2 row crops grown in large,borderless fields meant that certain components ofquail habitat were often missing, and the destructionof literally thousands of miles of brushy fencerowsand woody hedgerows not only eliminated criticalprotective cover, but often made remaining food sup-plies unaccessible to the birds. Finally, overall simpli-fication of the landscape itself not only eliminatedmuch usable space, but tended to fragment and isolatethat which remained.

Certain types of wildlife (e.g., waterfowl) can besuccessfully managed on relatively small public hold-ings such as refuges. This is not an option for quail. Tobe a viable game species, it is not sufficient for quail tobe only locally abundant – they must be reasonablyabundant over relatively large portions of the land-scape. Clearly, public areas such as refuges, wildlifeareas, or sanctuaries are too few and too small toimpact statewide or even county-level populations.Furthermore, such areas often provide inferior quailhabitat for a variety of reasons. Paradoxically, bob-white habitat can be adversely affected by either toomuch or too little human disturbance. The formerproblem usually exists on private lands whereas thelatter most often prevails on public lands such asparks, natural areas, or nature preserves. For this rea-son privately-owned farmland often has too littlecover for bobwhite whereas public land often has toomuch (98). When formerly agricultural areas are

acquired for conservation or recreational purposes,quail habitat is often initially improved as cultivatedfields are abandoned and early successional vegeta-tion predominates. However, if there is not a commit-ment to regularly maintain these areas by fire or soildisturbance, they quickly become too thick or rank forquail. Inmost cases, these types of management prac-tices are not used, or used too infrequently, to main-tain suitable quail habitat on state or federal holdings.Many public areas are intended for multiple uses, andearly successional vegetation and its management isoften not compatible with these uses (including man-agement of other species), or considered as aestheti-cally appealing as closed canopy forests or open grass-lands.

Local Extinction and RecolonizationAnother type of population fluctuation that can

be either short- or long-term is when local popula-tions go extinct. Plant and animal species have beengoing extinct for eons but modern-day conservation-ists warn that extinction rates have accelerated to analarming degree. We are talking here though, notabout species extinction, but about the temporary orpermanent extirpation of local populations, specifi-cally bobwhite populations. As we suggested earlier,isolated or semi- isolated populations are generally atgreater risk than those living in large blocks of con-tiguous habitat. This is especially true for bobwhitesbecause 1) their dispersal capabilities are limited, 2)they are subject to large numerical fluctuations owingto random weather events, and 3) they occupy prima-rily ephemeral habitats. Indeed, a recent study ofbobwhite counts from 81 North American BreedingBird Survey routes in Illinois (112) revealed that localpopulations along 12 of these 20-mile routes wentextinct, most following the severe winters of the late1970s. Five were later recolonized, but only 1 perma-n e n t l y .All of these populations were north of the primaryquail range in Illinois. Interestingly, recolonizations


29

took place only during years of below average snowfallsuggesting that such conditions may have increasedsurvival of immigrants.

The connection between bobwhite and agricul-ture is inescapable. How we humans use the land-scape to produce food and fiber determines its’ vege-tative composition and pattern, which in turn deter-mines the quantity and quality of bobwhite habitat.Because agriculture is the major force influencingland use in Illinois today, the bobwhite’s ultimate fateis inexorably linked with farm practices and policy.

Three-fourths of our State is farmland, so mostquail habitat must exist on agricultural lands if it is toexist at all. Furthermore, agriculture is the only landuse in Illinois capable of creating and maintainingquail habitat on any meaningful scale. Even on non-farmland, successful quail habitat management usual-ly employs agricultural methods. Unfortunately, thequail-friendly farming techniques of the past are forthe most part just that – a thing of the past. Present-day farming techniques and agricultural practicescommonly produce fields and landscapes that areinhospitable to quail. This section describes currentconditions across the Illinois agricultural landscapeand the problems and opportunities they present forbobwhite.

Regional DifferencesTraveling through the State of Illinois, even the

casual observer notices differences in topography andplant communities. Ecologists separate the state into14 natural divisions based on topography, soils,bedrock, climate, natural vegetation, and wildlife(103). The most rugged landscapes are generally cov-ered by deciduous forest, largely in private ownership.The land beneath these woodlands is too steep andinfertile to be used as cropland. Illinois forests are

concentrated on uplands along the Wabash, Ohio,Mississippi, and Illinois rivers and in the valleys oftheir major tributaries. In addition, there is a largeband of upland forest on the unglaciated hill countryextending across southern Illinois from the Ohio tothe Mississippi River (the Shawnee National Forest).In contrast, the flat, fertile lands formerly occupied byupland prairies and floodplain forests have beenalmost entirely converted to row crops (corn and soy-

beans). The Grand Prairie natural division occupieseast-central and north-central Illinois fromSangamon County east to Coles County on the southto Bureau and Kankakee counties on the north. Asidefrom some narrow corridors of forest along streams,

Bobwhite and Agriculture

Fig. 8. Land use changes in west-central (above) south-central (center) and south (bottom) quail managementregions in Illinois over the past 40 years.

30

this region is virtually 100% cropland and provideslittle or no quality bobwhite habitat. Between theseextremes are regions of the state with moderatelyrolling topography that support more diverse agricul-tural systems with intermediate amounts of forest.TheWestern Prairie Forest natural division runs fromthe Mississippi River east to the Illinois River andfrom Calhoun and Pike counties on the south toHancock,McDonough, Fulton, and Peoria counties onthe north. The Southern Till Plain natural divisionextends eastward from Madison County to CrawfordCounty on the north and south to the Shawnee Hills.The 3 bobwhite management regions in Illinois (Fig.7) generally correspond to natural divisions as fol-lows: Westcentral (Western Prairie Forest plus coun-ties immediately east of the Illinois river),Southcentral (Southern Till Plain), and South(Shawnee Hills). Overall land use (Fig. 8) is similar inthe Westcentral and Southcentral managementregions with 53-55% rowcrops, 18% forest, and 10-12% agricultural grasslands, but small grains are lesscommon (2%) in the former than the latter region(7%). In contrast, the rugged terrain of the Southregion is more heavily wooded (35%),with more agri-cultural grassland (15%) and less cropland (25%).

Changes Over TimeFarming practices and agricultural land use have

undergone major changes in Illinois since quail werelast abundant in the 1960s. Some of the changes areobvious while others are more subtle. All, however,have been detrimental to bobwhite. One obviouschange in the rural landscape is that there is less of it.As we noted earlier, some 2.7 million acres of Illinoisfarmland have been converted to other uses over thepast 40 years. Much of this conversion, though not all,has been to human dwellings or other developmentswhich has resulted in a net loss of quail habitat and/orhunting opportunities. Even when abandoned farm-land is used for recreational purposes, it is generallynot maintained in such a way as to provide optimumbobwhite habitat.

Another significant change has been a largeincrease in land devoted to corn and soybean produc-tion. In the 1960s, there were about 5.6 million acres

of rowcrops in theWestcentral and Southcentral quailrange; today there are more than 8 million (Fig. 8,Appendix B). Although more corn and soybeans aregrown now than in the past, the availability of thesepotentially important food sources to bobwhite hasactually declined. In the 1960s,most of the grain pro-duced in the quail range was available to bobwhitebecause it was grown in relatively small fields (20-40acres) bordered by brushy fencerows and woodyhedgerows that provided access for the birds. Today,most grain is produced in large (40-80+acre), border-less fields making it inaccessible to quail because thedistance to protective cover is too great. In addition,former harvesting and tillage practices left muchwaste grain and weed seeds in the fields for overwin-


In the past, waste grain and weeds in harvested crop-fields (top) provided a valuable overwinter food sourcefor bobwhite. Today’s cropfields (bottom) are generallyunusable by quail during the critical fall-wintermonths.

tering bobwhite, but today’s chemically-treated fieldsand efficient harvesting methods do not. On the pos-itive side, there has been a general increase in the useof “conservation tillage”methods which has increasedthe amount of overwinter cover on cropland(currently 28% of cropland is no-till and another 15%reduced tillage). However, large fields and absence ofprotective borders still rendersmost cropland inacces-sible to quail. Even where cropland is accessible, the 2-3 fold increase in insecticide and herbicide use haseffectively eliminated most of the weed seeds andinsects formerly available to bobwhite.

Veteran quail hunters recognize that there are farfewer woody hedgerows and brushy fencerows nowthan in the “good old days.” They realize too that thedestruction of just a few hundred yards of this type ofcover usually means the permanent disappearance of1 or more coveys of quail. Linear strips of woody orbrushy cover are critically important to bobwhite forseveral reasons. Not only do they provide valuableprotective cover themselves, but without them, other

habitat components such as feeding and nesting areasmay no longer be accessible to the birds. Although nodefinitive quantitative estimates are available, exami-nation of past and current aerial photographs andpersonal observations indicate that the loss of thistype of cover has been enormous over time and it con-tinues today. This, despite the fact that the totalamount of forested area has actually increasedthroughout the quail range (Fig.8). This increase doesnot, however, compensate for the aforementioned lossof hedgerow and small woodlot cover. Most gains inforested acreage result from advancing succession inplaces where there is already more than enough heavycover. In contrast, the destruction of small woodlots,or linear strips of woody vegetation along field edgesand riparian zones often represents loss of the lastvestiges of protective cover in a particular area.

Still another major landscape change with impor-tant ramifications for bobwhite involves agriculturalgrasslands, a critical component of habitat for groundnesting birds. Over the past 40 or so years, the com-bined acreage of hayfields, permanent and rotationalpastures, and diverted acres (fields temporarilyretired from crop production) in Southcentral andWestcentral quail regions has declined from almost 3million to just over 1.5 million acres (Fig. 8,AppendixC). Even more importantly, the composition andcharacter of these grasslands have changed fromquail-friendly vegetation to that which is mainlyunusable by quail. For example, hayfields formerlyprovided more usable habitat than they presently doowing to the use of different forage species. Cloversand lespedeza, which were the primary hay species inthe 1960s, grew more slowly and less densely thanmodern varieties of alfalfa, and were harvested later inthe summer (late June) and less frequently (twice pers e a s o n ) .Alfalfa, now the prevalent hay crop, is harvested asearly as May and may be cut 3 or 4 times during thegrowing season if rainfall is abundant. This level ofdisturbance makes it difficult if not impossiblefor bobwhite to successfully nest in such fields. In the1960s too, many permanent pastures consisted ofbluegrass, lespedeza, and various native forbs andgrasses, along with scattered shrubs and briars and

Brushy hedgerows (top) provide protective cover andaccess to fall/winter food supplies; unfortunately, thou-sands of miles of this valuable habitat component havebeen destroyed over the past 30-40 years (bottom).

