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KUCHLER TYPE

KUCHLER TYPE: Oak-hickory forest
KUCHLER-TYPE-NUMBER : K100 PHYSIOGNOMY : Medium tall to tall broadleaf forests [51]. Also, open woodlands with prairie forbs and shrubs in the understory. Typical deciduous trees on mesic, fairly rich soils are a maximum of 83 to 100 feet (25-30 m) tall with a nearly closed canopy. Trunk diameters may range up to 5 feet (1.5 m). There is usually a well developed shrub-sapling layer, but the herb and moss layer is variable, and is usually less than 100 percent cover [44]. Most deciduous forests have five well-developed vertical strata: the uppermost dense canopy, an open subcanopy of immature trees and mature small trees (such as flowering dogwood [Cornus florida] and eastern redbud [Cercis canadensis]), a shrub layer, an herb layer, and a surface layer of mosses and lichens [97]. This write-up includes some information on transitional vegetation (savanna) occurring at the interface between oak (Quercus spp.)-hickory (Carya spp.) forest and tallgrass prairie. United States Public Land Survey notes (1835) described these areas as "barrens." There is very little agreement, however, on what constitutes a barren and the term should be avoided [69]. Much debate continues over the degree to which oak savanna is "natural" [60,72]; however, most agree that it is a fire-dependent type (a fire disclimax that is stable over time given periodic fire). Clark [20] has suggested that closed oak-hickory forest is an artifact of fire suppression. Prairie and woodland burning by Native Americans had a great deal of influence on the extent and character of vegetation in the prairie-forest contact region, and in the eastern deciduous forest. The degree to which this activity altered the natural fire regime is largely unresolved [2,15]. OCCURRENCE : Oak-hickory forests occur in the central United States, ranging from central Iowa east to southern Michigan and south to Texas. These forests are extensive in some states, and have only local occurrences in others. In the Southeast, oak-hickory forests are a major type in Arkansas, Kentucky, and Tennessee and extend into Alabama, Mississippi, and Louisiana [15]. In the Southeast, at the beginning of the seventeenth century, there were approximately 1 million square miles (26 million sq km) of unbroken forest. By 1865, 65 percent of the surface area was forested; by 1923, less than 260,000 square miles (676,000 sq km) were left in second-growth merchantable timber [15]. The U.S. Forest Service indicated that there were 36.3 million acres (14.5 million ha) of oak-hickory forest in the north-central states in 1972 (this included Kuchler types 81, 82, 84, 89, 100, and 104) [64]. Oak savannas were an important feature of the midwestern landscape prior to European settlement. Nuzzo [69] estimated that at the time of settlement oak savanna covered some 28.6 to 33.8 million acres (11-13 million ha) of Minnesota, Wisconsin, Michigan, Iowa, Missouri, Illinois, Indiana, and Ohio. This estimate includes savannas occurring in Kuchler's oak-savanna (K081), oak-hickory forest (K100), and within the area mapped as bluestem (Andropogon and Schizachyrium spp.) prairie-oak-hickory mosaic (K082). There is some evidence that savannas existed in eastern Nebraska prior to settlement, some in areas previously thought to be exclusively tallgrass prairie [79]. Betz [8] estimated that presettlement savanna and associated sand prairie in Indiana covered 1.68 million acres (647,000 ha). Today only remnants of these types exist, most of which have been degraded due to various disturbances such as grazing and altered fire regime [7]. In 1985, 113 sites totaling 6,439 acres (2,607 ha) of relatively high-quality oak savanna remained, approximately 0.02 percent of the original extent. All but 100 acres (40 ha) were on sandy, rocky, or similarly droughty substrates [69]. STATES: AL, AR, IL, IN, IA, KS, KY, MI, MS, MO, NE, OH, OK, TN, TX COMPILED BY AND DATE : Janet Sullivan, 1995 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Sullivan, Janet. 1995. Oak-hickory forest. In: Remainder of Citation
Kuchler Type Index FEIS Home

KUCHLER TYPE DESCRIPTION


PHYSIOGRAPHY : Oak-hickory forests are found on all topographic positions including dry rocky ridges, deep coves, and well-drained valley floors [81]. West of the Appalachian Mountains, deciduous forests occupy areas geologically classified as plateaus, although erosion has produced rolling to mountainous topography [44]. CLIMATE : The climate of the north-central forest region (Minnesota, Wisconsin, Michigan, Iowa, Illinois, Indiana, Ohio, Missouri, and Kentucky) is continental and ranges from humid to subhumid. Winters are cold and summers hot. Temperatures differ greatly from north to south through the region, both in summer and winter. Temperature, growing season, and precipitation increase from north to south; at a given latitude, precipitation increases from west to east [44]. In the southeastern United States along the eastern edge of the oak-hickory forest, mean annual precipitation is approximately 45 inches (1,140 mm); at the western edge, annual precipitation averages 39 inches (980 mm). Lower precipitation in the western parts of this area is exacerbated by an episodic pattern of rainfall and more frequent drought. In southeastern oak-hickory forest, average annual precipitation increases from 42 inches (1,060 mm) in Indiana to 55 inches (1,400 mm) in Alabama [15]. At the southwestern limit of oak-hickory forest, central Texas has an average annual precipitation of 22 inches (560 mm) and mean annual temperatures around 65 degrees Fahrenheit (18.3 deg C). In eastern Texas mean annual temperatures are 62.6 to 66.2 degrees Fahrenheit (17-19 deg C), and the growing season is approximately 225 days [15]. SOILS : Oak-hickory forests occur on soils ranging from cool-moist Boralf and Orthod Spodosols to warm-dry Millisols and Alfisols. Soils range from clay to loam and are derived from glacial material, residual sandstones, shales, limestones, gneisses, schists, and granites [81]. In southeastern oak-hickory forests, soils encompass a wide range of physical and chemical features, ranging from shallow to deep, infertile to rich, and high clay fraction to very little clay. Soils are formed in residuum, primarily from limestone parent materials, but also from sandstones and shales. Soils in oak-hickory forests are fertile compared to other southeastern soils and represent some of the most productive agricultural land on these uplands [15]. Parent materials on the Ozark plateau are predominantly limestones, cherts, and dolomites. In Arkansas, soil parent materials in the eastern Ouachita province are mostly shales, sandstones, quartzites, and cherts, and in the western Ouachita province are mostly sandstones and shales [12]. VEGETATION : Classification: There is much debate over the most useful classification for the eastern deciduous forests including oak-hickory forest. Oak-hickory forest is not uniform across its range; dominants vary with climatic and edaphic conditions. The many differing interpretations of oak-hickory forest are probably a function of its vegetational complexity [15]. The most extensive area of unequivocal oak-hickory forest is found in the Ozark Mountains, where it covers most upland sites [15]. Hicks and Chabot [44] described oak-hickory forest that is largely equivalent to Kuchler's oak-hickory type, but Braun [12] characterized the central and eastern portions of K100 as western mesophytic forest. She defined the oak-hickory forest as the most westerly part of the deciduous forest region. Some authors use a definition of oak-hickory forest that is broader than either Kuchler's or Braun's, and include all forests dominated by oaks and having at least some hickory component as oak-hickory. These definitions usually include Kuchler's oak-hickory forest (K100), oak-hickory-pine (Pinus spp.) (K111), Appalachian oak forest (K104), and Braun's mixed mesophytic forest [37,81]. Monk and others [65] divided the eastern deciduous forest into three regions: the northern region, occupied by eastern hemlock (Tsuga canadensis), eastern white pine (Pinus strobus), and northern hardwood forests; the central region which included oak-hickory; and the southern region, occupied by pine and oak-pine forests. They defined oak-hickory as inclusive of Braun's oak-hickory, oak-chestnut (Castanea spp.), and about half of her western mesophytic forest; this definition is inclusive of Kuchler's oak-hickory forest but somewhat broader [65]. The Forest and Range Ecosystems of Garrison and others [37] include FRES 15, the oak-hickory ecosystem. This ecosystem includes Kuchler's oak-hickory but also the mosaic of oak-hickory and bluestem prairie (K082), Cross Timbers (K084), Appalachian oak (K104), oak-pine-hickory (K111), and oak savanna (K081) types [37]. This write-up includes mostly information about communities within Kuchler's [51] oak-hickory forest region as it was mapped, with the major focus on oak-hickory communities in the Ozarks. Communities further north (in Wisconsin and Michigan) are included where information is deemed relevant. An oak savanna type was mapped by Kuchler as a potential type for Wisconsin and Michigan (K081), but forests of similar structure and containing mostly the same species occurred widely in presettlement times throughout the prairie-forest interface. Savannas in the central states are therefore included because of their relevance to the fire ecology of oak-hickory forests. Other classifications describing oak-hickory forest include the following: The central hardwood forest [20] Plant geography--with special reference to North America [28] Ordination and classification of western oak forests in Oklahoma [31] A classification of the deciduous forest of eastern North America [65] The study of plant communities [71] Classification and evaluation of forest sites in the Cumberland Mountains [86] The natural vegetation of North America [97] Flora: Oak-hickory forests as described by Kuchler [51] are dominated by white oak, black oak, northern red oak, bitternut hickory (Carya cordiformis), and shagbark hickory (C. ovata). Other trees in alphabetical order of scientific name include pignut hickory (C. glabra), white ash (Fraxinus americana), black walnut (Juglans nigra), black cherry (Prunus serotina), chinquapin oak (Q. muehlenbergii), and basswood (Tilia americana). In the northern parts of the oak-hickory forest, other components are northern pin oak (Q. ellipsoidalis), and shingle oak (Q. imbricaria). In the southern parts of the oak-hickory type, forest tree species include black hickory (C. texana), mockernut hickory (C. tomentosa), southen red oak (Q. falcata), overcup oak (Q. lyrata), blackjack oak (Q. marilandica), Shumard oak (Q. shumardii), and post oak (Q. stellata) [51]. Occasional eastern white pine and eastern hemlock also occur [44]. Burkman and others [17] described oak-hickory forests as the largest and most diverse forest type in the eastern United States. Cover types include post oak-blackjack oak, bur oak (Q. macrocarpa), bear oak (Q. ilicifolia), northern pin oak, chestnut oak (Q. prinus), white oak-black oak-northern red oak, white oak, black oak, and northern red oak [17]. Oak-hickory forest sensu Kuchler is approximately equivalent to the Society of American Foresters white oak (Q. alba)-black oak (Q. velutina)-northern red oak (Q. rubra), white oak, and northern red oak forest cover types [39]. Braun's description of the oak-hickory forest flora resembles Kuchler's. In northern stands, bur oak has great prominence. Northern pin oak is confined to the northern division. Flowering dogwood is a common understory tree. Blueberries and huckleberries (Vaccinium spp.) are abundant shrub layer components, as is ceanothus (Ceanothus spp.). The herb layer often contains poverty oatgrass (Danthonia spicata), bird's-foot viola (Viola pedata), Appalachian mountainmint (Pycnanthemum flexuosum), stone-mint (Cunila origanoides), clasping aster (Aster patens), flax-leaved aster (A. linariifolius), and legumes including downy trailing lespedeza (Lespedeza procumbens), round-leaved tick-trefoil (Desmodium rotundifolium), wild sensitive plant (Chamaecrista nictitans), goat's rue (Tephrosia virginiana), pencil-flower (Stylosanthes biflora), and Sampson's snakeroot (Orbexilum pedunculatum) [12]. The Texas Plant and Wildlife Department published a list containing current information on the stability and distribution of plant communities occurring in Texas. They listed a post oak-black hickory series and post oak-blackjack oak series which occur within Kuchler's oak-hickory forest. In each of these community types, woody species diversity is highest to the east, where closed canopies may form. Post oak-black hickory forms open woodlands over deep sands in the western portion of its Texas range; species composition varies with soil texture and depth and herbs have higher species diversity than stands farther east. Components of this type include white oak, southern red oak, blackjack oak, pines, other hickories, flowering dogwood, yaupon (Ilex vomitorium), and beautyberry (Callicarpa americana). Common components of the post oak-blackjack oak series include cedar elm (Ulmus crassifolia), southern red oak, yaupon, beautyberry, water oak (Q. nigra), black hickory, eastern redbud, and deciduous holly (I. decidua), but composition is variable [89]. Savanna: Some authors define oak savanna as open-grown oaks with 10 to3 80 percent crown cover, with or without a shrub layer, and with a ground cover of grasses and forbs [23]. The understory vegetation of savanna is a mixture of both prairie and forest species, with prairie forbs and grasses more abundant in areas of high light, and forest forbs and woody species more abundant in areas of low light [13,69]. Early descriptions of Missouri savannas are of very open woodlands with ground covers of grasses and forbs and no underbrush or small timber [22]. The herb layer in oak savannas consists mostly of bluestem prairie species, namely big bluestem (Andropogon gerardii), little bluestem (Schizachyrium scoparium), Indian grass (Sorghastrum nutans), switchgrass (Panicum virgatum), and many prairie forbs [97]. In Indiana typical savanna herbs include little bluestem, butterfly weed (Asclepias tuberosa), flax-leaved aster, and yellow