North Central Forest Experiment StationForest Service - U.S. Department of Agriculture

1992 Folwell AvenueSt. Paul, Minnesota 55108

Published in 1997

General Technical ReportNC-195

Management of Early-Successional Communities inCentral Hardwood ForestsWith Special Emphasis on the Ecology and Managementof Oaks, Ruffed Grouse, and Forest Songbirds

Frank R. Thompson, III and Daniel R. Dessecker

TABLE OF CONTENTS

PageCHAPTER 1—INTRODUCTION ............................................................. 1

Basic Principles and Terms ...................................................... 3

CHAPTER 2—CENTRAL HARDWOOD FORESTS .................................. 4History and Ecology ................................................................. 5The Regeneration Ecology of Oaks ............................................ 6Oak Silviculture ....................................................................... 7

Even- versus uneven-aged management ......................... 7Clearcut method ............................................................ 7The shelterwood method ................................................... 7The seed tree method ..................................................... 8Selection methods ............................................................. 8Alternative methods .......................................................... 9

CHAPTER 3—HABITAT AND ECOLOGY OF THE RUFFED GROUSE .. 10Distribution ........................................................................... 10Status in Central Hardwood Region ........................................ 10Habitat Use and Annual Cycle ................................................ 11

Year-round adult cover ................................................. 12Nesting cover ................................................................... 14Brood cover ..................................................................... 14

Grouse Foods ......................................................................... 14

CHAPTER 4—EARLY-SUCCESSIONAL FOREST SONGBIRDS ............ 15Habitat Associations of Early-successional Forest Songbirds .. 15Residual Structure in Regenerating Stands ............................ 18Habitat Patch Size .................................................................. 18Importance of Landscape Context ........................................... 18

CHAPTER 5—FOREST MANAGEMENT PRACTICES FOREARLY-SUCCESSIONAL WILDLIFE ................................................... 19

Importance of Extensively Forested Landscapes ..................... 19Availability of Habitats Through Time ..................................... 19Habitat Patch Size .................................................................. 22Arrangement of Habitat Patches ............................................. 23Regeneration Methods ............................................................ 23Retaining Trees and Snags in Regeneration Cuts .................... 23Soft and Hard Mast Production .............................................. 25Consideration of Site Characteristics ...................................... 26Management of Early-successional Habitats Other Than Regeneration Forest ............................................................ 26

Diversity of Early-successional Communities ................. 26Old-Field Habitats ....................................................... 26Herbaceous Openings ..................................................... 27Conifers ...................................................................... 27

ACKNOWLEDGMENTS ...................................................................... 27

LITERATURE CITED .......................................................................... 27

CHAPTER 1—INTRODUCTION

In this paper we describe the ecology andmanagement of young forest communities in theCentral Hardwood Region. We refer to theseyoung forest communities as early-successionalforest because they are in the early stages offorest regeneration or growth following a distur-bance. We focus on early-successional specieshere for several reasons. In general there isconcern about the status of forest wildlife ineastern forests because of the loss, fragmenta-tion, or change in forest habitats as a result ofland-use practices such as land development,agriculture, and forest management. Some ofthe forest-dwelling species that are decliningspecialize in early-successional forest habitats.These include the ruffed grouse (chapter 3),several neotropical migratory songbirds (chapter4), and the American woodcock (Bruggink andKendall 1995).

A second reason to be concerned about thesespecies is that their habitats are dependent ondisturbance, and humans have altered historicdisturbance patterns and created new ones.Early-successional forest habitats are transitoryor ephemeral because they change over time asa result of forest growth and succession. Thismeans the community is dependent on repeateddisturbances (a disturbance regime) to create ormaintain habitat. Disturbance from fire, wind,insects and disease, timber harvest, and otherland-use practices have been an important partof the history of central hardwood forests and to

MANAGEMENT OF EARLY-SUCCESSIONALCOMMUNITIES IN CENTRAL HARDWOOD

FORESTS

With Special Emphasis on the Ecology and Management of Oaks, RuffedGrouse, and Forest Songbirds.

Frank R. Thompson, III and Daniel R. Dessecker

Frank R. Thompson, III, is a Project Leaderwith the North Central Forest ExperimentStation, Forestry Sciences Laboratory, Colum-bia, MO.

Daniel R. Dessecker is a Wildlife Biologist withthe Ruffed Grouse Society, Rice Lake, WI.

a large extent have determined the compositionand structure of present-day forests. Withinthe last 200 years, there have been significantchanges in the disturbance patterns of theseforests due to changes in land-use and forestmanagement.

Additionally, an understanding of the ecologyand management of these species is neededbecause of significant public interest in bothconsumptive and non-consumptive uses. Thereis also significant public and private interest intimber harvest, one of the primary disturbancefactors that creates early-successional forest.In addition, forest resource management andthe harvest of forest products receive a greatdeal of public interest, particularly on publiclands. Our focus on management for early-successional forest communities should notdetract from the need for other habitats orcommunities. For example, there is greatconcern for some species that live in matureforest as well as great concern for the extentand distribution of old-growth forest in thisregion. We offer some approaches to integratingconflicting habitat needs such as early- andlate-successional forest in landscapes andregions.

We focus on oak-dominated forests within thearea defined as the central hardwood forest(Clark 1989) (fig. 1). This area is within thearea of the oak-hickory forest type as defined bythe Society of American Foresters (Eyre 1980)and the eastern-broadleaf forest province of theNational Hierarchical Framework of EcologicalUnits (McNab and Avers 1994). Although, ourspecific focus is on oak-hickory forest, much ofthe information presented applies to otherforests in the Eastern U.S. in which oaks are animportant component.

A focus on management for oaks in centralhardwood forests is important from both a

wildlife and silvicultural perspective. Oaks havea fundamental role in central hardwood wildlifecommunities. Acorns are at the base of acomplex ecological web that affects the regen-eration and abundance of oaks, the abundanceof mast-consuming wildlife, the predators andparasites of mast-consuming species, and theabundance of defoliators and decomposers of

Figure 1.—Distribution of forest in the Eastern United States. Oak-dominated upland forest is shownas dark gray, all other forest as light gray. The extent of the Central Hardwood Region (asdefined by Clark 1989) is outlined in black.

oaks (Healy et al., in press). Acorns and otherseeds represent the most valuable and energy-rich plant food available to eastern wildlife inthe dormant season (Robbins 1993). Acornproduction is often 3 to 10 times greater thanbrowse production in oak forests (Rogers et al.1990, Segelquist and Green 1968). Oak hasincreased in importance as a food for eastern

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trees sufficiently uniform in species composi-tion, age structure, and condition to be distin-guished as a unit (Smith 1986). A landscapeconsists of a group or mosaic of habitat patchesand usually is a large area. Often landscapesare considered to be areas ranging from thou-sands to tens of thousands of acres. Forestmanagement is often regulated at scales re-ferred to as compartments, districts, or forests,which can often be considered landscapes.And, finally at a larger scale, we will discussregions. Regions are often defined geographi-cally, such as the Midwestern or SoutheasternU.S., or they are defined ecologically, such asthe Central Hardwood Region. A regional scaleis often important when talking about thegeographic distribution of a species or whencomparing patterns among landscapes.

For reasons already stated, landscapes are animportant scale at which to consider habitat forearly-successional forest wildlife. Habitatquality is affected by two general factors at alandscape scale, landscape composition andlandscape pattern.

Landscape composition is the amounts ofhabitats present in a landscape and can bethought of as habitat availability within alandscape. Forest managers often evaluatelandscape composition based on the distribu-tion of forest age classes. Landscape pattern isa result of the arrangement of habitats orspatial distribution in the landscape. Thespatial distribution of habitat patches in alandscape can affect the habitat quality of thosepatches or of the landscape as a whole. Land-scape pattern is determined by the amount,size, shape, and location of habitat patches.Another important component of landscapecomposition or habitat availability is temporalavailability. Ecological succession and distur-bances will cause landscape composition tochange through time so temporal availabilitymust be considered in habitat management.

Early-successional forest habitats are createdby natural or human-related disturbances.These disturbances, and the resulting habitat,create diversity or variety in a landscape. Adiverse landscape provides habitat for manydifferent species as well for species that requiremore than one type of habitat. Too muchhabitat diversity, however, can reduce thequality of some habitats for some wildlife spe-cies as habitat patches become small and

wildlife because of the elimination of Americanchestnut and decline in American beech. Afocus on oak management is important not onlybecause of its value to wildlife but also becauseof its dependence on disturbance for regenera-tion. Some form of active forest managementwill be required to maintain oak as an impor-tant component of future forests (Healy et al., inpress).

In chapter 2 we review the distribution, history,ecology, and silviculture of the central hard-wood forest. In chapters 3 and 4 we describeimportant aspects of the ecology and manage-ment of the ruffed grouse and early-succes-sional songbirds. In chapter 5 we suggest forestmanagement strategies for early-successionalcommunities within the context of maintainingthe native biological diversity of the CentralHardwood Region. In the remainder of theintroduction, we review relevant principles andterms of landscape ecology and forest manage-ment that are applicable to management ofearly-successional communities.

Basic Principles and Terms

Because of the ephemeral nature of early-successional habitats, their distribution in timeand space is important. Natural and human-induced disturbances and succession changethe availability and location of habitat for early-successional wildlife. This means that wildliferequirements need to be planned for over longtime periods and large areas. The distributionof regenerating oak stands is important from asilvicultural perspective as well. Sustainedyield of timber products and early-successionalhabitat is dependent on the frequency of timberharvest and its spatial distribution.