31

32

provided excellent nesting habitat for bobwhite. By1990, most remaining pastures had been cleared ofwoody plants, fertilized, and planted to durable, thickgrowing grass species such as smooth brome and tallfescue which are of little or no use to bobwhite.

Rotation pastures may still provide valuablebrood foraging and roosting habitat but are also har-vested more frequently at present or may be heavilygrazed thus reducing their utility for quail.

Another major change in the Illinois landscapemay not be obvious to the untrained eye, but has nev-ertheless been critical for bobwhite. We refer to thegeneral absence of early successional annual forbs andgrasses so necessary for quail abundance. In the past,this essential habitat component was routinely createdby timber clearing, wildfires, abandonment of mar-ginal farmland, and most importantly, the widespreadpractice of temporary fallowing cropfields for 1 or 2years. Now, however, cleared timber land is usually

converted directly to cropland or homesites, wildfiresare rare, and fallowing and crop rotation are no longerpart of normal farming operations. It is no accidentthat peak quail populations in the Midwest coincidedwith the early days of agriculture when periodic crop-


Diverse, mixed stands of grasses, forbs, and legumes(top) once offered prime nesting and brood rearinghabitat for bobwhite, but today’s agricultral grasslandsoften feature dense, monotypic stands of fescue (bot-tom) which are of little or no value to quail.

Habitat fragmentation (top) and human developments(bottom) have caused further loss of bobwhite habitat.

Early-successional vegetation, essential for bobwhiteabundance, is often lacking in present-day agriculturallandscapes.

33

land fallowing was a routine farming practice. Thisproduced large amounts of prime brood rearing coverand winter foraging areas for quail on a regular basis.Today, however, most of Illinois is either intensivelyrow-cropped, or developed for human habitation, orleft relatively undisturbed. There are no currentland use practices that result in sufficientamounts of early successional vegetation neces-sary to support quail at their former level ofabundance.

In addition to conditions at the field level such asdescribed above, changing agricultural practices andpolicies have altered the very composition and patternof the overall Illinois landscape itself. A comparison ofaerial photographs from the 1960s to the presentclearly reveal these changes. The earlier, less-intensiveagriculture created a mosaic of small, diverse fieldsinterconnected by a network of woody hedgerows,brushy fencerows, and vegetated waterways. Today’sagricultural landscape is simpler,more expansive, andhas a generally more open look from the air. The con-sequences of this for bobwhite are that not only hasthere been a net loss of habitat, but some of the habi-tat that does remain now exists in smaller patches thatare more or less isolated from neighboring patches.The negative implications this has for quail were dis-cussed earlier under the heading Metapopulations.

In summary, the Illinois landscape has undergonedramatic change since quail were last abundant, and itis these changes (listed below in no particular order)that account for most of the decline in quail numbersover the past 40 or so years:

1. Permanent destruction of woody hedgerowsand brushy fencerows.

2. Increase in rowcrops and correspondingdecrease in agricultural grasslands.

3. Conversion of remaining agricultural grass-lands to fescue.

4. Enlargement of cropfields and elimination ofwaste grain, weed seeds, and insects.

5. Lack of early successional vegetation.6. Human development of former farmland.7. Advancing plant succession in undisturbedareas.

8. Fragmentation and semi-isolation ofremaining habitat.

While these problems are general throughout theIllinois quail range, they do not all exist to the samedegree in every part of every region. For example, theloss of formerly rural quail habitat to human develop-ment has been greatest in the more metropolitancounties of St. Clair, Madison, and Peoria, althoughother counties have been affected too, especially thosewith high rural human population densities such asWilliamson and Franklin. Destruction of woodyhedgerows and brushy fencerows has been mostnoticeable in the Southcentral quail region while theshift from grassland to rowcrops has occurred in boththe Southcentral and Westcentral zones.A rather different set of problems exist in the Southregion. In many parts of this region, the landscapelacks the diversity and complexity characteristic ofgood quail range. Extensive tracts of mature forestblanket the hilliest terrain while large, borderless row-crop fields dominate the floodplains. In between,most of the small, irregularly-shaped cropfields for-m e r l ynestled in generally woody terrain have been replacedby fescue-dominated pastures or CRP fields. In eachof these settings, there is a notable lack of early-suc-cessional grasses and forbs necessary for quailabundance.

Federal Agricultural ProgramsHistory

Until the Great Depression in 1929, farm opera-tors produced livestock and crops that were bestadapted to their region and yielded the greatest returnfor resources invested. Their fortunes were made orlost depending on their skills, but they were totally atthe mercy of the economy and the weather.A protracted drought and the Great Depression forcedthe failure of a large number of farm operations in the1930s. This economic catastrophe was so widespreadthat Federal officials were concerned that the nation’sfood supply could be jeopardized. They determinedthat legislation was needed to protect farmers fromeconomic disasters and to conserve the productivityof the nation’s farm land.

Aside from natural forces such as climate, soils,and topography, Federal agricultural policy is the

34

major factor influencing land use in Illinois today. Inour nation’s history, some industries have come to beregarded as crucial to national survival. For this rea-son legislation has been enacted to secure adequatesupplies of energy, food, and other vital commodities.Federal agricultural policy is rooted in this conceptwith the goal of ensuring an abundant, reasonablypriced food supply by providing economic incentivesto farmers for the production of certain crops and forconservation of farm land. At various points in ourhistory, economic incentives have been used to stimu-late or reduce production of various crops based onmarket demands. Over the same period, a muchsmaller portion of the resources available have beenused to encourage practices that conserve soil produc-tivity on farm land.

During the mid-1950s to the mid-1970s, increas-ing agricultural production often surpassed demandfor many agricultural commodities. Market prices forfeed grains fell below the cost of production prompt-ing the Federal government to use economic incen-tives to encourage farmers to idle cropland to reduceproduction. These programs, which were operativefrom 1956 to 1970, involvedmillions of acres national-ly and were often beneficial for bobwhite. For exam-ple, the “Soilbank” program of the 1960s had optionsthat encouraged farmers to idle cropland for up to 10years and either plant it to grass/legume mixes orallow it to grow into annual grasses and forbs. The“Feed Grain” program of the 1960s and early 1970salso idled large acreages of cropland with annual con-tracts that required no planting of cover crops. Thenatural, early- successional grasses and forbs that vol-unteered in these fields provided excellent habitat forbobwhite and populations remained at relatively highlevels throughout the State.

By the early 1970s, domestic utilization was con-suming less than half of our total agricultural produc-tion, leading to a fundamental shift in the goals ofFederal farm programs. Government and agriculturalleaders decided to stimulate agricultural productionfor the rapidly growing world export market. Idle landwent back into production and nationally hundreds ofthousands of acres of rangeland, woodland, and wet-lands were plowed, cleared, or drained and convertedto production of corn, cotton, soybeans, and wheat.

This was the era of “fencerow to fencerow” farmingthat in Illinois alone, led to the destruction of literallythousands of miles of woody hedgerows and brushyfencerows that previously held bobwhite. As we saidbefore, the negative impact on statewide quail popula-tions was almost immediately evident (Fig. 1).

In less than a decade, the economic and environ-mental ramifications of maximum agricultural pro-duction for the export market became apparent.Foreign demand fluctuated widely from year to yearcausing large swings in prices that strained budgets ofboth producers and the government in years whenlarge income support payments had to be made.Artificially high prices for commodities drove up pro-duction costs and land prices. Farmers in other coun-tries increased production and began to compete withthe United States for export market share. On theenvironmental side,many thousands of acres of grass-lands, forests, and wetlands were converted to crop-land resulting in a tremendous loss of wildlife habitat.Water quality was negatively affected by the runoff ofsediment, fertilizers, and pesticides from marginalcropland and the conversion of highly erodible fieldsto production greatly increased soil erosion. By theearly 1980s a shift in the direction of agricultural pol-icy was being discussed among various governmentaland private groups, but it took a series of years withvery high world grain production and very lowdomestic grain prices to bring action.

The Food Securities Act (Farm Bill) enacted bythe Federal Government in 1985 was intended toaddress some of the problems created by full produc-tion agricultural policies. In particular, there wasrenewed effort to limit production of excess com-modities. In addition, greater emphasis was given tosoil conservation, water quality improvement, andwildlife habitat restoration. The major wildlife con-servation program of the 1985 Farm Bill was theConservation Reserve Program which encouragedlandowners to convert highly erodible cropland tosemi- permanent cover for a minimum of 10 years.Participants in the program were allowed to selectamong several optional cover types for initial plantingincluding various grass monocultures, grass/legumemixtures, or trees. The threefold objectives of CRPwere to control production of excess commodities;


35

reduce runoff of sediment, nutrients, and pesticidesinto waterways; and restore wildlife habitat. The pro-gram proved to be quite popular among landownersnationwide. In Illinois, enrollment reached a maxi-mum of about 880,000 acres by the mid-1990s. As of2003, enrollment was about 645,000 acres withapproximately three-quarters of the acreage locatedwithin the State’s quail range.

The next major change in agricultural policy thataffected bobwhite occurred in 1996.World grain supplies were at low levels and the 1996Farm Bill (aka. “Freedom to Farm”) provided largeeconomic incentives for full production farming. TheConservation Reserve Program was continued, but

enrollment requirements became more stringent and

some land was returned to crop production or usedfor hay or pasture. The 2002 Farm Bill continued thestrong economic incentives to maximize production,but also contained new programs and alterations toCRP that could potentially benefit bobwhite. Thesewill be discussed in the following section.

Opportunities for bobwhitePrograms that idle cropland to reduce grain pro-

duction have great potential to enhance habitat foropen land wildlife like bobwhite. Whether or not thispotential is realized, however, depends on a number offactors including amount and location of land divert-ed, the type of vegetation established on the retiredacreage, and its subsequent management.

Most authorities agree that CRP has been at leastregionally successful in reducing soil erosion andenhancing habitat for certain wildlife species (1).However, despite initial optimism (52), tangible bene-fits for bobwhite have been somewhat disappointing,both in Illinois (95) and nationwide (9). This is not tosay that individual CRP fields are not used by quail onoccasion - clearly they are (13). Furthermore, it is like-ly that certain local populations have benefitted fromCRP in terms of increased production. However, it isobvious that the program has not prevented regionaland statewide populations from continuing to decline.

To understand why the Conservation ReserveProgram has not had a more positive impact onIllinois bobwhite, we need to consider the nature ofindividual CRP fields and how they fit into the overallhabitat picture. Many CRP fields were of little or novalue to quail because of the type or structure of veg-etation they contained. This situation resulted from anumber of factors including initial establishment ofunfavorable vegetation such as fescue, excessive anduntimely mowing, and lack of provision for manage-ment practices to retard succession. During the 1st 5or so years of the program,87% of all CRP acreage wasplanted to introduced cool season grasses andlegumes (CP1) (20). Many of these fields quicklydeveloped into dense stands of fescue or orchard grassthat were of little or no value for quail. Other fieldswere rendered useless to quail by excessive and unnec-

The potential value of CRP fields to bobwhite dependson their vegetative composition and structure (top),and their location with regard to other habitat compo-nents (bottom).