sedge (Carex pensylvanica) [7]. Savanna trees are oak-hickory dominants; bur oak is common in northern savannas and post oak and blackjack oak are dominant in southern savannas. In Texas junipers (Juniperus spp.) and mesquites (Prosopis spp.) are common associates [97]. Indiana black oak sand savannas are dominated by black oak and white oak. Understory trees include black cherry, sassafras (Sassafras albidum), shining sumac (Rhus copallina), and smooth sumac (R. glabra) [7]. No species is known to be endemic to oak savannas, and relatively few species are modal (i.e., have their highest occurrence in this habitat) [69]. In Wisconsin only six species are considered modal in oak savanna (K081): sunflower-everlasting (Heliopsis helianthoides), kitten-tails (Besseya bullii), cancer-root (Orobanche uniflora), smooth phlox (Phlox glabberima), thick-root buttercup (Ranunculus fascicularis), and mountain deathcamas (Zigadenus elegans) [25]. Geographic Trends: Major vegetational trends from east to west include an increasing importance of oaks, particularly post oak, and a reduction in canopy tree species diversity. Mesic sites and vegetation are more restricted in the western part of the range, particularly with decreased precipitation and increased incidence of drought and fire. Pines increase in importance from north to south [15]. Oak-hickory merges with oak-hickory-pine (K111) in eastern Texas, grades into post oak-blackjack oak savanna (Cross Timbers [K084]) farther south and west in Texas, and alternates with tallgrass prairie on the western edges of its distribution (within the region mapped as mosaic of bluestem prairie and oak-hickory forest) [12]. WILDLIFE : Birds: Few bird species can be considered distinctive of oak-hickory forests. Species frequently encountered in oak-hickory forests include red-bellied woodpecker (Melanerpes carolinus) and cerulean warbler (Dendroica cerulea) [61]. Avian species richness is well documented within the southeastern portion of the oak-hickory forest. Breeding bird communities within southeastern oak-hickory forests range from about 120 species in western Tennessee to about 93 species in the southern extreme [38]. Dominant members of the avifauna of southeastern oak-hickory forests include downy woodpecker (Picoides pubescens), great-crested flycatcher (Myiarchus crinitus), red-bellied woodpecker, eastern wood-pewee (Contopus virens), tufted titmouse (Parus bicolor), Kentucky warbler (Oporornis formosus), red-eyed vireo (Vireo olivaceus), and summer tanager (Piranga rubra) in the Bluegrass region of Kentucky. On the Highland Rim and Knobs of Kentucky and Tennessee, dominant avifauna also include ovenbird (Seiurus aurocapillus), black-throated green warbler (Dendroica virens), black-and-white warbler (Mniotilta varia), blue jay (Cyanocitta cristata), red-eyed vireo, wood thrush (Hylcichla mustelina), Carolina chickadee (Parus carolinensis), hairy woodpecker (Picoides villosus), and eastern wood-pewee. In the Central Basin, Tennessee, dominant avifauna include most of the abovementioned species and yellow-billed cuckoo (Coccyzus americanus). In the Ozark Plateau blue-gray gnatcatcher (Polioptila caerulea) and northern cardinal (Cardinalis cardinalis) occur in addition to previously mentioned species; on the Coastal Plain additional species include Carolina wren (Thryothorus ludovicianus), northern parula (Parula americana), and hooded warbler (Wilsonia citrina). In lowland communities the avifauna is dominated by prothonotary warbler (Protonotaria citrea), American redstart (Setophaga ruticilla), wood duck (Aix sponsa), red-eyed vireo, American woodcock (Scolopax minor), yellow-throated vireo (Dendroica dominica), northern parula, hooded warbler, Acadian flycatcher (Empidonax virescens), pileated woodpecker (Dryocopus pileatus), white-breasted nuthatch (Sitta carolinensis), Carolina chickadee, and tufted titmouse [15]. Probst [75] summarized oak forest bird communities for the eastern United States, including both oak-hickory forests and oak-pine forests. The blue jay and the brown-headed cowbird (Molothrus ater) are species of generalized habitat requirements that are found in almost every stand, regardless of age or structure. However, most birds require particular habitat features. Five categories of birds that are present in oak-hickory forest, based on response to habitat feature include: 1) bark foragers such as northern flicker (Colaptes auratus), white-breasted nuthatch, downy woodpecker, and hairy woodpecker, 2) active gleaners feeding in the tree canopy such as chickadees (Parus spp.), tufted titmouse, and red-eyed vireo, 3) pursuers feeding in tree canopy with a sit-and-wait hunting strategy such as flycatchers (Empidonax spp.) and tanagers (Piranga spp.), 4) ground species associated with shrub and sapling layers such as thrushes (family Muscicapidae), rufous-sided towhee, and ovenbird, and 5) species associated with dense growth of saplings and small trees, mostly warblers (Emberizidae) [75]. Mammals: Southeastern oak-hickory forests generally are low in mammalian species richness with the exception of bats (Chiroptera). The limestone-based geology of the region has fostered a rich cave fauna including little brown myotis (Myotis lucifugus), big brown bat (Eptesicus fuscus), and pipistrelles (Pipistrellus spp.). The white-footed mouse (Peromyscus leucopus) is very common in southeastern oak-hickory forest; other common species include short-tailed shrews (Blarina brevicauda and B. carolinensis), eastern chipmunk (Tamias striatus), eastern gray squirrel (Sciurus carolinensis), eastern fox squirrel (S. niger), gray fox (Urocyon cinereoargenteus), raccoon (Procyon lotor), opossum (Didelphis virginiana), striped skunk (Mephitis mephitis), and white-tailed deer (Odocoileus virginianus) [15]. Reptiles: The garter snake (Thamnophis sirtalis) is very common in oak-hickory forests of the Southeast; other common reptiles include black racer (Coluber constrictor), black rat snake (Elaphe obsoleta), five-lined skink (Eumeces fasciatus), fence lizard (Sceloporus undulatus), rough green snake (Opheodrys aestivus), ring-necked snake (Diadophis punctatus), eastern hognose snake (Heterodon platyrhinos), and eastern box turtle (Terrapene carolina). Reptile species richness generally increases to the south and east of the southeastern oak-hickory region. Amphibians: Moist sites within oak-hickory forest support amphibians, most commonly slimy salamander (Plethodon glutinosus), dusky salamander (Desmognathus fuscus), American toad (Bufo americanus), Fowler's toad (B. woodhousei), spring peeper (Hyla crucifer), and gray treefrogs (H. versicolor and H. chrysoscelis) [15]. ECOLOGICAL RELATIONSHIPS : Species Distribution: Oak-hickory associates tend to clump together on ridges and upper slopes. In grassy openeings and on steep slopes and other dry exposures oak-hickory associates cooccur with plant species such as eastern redcedar (Juniperus virginiana) that are tolerant of xeric conditions [15]. On the Ozark Plateau dry ridges and south-facing slopes are usually occupied by open communities of post oak and blackjack oak. Locally, shortleaf pine (Pinus echinata) may form mixed stands with the