A habitat is an area with the appropriatecombination of resources (food, cover, water)and environmental conditions for the survivaland reproduction of a species. We refer toseveral spatial scales when discussing habitatsand land management planning. A habitatpatch is a contiguous block of habitat. Its sizeis relative based on the definition of habitat,which can be specific to a particular animal orplant. So, a habitat patch can be a group ofshrubs and saplings in a gap created by ablowdown or a 1,000-acre island of forestsurrounded by cropland. Habitat patches areoften considered analogous to stands. Standsare commonly defined as a contiguous group of

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fragmented. Habitat fragmentation is aprocess that results in increased habitat dis-continuity by breaking up blocks of habitat. Itranges from the creation of small disturbancepatches within a large block of habitat to wide-spread habitat loss resulting in small, isolatedpatches of habitat. Habitat fragmentation canhave positive or negative effects on wildlife,depending on the wildlife species of interest, theoverall landscape composition and structure,and the level or scale of fragmentation. Popula-tions in small isolated habitat patches or infragmented landscapes can be less viable thanthose in larger patches or more contiguoushabitat. Reasons for this include large arearequirements (home range), effects of smallpopulation size, and competition or predationfrom animals in adjoining habitats. Somespecies are considered area sensitive becausethey are not present in small patches of habitatdue to the effects of habitat fragmentation orbecause they avoid small habitat patches(Faaborg et al. 1995, Robinson and Wilcove1994).

Habitat fragmentation results in an increase inthe amount of edge. Edge is the border orecotone between adjacent habitats. Severalpatterns or processes, often referred to as edgeeffects, may occur at edges. Edge effects caninclude changes in animal and plant diversity orabundance, increased interactions amongspecies from adjoining habitats, (predators,competitors, parasites, and humans), andchanges in the micro-climate.

Forest fragmentation is a general term thatrefers to the fragmentation of forest habitats bynon-forest habitats. High levels of forest frag-mentation have negative consequences for manyforest wildlife species. Forest songbirds areoften absent from highly fragmented forests,and some species, while present in fragmentedlandscapes, have lower reproductive successthere. Although no studies have directly inves-tigated the effects of forest fragmentation onruffed grouse, these birds also presumablysuffer from the effects of fragmentation. Smallforest patches are not large enough to sustainpopulations of grouse, and grouse eggs andchicks are prey for many of the same predatorsthat depredate forest songbird nests in frag-mented landscapes. Also, grouse dispersing infragmented landscapes are likely to movethrough insecure cover and may be susceptibleto predation.

Forest habitat can be defined more finely basedon species requirements and can reflect differ-ences between successional stages, age classes,or forest types. Forest management activities,including the regeneration of forest stands,fragment forest habitats . Previously we dis-cussed fragmentation of forest by non-foresthabitats. Forest management practices main-tain forest habitat in general, but fragmentforest age classes, forest types, or habitats. Forexample, the use of regulated clearcutting willcreate patches of young forest mixed withmature forest, and forest habitats will be morefragmented than if the entire landscape werethe same age. Similarly, silvicultural practicescan change tree-species composition and foresttype, again fragmenting forest habitats. It isthis issue of habitat fragmentation resultingfrom the creation of early-successional foresthabitats that is often controversial with thepublic and land managers.

This type of habitat fragmentation creates foresthabitat diversity and can have positive andnegative consequences for forest wildlife. It haspositive effects for early-successional wildlife orspecies that require habitat diversity because itcreates patches of early-successional forestamidst older forest habitats. It has negativeeffects on late-successional wildlife because itresults in a loss of late-successional foresthabitat. Scientists have debated whether thislevel of habitat fragmentation results in some ofthe other negative consequences of forestfragmentation, such as edge effects. Mostevidence suggests the primary effect of this typeof habitat fragmentation is changes in theavailability of early- and late-successional forest(Thompson 1993, Thompson et al. 1996).Forest habitat fragmentation is generallyephemeral because of forest succession; forestfragmentation resulting from the conversion offorest to non-forest land uses is usually morepermanent.

CHAPTER 2—CENTRAL HARDWOODFORESTS

Oak-hickory forests cover approximately 127million acres (51 million ha) in the Eastern U.S.and make up 34 percent of eastern forests (fig.1). Approximately 82 percent of this forest landis owned by non-industrial private landowners,6 percent is in National Forests, 6 percent isheld by other public owners, and 6 percent isowned by the forest products industry (fig. 2)(Powell et al. 1993).4

Oak-hickory forests dominate the CentralHardwood Region but give way to mixed hard-woods in the north and in the AppalachianMountains, and to oak-pine forest to the south(Eyre 1980, Sander and Fischer 1989). Bottom-land hardwood forests extend into the region aswell. Dominant tree species in oak-hickoryforests are white, black, scarlet, and northernred oak. Other overstory trees are southernred, post, blackjack, chinkapin, bur, andnorthern pin oak; bitternut, mockernut, pignut,and shagbark hickory; black gum; yellowpoplar; red and sugar maple; white ash; elms;American beech; black walnut; and blackcherry. The most common understory trees orshrubs are flowering dogwood, sassafras,redbud, serviceberries, eastern hornbeam,American hornbeam, Witch-hazel, blueberries,mountain-laurel, rhododendron, and beakedhazel (Braun 1950, Eyre 1980, Sander andFischer 1989).

The oak-pine forest type is very similar to theoak-hickory type except shortleaf, loblolly,pitch, and Virginia pine make up 25 to 50percent of the forest. Mixed hardwoods aretypically found on moist, relatively productivesites primarily east of the Mississippi River.

Principal species are yellow poplar, white andnorthern red oak, and sugar maple. Manyother species are usually present includingwhite ash, black and chestnut oak, red maple,black gum, basswood, buckeyes, black walnut,black cherry, and hickories. Common under-story species include flowering dogwood, east-ern redbud, rhododendron, serviceberries,sourwood, and sassafras (Braun 1950, Eyre1980, Sander and Fischer 1989).

History and Ecology

The central hardwood forest has been changingsince the end of the last glaciation. Oaks,however, have dominated the region for the last6,000 years (Lorimer 1993). The rate of ecologi-cal change has accelerated since Europeansettlement. Deforestation and conversion oflands to agricultural land uses over the last 200years, the exclusion of fire during the 20thcentury, and the introduction of exotic diseasesand insects have greatly affected forest commu-nities (see review by Hicks 1997). The chestnutblight fungus eliminated American chestnutfrom eastern forests during the first half of thiscentury. American elm and American beechhave also been greatly reduced in abundance byinsects and pathogens (Healy et al., in press).Oaks have maintained their dominance over thelast 6,000 to 9,000 years, but they are nowlosing dominance in the overstory of manymesic (moist) sites whose understories areusually dominated by shade-tolerant species(Fralish et al. 1991).

Frequent and uncontrolled burning of oakforests ended less than a century ago in theMissouri Ozarks to several hundred years agoin the Eastern U.S. There is some debate aboutthe extent fire has affected oak forests(Beilmann and Brenner 1951, Steyermark1959). Some evidence suggests that historicdisturbance by Native Americans in the south-east was substantial (DeVivo 1991). Forestdisturbance by Native Americans throughoutEastern North America was greatly reducedafter their populations declined precipitouslyduring the 16th and 17th centuries (Hicks1997). In the Missouri Ozarks, fires occurredfrequently between 1785 and 1810 during aperiod of influx of Native Americans and Euro-pean explorers. This period was followed by aperiod of decreased fire frequency beginningwith an exodus of Native Americans around1815 (Guyette and Cutter 1991). Frequencyand extent of fires in Missouri forests are

Figure 2.—Trends in area of forestland byownership classes in eight states that makeup most of the Central Hardwood Region (IA,IL, IN, MO, OH, PA, TN, WV).

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further substantiated by evidence of vast oaksavannas, which historically covered an esti-mated one-third of the State (Nelson 1985).Repeated burning of oak forests in the morehumid regions to the east was also common(Day 1953, Komarek 1974, Little 1974, Lorimer1989). From an ecological perspective, thecurrent control of fire is likely the single mostsignificant human-induced alteration to thecentral hardwood forest landscape.

Logging had substantial impacts on the centralhardwood forests throughout the 1800’s. Forinstance, with the influx of settlers into theOzarks from 1850 to 1874 came the railroadand a period of accelerating logging during the1880’s (Smith and Petit 1988). The lumberboom came to an end in the Ozarks in the early1900’s and much of the land was sold by lum-ber companies sight unseen through the mailas farmland. During the period 1860-1899,more than 42 million acres of central hardwoodforest were converted to agricultural land use(Williams 1989). Widespread burning and openrange grazing of the forest occurred up until the1930’s in parts of the region.

The presettlement and early settlement historyindicate the Central and Eastern States wereheavily impacted by humans, and the wide-spread abundance of oak today is largely aresult of this disturbance history. Many oftoday’s oak-dominated stands are successionalin nature and will likely convert to other forestcommunities in the absence of continueddisturbance (Johnson 1993, Parker and Weaver1989).

Since the early 1900’s the amount of forest landin the United States has remained fairly con-stant. Within the Central Hardwood Region,forest land area has remained fairly stable sincethe 1950’s and has more recently shown aslight increase (fig. 2). To the south and eastthere are small declines in forest land (Powell etal. 1993). The increase in forest land is prima-rily the result of farmland and pasture revertingto forest. There are also trends in forest compo-sition indicating central hardwood forests arebecoming older and shifting to shade-tolerantspecies such as maple and beech (Raile andLeatherberry 1988, Smith and Golitz 1988).