36

essary mowing. Early in the program,as many as 40%of all CRP fields in Illinois were mowed even thoughthere was no requirement to do so (20). As timepassed, farmers began to use more quail-friendly veg-etation in initial CRP plantings. Currently, >25% ofCRP vegetation is native warm season grasses or“per-manent wildlife habitat” compared to only 3.5% dur-ing the early years of the program. Potentially benefi-cial introduced grass/legume mixes such as timothy,redtop, and lespedeza also are more commonly used.Nevertheless, even when preferred vegetation is ini-tially established, CRP fields can rapidly lose theirvalue to quail through the natural process of plantsuccession. During the 1st few years after planting,many CRP fields contain relatively open stands ofplanted grasses, legumes and volunteer annual forbswhich provide valuable brood foraging cover.As succession progresses though, fields become morerank, and grasses begin to crowd out the legumes andannual forbs making the fields less useful as broodhabitat. Once the legumes, forbs, and small patches ofbare ground disappear, the field’s value for quailbecomes virtually nonexistent.

Even when site conditions are favorable, typicalCRP fields cannot be expected to provide year-roundhabitat for bobwhite unless there is a woody/brushycomponent which is generally not the case. The bestthat can be hoped for is that the CRP acreage will pro-vide or enhance a habitat component that is otherwiselacking, usually nesting and/or brood foraging cover.Thus, the location of CRP fields with respect to exist-ing quail habitat becomes an important issue. Toinvestigate this, researchers from SIUC mapped everyCRP field (>8,800) in 11 representative counties with-in the Illinois quail range and examined how theyrelated spatially to other habitat components (119).The majority of CRP exists within landscapes that aregenerally suitable for bobwhite because both are large-ly confined to moderately rolling terrain.Nevertheless, researchers found that almost 25% ofthe CRP acreage occurred in areas with insufficientwoody edge to support high quail populations.Additional CRP acreage provided limited benefits to

quail because it was located in areas where nest-ing/brood rearing cover was apparently not the limit-ing factor. In some parts of hilly southern Illinois,CRP probably had a negative effect on bobwhitebecause it replaced small cropfields that were provid-ing vital food supplies.

It was perhaps unrealistic ever to assume thatCRP could return Illinois bobwhite to the “good olddays” because the acreage involved simply is not greatenough. The amount of agricultural grassland inIllinois is currently 1.1 million acres less than duringthe late 1960s when quail numbers were still high.Even doubling the acreage of existing CRP would notmake up this difference.

Despite the aforementioned limitations, there isstill some justification for optimism regarding theprogram. As noted in the previous section, the 2002Farm Bill contained new conservation-oriented meas-ures that could potentially benefit bobwhite. One ofthese is the Conservation Security Program whichrewards producers for adopting conservation prac-tices (viz., crop rotation) to reduce soil erosion,improve soil and water quality, and create wildlifehabitat. Farmers would receive payments for follow-ing corn or soybeans with wheat or oats and then amix of clovers, alfalfa, or lespedeza that would be leftunharvested for 2 years. Such a program could addmuch-needed diversity to the agricultural landscape.Another new initiative, the Grassland ReserveProgram would promote restoration of native grass-land and prevent conversion of existing grassland tocropland. Of particular importance to bobwhite,farmers would receive incentives to convert pasture-land from tall fescue to native grasses and forbs. Twochanges in the Conservation Reserve Program are alsobeing viewed with optimism by quail biologists (16).One is the addition of a new CRP practice commonlyreferred to as “bobwhite buffers” which would estab-lish much-needed grassy/herbaceous buffer zonesaround the perimeter of crop fields. The second,referred to as “mid-contract management”, is arequirement that 20- 30% of each CRP field be discedor burned annually to reduce litter, expose soil, and


37

stimulate growth of legumes, annual grasses, andforbs. If successful, such measures could alleviate thepreviously-described situation in which advancingsuccession progressively reduces the value of CRPfields to bobwhite.

Conservation implies preservation or “wise use”of a resource whereas management implies manipula-tion of that resource and/or its users for a specific pur-pose. Wildlife management generally attempts tomanipulate animal populations and/or their habitats.Objectives might include production of harvestablepopulations, control of nuisance situations, or restora-tion or protection of threatened populations and habi-tats. Traditionally,bobwhite management has focusedon maintenance of suitable habitats to produce andsustain huntable populations. As both populationsand habitats continue to dwindle, issues of conserva-tion and restoration also are beginning to emerge (9).In the remainder of this section, we will discuss vari-ous approaches to quail management as they relate tocurrent and projected conditions. We will attempt toexplain the theory behind the practice because, as arespected quail biologist recently said (40):“Management in a theoretical vacuum is successful[only] when it is lucky.”

StockingMany sportsmen ask:“Since we don’t seem to have

enough quail,why doesn’t the State just release (stock)them the way they do turkeys?” Before we address thisquestion and talk about what stocking can and cannotaccomplish, we should point out that there are 3 basictypes of stocking each with its own specific objectives.These are: 1) release of pen-reared birds for con-trolled shooting or field trial purposes (i.e.,“put-and-take”), 2) release of pen-reared birds to augmentnative wild populations, and 3) relocation of wild orsemi-wild birds to repopulate depleted range.

Put and takeTwo chronic, related problems associated with

using artificially-propagated (pen-reared) quail forrecreational purposes are low post-release survivaland lack of natural behavior. The degree of wildnessexhibited by “game farm” quail can vary greatly,depending primarily on their degree of isolation fromhumans during the rearing process. Behavior maybecomemore natural with time spent in the wild (96),but survivorship declines. For example, harvestreturns from >5,000 pen-reared quail in southernIllinois was <1% for birds released in spring, 8% forthose released 7-12 weeks before hunting, 13% forreleases made 2-3 weeks before, and 55% for birds putdown during the hunting season (62). Similar resultswere noted in Indiana where returns from 8,000released quail ranged from 3 to 16% depending ontime of release (81). A Georgia study reported 18%harvest return of 6,700 birds released 3 weeks prior tohunting (71). “Gentle releases” into heavy protectivecover provided with several days supply of grain maydiscourage premature scattering and enhance earlysurvival. Relatively elaborate “call back” systems havebeen developed for this purpose (45). Nevertheless,we agree with a southern colleague that to maximizereturns: “pen-reared birds should be released as closeto the beginning of the hunting season as possible and[because] year-to-year survival is minimal, [harvest-ed] without regard to ‘carry over’ for the followingyear” (71).

SupplementalWhereas, put and take stocking may be justified

under certain circumstances, there is virtual unani-mous agreement among quail experts that releasingpen-reared birds to augment wild populations is notonly futile (23, 96), but possibly harmful as well (9).The futility of such efforts stems from the extremelypoor survival of pen-reared quail that has been docu-mented again and again by scientific studies.Potential harm to resident wild stock may arise fromseveral sources including genetic contamination,

Conservation and Management

38

introduction of disease, attraction of predators, andgeneral disruption of routine activities. Althoughmore research of these issues is warranted, at least 2studies (in Kansas and Georgia) reported increasedmortality among wild quail following releases ofpen-reared birds (104, 106), and several groups ofleading quail biologists have recommended againstlarge-scale stocking of pen-reared quail because of thepotential adverse affects on resident wild birds(51, 66).

TranslocationAt this point, readers may be wondering if there is

a place for any kind of stocking in science-based quailmanagement – the answer is: maybe. Relocation ofbirds to accelerate recovery or reestablish populationsin depleted but suitable habitats might be a viableoption in local situations,butmore research is needed.In the past, it was assumed that suitable habitatswould naturally contain quail, but as we earlier noted,small populations in fragmented range can be deci-mated by severe winters with little or no opportunityfor recovery due to their isolation. In other instances,islands of newly created habitat might remain emptybecause distance to neighboring populations is toogreat for effective immigration. In such cases, a viablemanagement optionmight be to transplant or translo-cate wild birds into the area (9). Transplanted stockmight include captured wild quail or pen-reared off-spring from 1 or 2 wild parents. Pen- reared offspringof 2 wild parents reportedly performed better whenreleased than did standard “game farm” birds duringan Indiana study (4). However, researchers in Ohio(48), Illinois (96), and Texas (77) all found that pen-reared quail with wild parentage were inferior interms of survival and behavior to relocated wild quailthat never experienced prolonged confinement. Infact, the latter 2 studies found that wild or semi-wildquail reared in captivity performed no better thanstandard game farm birds.

And now, back to the original question of whystocking works for turkeys but not quail. The restora-tion of the eastern wild turkey across its former rangein North America is among the great success stories of

modern wildlife management. To accomplish this,wild (not game farm) turkeys are livetrapped wherethey are abundant, transported to areas of vacant butsuitable habitat, and released. Usually a release of 15birds (12 hens and 3 adult gobblers) is sufficient toestablish a new population given their relatively highannual survival rate and considerable ability to dis-perse. There are several factors that make bobwhite aless likely candidate for relocation. For one thing, theirsurvival is only about 1/2 to 1/3 that of turkeys andtheir dispersal ability is much less so initial stockingsof several hundred birds would probably be necessaryto have any chance of success. However, the main rea-son that turkey relocations were successful is thatbirds were released into suitable but vacant habitat. InIllinois, there is little, if any, vacant suitable habitat forbobwhite. Virtually all habitat that will support quailis currently occupied. If an area has no quail, it isalmost certain that it is lacking one or more criticalhabitat components, and relocating birds into such anarea would be doomed to failure.

Harvest ManagementTheory

An inherent capacity for rapid population growthpermits quail and other small game to sustain annualhunter harvests year after year without apparenteffect. Note the word apparent, however. Until per-haps 20-25 years ago, the“annual surplus” theory (69)formed the basis for most upland game huntingincluding bobwhite. Briefly, this theory held that gamepopulations generally produce more offspring thancan survive, and that regulated hunting merelyremoves this excess production without affecting thebreeding stock. More recent research, however, hasindicated that this view of hunting, and its impact onpopulations, is oversimplified,and can lead tomiscon-ceptions about harvest and harvest management.Field studies in Illinois and elsewhere have shown thatprotected quail populations in the Midwest normallysuffer about 50% reduction from fall to spring (98)whereas hunted populations may lose between 60 and80%, thus indicating that hunting can indeed increasetotal mortality and lower breeding densities. Fred


39

Guthery reached a similar conclusion for Texas bob-whites (35). But – and this is an important but – theloss of potential breeders caused by hunting is lessthan the actual number of birds shot; furthermore, amoderate reduction in breeding densities can actuallystimulate production.