oaks. White oak is usually more abundant on north-facing slopes, entering some ridge forests. In the western border area, deep ravines and gorges are occupied by mesophytic species; sugar maple (Acer saccharum), white oak, and northern red oak are abundant. Open forests of low-statured trees with sparse herbs occur on xeric plateaus [12]. On the Ozark Plateau in southeastern Missouri, ordination of forest stands showed a clear separation of upland forests into two groups. Acid upland stands were dominated by black oak, black hickory, and white oak. Other stands (intermediate in elevation and soil acidity) tended to be dominated by chinkapin oak, sugar maple, eastern redcedar, and basswood. White ash and slippery elm (Ulmus rubra) were sometimes present [101]. The eastern deciduous forest reaches its western limit in the central portion of Oklahoma as oak forest and oak savanna. Farther west, occasional stands occur in sandstone canyons in west-central Oklahoma and in the Wichita Mountains of southwestern Oklahoma [31]. These forests are attributed to climatic fluctuations which favored the westward migration of eastern species [12]. Prior to European settlement oak savanna was common in western Oklahoma; these stands now have a closed canopy developed with the advent of grazing and fire suppression. Ordination of forests in this area demonstrated that the vegetation pattern corresponded to a complex moisture gradient. Post oak and blackjack oak codominated forests occuping the xeric end of the gradient, post oak dominated stands located in the middle (slightly more mesic), and forests at the mesic end of the gradient included sugar maple, elm, and Shumard oak-dominated stands [31]. Post oak and black hickory appear to require relatively high levels of nutrients and moisture, while blackjack oak is tolerant of drier, less fertile sites [47]. Phenology: The understory herbs begin growth earliest in spring; the herb stratum is dominated by species that overwinter as rosettes or underground perennating organs [44]. Disturbance and Succession: Oak-hickory forests have a long history of disturbance; the presettlement pattern was a mosaic of seral stands, many due to Native American activities. Continuing and overlapping disturbances in the range of oak-hickory forest arise from the high value of the land for agriculture [15]. In some regions in the eastern United States, current oak dominance is similar to and is related to the importance of oaks in presettlement forests. In other regions, particularly in the Midwest, the current distribution of oaks exceeds that of presettlement vegetation [1]. Much development of oak forests has occurred through a variety of ecological pathways and disturbance patterns [2]. In northern Michigan sites on upland sandy soils that were dominated by eastern white pine, red pine, or mixtures were converted to northern red oak stands after disturbance by logging and fire [46]. In Missouri stands that were oak-pine or pine at the turn of the century are now dominated by scarlet oak (Quercus coccinea) and black oak following heavy cutting of the pine and post-harvest fires [24]. Most oaks are considered early to mid-successional species; there is recent evidence of a widespread potential for oak replacement by more shade-tolerant tree species in mature forests [53,68]. This phenomenon seems to be more prevalent on mesic than on xeric sites [1,2]. There is a strong successional trend to shade-tolerant species such as sugar maple and American beech (Fagus grandifolia) on moist sites [15]. The persistence of relatively shade-intolerant oaks in mesic forests may be related to cycles of synchronous tree death similar to the fir (Abies spp.) waves seen in New England, or shifting mosaic conditions caused by climate changes and/or large-scale disturbance [102]. Changes in fire frequency have had an impact on the species composition and structure of many forests [15]. For further discussion, see FIRE ECOLOGY AND MANAGEMENT. Old-field Succession: Species composition and the successional role of species may vary geographically. Generally, the first 3 years following the last crop are dominated by herbaceous members of the Asteraceae such as fleabanes (Erigeron spp.), asters (Aster spp.), ragweeds (Ambrosia spp.), and goldenrods (Solidago spp.), and grasses such as crabgrasses (Digitaria spp.), bristlegrasses (Setaria spp.), threeawns (Aristida spp.), and panic grasses (Panicum spp.). In many instances, woody species such as persimmon (Diospyros virginiana), elms (Ulmus spp.), hackberries (Celtis spp.) and junipers seed in but are not dominant for a number of years. Broomsedge (Andropogon virginicus) is a common dominant from the fourth year on, and may be a component for over 20 years. On limestone soils, eastern redcedar is a major old-field inhabitant. Oak-hickory and dry open forests of post oak, junipers, blackjack oak, and black hickory eventually develop on at least 50 percent of the relatively dry uplands. Other forest types develop in mesic coves and in bottomlands [15]. Old-field succession occurred on abandoned fields in western Illinois following a period in which the area was heavily cultivated and suffered severe erosion, then converted to pasturage and hay fields. Early succession was dominated by cool-season grasses including Kentucky bluegrass (Poa pratense). Early establishing woody species included boxelder (Acer negundo) and elms. Trees with heavy seeds only later invaded the edges of fields. The most rapid tree development took place in gullies [16]. Savanna Vegetation: Depending on local conditions including fire regime, climate, and settlement patterns, very different descriptions of the same area were sometimes recorded within a few years [69]. The structure of most savannas is highly dependent on fire frequency; savannas are converted to more closed forest in less than 50 years without fire. Further discussion of the relationship of savanna and fire is in FIRE ECOLOGY AND MANAGEMENT [15]. Nuzzo [69] distinguished between open savanna and scrub savanna. Open savanna is a parklike community with widely spaced trees, virtually no shrub layer, and an herbaceous ground layer. Scrub savanna is made up of moderate to dense thickets of oak sprouts within a prairie matrix, with a few fairly dwarfed open-grown trees. Open savannas usually occur on flatter, usually mesic areas, and scrub savannas are generally located on the dry to dry-mesic areas of steeper topography, particularly hillsides, dunes, and ridges. Both kinds of savanna vary in structure through time and space, depending on fire occurrence [69]. The persistence of some high-quality savannas may be related to the droughty soils on which they occur. Some savannas, in particular those located on thin soil or rocky substrates, have survived moderate grazing, exclusion of fire, and competition with alien and woody species. Light grazing may have helped maintain the savanna. Some stands have survived because they have been burned relatively frequently [69]. The roots of savanna trees may show morphological responses to growing in dry soils; bur oak taproots are at least 13.2 feet (4 m) deep in dry soils but are not as deep in moist soils [103].