The Regeneration Ecology of Oaks

Oaks are generally intolerant to moderatelytolerant of shade. Many of their associates

such as maples, blackgum, elms, beech, dog-wood, and redbud are much more shade toler-ant. The dominance of oaks in many forests istherefore dependent on canopy disturbancesthat allow light to reach the forest floor as wellas on the ability of young oaks to quickly gain acompetitive position in the canopy.

In most years nearly all oak flowers and oakacorns are destroyed by insects, fungi, oranimals. Occasionally, usually following abumper acorn crop, enough seed survives toproduce seedlings. Oak seedlings grow veryslowly and, unless released by a canopy distur-bance, cannot compete with other species. Inxeric (dry) ecosystems, oak reproduction tendsto accumulate in the understory of the parentstand. The repeated dieback and sprouting ofoak seedlings appears to be important to oaksbecause it allows seedlings to survive anddevelop a large root mass. Accumulation ofyoung oaks may occur over several decades,resulting in populations of seedlings and seed-ling sprouts that originated from several spo-radic acorn crops. These seedling sprouts arecapable of rapid and competitive growth whenthere is a significant reduction in the density ofthe canopy (Johnson 1979, Johnston 1941).Such events can result from fire, windthrow,insect- and disease-related mortality and defo-liation, drought, and timber harvesting.

Accumulation of oak reproduction under theparent stand is one of the most importantaspects of the regeneration ecology of oaks.This accumulation of small oak trees is called“oak advance reproduction” because it ispresent in advance of a regeneration harvest.Oak advance reproduction largely determinesthe composition of the next stand and thecombination of advance reproduction, seedlings,and stump sprouts represent the oak “regenera-tion potential” of a stand (Sander et al. 1984).Sustaining oak-dominated forests thus dependson perpetuating preestablished reproductionfrom one generation to the next. Oak reproduc-tion does not need to be present at all times in astand but should be present when the stand isregenerated.

Recurrent fire promotes the accumulation ofoak reproduction by eliminating or reducing thenumber of fire-sensitive understory competitors,including shrubs and shade-tolerant trees, andby killing overstory trees with thin, fire-sensitivebark. In addition, fire kills stems of oak repro-duction, which increases the root:shoot ratio of

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those that survive by resprouting. Oak repro-duction is well adapted to surviving fire becauseof the concentration of dormant buds near theroot collar. These buds often remain an inch ormore below the soil surface where they areprotected from potentially lethal temperatures(Korstian 1927).

Not all oak-dominated ecosystems require fireor disturbance to be sustained. Many xeric oakforests appear to be relatively stable communi-ties that show little evidence of succession tomore shade-tolerant or mesophytic species.Probably the largest North American ecosystemof this type occurs in the Ozark Plateau inMissouri where oak reproduction may accumu-late in the understory for 50 years or longer.However, even in this ecosystem, the capacity ofoak reproduction to accumulate varies withtopographic features such as slope position andaspect, which collectively control light, heat,and soil moisture (Sander et al. 1984).

There is a general relation between site qualityand regeneration success: ironically, the betterthe site the more difficult it is to regenerateoaks (Arend and Scholz 1969, Loftis 1990b,Lorimer 1989, Trimble 1973). Obtaining theaccumulation of oak reproduction necessary forsuccessful regeneration on highly productivesites requires recurrent disturbance. Histori-cally, fire created the necessary conditions.Thus, one potential solution to sustaining oak-dominated forests on productive sites is pre-scribed burning. Based on a review of researchresults on prescribed burning in eastern hard-woods and southern mixed pine-hardwoodstands, Van Lear and Waldrop (1988) concludedthat fire, if correctly employed, has the potentialto create the necessary conditions for sustain-ing oak-dominated forests on productive sites.

Oak Silviculture

Many forests in the Central Hardwood Regionwill likely succeed from oak-dominated foreststo forests comprised primarily of shade-tolerantspecies. Most silvicultural systems whenapplied to oak-dominated forests will maintain ahardwood forest of which oak is a component.The cutting or regeneration method used,however, will determine to what degree oaks arereplaced by other species. Generally, where theobjective is to perpetuate oaks, even-agedmanagement has been accepted as the mostappropriate regeneration method. Group-selection cutting, however, is increasingly being

used where smaller canopy disturbances andpartial cutting methods are desired.

Even- versus uneven-aged management

Regeneration methods can be classified as thosethat produce either even-aged forests or thosethat produce uneven-aged forests. Even-agedstands typically have one dominant age-class oftrees (which may vary in diameter) and gener-ally a level, closed canopy. An uneven-agedstand contains three or more age classes andtrees of various diameters and height (fig. 3).The canopy of an uneven-aged stand is dis-tinctly irregular in height. The clearcutting,seed-tree, and shelterwood methods are used toregenerate even-aged stands, and the single treeor group selection methods are used to regener-ate uneven-aged stands (Smith 1986). Recentlythere has been increased interest in the man-agement of two-aged stands, and silviculturalmethods for two-aged stands are under develop-ment.

Clearcut method

Where adequate advance oak reproductionexists, clearcutting will produce rapidly growingfully stocked stands containing oaks, hickories,and other shade-intolerant species as well assome tolerant species. The better the sitequality, the more difficult it will be to regenerateoaks. Clearcutting is most successfully used onrelatively dry sites where oak reproductionnaturally accumulates in the understory. It isoften unsuccessful in regenerating oaks onmesic sites (Beck and Hooper 1986, Gammon etal. 1960, Johnson 1976, Loftis 1983b). Failuresare largely because oak advance reproductiondoes not accumulate under the heavy shade ofthe parent stand. Where clearcutting fails toregenerate oaks, other commercially valuablespecies often become established. In manymesic ecosystems where adequate advance oakreproduction is not present, clearcutting willaccelerate the succession of oak-dominatedforests to mixed-mesophytic forests (Johnson1993).

The shelterwood method

The shelterwood method may be useful on siteswhere advance oak reproduction is not present.The shelterwood method involves the removal ofa stand in a series of partial cuttings. It isuseful in regenerating oaks on mesic sitesbecause it controls stand density near the end

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of the rotation when oak reproduction needs toaccumulate. Natural reproduction or planted-oak seedlings maintain their viability under thepartial canopy of the parent stand and arereleased by the final harvest.

The shelterwood method has been moderatelysuccessful in regenerating oaks. The initialshelterwood cut reduces the basal area byabout 70 percent. Herbicides or prescribedburning may be needed to reduce competitionfrom shade-tolerant trees in the understory andto increase survival of oak reproduction (Loftis1990b). When adequate oak advance reproduc-tion is present, a final cut removes the remain-ing overstory. The accumulation of adequateoak advance reproduction may take anywherefrom 1 or 2 years on dry sites to 10 or moreyears in mesic forests (Johnson 1993, Loftis1990a, Sander 1989). Planting of oak nurserystock can also be used in conjunction with theshelterwood method to make up for a deficit innatural oak reproduction (Johnson 1989).8

The seed tree method

The seed tree method is an even-aged methodthat usually leaves 10 or fewer seed-producingtrees on every acre (Smith 1986). The seed treemethod is not appropriate for regenerating oaksbecause successful regeneration is dependenton the amount of advance reproduction and notseed production after the harvest. Seed treesleft after the tree harvest would contribute verylittle to the regeneration of oaks but may bedesirable if sustaining acorn production (forwildlife) on harvested areas is important. Theregeneration method in this case is the clearcutmethod with trees retained for values otherthan regeneration.

Selection methods

Selection methods maintain a range of tree-ageclasses and diameters in a stand. Selectioncutting is regulated by a diameter distributioncurve. The target diameter distribution is

Figure 3.—Cross-sections of A) a mature even-aged stand, and B) an uneven-aged stand and theirrespective tree-diameter distributions.

determined by the largest diameter tree to beretained in the stand, the q-value, and thedesired residual basal area or stocking level.The q-value represents the average quotientbetween the number of trees in consecutivediameter classes and essentially determines thesteepness of the curve. A stand with a q-valueof 1.3 will have more large trees than a standwith a q-value of 1.7. Generalized guides existif an inventory cannot be conducted and adiameter distribution curve constructed (Lawand Lorimer 1989). Stands can be harvestedwhenever stocking exceeds 80 percent. Onmost average to good sites, this should be every15 to 20 years. Stands regenerated by selectionmethods retain many more large trees thanstands managed with even-aged regenerationmethods, and maintain some of the attributes ofmature forest. Because they retain significantamounts of canopy, however, they do not pro-vide many of the attributes of early-successionalcommunities.

The group-selection method—The groupselection method can be used to regenerateoaks. Trees are marked for harvest with atarget diameter distribution in mind for thestand. Some trees are harvested in smallgroups to create small regeneration openings.Just as with clearcuts, openings should belocated where there is sufficient advanced oakreproduction. The openings provide the neces-sary light for intolerant species like the oaks togain a competitive advantage over more shade-tolerant species. Recommended opening sizeranges from 1/10 acre to 1/2 acre (Law andLorimer 1989, Marquis 1989, Roach 1963).Larger openings should be referred to as patchcuts or clearcuts if the objective is to perpetuatethese as even-aged units through successiverotations (Marquis 1989). As with clearcut orshelterwood treatments, the reproduction thatdevelops in the openings depends on the ad-vance reproduction present at the time theopening is created (Sander and Clark 1971).

The single-tree selection method— Trees areharvested across all diameters to maintain adesired diameter distribution, but there is nointentional effort to create regeneration open-ings as in group selection. The single treeselection method generally has been consideredinappropriate for managing oak forests (Sanderand Clark 1971). Canopy openings the size ofindividual trees presumably provide inadequatelight for the accumulation of oak reproduction.