It is important to understand how these 2 thingscan be, so let’s examine them separately. First of all,hunting is not completely additive to natural mortali-ty because some of the birds shot would have diedanyway just as the old “annual surplus” theory held.Biologists refer to this as competing risks. It’s just afancy way of saying that even though an individualhas a chance of dying more than one way, it can in factonly die once. This relationship can be approximatedby the simple equation: T =H+N - (HN) which statesthat total fall-spring mortality rate (T) is equal to theharvest rate (H) plus the natural mortality rate in theabsence of hunting (N) minus that portion of the har-vest that would have died anyway (HN). In additionto this purely numerical compensation, birds that sur-vive the hunting season tend to suffer slightly less late-winter mortality than nonhunted birds (98), probablybecause as hunting thins out the population, morefood and cover is available to the remaining birds thusincreasing their chances for survival. The factremains, however, that everything else being equal,hunted populations generally have fewer birds left atthe end of winter than non-hunted populations. As itturns out though, this is not necessarily a bad thingbecause of the phenomenon of density-dependentreproduction which we talked about in an earlierchapter. When breeding densities are temporarilyreduced below carrying capacity, as in the case ofhunting, they tend to respond with increased individ-ual production. This increased reproductive outputpartially mitigates the fact that there are fewer breed-ers in the hunted population.

To put it another way, populations at carryingcapacity tend to be sluggish with lower annual turn-over, higher proportion of older individuals, andreproductive output just enough to match mortality.In other words, there’s little or no excess productionbecause the population is no longer growing. By con-trast, populations below carrying capacity tend to bevigorous and productive, with a high potential for

growth. By forcing breeding densities to this lower butmore productive level, hunting actually stimulatesexcess production, thereby producing a “harvestablesurplus.” The relationship between annual harvestrate, population size, and total yields can be visualizedin Fig. 9. If hunting pressure is very low, the standing

crop remains high but yield suffers because only asmall proportion is taken each year. Conversely, exces-sive hunting pressure can reduce standing densities tothe point where yields suffer. An intermediate harvestrate that promotes high individual productivity whilemaintaining an adequate number of breeders general-ly produces the greatest long-term yields and is theaim of management.

The next logical question is what constitutes areasonable harvest rate and how can it be achieved? Arecent book on quail management (40) recommendedharvesting to reduce fall populations to spring levelsthat would maximize percent summer gain.Theoretically, such an approach would achieve maxi-mum sustained yields (MSY), i.e., permit the largestnumerical harvest year after year by keeping the pop-ulation at its most productive level (88,89). In Illinois,MSY could probably be achieved by removing abouthalf the fall population each year. However, this levelof harvest would leave little margin for error, especial-ly when crippling losses and periodic hard winters areconsidered A more prudent harvest might be around40-45% (90, 92). Even this more conservative offtakeis somewhat higher than generally recommended by

Fig. 9. General relationship among harvest rate, yieldand population size.

40

quail biologists in the Southeast (102) and Southwest(105). Actually, there is no single rate of harvestappropriate for all situations or places. Statewide, thegoal is generally to provide some recreation for themaximum number of hunters, whereas on private orcommercial areas, the goal is often to provide qualityhunting for fewer individuals. The latter situationclearly calls for lower hunting pressure and harvestrates. Allowable harvest also can vary geographically,e.g., northern populations can generally sustain high-er rates of harvest than southern populations (36).This may seem counterintuitive given that naturalmortality rates are higher for northern birds.However, as we can easily demonstrate with the previ-ous equation T = H + N - (HN), the higher the natu-ral mortality rate, the less additive hunting losses areto it. Furthermore, northern populations, with theirnormally higher annual turnover, have naturally high-er reproductive rates allowing them to better compen-sate for the additional mortality caused by hunting.

PracticeAs a practical matter, statewide quail populations

cannot be estimated, nor hunting regulated, closelyenough to achieve specific harvest rates. The goal ofstatewide harvest management is simply to providereasonable recreational opportunities for sportsmenwhile ensuring the continued welfare of the resource.Depending on the species, recreational hunting inIllinois is generally controlled by regulating a) num-

ber of hunters afield, b) daily or seasonal bag limits,and/or c) season length. With regard to quail, only thelatter 2 apply.Over the past 50+ years, the bag limit forbobwhite in Illinois has gone from 10 to 8 to 6 and(since 1988) back to 8 (Appendix D). According tosurveys taken in the 1990s (2,3) approximately 60% ofIllinois quail hunters would like to see the bag limitreduced to 6, while only 6% would like it increased to10. In reality, adjusting the current bag limit wouldprobably have negligible effect on overall harvest. Forexample, even if 1/4 of the total harvest resulted fromhunters taking a limit of quail (8), reducing the limitto 6 birds would lower the total harvest by only 6%.

A more viable approach to regulating the totalharvest is by adjusting the length, and to a lesserextent, the timing of the open season. Bobwhite hunt-ing seasons in Illinois have gotten progressively longerover the years. They were only 30-40 days long duringthe 1950s but currently range from 63-69 days in thenorthern zone and 70-76 in the southern zone(Appendix D). Since 1989, opening day has been the1st Saturday in November. A majority of hunters aresatisfied with this date, but some (16-25%) feel it istoo early (2, 3). They argue that early Novemberweather can be too hot and dry for good dog work,some birds aren’t yet fully grown, and crops are oftenstill in the fields making it difficult to locate birds.Records show, however, that over 85% of all quail areat least 4/5 grown by November 1. Furthermore, thenumber of birds available for harvest is higher than itwould be later in autumn. Whereas, most hunters aresatisfied with an early November starting date, 56% ofthose surveyed in 1991-92 wanted regional seasonsextended beyond January 1 and January 7 (2). Evenwhen closing dates were subsequently extended toJanuary 8 and 15, 37% still thought the season endedtoo early (3). Despite popularity with some hunters,late-season hunting has negative biological implica-tions prompting several quail biologists to go onrecord against it (17, 92). The main problem is thatthe later in the year that hunting losses occur, themorethey reduce the breeding stock (80, 84). To put it sim-ply, a bird bagged in November probably wouldn’thave survived to breed anyway, whereas a bird shot inlate January probably would have. Some would arguethat hunting pressure typically declines as the season


The goal of harvest management is to provide reason-able recreational opportunites for sportsmen whileensuring the continued welfare of the resource.

41

progresses, thus providing a built-in safeguard againstoverharvest. However, those avid hunters who do per-sist are often the most experienced and skilled.Furthermore, quail are more vulnerable to hunting asthe season progresses because escape cover is beingcontinually reduced by crop harvesting and break-down of natural vegetation. Finally, late-season hunt-ing may affect even those birds that escape the gun,i.e., the breeding stock. Waterfowl biologists recognizethe importance of sending females to the breedinggrounds in the best possible physical condition. Thesame holds true for bobwhites – they must comethrough the winter in good shape if they are to repro-duce at maximum capacity, a necessity for heavilyhunted populations. Some biologists worry that late-season disturbance by hunters and dogs can disruptnormal feeding patterns during a critical time of theyear. Such disturbance can be particularly damagingduring severe weather or periods of prolonged snowcover. Given all these considerations,we do not believeit wise to extend quail hunting in Illinois much pastthe 1st week in January.

It has been suggested that perhaps quail huntingseasons should be adjusted according to the anticipat-ed size of the fall crop, i.e., liberalizing the harvest dur-ing good years, and reducing it during poor ones (35).Computer modeling in Illinois showed that such astrategy might be marginally beneficial (88). In actu-ality, hunting pressure and thus harvest intensity isprobably at least partially self-regulatory, i.e., interestand participation may rise and fall with quail abun-dance, both short- and long-term (56, 79). However,management cannot and should not rely solely on vol-untary actions to safeguard the resource (90).

It is becoming increasingly apparent that long-held assumptions and traditions regarding quail hunt-ing may have to be reevaluated in the face of decliningpopulations and deteriorating habitat (9, 12, 93).Much of the natural resilience that quail populationshave to hunting, predation, and weather, results fromthe numerical and physical security that characterizelarge, thriving populations in prime habitat.Unfortunately, wildlife managers no longer have theluxury of always working with such populations. Theynow find themselves dealing more and more withbirds living, as the late long-time Missouri quail biol-

ogist Jack Stanford (108) put it, “closer to the edge.”Quail in smaller, more isolated pockets of habitat aresimply not as resilient to routine pressures from hunt-ing, predation, and weather as are larger populationsoccupying extensive tracts of prime habitat. In recog-nition of this fact, some quail biologists have suggest-ed that specific harvest regulations be applied at geo-graphic scales smaller than statewide (121), or thatmore restrictive regulations be employed for at-riskpopulations that do not greatly impact hunting oppor-tunities elsewhere (42).

Predator ControlTheory

Predation, and the effect it may or may not haveon quail populations, is an emotionally charged issuethat evokes much debate among quail enthusiasts (12,74). Many hunters operate on the assumption thatsince predators kill quail, fewer predators would meanmore quail. Biologists argue that it is not that simple.Complicating the issue further is that many non-hunters do not believe that predator control is justifiedregardless of whether or not game populations wouldbenefit. In any case, we should not lose sight of thefact that bobwhites have successfully coexisted withpredators throughout their entire evolutionary histo-ry, and in the process have developed effective life his-tory strategies (large clutches, renesting, etc.) to copewith routinely heavy annual losses (10). Furthermore,many of the very behavioral traits that make bobwhitea prized gamebird (holding for pointing dogs, explo-sive flushes, rapid flight, etc.) undoubtedly evolved asresponses to predation (107).

In the classic predator-prey relationships of thefar north (e.g., lynx/hare, wolf/deer) the prey general-ly represents the predator’s main food source while thepredator is the prey’s principal enemy. Such is not thecase with bobwhites, however. Quail are preyed uponby a wide variety of birds, mammals, and even rep-tiles. Predation of adult bobwhites during fall andwinter is mostly from hawks and owls, whereas mam-mals of one kind or another are primarily responsiblefor most nest predation. None of these, however, relyon quail as their primary food source. Quail preda-tors, especially the mammals, tend to be generalists intheir feeding habits and quail are normally just inci-

42

dental (and often rare) items in their diet. As we shallsee later, these are important considerations whenassessing the pros and cons of predator management.