KUCHLER TYPE VALUE AND USE

KUCHLER TYPE: Oak-hickory forest
FORESTRY VALUES : Forests dominated by oaks comprise the largest type of commercial hardwood forest land in the United States. Collectively the red and white oaks comprise 38 percent of the total hardwood sawtimber volume in the United States. Oak wood is strong, hard, and tough. It has good working characteristics and is used extensively for furniture, flooring, paneling, ties, and cooperage. Oak manufacturing residues and low-grade stems not suitable for lumber have been increasingly used for pulp production [81,83]. RANGE VALUES : Livestock grazing is incompatible with timber production in most of the north-central states; forage production is low and cattle damage tree reproduction and compact forest soils. Grazing on Federal land in this region is minimal. Forest range suitable for grazing is grassland adjacent to or within forested areas, or savanna [64]. WILDLIFE VALUES : Oaks are important to wildlife species for both cover and food. Young oaks with branches close to the ground provide foliage browse long into the winter months, and often provide the only brushy cover in dense pole stands. Dried oak leaves are important in the winter diet of white-tailed deer in some areas. Different parts of oak trees are consumed by 186 different kinds of birds and mammals; the geographic distribution of many animals coincides with or depends on the range of oaks [84,94]. Acorn production is of primary importance to many birds and mammals. For example, in the Ozarks of Missouri acorns comprise 37 percent of the yearlong diet of wild turkey (Meleagris gallopavo) and 54 percent of white-tailed deer diets [84]. Many mammals consume acorns, including white-footed mouse, eastern chipmunk, eastern fox squirrel, eastern gray squirrel, red squirrel (Tamiasciurus hudsonicus), flying squirrels (Glaucomys spp.), and deer mouse (Peromyscus maniculatus). Acorns (particularly of northern red oak) are an important food for northern bobwhite (Colinus virginianus), red-headed woodpecker, red-bellied woodpecker, blue jay, tufted titmouse, common grackle, white-breasted nuthatch, sapsuckers (Sphyrapicus spp.), quail (Phasianidae), ruffed grouse, and various waterfowl including common golden-eye (Bucephala clangula), gadwall (Anas strepera), mallard (A. platyrhyncos), wood duck (Aix sponsa), hooded merganser (Lophodytes cucullatus, and others [57,94]. Black oak has high cavity value for wildlife [26]. Cavity nesters are an important component of oak-hickory forests. Cavity nesters comprise the 10 most frequent bird species in all 7 sampled vegetation types, most of which are oak-hickory forest types in the Missouri Ozarks [41]. Current land use patterns that decrease forest cover and increase herb cover result in an increased abundance of grassland bird species such as dickcissel (Spica americana) and horned lark (Eremophila alpestris) [15]. Many animals that depend on Illinois savanna or open woodlands are decreasing in abundance due to forest closure caused by fire exclusion. Examples include Kirtland's snake (Clonophis kirtlandii), Cooper's hawk (Accipiter cooperii), and silvery blue butterfly [72]. Sharp-tailed grouse (Tympanicus phasianellus) habitat in the eastern United States includes oak savanna and recently burned areas [33]. Nine species of terrestrial vertebrates on the Endangered and Threatened Wildlife and Plants list occur in oak-hickory forest: gray bat (Myotis grisescens), Indiana bat (M. sodalis), Ozark big-eared bat (Plecotus townsendii ingens), eastern cougar (Felis concolor couguar), Bachman's warbler (Vermivora bachmanii), bald eagle, peregrine falcon (Falco peregrinus ssp. anatum and F. p. ssp. tundrius), and Kirtland's warbler (Dendroica kirtlandii) [112]. Branson and others [11] listed 81 species of terrestrial vertebrates with ranges in oak-hickory forests in Kentucky that are considered threatened, endangered, of special concern, or of undetermined status because of lack of information. A summary of state lists of rare, threatened, or of special concern was prepared by Meredith [62] and included many species occurring in oak-hickory forests. OTHER VALUES : Forests in the north-central states offer a wide variety of recreational opportunities. All nine north-central states have both State and Federal forest-oriented recreation facilities. Activities include picnicking, camping, hiking, hunting, and fishing [64]. Forests in the north-central states provide habitat for most big game, and many small game animals including game fish. There were 26,400 deer, 474 bear, 151 moose, and 1,927 wild turkey taken on National Forest land in 1974 (National Forest land is only a small portion of the available hunting areas in this region) [64]. MANAGEMENT CONCERNS : Diverse vegetation is important for maintaining wildlife species diversity in oak-hickory forests. Mature stands are high in cavity and den value. Regeneration cuttings in even-aged stands create openings and edge habitat. Regeneration areas provide browse, sapling stands are good habitat for certain bird species, and pole, immature sawtimber, and sawtimber stands are good habitat, providing mast and cover for many species [11]. Removal of culls and snags from oak-hickory forest stands has detrimental impact on cavity nesting species. Even-age management of eastern deciduous forests creates cull-free, young, fast-growing stands that have very few cavities available to cavity nesting species. For high quality bird habitat clearcuts should be kept small and planned so that each management unit contains diverse stand age classes. Leaving dead snags and trees with heart rot during regeneration cuts and subsequent thinnings may maintain habitat for cavity nesting species. On medium-quality sites, killing unwanted trees and leaving them standing also provides habitat for cavity nesters [41]. Oak regeneration has become a subject of concern in oak-hickory forest management; in many areas oaks are being replaced after harvest by mesic hardwoods such as sugar maple [20,54,55,81]. Oak stands on sites with adequate soil moisture usually have high densities of mesophytic species in the understory, which often outcompete oak regeneration, even after timber harvesting [15,55]. In Wisconsin planted oak seedlings showed excellent (>90%) survival on plots in which tall (>5 feet [1.5 m]) understory vegetation was removed; on control plots where tall understory vegetation was left intact more than 70 percent of planted oak seedlings died within 5 years. Natural seedlings were also more abundant on opened plots [55]. Merrit [63] indicated that the present mature oak stands originated when woodlands were subjected to severe cutting, grazing, and fire; these conditions have largely been eliminated from current management practices [15]. Northern red oak was favored by frequent fires and/or heavy cutting because of its sprouting ability. In