The method also encourages the development ofa shade-tolerant understory. The ecological andsilvicultural literature generally substantiatesthese assertions, although virtually all therelated studies were in mesic ecosystems whereoak reproduction apparently did not accumu-late (Della-Bianca and Beck 1985; Schlesinger1976; Trimble 1970, 1973).

The method has nonetheless been used suc-cessfully for 50 years on a large industrial forestin the Missouri Ozarks (Loewenstein et al.1995), and the diameter distribution of thisforest approximates the classical reverse J-shaped distribution characteristic of an all-agedforest (Loewenstein 1996). The key to thesuccess of this method may lie in the largenumber of small diameter oaks typically foundin xeric ecosystems such as the Ozark High-lands (Johnson 1993). In contrast, the smallerdiameter trees in mesophytic oak forests arelikely to be shade-tolerant non-oaks; smalldiameter oaks are typically absent (Loftis1983a). Guidelines are currently being devel-oped for use of the selection method in theOzark Highlands (E. Loewenstein, personalcommunication), but it is not recommended formore mesic, eastern parts of the Central Hard-wood Region.

Alternative methods

A number of alternative silvicultural systemsand revised practices are currently underdevelopment in response to increased publicconcern over the ecological effects (whether realor presumed) of traditional forestry practices.This includes the use of shelterwoods withoutfinal removal cuts as well as management oftwo-aged stands, which retain more residualstructure than clearcutting, but usually fewerresidual stems than the traditional shelterwoodmethod.

Diameter-limit cuts are frequently used onprivate non-industrial forest lands. Individualtrees of marketable species that exceed a cer-tain diameter are harvested. This methodretains forest cover on a site while removingmarketable trees. Unlike the previous silvicul-tural methods, however, the focus is on theharvest and not on the regeneration or struc-ture of the future stand. Because the largesttrees of desirable species are removed, theslower growing trees and undesirable speciesmake up the residual stand. This may have

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serious negative impacts on forest compositionand structure. Diameter limit cuts may exacer-bate oak regeneration problems and typically donot provide habitat for most early-successionalspecies.

CHAPTER 3—HABITAT AND ECOLOGY OFTHE RUFFED GROUSE

Distribution

The ruffed grouse (fig. 4) is North America’smost widely distributed gallinaceous bird(Johnsgard 1973). Ruffed grouse can be foundthroughout a significant portion of the CentralHardwood Region, yet are common only whereextensive tracts of forest dominate the land-scape (fig. 5). The predominately agriculturallandscapes in the Farm Belt that extends fromcentral Ohio westward through Indiana, Illinois,Missouri, and Iowa are largely inhospitable toruffed grouse. Here, scattered populations existonly along wooded river drainages or otherareas where sufficiently large tracts of forestexist. Ruffed grouse are found in central hard-wood forests throughout the southern parts ofthese States as well as Kentucky, Tennessee,parts of Arkansas, and the central and south-ern Appalachian Mountains. The southernextreme of the range of ruffed grouse is consis-tent with the southernmost edge of the Appala-chians in northern Georgia. Ruffed grouse aregenerally uncommon below 1,500 feet (460 m)in the extreme southeastern part of their rangeeven though habitats that appear suitable existin the Piedmont from Louisiana east to Georgia

and north through Virginia. In Kentucky,central Virginia, and further north, the impactof elevation on ruffed grouse distribution be-comes less pronounced than in more southerlylatitudes. This may indicate that warm south-ern climates are inhospitable to ruffed grouse.

Status in Central Hardwood Region

The ruffed grouse has a fragmented distributionthroughout the Central Hardwood Regionexcept in the largely contiguous forests of theAppalachian Mountains. This distribution is

Figure 4.—The ruffed grouse,perhaps more than anyother resident wildlife, isan early-successionalhabitat specialist. Whileoften thought of as a bird ofnorthern forests, its rangeextends throughout theCentral Hardwood Regionand southern AppalachianMountains.

Figure 5.—Distribution of the ruffed grouse in theEastern United States. Modified fromJohnsgard (1973) with input from Statewildlife management agencies.

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largely the result of land-use patterns andactive efforts to restore ruffed grouse popula-tions. Efforts to restore grouse populationsbegan in the late 1940’s and have occurredthroughout the Central Hardwood Region.These efforts have successfully restored somepopulations, however, local extirpations ofruffed grouse populations are likely to occurperiodically throughout this region because ofthe fragmented habitat, its location near theedge of the grouse range, generally low grousedensities, and the successional nature of re-quired habitats.

There is no range-wide population survey ofruffed grouse, but many States monitor popula-tions or harvest rates. Surveys of drummingmale grouse show some distinct patterns ingrouse numbers. Densities of drumming malegrouse are lower in the Central HardwoodRegion, and the Southeast in general than inthe northern portion of the grouse range (figs. 6,7). Approximately 4 to 6 birds can be expectedin the fall population for each drumming maleidentified in the spring. Central hardwoodpopulations do not show the strong cyclicfluctuations exhibited by northern populations

(fig. 7). Central hardwood populations inIndiana and Ohio may have declined overseveral decades (fig. 7). Hunter flush-rate datafrom Ohio, Tennessee, and Virginia also indi-cate long-term declines in these States’ ruffedgrouse populations, while West Virginia andNorth Carolina show periodic fluctuations butno consistent long-term trends.

Ruffed grouse population densities in thecentral hardwood forests seldom reach levelsattained in the aspen forests of the Great Lakesregion (fig. 6). The reasons for this pattern ofabundance are likely related to differences inthe forest composition and climate. Aspen, animportant component of northern forests,provides a unique combination of food andcover that is not provided by central hardwoodspecies. Note that, in general, populationdensities are greater in the northern portion ofthe Central Hardwood Region than in the south.Ruffed grouse can, however, be locally abun-dant in this region where quality habitats exist.

Habitat Use and Annual Cycle

Ruffed grouse are habitat specialists. Althoughthey can survive in various forest communities,they are common only on extensively forestedlandscapes that include numerous young (< 15years old), even-age hardwood stands. Opti-mum ruffed grouse habitat is most often cre-ated through the drastic disturbance of matureforest stands by processes such as timberharvest, fire, blowdown, or by succession ofopen lands back to forest. The high stemdensities characteristic of these stands (5,000to 8,000 or more stems/acre) protect ruffedgrouse from predators and enable local popula-tions to attain levels substantially higher thanon landscapes dominated by mature forest(Gullion 1984a, Kubisiak 1985, Stoll et al.1979). Perhaps more than any other year-round resident wildlife, ruffed grouse are early-successional forest specialists.

Early-successional forest provides severalimportant habitat components. Most importantis the low-dense overhead cover provided byhigh densities of saplings and tall shrubs.Overhead cover protects grouse from their mostimportant source of mortality—avian predators.The shade cast by dense overhead cover pre-cludes the growth of a dense herbaceous under-story, providing grouse with both unobstructedmovement and the visibility needed to detectapproaching predators.

Figure 6.—Densities of drumming male ruffedgrouse in nine Eastern States. Data sources:Georgia, Hale et al. (1982); Tennessee, Boyd(1990); Kentucky, J. Sole (unpubl. data);Missouri, Hunyadi (1984) and F. Thompson(unpubl. data); Indiana, Backs (1984); Ohio,Stoll and Culbertson (1995); Iowa, Porathand Vohs (1972); southern Wisconsin,Rodgers (1981); central Wisconsin, Kubisiak(1985); northern Minnesota, Gullion (1984).

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Ruffed grouse habitat needs vary during theyear and between males and females and youngand adults. It is easiest to understand thesedifferences by following the birds through theirannual cycle. The breeding season occurs inthe spring. Male grouse drum to advertise theirpresence to females and male competitors. Inthe Central Hardwood Region, drumming isusually at its peak during March and April.Hens visit males during this period and initiatenesting in early or mid-April. Broods hatchthroughout May and into early June. Hensspend the summer with their broods, andjuvenile birds disperse in the early fall. Grouseare primarily solitary during the winter, al-though they will sometimes concentrate in goodhabitat to take advantage of locally abundantfood sources.

Subtle differences sometimes exist betweenquality brood habitat and habitats frequentedby adults throughout much of the year. Ingeneral, habitat requirements for nesting ap-pear to be imprecise; nesting hens can be foundin a wide variety of habitats.

Year-round adult cover

Optimum adult cover usually consists of youngregenerating forest or shrub cover. This dense,

almost impenetrable cover protects grouse fromthe host of predators with which they mustcontend. The dense overhead cover providesprotection from hawks and owls and shades outground cover, enabling grouse to detect mam-malian predators. Dense, young forest standsare especially important to drumming malesadvertising their presence to nearby females.The visual and auditory displays of a perform-ing male are designed to draw attention to thebird. Drumming habitat must provide denseoverhead cover for protection from avian preda-tors while providing good visibility at groundlevel so a displaying male can detect approach-ing mammalian predators or other grouse. Amale’s activity center, the area that encom-passes his drumming platform or platforms, istypically the most structurally dense and,therefore, the most secure site available.

Grouse are sometimes observed in more opencover, usually because they are traveling be-tween habitat patches or feeding on a particu-larly attractive food. Grouse are more suscep-tible to predation in open cover, and on average,grouse densities will be higher where all habitatcomponents are provided in close proximity todense protective cover.

Figure 7.—Trends in ruffed grouse abundance determined by surveys of drumming males. Dataprovided by State wildlife management agencies. Data may not be comparable between Statesdue to differences in survey methods.