The renowned pioneer of quail research, HerbertStoddard, felt that predation played a significant rolein quail population dynamics (109), but it was thewritings and opinions of his even more famous con-temporary, Paul Errington, that influenced quail biol-ogists for decades. Errington,working with quail pop-ulations inWisconsin and Iowa, advanced the“thresh-old of security - doomed surplus” theory which tend-ed to minimize the impact of predation (and hunting)on breeding populations. Errington believed that latewinter habitat conditions acted as a sort of bottleneckso that the number of birds surviving to spring wasrather uniform regardless of how many entered thefall (31). He reasoned that predation seriously affect-ed only “surplus” birds that exceeded their “thresholdof security”(i.e., carrying capacity), and that breedingdensities were therefore regulated by habitat condi-tions and not predator populations (30).

This theory of game bird population dynamicsremained popular and accepted for decades, eventhough certain aspects were being challenged by long-term population studies of both ring- necked pheas-ants in Wisconsin (117) and bobwhite in southernIllinois (98). Specifically, these studies found thatrelationships between fall densities and rates of winterloss and subsequent spring populations did not neces-sarily support the idea of a“threshold of security”and“doomed surplus” of birds. The implications of thiswas that both hunting and predation may exert agreater effect on quail abundance than was previouslysuspected. Furthermore, Illinois researchers (63, 98)as far back as the early 1980s were warning thatchanging landscape and habitat conditions might bealtering relationships between quail and their preda-tors (both animal and human). More recently, otherresearchers, especially in the Southeast andSouthwest, have proposed that under certain condi-tions, nest predation can limit bobwhite recruitmentand thus negatively affect fall abundance (50, 75, 85).

Various factors (all human-related) have alteredbobwhite/predator relationships over the years. Forone thing, certain predators are more abundant nowthan 30-40 years ago. This is particularly true for

medium-sized mammals (potential nest predatorssuch as racoons, skunks, and opossums) probablybecause of declining fur prices, less recreational trap-ping and hunting, and possibly landscape changes.Raptor numbers also have increased due to elimina-tion of organochlorine insecticides, increased lawenforcement, and public education.

More important still are landscape changes thathave drastically altered the amount and spatial pat-terns of remaining bobwhite habitat. Nesting cover inparticular has been greatly reduced and now oftenoccurs in small, isolated patches or narrow strips thatmake for easier hunting by nest predators.Additionally, fragmentation of much year-round habi-tat has likely increased the vulnerability of residentbirds to all forms of mortality including weather,hunting, and predation.

PracticeIn the view of many hunters, the most direct and

logical method of controlling predation would be toreduce the number of predators. However, there areseveral problems with this approach. First, there is nosingle, dominant quail predator on which to focuscontrol efforts. As we said, quail are taken on a year-round basis by a wide variety of animals, none ofwhich“make their living” from quail. Secondly, killinghawks and owls is not even an option. It is sociallyunacceptable and it is illegal! Thirdly, high popula-tions of certain mammals may be related more to eco-nomic and social than to biological factors, and thusnot readily addressable on a regional or statewidebasis by natural resource agencies. Individuals orgroups of landowners might consider encouraginglegal trapping and furbearer hunting on their proper-ties although there is no guarantee that quail numberswould increase. Furthermore, reductions of certainpredators might actually produce negative results.Coyotes, for example, are often blamed for low quailnumbers when, in fact, they may tend to suppressmore effective quail predators such as red foxes, rac-coons, and skunks (10, 47, 85) .

Alternatives to reducing predator abundancemight involve habitat manipulation to limit theireffectiveness. For example, increasing the size andnumber of nesting habitat patches may help reduce


43

nest predation, while increasing understory coverwithin hedgerows and providing brushy buffer zonesbetween mature woods and cropfields may reducepressure from avian predators. The most effectivepredator management, however, would simply be toincrease the amount of bobwhite habitat in general.Predation normally poses little threat to large, vigor-ous quail populations inhabiting extensive tracts ofquality habitat; it becomes a potential problem only insituations of reduced habitat quality, fragmentation,and population isolation (107). Past research hasclearly shown that good habitat can produce goodquail populations even when predators are abundant.Conversely, quail cannot exist in the absence of habitatregardless of the predator situation. In other words,predator management cannot be substituted for habi-tat management (10).

Habitat ManagementThroughout this booklet we have attempted to

stress one simple, but all-important, fact: the key toquail abundance is habitat. For bobwhite, as for allother wildlife, habitat is not a luxury, it is an absolutenecessity. There can be no substitute – not stocking,not closed seasons, not predator control. If we want tohave quail, we have to provide them with a place tolive, and if we want to have lots of quail, we have toprovide them with lots of places to live. And that in anutshell, is why we don’t have as many quail as weused to – there simply aren’t as many places for themto live as there used to be. The heyday for bobwhite inthe Midwest was when they existed as a normalbyproduct of routine farming operations. In thosedays, much of the rural landscape provided excellentquail habitat, not because it was managed for quail,but because the diverse, low-impact, chemical-freeagriculture provided a naturally ideal setting for quail.Today, however, most high-quality quail habitat mustbe created and maintained via specific managementefforts that rely on 3 key elements: knowledge, com-mitment, and opportunity.

Because bobwhite/habitat relationships varyacross spatial scales (i.e., field, farm, landscape,region, state), management must do likewise. At thefield level,management usually involves manipulatingplant succession to create andmaintain favorable con-

ditions. Almost invariably, this means retarding orsetting back succession to an earlier stage either byfire or some type of soil disturbance. The knowledgeand expertise to do this is readily available from vari-ous natural resource agencies. Commitment becomesan issue here because the dynamic nature of succes-sional vegetation requires a concerted effort to main-tain it at the proper stage. Quail habitat cannot be cre-ated and then ignored – it will inevitably progress tosomething less desirable, and this progression can beremarkably quick, just a few years in some cases.

Management at the next larger scale (i.e., farmlevel) obviously includes field-level habitat manipula-tion, but must also incorporate the concept of limitingfactors. Just as a chain is only as strong as its weakestlink, habitat is only as good as its poorest (most limit-ing) component. For example, a farm lacking ade-quate winter cover cannot be improved for quail byadding food plots. Here again, knowledge of whathabitat component is most lacking and how it can beprovided is generally available from natural resourceagency professionals.

Despite its importance, management at the farmand field level may not always be sufficient to produceviable quail populations in today’s landscapes.Management at broader geographic scales involvinggroups of adjacent landowners may be necessary toaddress current bobwhite needs (121).As we said earlier, quail populations may require asmany as several thousand acres of habitat to ensuresuccessful long-term persistence. It is at this land-scape level where opportunity (or lack of) formanage-ment becomes paramount. Clearly, if habitat is to beprovided at this spatial scale, the bulk of it must occuron privately-owned farmland which makes up 75% ofIllinois. As we have repeatedly pointed out, modern-day agriculture no longer routinely provides a favor-able setting for bobwhite as it once did. Quail habitatmust now be specifically planned and provided forwithin the context of production agriculture, and to alesser extent, forestry practices. Those portions ofIllinois that are flat and fertile will likely remaininhospitable to bobwhite barring major, unforseenchanges in land use. Hopefully though, there will beopportunity to improve conditions within less-inten-sively farmed landscapes with existing or emerging

44

conservation-oriented agricultural programs, contin-gent of course, on the willingness of landowners tocooperate in these efforts.

Whereas much practical knowledge exists regard-ing the mechanics of bobwhite management at thefield and farm level, less is known about habitat rela-tionships and management needs at larger spatialscales (93). For example, it is not entirely clear howlarge contiguous habitat tracts must be to ensure pop-ulation viability, or at what distances do populationsbecome dangerously isolated, or how landscape struc-ture impacts population responses to hunting andpredation (12).

Questions also arise as to where managementefforts should be concentrated within landscapes.One strategy would be to focus on rapidly decliningpopulations to prevent possible extirpation; an alter-native would be to focus on currently stable popula-tions to maintain their strength and prevent futuredeterioration (113). Most wildlife professionals agreethat management should concentrate on the currentIllinois quail range. The logic here is that the inten-sively row-cropped east-central portion of the State isso lacking in basic necessities for bobwhite that isolat-ed management efforts would be futile. Even withinthe established quail range, there are areas so inten-sively farmed or heavily wooded that attempts tomanage for quail would probably be a waste of time.Researchers at SIUC (100, 111), using habitat model-ing and satellite imagery, recently produced statewidemaps identifying those landscapes potentially suitablefor bobwhite (Fig. 2). It is hoped that results from this,and similar, research will enhance and focus future

bobwhite management in Illinois and elsewhere.Finally, there is the question of whether manage-

ment should emphasize improvement of existinghabitat or creation of new habitat. It has long beenheld that bobwhite habitat quality represents a contin-uum from excellent to average to poor. Recently, how-ever, a respected quail biologist (37) advanced theargument that if a particular piece of land is alreadyoccupied by quail, then it cannot actually beimproved, and that the best way to “improve” an areais to make additional space usable by quail, eithermore of it, or for longer periods of the year.Regardless, creation of new habitat is clearly desirableso long as the addition is not so small or remote that itcannot attract and support viable quail populations.In this regard, we recommend new habitat be estab-lished adjacent, or in very close proximity, to existinghabitat, thereby enhancing the value of both. Keep inmind, however, that even if occupied habitat cannotbe improved, it still must be maintained, i.e., theremust be frequent management attention to controlsuccession.

To summarize, the presence or absence of plantcommunities that make up bobwhite habitat reflectsthe interaction of climate, topography, soils, andhuman use of the land. Currently, decisions aboutland use are the most important forces determiningthe distribution and abundance of such habitat. Of themajor categories of land use, agricultural enterpriseoccupies the greatest land area and has the mostpotential to produce usable habitat for bobwhite. Theprimary objective of most agricultural land ownersand land users is tomanipulate that land in such a way


45

as to produce an income sufficient to support them-

selves and their families.Fortunately,bobwhite habitat

and profitable, sustainable agricultural land use arenot mutually exclusive. The preceding few paragraphsdescribed some of the theoretical considerations ofrestoring and maintaining habitat for bobwhite. Thenext section describes, in a more practical way, justhow this can be accomplished in today’s landscapes.

Private LandsBecause bobwhite likely occur in nearly every

county in Illinois, landowners could theoreticallyattempt habitat restoration anywhere in the State.However, probability of success would be much high-er in the milder climates of central and southernIllinois where bobwhite are more uniformly distrib-uted over the landscape. Tracts isolated from existingpopulations may be hard to improve. A covey or 2might get started, but if they were eliminated by asevere winter storm there would be no opportunity fornew birds to move onto the site. It is nearly impossi-ble to start from nothing and build a sustainable bob-white population unless birds are able to move fromsurrounding areas to the tract selected for habitatimprovement. Furthermore, bobwhite management ismuch more likely to succeed in landscapes alreadycontaining a suitable mix of woody edge and openfields as opposed to vast unbroken tracts of croplandor closed canopy forests. In other words, start byselecting a suitable area for management. Do notattempt to turn a forest or wetland into habitat forbobwhite.