Indiana, however, both even- and uneven-aged silvicultural practices have converted many stands containing large proportions of oak sawtimber-sized trees to species such as sugar maple and yellow-poplar (Liriodendron tulipifera) [23]. Advance regeneration in oak-hickory forest is poor [19,20,23,82]. Underlying causes may be related to fire exclusion. Fire may have a beneficial influence on oaks by reducing competition from more fire-sensitive tree species in the sapling layer [54]. Fire reduces the amount of litter under a stand, which, according to Lorimer [54], may discourage rodent predation of acorns. Fire may indirectly influence rodent populations as well, by reducing available nest sites and food availability. Oak-hickory forests in the northeastern United States have notable pest problems. Gypsy moth larvae have caused widespread defoliation contributing to oak decline and mortality in many areas, and oak wilt is widespread in the central states and in some eastern states [83]. Butternut canker has caused widespread losses of butternut (Juglans cinerea), and dogwood anthracnose has caused serious losses throughout its range [17]. A widespread decrease in oak vigor and growth rate and an increase in mortality have recently been attributed to high atmospheric levels of sulfur dioxide and other pollutants [15]. In the central United States, savanna has declined drastically since settlement. In Wisconsin and other northern areas, most savanna trees were cut within 25 to 30 years after settlement. Savanna that was protected from fire developed into dense forest [15].

KUCHLER TYPE FIRE ECOLOGY AND MANAGEMENT

KUCHLER TYPE: Oak-hickory forest
FUELS, FLAMMABILITY, AND FIRE OCCURRENCE : Fuels and Flammability: In the Sand Prairie Scrub Oak Nature Preserve, Illinois, estimated fuel loads were highest in closed forest, moderate in savanna, and lowest in prairie. When blackjack oaks in savanna reach sufficient size, grass growth under their crowns is reduced and wind action sweeps away litter; this reduces the amount of fuel near the tree trunk and prevents fire from reaching the tree base. Following a spring prescribed fire, a study of patterns of fuel consumption around isolated blackjack oak savanna trees showed that fire never reached their bases. In closed forest, tree mortality was high, and trees experienced more scorch at stem bases than trees on the forest-prairie edge [5]. In hardwood stands in the north-central states, fire danger is high in both the spring and fall and low in the summer months when the vegetation is green [64]. At Indian Creek Nature Center, Iowa, prescribed fires were conducted in spring under moist conditions in grassland and adjacent oak-hickory woodlots. In the woods, fires were generally patchy and left large areas unburned [73]. Oak leaf litter affects the likelihood of fire and increase fire intensity [54]. Oak leaves crinkle up when they dry and are therefore held above the ground surface creating a loose, porous fuel bed that carries fire easily [54,73]. In contrast, leaves of locust (Robinia spp.), maple, and elm lie flat, stay damp, and do not burn well [73]. Oak leaf litter is easily carried by wind; burning leaves can ignite spot fires and increase fire spread. Dead leaves of black oak often remain on the tree, creating deep layers of dry fuels in young stands [54]. Wildfire hazard has increased due to fuels buildup and fire exclusion on the Hercules Glades Wilderness Area in southwestern Missouri since its designation as wilderness. Management-ignited fire and prescribed natural fire have been recommended [56]. Fire Occurrence: It is widely accepted that along the western boundary of the deciduous forest Native Americans set many fires that kept oak-hickory forests open, helped determine the grassland-forest boundary, and limited the range of maple-basswood forests [73,97]. Nearly every acre in the Midwest probably experienced at least occasional fire [73]. There are numerous reports of Native Americans burning Missouri savannas and grasslands on an annual basis, usually in the autumn after leaf-fall [22,56]. Northern Missouri was virtually fire-free soon after settlement; southern Missouri continued to be burned by early settlers, but fires were of lower severity than presettlement, aborigine-set fires. In 1936, the state Conservation Commission began a concerted effort to suppress fire throughout the state [22]. In the Missouri Ozarks, fire history of a post oak savanna growing on a ridge was reconstructed for 1734 to 1991 using tree ring analysis. The mean fire return interval from 1740 to 1850, the period of greatest fire frequency, was 2.8 years. After 1850, the fire return interval increased to 24 years [26]. In southern Wisconsin (the area mapped by Kuchler as oak savanna [K081] but containing virtually the same species as oak-hickory forests and bluestem prairie-oak-hickory mosaic), oak savannas were widespread and common in presettlement times and disappeared rapidly as soon as fires ceased. It was therefore estimated that presettlement fire frequency was very high and in some areas fire occurred annually [54]. In the north-central states, there were more than 11,000 fires reported between 1970 and 1972. Many fires were caused by debris burning (approximately 33%), many were incendiary (20%), and only a very few (1%) were started by lightning. More than 50 percent of the fires burned in hardwood stands [64]. Boerner and Cho [10] estimated fire return intervals for heavily forested sites in southeastern Ohio from 1923 to 1978. They calculated an average fire return interval of 643 years given current conditions. Average fire intervals of 900 years were estimated for the Shawnee National Forest, Illinois [54]. FIRE EFFECTS ON SITE : In Missouri seasonal rates of nonsymbiotic nitrogen fixation in surface soils of oak-hickory forests were measured on plots exposed to 30 years of annual or periodic prescribed fire. Fire treatments had no influence on fixation rates, which averaged 0.1 kg/ha/year, or on the proportion of samples showing nitrogen fixing activity [93]. In Missouri annual fire in oak-hickory forests may reduce nutrient availability and water infiltration. Burning significantly reduced quantities of extractable ammonium in the soil; annual burning resulting in lower amounts than burning every 4 years [92]. In Missouri, oak forests were prescribed burned annually or periodically (every 5th year) in spring. Annual fires were patchy and of low severity due to lack of surface fuels. Soils of annually burned plots became increasing compacted and had little or no earthworm and insect activity compared to unburned soils and litter. After seven annual fires, water infiltration rates were 4.5 times faster on unburned soils than on burned soils [73]. Arend [111] reported reductions in infiltration rates after prescribed fire for some soils under mixed upland oaks in Missouri. FIRE EFFECTS ON