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In the Central Hardwood Region, adult cover isprovided by habitats such as regenerating oakor mixed hardwood forests, advanced stages ofold-field succession with sapling and pole-sizehardwoods and eastern redcedar, and older oakor mixed hardwood stands with dense under-stories of shade-tolerant shrubs or tree repro-duction (Atwater and Schnell 1989; Dimmick etal., in press; Stoll et al. 1979; Thompson et al.1987; Triquet et al. 1988).

Regenerating hardwood stands provide thehighest stem densities and most secure cover.Six- to 15-year-old stands generally provideexcellent adult cover (fig. 8). A 10-year-oldupland-oak stand on average will have from2,400 to 6,800 tree stems/acre, 1 to 2 inchesd.b.h. (diameter at breast height), and 7 to 21feet tall (Schnur 1937). The variation is largelydue to site quality (site index); poor sites willhave higher densities of smaller stems thanbetter quality sites. Total stem densities,including all shrubs and trees >1m tall, oftenrange from 6,000 to 25,000 stems/acre(Kurzejeski et al. 1987, Laubhan 1987, Stoll etal. 1979). Resource Managers are often inter-ested in regenerating a high density of oaksbecause of their commercial and wildlife value.From the perspective of grouse cover, treespecies composition is less important than afully stocked sapling stand of hardwoods.Oaks, however, also provide an important food(acorns) in the Central Hardwood Region. Inthe Great Lakes Region, adult cover is usuallyprovided by dense stands of aspen saplings 6 to15 years old, or older stands with dense under-stories of alder or hazel.

Old-field habitats are the result of land aban-donment. These habitats are composed ofinvading hardwood trees such as oaks, sassa-fras, persimmon, yellow poplar, hickory, ashand maple; shrubs such as dogwoods, multi-flora rose, blackberry, hawthorn, and honey-suckle, as well as conifers such as easternredcedar. Old fields usually do not have theuniformly distributed high stem density thatregenerating forest does, but this may be par-tially compensated for by the low dense ever-green growth of eastern redcedar or low cano-pies of trees such as hawthorn and dogwood.Redcedar has unique value as winter cover inportions of this region and is found almostexclusively in this habitat. Old-field habitatsalso have a diversity of fruit-producing shrubsand trees that provide important foods forgrouse and other wildlife.

Ruffed grouse populations in the Central Hard-wood Region seldom are confronted by pro-longed, severe winter weather conditions char-acteristic of northern latitudes. However, briefperiods of ice, snow, and extreme cold can beexpected during most winters. Contrary towhat might be expected, it is the lack of deepsnow during most winters throughout theCentral Hardwood Region that can stress ruffedgrouse. As temperatures drop below freezing inthe north, ruffed grouse escape the cold andpotential predators by burrowing into the snowuntil they are completely covered. Snow condi-tions are rarely suitable for snow-roosting in theCentral Hardwood Region. Conifers with adense canopy such as spruce, fir, or redcedarcan provide thermal protection for ruffed

Figure 8.—An example of goodyear-round adult coverand a drumming log in a10-year-old oak-hickorystand in central Missouri.

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grouse, as can evergreen shrubs - rhododen-dron and mountain laurel. In Missouri ruffedgrouse strongly prefer redcedar trees as awinter roost, and birds roosting in redcedar lossless heat and save more energy than those inhardwood cover (Thompson and Fritzell 1988).

Another component of adult cover is the pres-ence of suitable drumming platforms. Preferreddrumming platforms are typically large-diam-eter (>10 inches [25 cm]), partially decayedfallen logs > 10 feet (3 m) in length. Rocks,stone fences, upturned roots, and other el-evated perches are also used as drummingplatforms. In hilly terrain, logs oriented alongthe contour provide a relatively level platformand are most likely to be used by displayingmales. Although ruffed grouse activity centerscan be located anywhere on a hillside, quiteoften males select upper slopes or benches nearthe top or the base of the ridge where the slopeis < 25 degrees (Boag and Sumanik 1969,Porath and Vohs 1972, Rodgers 1981, Stoll etal. 1979, Thompson et al. 1987, Triquet et al.1988).

Nesting cover

A hen grouse sitting on a nest is well camou-flaged and predators are likely to pass bywithout detecting her. As a result, a key habitatfeature is visibility. This allows hens to watchpredators and determine when or if she needsto flush and potentially expose her nest. Nestsare usually next to a tree or log, perhaps toprovide cover in the one direction the hen can’tsee. Other than visibility and proximity tobrood and adult cover, there are no otherapparent requirements of nesting cover. Nest-ing cover is frequently located in pole-size, orlarger, hardwood stands. These older standsgenerally have good ground-level visibilitybecause their closed canopies shade out under-story and ground vegetation. In most forestedareas nesting cover is readily available but it isimportant that it be close to secure adult coverand brood habitat.

Brood cover

Broods, like adults, need suitable cover forprotection from predators. They also need anabundant high protein food source to supportrapid growth. Grouse chicks, like the young ofmany other birds, feed heavily on insects duringthe first few weeks of life. Throughout much ofthe grouse range, broods are seldom found far

from dense protective cover characteristic ofregenerating forest stands or shrub-dominatedold-field habitats. Broods are often found in 3-to 7-year-old regenerating stands that still havea significant herbaceous component. Thesehabitats are often more “patchy” and supportmore herbaceous ground cover than adultcover. Brood habitat is often more mesic thanadult cover, often along creek bottoms, alderswales, or lower north or east slopes (Thompsonet al. 1987). Small herbaceous openings with asignificant shrub component provide an impor-tant source of insects for developing chicks.

Grouse Foods

Grouse eat a wide variety of buds, catkins,fruits, and leaves. Their diet varies from regionto region. Grouse populations in the CentralHardwood Region are more likely to be limitedby a lack of quality winter forage than northernpopulations (Servello and Kirkpatrick 1987).Catkins and buds of yellow and paper birch,hazel, and especially quaking and bigtoothaspen are important winter foods throughoutthe northern part of the grouse range. Centralhardwood forests contain few species of trees orshrubs that support catkins that are readilyavailable to feeding grouse. Aspen is an impor-tant winter food because grouse can quickly filltheir crops foraging in trees and then return tosecure cover or snow burrows.

Grouse depend more on soft and hard mast andleaves for their fall and winter diet in centralhardwood forests than in northern forests(Korschgen 1966, Norman and Kirkpatrick1984, Thompson and Fritzell 1986). Grousemay spend considerable time and energy feed-ing on these foods because of the greater diffi-culty of foraging in fruiting shrubs or searchingfor succulent vegetation and other food on theforest floor. Also, important foods such as hardmast (acorns, hickory, and beech nuts) are notoften found in dense, young forest stands.Therefore, to take advantage of these foods,grouse must forage in relatively open, maturestands and increase their exposure to potentialpredators. Soft mast, the fleshy fruits fromdogwoods, grapes, greenbrier, hawthorn, servi-ceberry and many other fruiting plants, isreadily consumed by ruffed grouse.

Insects make up the majority of the diet ofruffed grouse chicks during their first 4 to 6weeks of life. Insect abundance is greater inopen habitats dominated by herbaceous vegeta-tion than on the forest floor beneath a closed14

canopy (Bump et al. 1947, Hollifield 1992).These herbaceous openings can provide anabundant source of insects for young chicks.Unfortunately, foraging in an opening is unsafefor ruffed grouse, and the potential benefits ofherbaceous openings must be weighed againstthe likelihood of increased predation. Youngforest regeneration cuts or old fields that con-tain patches of herbaceous ground cover as wellas dense shrub and sapling cover are idealbrood habitat because they provide both foodand cover.

CHAPTER 4—EARLY-SUCCESSIONALFOREST SONGBIRDS

Early-successional forest habitats provide food,cover, and nest sites for a variety of songbirds.Some of these species are specialists thatdepend on dense shrub-sapling cover, whileothers are generalists that use a wide range ofhabitats. Some are abundant, while others aredeclining and there is a high level of concern fortheir conservation.

Birds that nest in shrubland or young-foresthabitats make up an important component ofthe midwestern avifauna. Probst and Thomp-son (1996) compiled information on midwesternneotropical migratory birds. They identifiedspecies for which there was a high degree ofmanagement concern and species that weredeclining in numbers. Of 187 species thatbreed in Midwestern North America, 95 useshrub-sapling or young-forest habitats to somedegree during the breeding season (fig. 9).

Population trends of some common early-successional forest birds are shown in figure10. Of the 12 species depicted in figure 10, thebrown thrasher, prairie warbler, yellow-breastedchat, indigo bunting, and rufous-sided towheeshow significant long-term declines.

Habitat Associations of Early-successionalForest Songbirds

Midwestern, early-successional birds use a widevariety of shrubland or young-forest habitatsranging from semi-forested grasslands, such asglades, barrens or savannas; old fields; andyoung forest resulting from silvicultural treat-ments. General habitat requirements of somecommon birds in central hardwood forested andsemi-forested habitats are shown in figure 11.

Several distinct habitat components of early-successional forest are used by songbirds.

Figure 9.—Number of neotropical migratorybirds, priority neotropical migratory birds,and declining neotropical migratory birds inthe Midwestern United States grouped bymajor breeding habitats. Priority species arespecies considered a high managementconcern based on abundance, distribution,population trend, and known threats. Declin-ing species had a significant (P<0.1) negativepopulation trend based on the Breeding BirdSurvey. Adapted from Probst and Thompson(1996). 15

Figure 10.—Population trends of common breeding songbirds in early-successional forest habitats inNorth America. Data are from the North American Breeding Bird Survey (Bruce G. Peterjohn,National Biological Service, Patuxent Environmental Center, Laurel, Maryland).