Another important consideration is theprimary objective of the land. Most often the produc-tion of grain, forage, or timber is the reason for landownership, although a growing number of acres arebeing purchased for recreational use including hunt-ing for bobwhite. The primary ownership objectivesdetermine if habitat restoration for bobwhite is feasi-ble and the amount of land that can be devoted tomanagement. Many guides to management state that20 to 40 acres of habitat will support 1 or 2 coveys ofbobwhite. This should not be interpreted as meaning

Habitat Restoration

Bobwhite management is much more likely to suc-ceed in landscapes featuring a mix of woody edgeand small, open fields (top) as opposed to unbrokentracts of cropland (middle) or forest (bottom)

46

that 20 to 40 acres of farmland must be retired fromproduction for each covey desired by the landowner.Bobwhite will utilize many of the plant communitiesexisting on the farm. In some cases, habitat restora-tion may be accomplished without removing a singleacre from income producing uses. Often all that isneeded are minor adjustments in farming practices ortiming of agricultural activities.

As an example, consider an average 300-acre farmin southcentral Illinois with 160 acres of row crops, 30acres of small grains, 10 acres of permanent pasture,20 acres of rotation pasture, 20 acres of hay, 15 acres ofCRP grassland, and 40 acres of woodland plus 5 acresoccupied by the farmstead, fencerows, ravines, andfield roads. Land in row crops (corn, soybean, grainsorghum) will provide shelter and escape cover duringthe growing season and a source of waste grain andweed seeds after harvest if a no-till or reduced tillagesystem is used. The winter food source could beenhanced by deferring harvest of a few rows of grainadjacent to woodlots or shrubby fencerows untilspring. Small grain fields also provide shelter andescape cover during the growing season and a sourceof insects for foraging broods. After harvest, untilledstubble provides waste grain, weed seeds, insect for-age and roost cover. Permanent, monotypic fescuepastures provide little or no usable habitat for bob-white. Such pastures can be improved as forage andbrood habitat by interseeding with leguminous foragespecies such as red clover or ladino clover. Rotationpastures provide an additional source of insects forbroods even if hayed or grazed. Legume hayfields alsoprovide brood foraging areas after harvest, but fescuehayfields are of no value to bobwhite. ConservationReserve Program grassland will provide nest cover forbobwhite if planted to native warm season grasses andforbs or thin stands of fine-stemmed cool seasongrasses and legumes such as redtop and Korean les-pedeza. Woodlands and fencerows with brushyunderstory will provide shelter from weather andpredators.

From this example, it is obvious that the“buildingblocks”of bobwhite habitat are present onmany farmsin Illinois. Small adjustments in agricultural opera-tions are usually all that is needed to significantlyincrease habitat for bobwhite on the farm. If land

must be retired from crop production, it is often eligi-ble for CRP which will compensate the landowner forincome lost from the cessation of farming. Costs ofplanting grasses, forbs, and shrubs will be shared bythe U.S. Department of Agriculture, the IllinoisDepartment of Natural Resources, or conservationgroups such as Quail Unlimited (QU). The importantmessage is that habitat restoration for bobwhitecan be both economical and compatible with nor-mal farming operations in most regions of Illinois.

Perhaps you are fortunate enough to own landthat can be devoted completely to habitat restoration.What should you do? You might start by subdividinglarge blocks of open land into fields 10 to 20 acres insize. Field borders should be designed to provide acombination of shrub habitat, native warm seasongrass/forb nest cover, and legume brood foraginghabitat. For example, the center could contain a dou-ble row of native shrubs 10 feet wide. On either side ofthe shrubs, there could be a 10-foot wide strip ofclover or Korean lespedeza which, if mowed in Augustof each year,would provide brood foraging habitat anda protective firebreak for the center strip of shrubs.The outside edges of the field border could be 35-footwide strips of nesting cover in the form of nativewarm season grasses and forbs. Such a field borderwould be 100 feet wide and provide 12 acres of habitatper mile (1 acre per 435 feet). If agricultural fields arefarmed in continuous row crops, the cropping systemshould be changed to a rotation of corn or grainsorghum followed by oats or wheat followed by cloveror lespedeza for 2 years. If legumes are harvested forhay, 50-foot borders should be left unharvested forescape cover for broods. About half of the remainderof the field could be harvested alternating harvestedstrips with unharvested strips. Areas of permanentpasture should be converted to a mix of native warmseason grasses and forbs.After the planting is established, about 1/3 of theacres should be burned each year in March or discedin October if burning is not feasible. Woodlots shouldbe thinned to at least 50% canopy closure, removingshade tolerant species such asmaple and ash. Existingfencerows should be allowed to grow up in shrubs,vines and briars, and strips of lespedeza and nativewarm season grasses, and forbs should be planted on


each side of already established fencerows as previ-ously described. Large trees should be removed fromexisting fencerows.

Be sure to inventory the plant communities on theparcel selected for management. The “buildingblocks”of bobwhite habitat are woodland and woodededges, grassland (in a variety of growth forms andstructures), and cropland or early succession idle landthat provides a source of plant and insect foods. All 3“building blocks”must be present. If any 1 is missing,the parcel is not bobwhite habitat. Also take note ofany useful plant communities on adjacent lands. Ifclose enough, bobwhite may utilize habitat on neigh-boring lands making it unnecessary to duplicate thehabitat on the parcel proposed for management. Theimportance of inventorying existing plant communi-ties cannot be overstated. It is not unusual to see amilo food plot planted on the edge of a corn field.Wouldn’t it have been more economical to leave a fewrows of corn unharvested at the edge of the field?Following is a discussion of the “building blocks” ofhabitat for bobwhite.

Woodlands and wooded edgesIn terms of plant communities, the presence of

woodlands and wooded edges seems to be the mostimportant in determining the distribution of bob-white in Illinois. Quail are not forest dwellers, but theyseldom stray far from forest edges,wooded ravines, orshrubby fencerows. Woodlands and wooded edgesprovide shelter from severe weather and escape coverfrom predators as bobwhite move about the landscapeto feed,nest, roost, and raise their broods. These plantcommunities are vital, but they should not occupy alarge portion of the area available for management.For winter cover in Illinois, a 2 to 5 acre woodlotshould be adequate.As previously mentioned,woodlots should be open incharacter (50% or less canopy closure) with a groundcover of grasses, forbs, and shrubs. The common mixof upland hardwood species (oaks and hickories) issuitable.

Many woodlots and forests in Illinois are over-stocked with large, shade tolerant tree species such ashard maple, ash, and hackberry as a result of manyyears of fire suppression. Due to heavy shading, these

forests usually have sparse, poorly developed shrublayers and ground cover, the very zones most impor-tant to bobwhite. Forests in this condition may begreatly improved by thinning. Species such as ash andmaple should be removed in the thinning process.Reducing the canopy to 50% coverage or less willallow development of shrubs, grasses, and forbs nec-essary to create wintering areas for bobwhite. In someinstances, planting native warm season grasses andforbs adapted to open forest or savanna conditionssuch as little bluestem (1 pound of pure liveseed/acre), sideoats grama (1 pound of pure live seed/acre), and partridge pea (2 pounds of pure live seed/acre) may be desirable (plant in April or May with ano-till native grass drill). In other situations, the for-est floor may already contain a seed bank of desirable

Woodlots must have suitable understory (top) to beconsidered potential quail habitat. Closed canopyforests with little or no ground cover (bottom) are of novalue to bobwhite.

47

48

plants adequate to re-establish ground cover naturally.Many farms are comprised almost entirely of croplandand woodlots. Thinning woodlands on these farmsmay be the best way to create additional habitat forbobwhite.

When woodlands are returned to the desirablecondition of scattered large oaks and hickories withan understory of shrubs, grasses and forbs, some formof periodic disturbance will be necessary to maintainthis ecological stage. Prescribed burning in lateFebruary or early March every 3rd year should be ade-quate. Burn only 1/3 of the area each year to leaveresidual cover for nesting and shelter. Discing ormowing in October may be substituted for prescribedburning. Rotate treatments so that only 1/3 of the areais treated each year.

Wooded areas utilized for shelter and escape cover

must be connected to cropland and grassland by fieldborders of shrubs that provide overhead protectionwith an open understory to facilitate movement bybobwhite. Many species of shrubs are suitable forestablishing field borders. Some of the common“fencerow” species include crab apple, hawthorn, wildplum, gray dogwood, silky dogwood, and hazelnut.Shrub field borders are usually established by plantingsmall, bare root seedlings in single or double rows ona 5-foot by 5-foot spacing in March or April. In sever-al years, shrub rows will begin to widen as additionalstems grow up from the roots. Shrub clumps provideexcellent resting areas for coveys of bobwhite. Shrubfield borders are preferable to large trees because theyare low growing (10 to15 feet tall) and less competitivewith adjacent crops for moisture and nutrients. Inaddition, they do not provide hunting perches foravian predators. Shrub field borders require somemaintenance. Periodically inspect them and cut outseedlings and saplings of large trees such as oak, ash,hackberry, mulberry, and osage orange.Also important, is the distribution of wooded edge. Itshould be adjacent to every field that contains food orcover that bobwhite need to use. Research in Illinoisfound few bobwhites present in areas with less than 40feet per acre of wooded edge (100). On a larger scale,that translates to 1,600 feet per 40 acres, a little morethan one side of a square 40-acre field.

Agricultural grasslandsThe distribution of bobwhite is primarily depend-

ent on woody cover, but their abundance seems main-ly determined by the amount and quality of nest coverand brood habitat. Bobwhite are a short-lived specieswith an annual mortality rate of 75 to 80%; therefore,long-term survival is predicated upon high rates ofp r o d u c t i v i t y .Without the plant communities necessary for success-ful nesting and brood rearing, bobwhite cannot per-sist. Some agricultural grasslands have the potentialto provide safe nest cover while others are better suit-ed to providing brood foraging areas. Most of Illinois’bobwhite range is deficient in both.

Portions of agricultural landscapes that havepotential to provide nest cover include grass field bor-ders, roadsides, pasture (active and idle), and fields,


Prescribed burning is an excellent tool for maintainingquality grass/shrub cover in open woodlands.

Shrubby fencerows allow bobwhites access to feedingand nesting areas (the tall trees in this example shouldbe removed).

contour buffers and filter strips enrolled in theConservation Reserve Program. Nest cover shouldoccupy 20 to 30% of the farm because nest densitiesseldom exceed 0.5 to 1 nest per acre, and hatching suc-cess seldom reaches 50% (64). It is encouraging thathighest nest densities are usually found in strip coversuch as field borders and roadsides (64) because stripcover is far more common in modern agriculturallandscapes than large grass fields. To ensure maxi-mum nest production, nest cover should remainundisturbed from April 1 to August 1.