VEGETATION : Fire is an important factor in establishing and maintaining vegetation patterns in the Ozark Mountains [2,74]. The closed-canopy oak-hickory forest that now predominates in the Missouri Ozarks is thought by some authors to be an artifact of fire suppression and other anthropogenic disturbances. People travelling in the area in presettlement times described a wide variety of plant communities ranging from grassland, to oak savannas, to oak-pine forests [26]. Repeated fires create open, parklike stands of oak [90]. Drought is the primary agent controlling the frequency and severity of fire and the impact of fire on trees in oak-hickory forests. Drought contributes to low fuel moistures and therefore high fireline intensity, resulting in more severe fire. Individual trees stressed by drought are more susceptible to pathogens after fire wounding and sprout less vigorously after top-kill. Stands affected by insects or pathogens also have high amounts of fine fuels [17]. Species Responses: Most oaks are resistant to fire to some extent; fire resistance generally increases with stem diameter [54,78]. Most oaks sprout from dormant basal buds after top-kill. Sprouting species are favored over other hardwoods by occasional fires [23]. Boerner and Cho [10] hypothesized that large, emergent oaks in an old-growth forest in southeastern Ohio were established following one or more large-scale disturbances such as fire or windstorm. After oak establishment on good sites, severe fire maintains oak dominance [54]. Rouse [78] listed the following oaks in order of decreasing bark thickness: chestnut oak, black oak, northern red oak, and white oak. Lorimer [53] rated upland oaks slightly differently in order of decreasing bark thickness: bur oak, black oak, white oak, and northern red oak. The thinner bark of northern red oak may acount for its susceptibility to top-kill by fire. Bur oak and white oak are often maintained in oak savannas, whereas frequent fire reduces northern red oak to shrubby clumps of sprout origin. Black oak is a short-lived (less than 150 years) tree, intermediate in shade tolerance. It sprouts after injury or top-kill. Susceptibility to fire is intermediate in larger individuals and high in young seedlings and sprouts [42]. Both mature [110] and young black oak seedlings sprout [54]. Fire-caused basal wounding provides entry for wood-rotting fungi and insects [74]. Loomis [52] published a method for predicting tree mortality and estimating basal wound size for surviving trees. Mortality equations were derived for black oak, white oak, post oak, and scarlet oak. The black oak formula also applies to hickories and to northern red oak. Mortality depends on tree species, diameter at breast height, height of bark blackening, and the width of bark blackening at 1 foot above the ground [52]. Prescribed spring fire in grasslands and adjacent woods in Iowa resulted in a great increase in flowering of forbs, and reduced woody species such as common prickly-ash (Zanthoxylum americanum), black locust, barberry, buckthorns, and maples [73]. Survival of lichens after fire in a Michigan oak savanna was summarized by Wetmore [105]. Many species showed lower cover after fire on the part of tree trunks closest to the soil; cover reduction was less pronounced high on the tree trunks, even at high fire frequencies [105]. Effects of Fire on Species Composition: Fire influences species composition towards oak-hickory and away from maple-beech dominated forest [15]. After a severe wildfire in West Virginia, oaks increased from 9.3 percent of the dominant trees to almost 30 percent of the total, mostly due to top-kill of mesic hardwoods [54]. Frequent fire reduces tree and shrub cover in favor of prairie species. In the Driftless Area (Wisconsin and Minnesota), almost pure stands of northern red oak occur on moist sites that would be suitable for sugar maple. These are even-aged stands that probably established after severe fires [23]. In many oak-hickory forests, sugar maple and other shade-tolerant and fire susceptible species are increasing in number and cover. Evidence that this is a result of fire exclusion includes the tendency of sugar maple and other fire-susceptible species to co-occur on protected sites that show no evidence of past fire [67]. In southern Wisconsin oak forests (which experienced fire in presettlement times but have not burned recently), oak regeneration is scarce. McCune and Cottam [59] suggested that since none of the understory species in southern Wisconsin (hackberry, boxelder, American elm, butternut, black cherry, and red maple) are true climax species, the absence of disturbance is an historic anomaly and the future composition of these forests is unclear. Henderson [42] compared two black oak stands in Indiana with differing fire histories. The stand which had experienced frequent, low-severity surface fires was dominated by black oak and a few northern red oak, and there were few other woody species. The stand that had experienced more severe fire less frequently was also dominated by black oak, but had less fire-tolerant species present as well: red maple (Acer rubrum), sassafras (Sassafras albidum), and black tupelo (Nyssa sylvatica) [42]. In Illinois grassy openings with few trees were burned annually with prescribed fire from 1969 to 1973. They were not burned again until 1990, 1991, and 1992. With annual fire, prairie forbs increased and woody species declined. In the 15-year fire-free interval, the number of shrub species increased. When prescribed fire was resumed, prairie willow (Salix humulis) and silky dogwood (Cornus amomum) sprouted vigorously after fire; prairie forbs also increased [3]. Effects of Fire on Stand Structure: Fire was important in maintaining oak savannas at the forest-prairie ecotone. There are numerous historical references to the rapid conversion of savanna to oak forest following European settlement and cessation of fire [23]. Anderson and Brown [5] proposed that the mosaic of presettlement vegetation types was strongly influenced by fire patterns that tended to limit closed forests to areas protected from most fires. In Tennessee annual and periodic prescribed surface fires were conducted in upland oak-dominated vegetation. Upland stands were mostly post oak-blackjack oak, southern red oak-scarlet oak, or white oak [30]. After 25 years, understory cover of woody plants was irregular but more or less constant in annually burned plots. Overstory cover was open, with few trees per acre. In periodically burned plots, woody understory cover increased irregularly. In unburned control plots, woody understory plant cover first decreased but then increased over the 25-year period [29]. In presettlement Indiana grass-dominated openings were maintained by annual burning after autumn frost. These frequent fires prevented the growth of trees and "permanent vegetation". In less than 50 years