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During the early stages of forest regeneration (1to 3 years after canopy removal), these habitatshave dense clumps of grasses and herbaceousvegetation that are used for nesting and forag-ing by species such as the prairie warbler andfield sparrow. As seedling-sapling-size treesand shrubs grow, they begin to shade outground cover. Regenerating stands support avery diverse and abundant bird communityduring this stage because there is a mix ofgrasses, forbs, shrubs, and tree reproduction.Species such as the yellow-breasted chat andblue-winged warbler are common in suchstands. This stage of stand development,however, lasts only a few years (approximately 3to 9 years after regeneration) before the canopycloses and much of the groundcover succumbsto shade. Species such as the wood thrush,ovenbird, and worm-eating warbler begin to usethese stands within 10 to 20 years after theyare regenerated.

Residual Structure in Regenerating Stands

Snags, slash, and residual live trees that werenot removed during timber harvests are alsoimportant habitat components in early-succes-sional forests. Snags, woody debris, and somelarge live trees from the previous stand providehabitat features needed by certain species andincrease within-stand diversity in deciduousand coniferous stands (Balda 1975, Dickson etal. 1983, Niemi and Hanowski 1984, Scott1979). Snags are used for nesting and feeding.Primary cavity nesters such as woodpeckersexcavate their own nest cavities. Secondarycavity nesters such as bluebirds and great-crested flycatchers use existing natural cavitiesor cavities excavated by primary cavity nesters.A wide variety of species feed on snags or usethem as perch sites. Residual live trees inregeneration cuts provide structure for opencanopy species such as blue-gray gnatcatchers,northern orioles, and yellow-billed cuckoos, orspecies from surrounding forests such as red-eyed vireos and some flycatcher species (Probstet al. 1992). However, tree regeneration andshrub stem densities can be reduced if toomany live residual trees are left standing,thereby reducing habitat quality for speciesdependent upon structurally dense habitats.

There has been some speculation that snags orresidual trees might provide perches for brown-headed cowbirds (a brood parasite) and nestdepredators (such as blue jays and crows). Nestpredation and parasitism can be high for some

birds nesting in early-successional forest(Annand and Thompson 1997), so this could bea real concern. Clustering residual trees orleaving them near the periphery of the standcould reduce the effective use of these by cow-birds and predators. Large stands with ahigher area to perimeter ratio will also have lessforest edge and fewer potential perch sites forcowbirds and predators.

Habitat Patch Size

In addition to habitat structure, the size of apatch of early-successional forest influences thebird community. Some species of songbirdsthat breed in early-successional central hard-wood forest have strong preferences for specificpatch sizes. Patch-size preferences generallyfall into two categories: species that prefersmall patches created by small disturbancesthat remove one to several trees, or species thatprefer large contiguous patches of at least 3 to 5acres. Small gaps can be created by windthrow,single tree deaths due to insects or disease, orby selection cutting. Hooded and Kentuckywarblers prefer small gaps created by single-tree or group selection methods, and manyother late-successional forest species are alsoabundant in these stands. Large patches of atleast several acres are created by large-scalewindthrow, high intensity fires, insects anddisease, and even-aged regeneration methods.Yellow-breasted chats, prairie warblers, blue-winged warblers, and white-eyed vireos aretypically found in stands regenerated by theclearcut or shelterwood method but not in thoseregenerated by selection methods (fig. 11,Annand and Thompson 1997).

Importance of Landscape Context

In addition to habitat structure and patch size,landscape context may also be an importantfactor affecting the quality of early-successionalforest habitat for songbirds. Landscape contextrefers to the overall landscape composition andpattern within which a specific habitat is lo-cated or being considered. Forest songbirds,including those that nest in mature or youngforests, have higher reproductive success inlandscapes that are predominately forested.Patches of early-successional forest habitatlocated within heavily forested landscapes maybe of more value to these species than habitatsin agricultural landscapes. This may seemcontrary to many managers’ ideas of habitat forthese species because they are often associated

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with forest edges bordering agricultural fields,and are sometimes referred to as “edge species.”Species nesting in agricultural landscapes,however, have a greater risk of brood parasitismby brown-headed cowbirds or nest depredationby predators (Donovan et al. 1995a, 1997;Robinson et al. 1995; Thompson et al., in press).

CHAPTER 5—FOREST MANAGEMENT PRAC-TICES FOR

EARLY-SUCCESSIONAL WILDLIFE

Resident and migratory birds use a wide rangeof forested and semi-forested habitats in centralhardwood landscapes. Even when managementactivities are focused on a reduced set of highpriority or featured species, these species stillspan a wide range of habitats. Conservationplanning is further complicated when lands aremanaged for multiple use, including recreationand forest products. Land managers and thepublic should recognize that management of anyparticular landscape will benefit some speciesand harm others, but that across landscapesthe needs of all species can be met. Thompsonet al. (1996) suggest that a mix of even- anduneven-aged silvicultural practices, designatedreserve areas, and use of prescribed fire will berequired within the Central Hardwood Region tomeet bird conservation objectives and otherobjectives for forest lands. Even if we narrowour focus to species found in early- successionalforest created by silvicultural practices, a mix ofregeneration methods will be needed to meetspecies habitat needs (Annand and Thompson1997). Thompson et al. (1996) used a simplelandscape model to demonstrate how mostregulated, sustainable forest managementpractices likely will sustain most species acrossa landscape. Species abundances, however,differ greatly among alternatives.

Depending on forest ownership, legal mandates,historical landscape composition, and manage-ment objectives, practices that create early-successional habitats or produce forest productswill be appropriate. Regulated forest manage-ment with standards and guidelines for wildlife,practiced within an adequate forest landbase,can accommodate the needs of central hardwoodforest songbirds.

Importance of Extensively ForestedLandscapes

Fragmentation of forest by non-forest land-uses,such as urban and agricultural uses, has nega-tive consequences for many species of forest

wildlife. While the total amount of forest area inthe Central Hardwood Region has in recentyears been relatively stable, few areas areextensively forested or minimally fragmented.We believe an important conservation need is tomaintain landscapes in these regions in anextensively forested condition. Active reforesta-tion, or encouraging succession of non-forestlands to old fields and forest, could minimizefragmentation in some landscapes and benefitforest wildlife (though at the expense of farm-land wildlife).

Availability of Habitats Through Time

Early-successional forest-wildlife will be mostabundant through time in landscapes managedfor the greatest amount of early-successionalforest habitat that is sustainable. The appropri-ate amount of early-successional forest habitatin the landscape will depend on wildlife objec-tives (particularly the balance between early-and late-successional wildlife), silviculturalpractices and objectives, and ecological landtype or suitability of the site for these communi-ties (particularly glades, barrens, and savan-nas).

Because early-successional forest habitat isephemeral, its availability varies by location(space) and through time. This means that landmanagers must carefully plan managementactivities over long time periods and large areasto ensure the availability of these habitats. Forinstance, if even-aged forest management is theprimary method being used to create early-successional forest, regulating harvest activitiesto provide sustained yield over the commercialrotation will result in a balanced forest age-class distribution and consistent levels of early-successional habitat through time. This pro-vides a consistent level of habitat availability forlate-successional forest wildlife as well. Forexample, in central hardwoods, generally 10 or12 percent of forest land can be regeneratedper decade under regulated forest managementwith a 100- or 80-year rotation, respectively,and would sustain 10 to 12 percent of thelandscape in 1- to 10-year-old forests. Moreforest can be regenerated in any one decade,but a higher level of regeneration cannot besustained throughout the rotation and wouldeventually result in widely fluctuating amountsof habitat and population levels.

Shorter rotations will result in more youngforest in a landscape than longer rotations.

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Figure 11.—Habitat associations of some common breeding songbirds in the Central HardwoodRegion. Glades and savannas are semi-forested habitats managed by prescribed fire. Clearcut,shelterwood, group selection, and single tree selection refer to stands treated within 10 years bythese silvicultural methods. Mature forest was even-aged forest 60 to 100 years old. Adaptedfrom Thompson et al. (1996) and Annand and Thompson (1997).

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Because most oak forests are regenerated bycommercial harvest for saw logs, minimumrotation ages are usually dependent on the ageat which oaks reach saw log size and are typi-cally around 80 years old. Conceivably, ifmarkets existed for smaller diameter trees,shorter rotations could be used as long as thecombined regeneration potential of stumpsprouts and advanced reproduction was ad-equate to regenerate the stand. Other manage-ment objectives should also be considered whensetting rotation ages. For instance, extendingrotation ages beyond a typical commercialrotation will result in more habitat for late-successional forest wildlife, more old-growthcharacteristics, but less early-succession foresthabitat.

Regulating harvest activities to maintain abalanced age-class distribution with a 80-yearrotation will ensure relatively consistentamounts of habitat for all forest wildlife throughtime. Regulated harvest on a 80-year rotationwould maintain about 15 percent of the forestin ruffed grouse brood or adult cover (forest 3 to15 years old). Other direct management ofhabitats such as old fields or riparian forest(discussed below) could increase this area tomore than 15 percent of the landscape. Theamount of the landscape that can be main-tained in prime grouse habitat by regulated-commercial timber harvest is notably smaller incentral hardwood forests than in aspen forests.Twenty-five percent or more of northern aspenforests can be maintained in brood and adultcover, largely because of younger rotation ages.