Unfortunately, very few existing grasslands pro-vide nest cover suitable for bobwhite. Grass must havethe proper density and structure or it cannot be used.The most common problems are excessive thickness,lack of diversity, and disturbance (mowing, haying,grazing) during the nesting season. Dense,monotyp-ic stands of fescue, smooth brome, or reed canarygrass simply cannot be used by bobwhite. Fescuefields in CRP that are mowed like lawns every year areof no value to bobwhite. Stands such as these must bekilled before more desirable species can be estab-lished. To convert existing fescue sods in strips or

fields, it is necessary to kill the sod with herbicides(usually fall and spring applications of Roundup® or afall application of Roundup® and a spring applicationof Plateau®). Spraying should be preceded by closemowing. Allow the stand to grow a few inches beforespraying to improve uptake of the herbicide.Following herbicide treatments, a light seeding ofnative warm season grasses and forbs such as littlebluestem (2 pounds of pure live seed/acre), sideoatsgrama (2 pounds of pure live seed/acre), and forb mix(1 pound of pure live seed/acre) should be plantedwith a no-till native grass drill in April or May.

Once established, the stand should be burnedonce every 3 years in March to maintain the properdensity and structural diversity. Remember to treatno more than 1/3 of the nest cover each year.Bobwhite build their nests from residual plant materi-als from the previous year’s growth. Stands of nestcover may be disced in October if burning is not feasi-ble. A few species of introduced grasses and legumessuch as redtop (1 pound of pure live seed/acre) andlespedeza (3 pounds of pure live seed/acre) also canprovide suitable nest cover. Planting should be done

Grasslands must have proper density and structure to be used by quail. A mix of bunch grasses and forbs (top) isideal, but dense monocultural sods (bottom) are of little or no value.

49

in March orApril. Disturbance is also necessary every3rd year in the form of light discing in October.

Larger fields (>10 acres) of native warm seasongrasses and forbs or introduced grasses and legumesshould be subdivided into 3 equal sized units by plant-ing 30 to 50-foot wide strips of legumes (clover, les-pedeza or alfalfa) to serve as firebreaks when pre-

scribed burning is used to maintain fields. Strips alsoprovide brood foraging habitat. In another variation,the strips may be used as foodplots as well as fire-breaks. The strip should be disced in October beforea burn the following March. After the burn, corn ormilo may be planted in the firebreak. The followingspring in February or March, lespedeza may be broad-cast onthe corn or milo residue. The plot should be leftundisturbed during the 3rd spring to permit thegrowth of annuals and lespedeza. Fields, field borders,

and filter strips in CRP may be established and man-aged in a similar fashion. For CRP grasslands, a writ-ten plan will be prepared by U.S. Department ofAgriculture (USDA) Natural Resource ConservationService (NRCS) or Illinois Department of NaturalResources (IDNR) personnel for each field under con-tract. Be sure to inform the individual preparing theplan that your objective is to provide the best possiblenest cover for bobwhite. Active pastures may providesome additional nest cover if 30 to 50-foot wide stripsof native warm season grasses and forbs (littlebluestem, sideoats grama, and broad leaf mix) areplanted around the edges of grazing lands. A 40-acrepasture with 50-foot borders on all sides would pro-

vide 6 acres of nest cover. Prescribed burning, neces-sary to maintain the grass/forb stand, would alsoreduce the invasion of pasture edges by trees andshrubs.

Brood habitat may be similar to nest cover but it is


50

Herbicides must be used to kill fescue (top) beforewarm-season grasses can be successfully establishedusing specialized grass drills (bottom)

Legume fields (top) provide an excellent source ofinsect foods for bobwhite chicks (bottom).

51

usually thinner with more broad-leafed plants,legumes and annual grasses, and less perennial grass.A number of common crops can provide brood forag-ing areas with little or no modification. Hayfields ofalfalfa, clover, or lespedeza usually support high insectpopulations. After cutting, stands are usually openedto the point where chicks can move through the fieldand locate insect foods. Small grain fields after har-vest also provide suitable brood habitat, especiallywhen seeded with a companion crop of red clover.Even soybeans, no-till planted into wheat stubble, canprovide some brood foraging areas. In larger CRPgrasslands, brood habitat can be created by discingstrips in the grass stand in October. Disced stripsshould be about 50 to 75 feet wide and cover about30% of the field. Discing should be no deeper than3 inches and should leave 50% of the ground coveredby plant residue. Untreated areas of grass andlegumes twice as wide as the treated areas (100 to 150feet) should be left between disced strips for nestingcover. Each year discing should occur on the stripsthat werenot disced the year before. If desired, Korean les-pedeza may be broadcast on the disced strips in latewinter at a rate of 5 lbs/acre. In 3 years, the entire fieldshould have been covered.

CroplandIn Illinois, row crops (corn, soybeans, and milo)

and the weed seeds associated with their production

are the primary sources of plant foods for bobwhite.During the summer, row crops also provide consider-able shelter from severe weather and predators. Withcurrent farming practices, there is an excess of row

crops in most landscapes, but they are under-repre-sented in some areas of the Shawnee Hills where for-est and pasture have replaced general farms. Somecounties with high enrollments of CRP have areaswhich may be deficient in plant foods because of theretirement of row crop fields. The best way to managecropfields to provide over-winter food for bobwhite isto leave a few rows of unharvested grain next to wood-lands or wooded fencerows. Leaving an undisturbedfallow strip at least 30 feet wide between crops andprotective woody cover also is very beneficial. Finally,adopt a no-till farming system if possible. If no-till isnot feasible, use conservation tillage or at the veryleast, avoid fall plowing.

Where row crops are unavailable, establish a sys-

Strip discing is an excellent way to restore legumes andannual seed producers in fields with dense grass sods.

Leaving a few rows of unharvested crops next to securewinter cover is the best way to provide an emergencyfood supply for bobwhite,.

Where access to grain fields is limited, it is importantto plant annual foodplots adjacent to good wintercover.

52

tem of rotated foodplots on the best soils available.Five percent of individual CRP fields may be plantedto foodplots, which is usually adequate if care is exer-cised in locating the foodplots. Use foodplots to dividefields into 3rds for prescribed burning or discing aspreviously described.Always plant foodplots on contour in fields with >5%slopes. Planting rates vary by method of planting.When a corn planter is used, plant seed at a rate of20,000 per acre. Milo should be drilled at 10 poundsper acre. Use higher rates if seed is broadcast and cov-ered with a disc or harrow. Apply a balanced fertilizerat a medium rate and plant in April or May.Sources of assistance

Assistance is available for restoration of bobwhitehabitat. The NRCS can provide technical assistanceand financial assistance for planning and implement-ing habitat restoration for bobwhite. The FarmServices Agency is the unit of the USDA that operatesthe CRP. This program provides both land rental andcost sharing for habitat development. Offices of bothUSDA agencies are usually found at a common loca-tion in each county. You should contact the countyoffice in which your property is located. The IDNRDivision of Wildlife Resources also provides technicalassistance, limited free plant materials, and some cost

sharing. District Wildlife Biologists usually serve a 3or 4-county district.You can locate your district biologist by calling theDivision of Wildlife Resources office in Springfield(217) 782-6384.

Assistance also is available from QU, a privateorganization devoted to the conservation of quail

throughout North America. Currently, there are 18active chapters located throughout westcentral, south-central, and southern Illinois. These groups may alsoprovide some financial assistance with bobwhite habi-tat restoration projects. To locate the chapter servingyour area call (812) 536-2272.

Public LandsThe IDNR currently owns or leases approximately

356,000 acres at 391 sites of which 116 (30%) offerpublic hunting for bobwhite. Unfortunately, popula-tion trends for bobwhite on Illinois’ public wildlifemanagement areas are very similar to those on privateland over the last 30 years. With a few exceptions,management efforts on public land in Illinois havebeen largely unsuccessful in establishing and main-taining high numbers of bobwhite. Research biolo-gists and wildlife managers believe this lack of successis attributable to the inability to maintain large por-tions of sites in the early successional habitatsrequired by bobwhite for nesting, brood foraging, androosting. Establishment and maintenance of thesehabitats require intensive application of a variety ofpractices including native warm season grass/forbplanting, prescribed burning, strip discing, establish-ment of wooded field borders to provide corridors forsafe movement, as well as chemical and mechanicalbrush control in oldfields. Cultivated land with anappropriate crop rotation (row crops, small grains,legumes) also is an important component of bobwhitehabitat. Cropping systems that contain only row crops


The Illinois Department of Natural Resources providesassistance in restoring habitat for bobwhite.

Large amounts of early successional vegetation are nec-essary for successful bobwhite reproduction.

53

provide a very limited spectrum of benefits to bob-white.

In the mid-1960s, Illinois researchers (27) con-ducted a classic habitat improvement demonstrationon Sam Dale Lake Conservation Area in WayneCounty and Stephen A. Forbes State Park in MarionCounty. High quail populations and sustained har-vests were maintained through a program of herba-ceous vegetation management that featured crop rota-tion via sharecropping and prescribed burning.Despite the success of this approach, most manage-ment efforts on Illinois public lands continued tofocus on increasing escape cover and over-winter foodsupplies. Management to provide herbaceous habitatsessential for reproduction has been largely ignoredresulting in grasslands on public areas that are often oflittle or no value to bobwhite. Most consist of thickstands of introduced species such as smooth brome,fescue, or reed canary grass. Even where native warmseason grasses have been established, the stands areoften too thick for bobwhite because they were plant-ed at high seeding rates and without forbs or legumes.These grasslands generally receive little or no distur-bance after planting in the form of burning, discing orherbicide treatments to reduce grass density, exposebare soil and increase forbs and legumes. Ideally, 30%of each unit of grassland should receive appropriatelytimed (fall discing or spring burning) disturbanceeach year.

For bobwhite, site management objectives shouldbe to increase and better distribute nest cover, broodhabitat, shelter/escape cover, and winter food supplies

over as much of the site as is feasible. Habitatimprovement should focus on renovation of largeblocks of grass sods and better utilization of croplandon site by dividing large fields (>40 acres) with shrubfield borders and adopting a crop rotation includingcorn or milo, oats or wheat, and 2 years of legumesthat are unharvested or partially harvested. A fewrows of corn or milo should be left standing adjacentto all woodland edges and shrub field borders. Iflegumes are harvested, a 50-foot border should be leftaround all field edges. In addition, half of the hayfieldcould be harvested by alternating 50-foot wide har-vested strips with 50-foot wide unharvested strips.