without fire, former openings were converted to forest [49]. Repeated fires are thought to be responsible for scrub oak lands (stands of scrubby oaks of sprout origin) in the Midwest [23]. In the Sand Prairie Scrub Oak Nature Preserve, a March 1977 prescribed fire in closed forest consumed 95 to 99 percent of the surface litter in study plots. Numerous trees had fire scars up to 3.3 feet (1 m) above the ground. In the savanna the fire did not reach within 1 foot (30 cm) of the base of any of the 14 trees. Surface area burned ranged from 7.1 percent to 52.4 percent in different plots. None of the savanna trees showed any deterioration in 3 years after the fire even though some had scorching of the lower branches [4]. Black oak sand savannas at Hoosier Prairie Nature Preserve, Indiana, are maintained by prescribed fire. One area has been burned four times in 8 years, three times in the spring, and once in the fall. The number of woody stems increased after the first fire, but decreased thereafter. Postfire sprouting of smooth sumac occured at the same rate through four fires, but black oak and prairie willow sprouting rates declined. Another area of oak savanna at Hoosier Prairie was burned five times in 8 years. Allowing plots to rest from fire for least 2 years resulted in an increase in the numbers of woody stems, but the number of woody stems decreased after two consecutive fires [7]. FIRE EFFECTS ON RESOURCE MANAGEMENT : Few instances of direct mortality from fire have been recorded for wildlife species in oak-hickory forests. In Oklahoma, Bigham and others [9] observed many dead eastern box turtles immediately after severe fires in post oak-blackjack oak woods and adjacent pasture and range. They also observed living turtles, snakes, northern bobwhites, and a rabbit on the burned area 4 days after the fire [9]. FIRE USE CONSIDERATIONS : Moore [66] published a method for field measurement of fuel moisture to assess burning conditions in oak-hickory forests. FIRE MANAGEMENT CONSIDERATIONS : Fire was once an important factor in the oak-hickory ecosystem and maintained the open character of most stands in the Midwest [15,54]. Many historians and ecologists feel that the midwestern oak-hickory forests evolved with frequent fire and remain fire-dependent [73]. Prescribed fire is useful for maintaining the diversity of oak-hickory forests, and is being used in Indiana, Illinois, Missouri, Wisconsin, and Iowa with good results. Prescribed fire use continues in hardwood and mixed forests as a wildlife management tool [106] and to modify understory composition or size class structure [35,99]. Unfortunately, few short- or long-term ecological studies of fire perturbations have been conducted on oak-hickory forests [15]. Implications for fire in management of currently existing closed-canopy forests are unclear. Regeneration problems associated with these forests are similar to problems occurring in forests farther east. Prescribed fire is being considered as a silvicultural tool in oak forests. It seems likely that several low- to moderate-severity prescribed fires will reduce the number of shade-tolerant competitors, and it is possible that such fires may encourage establishment of oaks. A single fire will have little lasting impact, and in some cases may be detrimental to oak regeneration. Several fires are probably necessary to select against less fire-resistant competitors [54]. Lorimer [54] commented that experiments with medium-severity fire are to be encouraged; low-severity fire may not kill competing species at rates high enough to be beneficial to oak regeneration. In some areas silvicultural approaches may be as least as successful as prescribed burning programs [54] because basal wounding by fire allows entry of fungi and insects and reduces the value of timber trees [74]. In an attempt to improve oak regeneration in an upland hardwood forest in northern Alabama, a prescribed fire was conducted in 4- to 6-year-old oak stands that appeared to be in danger of replacement by other species. One fire in these stands did not substantially alter the trend to replacement of oaks by other species. Stands recovered rapidly from fire; density of browse species and herb cover increased, resulting in improved wildlife habitat [45]. With increasing age of forests, regeneration of oaks after disturbance tends to decrease. Sprouting tends to decrease with age. Established shade-tolerant species in the understory and subcanopy are more competitive than low-vigor oak sprouts [15]. In the Midwest many savanna and grassland areas have been prescribed burned on a regular basis. These fires have been prevented from spreading into adjacent forests, with the result that there are no transitional areas. Under presettlement conditions, grassland fires sometimes spread into adjacent oak-hickory forests, depending on local conditions. These fires maintained savanna and ecotonal communities in irregular mosaics. It has been recommended, therefore, that prescribed fires for grassland maintenance be allowed to burn into adjacent forest and reestablish savanna vegetation and ecotones [73]. In Indiana fire management policies are based on the premise that oak savanna and prairie communities will best be maintained by a fire interval of 2 to 8 years, depending on unit history and specific goals. Annual fires are not recommended for either restoration or maintenance of oak savanna because there is usually not enough fuel to carry fire, and annual fires may damage some herbs. Remnants of oak savanna prairie need to be burned at least every 15 years or they will develop into dense forest. Spring fires are thought to be the most effective [32]. In Indiana black oak sand savannas, maintenance and restoration of vegetational mosaics was initiated with prescribed fire. Plots were burned biennially, three times in the spring and once in the fall, for a total of four times in 8 years [7]. On the Grand Prairie, Illinois, U.S. Forest Service personnel are attempting to restore the transition landscape (as savanna) by allowing firewood removal until approximately 10 trees are left per acre (25 trees/ha); tree removal is followed by annual prescribed fire [58]. There has been some debate over the choice of sites and appropriate methodology for restoration of presettlement vegetation including savannas [60,72]. Mendelson and others [60] contend that savanna vegetation represents an ecotonal and therefore not unique vegetation type and should not be created by artificial means. Packard [72] and his colleagues believe that, since the presettlement vegetation has been grossly disturbed by humans, drastic measures (i.e., prescribed fire and tree thinning) are necessary to restore savanna and other presettlement vegetation types. REHABILITATION OF SITES FOLLOWING WILDFIRE : NO-ENTRY

REFERENCES

KUCHLER TYPE: Oak-hickory forest
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