Habitat Patch Size

Songbirds that breed in early-successionalforest and ruffed grouse will benefit most frompatches of regenerating forest greater than 5acres and preferably 10 to 40 acres. Largerpatches have less edge per unit area and aremore efficient to manage. Forest wildlife ingeneral, however, may benefit more from arange of patch sizes. Some species, such as thehooded warbler, are primarily found in habitatpatches <1 acre. For example, a mix of even-aged and uneven-aged regeneration methodsapplied at different intensities across the land-scape will maintain diversity at a large scalebetter than one method uniformly appliedacross the region (Thompson et al. 1996). Also,the variation in habitat-patch size created by amix of even- and uneven-aged management

more closely mimics historic disturbance pat-terns. Fire and wind tend to create many smallhabitat patches (<1 acre) and fewer largepatches (>10 acres), resulting in a reverse J-shaped distribution in the frequency of differentsize gaps. This distribution can be approxi-mated using even-aged and uneven-aged meth-ods on approximately equal areas of the forest.In even-aged management, the size of standsand the rotation lengths can be varied.

Research in aspen forests of the Great LakesRegion shows that small harvest units (3 to 5acres) are of greater benefit to ruffed grousethan larger harvest units (Gullion 1984a). Thesmall harvest units are designed to provideruffed grouse with patches of protective cover(6- to 15-year-old stands) interspersed withmature stands where grouse find their principalwinter food, the flower buds from male aspentrees. The small harvest unit size ensures thatstands of varying ages are close to one another,thereby promoting maximum ruffed grousedensities.

Grouse in central hardwood forests will alsobenefit from interspersion of habitats, but theymay not benefit from a pattern of small-blocktimber harvests to the same degree as in aspenforests. We recommend regenerating stands 10acres or larger because the level of forest age-class interspersion required in aspen forestsmay not be required in central hardwood for-ests. Throughout much of the Central Hard-wood Region, there is no universally available ordominant food comparable to mature aspen innorthern forests. Many foods are abundant inrecently regenerated stands and mature stands.However, ruffed grouse that are forced to feed inthe relatively open understory of mature foreststands are susceptible to predation. Interest-ingly, small-block cutting units (2.5 acres) inmixed oak stands in central Pennsylvaniasupport surprisingly high ruffed grouse densi-ties (Storm, unpubl. data). Precise densityestimates for these mixed oak stands are diffi-cult to determine due to the presence of adja-cent small-block cutting units in aspen andscrub oak (Quercus ilicifolia) communities.Scattered small-block harvest units in land-scapes dominated by mature forest stands canprovide quality habitat for ruffed grouse, butthese isolated islands likely provide only limitedsecurity from predators. Although there is noconclusive research, we believe cutting units> 10 acres, and perhaps 20 to 40 acres, may

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provide better security for ruffed grouse incentral hardwoods. Most of the resources theyneed will be within or immediately adjacent tothe cutting unit.

Arrangement of Habitat Patches

Just as habitat use varies among wildlife spe-cies, so do the benefits derived from differentspatial arrangements of habitats. Some spe-cies, such as songbirds, generally restrict theirbreeding activities to a single habitat patch.Generally the larger the habitat patch, thelarger and more secure the local population willbe. For early-successional songbirds in man-aged-forest habitats, this means the larger,regenerating stands are better than smaller.Regenerating adjacent stands will also create alarger expanse of contiguous habitat. Regener-ating large stands or regenerating groups ofstands will also minimize any potential negativeedge effects for early-successional and late-successional species. Migrant songbirds mayuse several different habitats during the yearbut because of their great mobility they readilytravel between them. In fact, neotropical migra-tory birds travel among several countries andtemperate and tropical zones to meet theirannual habitat needs.

By contrast species such as ruffed grouse thatuse several habitats during a day, season, oryear; and that are less mobile, will benefit fromthe proper interspersion or local diversity ofhabitats. The optimum habitat arrangement forruffed grouse should provide all their annualhabitat requirements adjacent to 10 acres ormore of dense protective adult cover. Forinstance, a good arrangement of habitats forruffed grouse is a 10-acre or larger 5- to 15-year-old regeneration cut on a northeast slope,an old field on the adjacent ridgetop, andriparian forest along the stream at the base ofthe slope managed by group selection method(fig. 12).

Regeneration Methods

Even-aged methods (clearcut, seed-tree, andshelterwood) are the most appropriate methodsfor creating habitat for early-successional forestspecialists like ruffed grouse, prairie warblers,and blue-winged warblers. These methodsremove sufficient canopy from the parent standto result in enough understory development toprovide protective cover for ruffed grouse and

foraging and nesting cover for songbirds. Selec-tion methods seldom remove sufficient overstoryto allow the development of a sufficiently denseunderstory, or create large enough openings formany early-successional forest species. Groupselection methods can produce stem densitiescomparable to clearcuts in central hardwoodforests (Weigel and Parker 1995), but regenera-tion patches are generally too small to providelarge enough patches of contiguous habitat.Uneven-aged management should not be pre-scribed where early-successional forest speciesare the primary management objective. Inareas where regeneration methods are limited toselection cutting because of other objectives orregulations, group selection should be used,groups should be made as large as possible,and group openings should be clustered. Selec-tion methods do provide habitat for gap speciessuch as the hooded warbler and indigo bunting,and they provide other habitat values that maywarrant their inclusion in forest landscapes.Selection methods may be used in riparianzones where clearcutting may be inappropriateand understory and groundcover developmentis desired for ruffed grouse and other species.

Although not a regeneration method, crop treerelease deserves some mention as a manage-ment consideration. Throughout the CentralHardwood Region, likely crop trees exhibitdominance and can be identified as early as 5years after the parent stand has been regener-ated (Marquis and Jacobs 1989). The growth ofthese crop trees can be enhanced by felling alladjacent stems. Initial crop tree release typi-cally is conducted in stands < 15 years old.This 10-year window from age 5 to age 15 isprecisely that period in stand developmentwhen habitat quality for ruffed grouse is opti-mum. The release of crop trees in 5- to15-year-old stands can significantly reducestem densities and greatly diminish the protec-tion afforded ruffed grouse by these stands.Such treatments are inconsistent with ruffedgrouse management and should not be pre-scribed where the development of quality habi-tat for ruffed grouse is an objective.

Retaining Trees and Snags inRegeneration Cuts

Retention of mature live trees in regenerationcuts provides potential cavity and den trees,mast production for grouse and other wildlife inthe regenerating stand, song perches for song-birds, structural diversity in even-aged stands,

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potential large den trees and snags in develop-ing stands, and the potential for some largetrees that might not develop under intensiveeven-aged management. Many of these charac-teristics are not of direct value to early-succes-sional songbirds but are easily accommodatedin management plans and benefit other wildlifespecies.

To accommodate cavity and snag-using wildlifein young even-aged stands, four or more livecavity trees and three or more snags should bemaintained across a range of tree diametersfrom 6 to >19 inches d.b.h. (Titus 1983). Re-taining individual trees throughout a standwould result in a uniform distribution, butclustering retained trees may increase theirsurvival because they may be less susceptible to

Figure 12.—An example of forest management for early-successional wildlife. The habitat composi-tion of this landscape represents a balanced age-class distribution for regulated timber harveston a 80-year rotation. Young forest stands on side-slopes, adjacent to old fields on ridgetops andselectively cut riparian forest, provide a good arrangement of habitats for ruffed grouse. A diver-sity of early-successional habitats provides habitat for many songbirds. Grouping harvestactivities in one portion of the landscape maximizes habitat quality for early successional wildlifein that portion of the landscape while providing a large block of late-successional forest for otherwildlife.

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wind, lightning, or dieback after cutting. Thereis also concern that large trees or snags inearly-successional habitats create perches forpredators and cowbirds (a brood parasite);clustering retained trees will reduce the disper-sion of potential perches. When clusteringretained trees and snags in stands regeneratedwith even-aged methods, consider retaining two1/6-acre clumps or one 1/3-acre clump per 5acres. A range of tree sizes exists in uneven-aged stands but a conscious effort may beneeded to be sure that a minimum number ofsuitable cavity trees and snags is present.

Retaining some mature oaks in regenerationcuts can provide for substantial acorn produc-tion in the regenerating stand. Substantialgains in acorn production can be obtained byselecting good acorn producers in a stand andconsidering the d.b.h. at which acorn produc-tion peaks for different species (see Johnson1993 for guidelines). Identifying the seedproducers, however, requires long-term recordson seed production, which few forest managersare likely to have at their disposal. Sharp(1958) observed that fewer than 30 percent ofwhite oaks produced acorns and many of thosewere poor producers. If this is typical of mostoak species, leaving 10 good acorn producersper acre would retain 40 percent or more of theacorn-producing capacity of the original stand.This assumes that about 75 oaks are in theoverstory at the end of the rotation. Moreover,under the open-grown conditions created by theseed tree method, seed tree crowns can expandto their maximum, thus increasing acornproduction per tree. The crowns of some seedtrees may degenerate from crown diebackresulting from their sudden exposure (Smith1986). However, the snags that ultimatelydevelop may provide valuable habitat for cavitynesting birds and the standing dead woodessential to preserving components of biologicaldiversity. Moreover, seed trees can be quicklyconverted to dead snags by girdling once it isdetermined they have little value for acornproduction or other purposes.

In general, the greatest amount of overstoryremoval will yield the greatest degree of under-story development and cover for ruffed grouseand other early-successional wildlife. Retentionof a limited number of mature trees may havelittle impact on stem densities in the developingstand. Smith et al. (1989) found similar stemdensities 5 years post treatment in clearcutstands and stands with < 20 ft2/acre of residual

basal area. Regeneration treatments with lowlevels of basal area retained have been calleddeferment cuts or modified shelterwood har-vests and can lead to a stand dominated by twodistinct age classes. If many trees are retained,however, stem densities in the developing standwill be significantly reduced. The diameterdistribution of residual trees can also have asignificant impact on regeneration stem densi-ties. For example, 20 ft2 of residual basal areaper acre is provided by retaining approximately16, 15-inch diameter trees or approximately150 5-inch diameter trees. Although thecrowns of the larger diameter trees are far moreexpansive than those of the 5-inch trees, thelatter would quickly respond to release, and theshade cast by the combined crowns of thesesmall-diameter trees would likely have a greatereffect on the developing regeneration thanwould the shade cast by the larger trees.