Three IDNR sites currently have relatively suc-cessful bobwhite management programs. These arethe Jim Edgar Panther Creek State Fish and WildlifeArea in Cass County, Pyramid State Park in PerryCounty, and Ten Mile Creek State Fish and WildlifeArea in Hamilton County. These sites have at least 3things in common: relatively large size (5,800 - 19,400

acres), an emphasis on management for nesting andbrood rearing habitat, and hunting programs thatlimit pressure to some degree. The latter factor isimportant because the abundance of bobwhite onpublic land may be adversely affected by excessivehunting pressure. Because many public hunting areasexist as islands of suitable habitat in a matrix of rowcrops, there is little opportunity for repopulation ofexcessively exploited sites through ingress from sur-

Without periodic disturbance of grasslands, woodyinvasion will ultimately eliminate its value as nestingand brood rearing habitat.

Edges of legume fields adjacent to protective cover pro-vide excellent brood habitat for bobwhite.

54

rounding areas. In most cases, sites are available forbobwhite hunting throughout the season by unlimitednumbers of hunters. Hunting pressure undoubtedlyexceeds levels consistent with maximum sustainedyield of bobwhite at most sites. Depending on size ofthe area, the number of hunting parties per day shouldbe limited to perhaps 1 party of 4 per 300 acres ofopen land. In addition, the number of days open to

hunts should be limited to 2 or 3 per week.The Shawnee National Forest, managed by the

U.S. Forest Service, has exciting potential for restora-tion of bobwhite habitat if management efforts can beupdated and improved. On many national forestsincluding the Shawnee, decades of fire suppression,planting of exotics, and severely limited timber har-vest has resulted in a buildup of fuel similar to those


Native warm season grass field borders provide excellent nesting areas for bobwhite.

55

that led to recent catastrophic wildfires in the westernUnited States. Currently, the Forest Service is reexam-ining its policies and prescribed fire and increasedharvesting are being considered to reduce fuel loadson national forests. These activites have the potentialto restore thousands of acres of habitat for bobwhite.Thinning or eliminating stands of exotic pines plant-ed in the past would provide large openings usable bybobwhite. Prescribed burning would stimulate rem-nant stands of native warm season grasses and forbsand oaks and hickories known to have occupied theIllinois Ozarks at the time Illinois was settled.

Humans dominate the landscape in a populous,agricultural state like Illinois, and how we humans usethe land will ultimately determine the bobwhite’sfuture. To be honest, this future does not look partic-ularly bright at the present time. To suggest otherwisewould be to ignore current facts and trends. As weearlier noted, if downward population trends contin-ue, bobwhites may be all but eliminated from theSoutheast and Midwest in the next 30-50 years. Thisis a sobering projection to be sure, but is it inevitable,or can it be reversed, or at least slowed?

To address these questions,we must first examineprojected trends in those factors that are contributingto the bobwhite’s decline.

One trend that certainly does not favor bobwhiteis the continued growth and expansion of the humanpopulation, coupled with our seemingly insatiabledemands for more and more living space (94).Illinois has lost nearly 2.7 million acres of potentialquail habitat to non-agricultural uses over the past 40years and this trend can only be expected to continue.Quail obviously cannot use land covered by concreteor asphalt, but vegetated suburbs also are usuallyinhospitable to quail. Even where usable habitatremains near expanding developments, huntingopportunities are often lost.

Despite continued conversion of rural land tourban uses, Illinois’ land base remains predominant-ly agricultural, and the bobwhite’s fate will mainlydepend on how this land is farmed (9). Future farm-

ing practices and agricultural policies, in turn, willdepend on a number of factors including global eco-nomics, federal policies, technological advances, andpossibly even climate change. Although changes inagriculture are almost inevitable, the nature of thesechanges, and their potential effect on bobwhite arevery speculative (19). As the proportion of row cropsincreases, the quality of bobwhite habitat generallydeclines, although an increase in row crop productionmight actually benefit quail in the more heavily wood-ed, less intensively cultivated portions of southernIllinois. Quail in our State seem to prefer landscapeswith about 30-65% rowcrops. It is hard to envisionscenarios that would reduce the intensity of agricul-ture to this level on the flat, highly fertile portions ofthe State. Thus, the bobwhite’s potential future will bedetermined by what happens on farmland of interme-diate to low quality on slightly rolling topography (i.e.,the current quail range; Fig. 2). Undoubtedly, thegreatest pressure for increased row crop productionwill come from an ever-expanding human population.As the world population continues to grow, and lessdeveloped nations continue striving to raise theirstandard of living which often includes increasing theamount of red meat in their diets, there will beincreased demand for feed grains such as corn, soy-beans, and grain sorghum. Expanded use of alterna-tive fuels such as ethanol also could stimulateincreased corn production, although perennial plantssuch as native warm season grasses could be used forbiomass production of ethanol without the expense ofannual planting.

There are other factors, however, that might workto reduce corn/soybean acreage, at least on less pro-ductive areas of the state. For example, an increasingnumber of foreign countries are beginning to exportthese products and there is growing opposition fromthese producers to the U.S. government subsidizinggrain production for the export market. Future trendstoward larger, corporate farms also might reduce agri-cultural use of “marginal” lands. The opposite trendtoward more diversified, “sustainable” agriculturecould have the same effect (19). The latter could be

The Future

56

promising for bobwhite, especially if it meant return-ing to such quail-friendly practices as crop rotationthat included small grains, legumes and regular, peri-odic short-term fallowing. However, if fescue orsomething similar continues to dominate agriculturalgrasslands, then benefits would be minimized.

There will continue to be scientific advances lead-ing to higher yields and more efficient production ofcrops. Just how this will impact bobwhite is unclear.Conceivably, better efficiency could allow greater pro-duction on the same or less acreage; conversely, itcould result in even more acres devoted to agriculture(19). Likewise, increased development and use ofgenetically modified crops (e.g., Bt corn and RoundupReady soybeans) could potentially benefit quail byreducing use of insecticides and herbicides. However,such advances could also promote additional grainproduction which might not always favor bobwhite.

Another factor that could impact quail directly orindirectly is climate change. Several biologists havealready speculated on the potential biological effectsof global warming on bobwhite distribution andabundance (43), but perhaps the greatest impact onquail would come from land use and agriculturalchanges resulting from altered climatological patterns(91).

The environmental/land use changes we havebeen talking about could conceivably affect quail overthe next 20-50 years or so. More important for theimmediate future are the kinds of federal agriculturalprograms that are initiated or maintained. As we havenoted, land-retirement programs such as CRP canbenefit bobwhite, but only if there are specific provi-sions designed to maintain early successional vegeta-tion. Recently added, potentially quail-friendly CRPpractices such as strip discing, controlled burning,field borders, and incentives for mid-contract man-agement are encouraging. However, CRP regulationsare subject to political pressures and will require con-tinued effort by quail enthusiasts to promote benefi-cial practices.

This leads us to the troubling question of just whowill speak for quail interests in the future? Census fig-ures tell us that the United States population is becom-

ing increasingly more urban and less rural. And whilethere may be a superficial increase in public aware-ness and concern about conservation issues (12), thenumber of people with direct experience or knowl-edge of bobwhite and their habitat needs is declining(19). The traditional base of public support forwildlife conservation and management has comefrom hunters. However, the proportion of UnitedStates citizens that hunt is dwindling, with the trendespecially pronounced among upland game hunters.In Illinois, for example, the number of quail huntershas declined from >155,000 in the late 1960s to<35,000 currently. Equally troubling is the fact thatuniversity students, our future wildlife managers andresearchers, increasingly do not hunt and thus willhave less interest in game bird management and hunt-ing issues (19). How bobwhites will fare in the emerg-ing “ecosystem/biodiversity” conservation agendas ofthe future is not yet clear (93).

On the positive side, organizations such as QUremain popular and active. Currently, QU has approx-imately 300 chapters and 48,000 members nationwide(18 chapters and 1,900 members in Illinois). Theseorganizations are not only actively involved in habitatrestoration, but provide a base of public support forstate and federal programs affecting quail (120). Alsoencouraging is the fact that there is now in place anationwide plan for quail population recovery calledThe Northern Bobwhite Conservation Initiative (25).This plan seeks to promote cooperation between pri-vate land owners, state and federal agencies, and otherconservation organizations for the purpose of restor-ing bobwhite habitat on a landscape scale throughoutthe species’ range. Plans for implementing suchefforts in Illinois are currently being developed (16).

Despite a rather pessimistic outlook by some (41,94), the future can still provide opportunities toincrease or at least stabilize bobwhite abundance inIllinois. But unlike the past, it will require strong,focused,well-directed effort instead of benign neglect.We cannot help bobwhite if we are unwilling to recog-nize the reasons for population declines or the actionsnecessary to reverse these declines. We must stopblaming predators, turkeys, or the closure of game


57

Photo Credits

Adele Hodde – page 6 (left,right)

Dave Howell – page 18 (bottom); page 19 (bottom right); page 21 (center left); page 22 (bottom); page 32 (bottom left,bottom right), page 48 (bottom); page 50 (top left, bottom left, top right); page 51 (left); page 52 (right); page 53(right); page 54 (left, right)

John Roseberry – page 12; page 17 (top,bottom); page 18 (top); page 31 (top); page 49 (top left, top right, bottom right); backcover

Paul Willms – page 10; page 11 (top, bottom); page 21 (top left, bottom left, bottom right); page 22 (center); page 30 (top);page 31 (bottom); page 35 (top); page 40; page 45 (top); page 47 (top); page 48(top); page 51 (top right, bottomright); page 53 (left)

Scott Yaich – front cover

CooperativeWildlife Research Laboratory, SIUC – page 8; page 19 (top left, top right); page 21 (top right, center right); page22 (top); page 30 (bottom); page 32 (top left, top right, center right); page 35 (bottom); page 45 (center, bottom);page 47 (bottom); page 49 (bottom left)

Illinois Department of Natural Resources – author page (left, right), page 52 (left)

Nebraska Game & Parks Division – page 19 (bottom left)

Virginia Department of Game and Inland Fisheries – page 50 (bottom right)

The bobwhite wing drawings on page 58, by Richard Parks, were reproduced with the kind permission of James M. Rosene.

59

Juvenile with molt in progress

Adult with molt in progress Adult with molt complete

Juvenile with molt complete

Equal opportunity to participate in programs of the Illinois Department of Natural Resources (IDNR) and those funded by the U.S. Fish andWildlife Service and other agencies is available to all individuals regardless of race, sex, national origin, disability, age, religion or other non-meritfactors. If you believe you have been discriminated against, contact the funding source’s civil rights office and/or the Equal EmploymentOpportunity Officer, IDNR, One Natural ResourcesWay, Springfield, Ill. 62702-1271; 217/785-0067; TTY 217/782-9175.

Printed by the authority of the State of IllinoisPRT4031234 - 5M - 6/06

IISG400

the bobwhite in illinois:

Documents