The spatial distribution of residual trees withina harvest unit can also have a significantimpact on regeneration stem densities. If weuse the above example, 150 5-inch diametertrees equally spaced across 1 acre would re-quire that the trees be only 17 feet apart. Thisspacing would lead to substantial crown closureover the developing regeneration and wouldhave a significant negative impact on regenera-tion stem densities. Residual basal area main-tained in discrete patches will minimize shadingof regenerating hardwoods and, therefore, theeffects of this shade on regeneration stemdensities. Residual basal areas > 20 ft2/acrecan significantly reduce regeneration stemdensities and should not be maintained withinharvest units designed to provide quality habi-tat for ruffed grouse and other early-succes-sional species. In addition, residual basal arealevels < 20 ft2/acre can under some circum-stances reduce stem densities and habitatquality for ruffed grouse.

Soft and Hard Mast Production

Acorns are an important resource in oak-dominated ecosystems because of their role intree reproduction and as a key component incomplex food webs. They are one of the mostimportant fall and winter foods of grouse in theCentral Hardwood Region (Korschegen 1966,Thompson and Fritzell 1986). Acorn productionvaries greatly among trees, but in general large-diameter trees produce more acorns than small-diameter trees (Johnson 1994). Acorn produc-tion can potentially be increased in adult grouse

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cover by retaining some large-diameter, high-producing trees in regeneration cuts, as previ-ously discussed. This ensures acorn produc-tion within the secure grouse cover of thedeveloping stand. An important general guide-line for ensuring mast production is to usesilvicultural methods that will regeneratestands with a large percentage of oaks. Theimportance of mast as a grouse food is thepredominant reason for interspersing regenerat-ing and mature oak stands in areas managedfor grouse. Regulating even-aged managementon a 80-year or greater rotation should meet theneeds of mast-consuming wildlife at a land-scape scale.

Soft mast is another important wildlife food incentral hardwoods. Fruits of shrubs and treesare important summer, fall, and winter foods ofbirds. Regeneration cuts can increase produc-tion of these foods because they remove theoverstory. Mast-producing species are some-times controlled by felling or herbicide treat-ments when stands are regenerated. Wherewildlife habitat is an objective, control of mastproducing species should be minimized.

Direct planting of soft-mast producing trees andshrubs can benefit ruffed grouse, but it isexpensive and usually not cost-effective at alarge scale. Deciduous plantings are ofteneaten by deer unless protected by fencing ortree shelters, which can add significantly toplanting costs. Species selected for plantingshould be well suited to local soil and climaticconditions. Examples of trees and shrubs thatprovide fruits eaten by ruffed grouse includehawthorns (Crataegus spp.), Viburnum spp.,dogwoods (Cornus spp.), and cherry (Prunusspp.).

Existing sources of soft mast can also be main-tained and enhanced in established stands.Fruiting trees and shrubs and grape arbors canbe released from the competition of surroundingoverstory trees. These food sources are mostbeneficial to ruffed grouse when they are lo-cated in close proximity to dense protectivecover.

Consideration of Site Characteristics

Microclimate is an important parameter toconsider when identifying specific sites forruffed grouse habitat development efforts.Typically, relatively moist sites produce agreater abundance of soft mast and succulent

herbaceous vegetation than do droughty sites.In hilly or mountainous terrain, clearcut regen-eration harvests on north- and east-facingslopes are more likely to provide quality foragefor ruffed grouse than are similarly treatedstands on south or west exposures. However,the availability of dense woody vegetation onrelatively warm south and west exposures canbenefit ruffed grouse during inclement winterweather. Habitats positioned at or near thebase of a slope often provide a more cool, moistmicroclimate than those near the top of a ridgeand are typically preferred by ruffed grouse.Severely restricting silvicultural treatmentswithin varying distances from small streamsexacerbates efforts to establish early-succes-sional communities on these inherently produc-tive sites. Clearly, riparian corridors warrantspecial consideration before any timber harvestoperation is implemented, but here too distur-bance must be allowed to play a role. Sufficientoverstory must periodically be removed fromsites where conditions allow to promote aresponse from understory vegetation and atleast marginal habitats for ruffed grouse.

Management of Early-successional HabitatsOther Than Regenerating Forest

Diversity of early-successional communities

Management for early-successional forestcommunities should provide a range of habitatsincluding regenerating forest, glades, barrens,savannas, and old fields. Glades, barrens,savannas, and old fields generally have lowerwoody-stem densities and a greater grass andforb component than regenerating forests. Birdspecies abundances vary greatly among thesehabitats, so a mix of these habitats is mostlikely to meet the needs of all early-successionalforest birds.

Old-field habitats

Ruffed grouse habitat can develop as agricul-tural cropland or pasture is retired from pro-duction and allowed to revegetate naturally.Succession on recently abandoned agriculturallands can be a slow process. Decades may passbefore woody vegetation of sufficient heightexists in sufficient densities to support ruffedgrouse. The rate of establishment of woodyvegetation is dependent upon many site factors.A dense sod layer can limit seed germinationand seedling development. The thick humuslayer and the interwoven root systems of

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densely matted herbaceous vegetation canphysically preclude seeds from shrubs or treesfrom making contact with moist mineral soils, arequirement of many species for successful seedgermination. The natural regeneration of woodytrees and shrubs can be accelerated by me-chanical treatments. Mineral soil can be ex-posed by physically disrupting the humus andthe established root systems. A shallow-run-ning plow or disk can provide the physicaldisturbance required, but mechanical treat-ments can be expensive and they are impracti-cal on steep terrain. Mechanical treatments toexpose mineral soil should be timed to providean appropriate seedbed in the spring when soilmoisture is high. This means site preparationshould occur in spring or the previous fall afterherbaceous plant growth and seed dispersalhave largely ended for the growing season.

Existing old fields will require periodic treat-ments to set back succession. Managementpractices could include felling of pole-size orlarger trees, prescribed burning, and mowing orbrush cutting. The objective is to maintainsecure woody cover (including redcedar), adiversity of fruit-producing shrubs and vines,and patches of herbaceous cover. If the site isopened up too much, grasses and forbs willdominate and make the site less valuable toruffed grouse.

Herbaceous openings

Maintained herbaceous openings can provideruffed grouse with a readily available source ofquality forage. These openings also supportgreater arthropod densities than do habitatsdominated by woody vegetation (Bump et al.1947, Hollifield 1992). These openings are oftenplanted with several grasses and a legume suchas clover. Annual mowing helps to maintain thelegume component of these openings. Legumesremain relatively succulent throughout much ofthe year and provide ruffed grouse with a nutri-tious source of forage during winter when littleother herbaceous vegetation is available. Theabundant arthropods supported in maintainedopenings can be an important source of protein-rich food for developing chicks (Hollifield andDimmick 1995). The high-contrast edge formedby the maintenance of herbaceous openingsadjacent to forested habitats, however, mightlead to increased predation. Increased preda-tion can be minimized by locating herbaceousopenings adjacent to dense, young forest habi-tats. Dimmick et al. (in press) recommend

seeding landings and logging roads with amixture of clover and orchard grass to accom-plish this objective.

Conifers

Snow depths sufficient to allow snow roostingare uncommon in the Central Hardwood Re-gion. Establishing small patches of conifers canprovide ruffed grouse with protection frominclement winter weather. Conifer cover forruffed grouse should be dense and low to theground. Eastern redcedar is perhaps the bestnative conifer for grouse cover in the region,while species such as short-leaf pine are notsuitable. Species exotic to the region, such aswhite spruce (Picea glauca) or Norway spruce(Picea abies), also provide good cover. Norwayspruce is relatively unpalatable to white-taileddeer and, therefore, is the easiest to establish inareas that support high deer densities. Plant-ings should be > 0.5 acres in size to provide aneffective refuge from inclement weather. Thisrefuge effect can be enhanced by using a rela-tively tight spacing (6 ft X 6 ft) during planting.

ACKNOWLEDGMENTS

We thank Steve Backs, Al Bourgeois, SamBrocato, Dirk Burhans, Ralph Dimmick, ErikFritzell, Mark Gudlin, Eric Kurzejeski, EdLoewenstein, Jeffery Sole, and Robert Stoll fortheir comments on this manuscript. Thepublication is in large part a product of theMissouri Forest Wildlife Project, which wasinitiated and administered by Erik Fritzell andlater by Frank Thompson. We especially thankThe Ruffed Grouse Society for their long-termsupport of this project. Additional funds for theMissouri Forest Wildlife Project were providedby the University of Missouri, the USDA ForestService’s North Central Forest ExperimentStation, the Missouri Cooperative Fish andWildlife Research Unit, and the Missouri De-partment of Conservation.

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Thompson, Frank R., III; Dessecker, Daniel R.1997. Management of early-successional communities in central

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Describes the history, ecology, and silviculture of central hardwoodforests and the status and ecology of early-successional forest song-birds and ruffed grouse. Concludes with management guidelines forearly-successional communities in central hardwood forests.

Key Words: Early-successional forest, ruffed grouse, songbirds,wildlife habitat, central hardwoods, silviculture, oaks.