|
|
FEIS Home Page |
SPECIES: Quercus alba
SPECIES: Quercus alba Introductory
AUTHORSHIP AND CITATION : Tirmenstein, D. A. 1991. Quercus alba. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: https://www.fs.usda.gov/database/feis/plants/tree/quealb/all.html [].
ABBREVIATION : QUEALB SYNONYMS : NO-ENTRY SCS PLANT CODE : QUAL COMMON NAMES : white oak stave oak ridge white oak forked-leaf white oak fork-leaf oak TAXONOMY : The currently accepted scientific name of white oak is Quercus alba L. [69]. It is a member of the order Fagales and has been placed within the white oak subgenus (Lepidobalanus) [16]. Three varieties of white oak are commonly recognized [148]: Quercus alba var. alba Quercus alba var. repanda Michx. Quercus alba var. latiloba Sarg. Some authorities recognize these entities as forms rather than varieties [108,117,131]. White oak is highly variable genetically [58], and many forms and ecotypes have been described. According to Fowells [47], "no definite races have been defined, but within such a tremendously diverse habitat, climatic races undoubtedly exist." White oak readily hybridizes with many other species within the genus Quercus [58], including swamp white oak (Q. bicolor), bur oak (Q. macrocarpa), chinkapin oak (Q. muehlenbergi), dwarf chinkapin oak (Q. prinoides), overcup oak (Q. lyrata), swamp chestnut oak (Q. michauxii), sandpost oak (Q. margaretta), chestnut oak (Q. prinus), English oak (Q. robur), Durand oak (Q. durandii), and post oak (Q. stellata) [58,83]. Hybrids, their common names, and purported origins are listed below [69,148]. Beadle oak X beadlei Trel. (Quercus alba x michauxii) Bebb oak X bebbiana (Q. alba x Q. macrocarpa) X bimundorum Palmer (Q. alba x Q. robur) Deam oak X deamii (Q. alba x Q. muehlenbergi) Faxon oak X faxonii Trel. (Q. alba x Q. prinoides) Fernow oak X fernowii Trel. (Q. alba x Q. stellata) Jack oak X jackiana Schneid. (Q. alba x Q. montana) Saul oak X saulii Schneid. (Q. alba x Q. prinus) Saul oak was formerly known as Q. alba f. ryderii but is now considered a heterozygous hybrid form of white oak [3]. Introgressive populations are locally common throughout much of the range of white oak. Hybrid swarms derived from complex mixtures of parental forms are particularly common on disturbed sites, at the margins of white oak's range, and where several oak species occur sympatrically [58]. LIFE FORM : Tree FEDERAL LEGAL STATUS : No special status OTHER STATUS : NO-ENTRY
SPECIES: Quercus alba DISTRIBUTION AND OCCURRENCE
GENERAL DISTRIBUTION : White oak grows throughout much of the eastern United States from southwest Maine to northern Florida, Alabama, and Georgia [53,83,148]. It extends westward throughout southern Ontario and Quebec into central Michigan, northern Wisconsin, and southeastern Minnesota and south to southwestern Iowa, eastern Kansas, eastern Oklahoma, and eastern Texas [55,83]. Little [83] reported that white oak may have been eliminated from southeastern Nebraska. The best growing conditions for white oak occur on the western slope of the Appalachian Mountains and in the Ohio Valley and central Mississippi Valley [148]. White oak is mostly absent from conifer-dominated stands at higher elevations within the Appalachian Mountains and from the lower Mississippi Delta and coastal areas of Texas and Louisiana [148]. The variety latiloba occurs at the northern edge of the species' range [47]. The range of var. repanda is poorly documented, but it has been reported in parts of New England [117]. ECOSYSTEMS : FRES10 White - red - jack pine FRES12 Longleaf - slash pine FRES13 Loblolly - shortleaf pine FRES14 Oak - pine FRES15 Oak - hickory FRES16 Oak - gum - cypress FRES18 Maple - beech - birch FRES19 Aspen - birch STATES : AL AR CT DE FL GA IL IN IA KS KY LA ME MA MI MN MS MO NE NH NJ NY NC OH OK PA RI SC TN TX VT VA WV WI ON PQ BLM PHYSIOGRAPHIC REGIONS : NO-ENTRY KUCHLER PLANT ASSOCIATIONS : K081 Oak savanna K089 Blackbelt K095 Great Lakes pine forest K100 Oak - hickory forest K103 Mixed mesophytic forest K104 Appalachian oak forest K111 Oak - hickory - pine K112 Southern mixed forest K113 Southern floodplain forest SAF COVER TYPES : 14 Northern pin oak 15 Red pine 19 Gray birch - red maple 21 Eastern white pine 22 White pine - hemlock 23 Eastern hemlock 26 Sugar maple - basswood 27 Sugar maple 40 Post oak - blackjack oak 42 Bur oak 43 Bear oak 44 Chestnut oak 45 Pitch pine 46 Eastern redcedar 51 White pine - chestnut oak 52 White oak - black oak - northern red oak 53 White oak 55 Northern red oak 57 Yellow poplar 58 Yellow poplar - eastern hemlock 59 Yellow poplar - white oak - northern red oak 60 Beech - sugar maple 61 River birch - sycamore 64 Sassafras - persimmon 65 Pin oak - sweetgum 75 Shortleaf pine 76 Shortleaf pine - oak 78 Virginia pine - oak 80 Loblolly pine - shortleaf pine 81 Loblolly pine 82 Loblolly pine - hardwood 91 Swamp chestnut oak - cherrybark oak 110 Black oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : White oak grows as a dominant in many communities and as a major species in several cover types [95,96]. Common codominants within the overstory include northern red oak (Quercus rubra), scarlet oak (Q. coccinea), northern pin oak (Q. ellipsoidalis), black oak (Q. velutinus), beech (Fagus spp.), sweetgum (Liquidambar styraciflua), chestnut (Castanea dentata), red maple (Acer rubrum), sugar maple (A. saccharum), and hickories (Carya spp.). Understory dominants or codominants include deerberry (Vaccinium stamineum), leadplant (Amorpha canescens), trailing arbutus (Epigaea repens), huckleberries (Gaylussacia spp.), meadow-rue (Thalictrum spp.), and false Solomon's-seal (Smilacina racemosa). Published classifications listing white oak as an indicator or dominant in habitat types (hts) are presented below: Area Classification Authority AL general veg. cts Golden 1979 s IL general veg. cts Fralish 1976 IN general veg. cts Keith 1983 ne IA general veg. cts Cahayla-Wynne & Glenn-Lewin 1978 MI general veg. cts Hammitt and Barnes 1989 general veg. eas Pregitzer and Ramm 1984 n MI, ne WI forest hts Coffman and others 1980 sw OH general veg. cts Braun 1936 e TN general veg. cts Martin and DeSelm 1976 n WI forest hts Kotar and others 1988 Smoky Mtns general veg. cts Whittaker 1956
SPECIES: Quercus alba MANAGEMENT CONSIDERATIONS
WOOD PRODUCTS VALUE : White oak wood is heavy, hard, strong, and durable [131]. When properly dried treated, oak wood glues well, machines very well and accepts a variety of finishes [97]. White oak is the most important timber oak and is commercially important throughout much of the South and East [35,141,148]. White oak is an important source of wood for furniture, veneer, paneling, and flooring [28,95,101]. It has been used to make railroad ties, fenceposts, mine timbers, ships, and caskets [95]. White oak has long been used in cooperage [125] and is currently the major source of wood for whiskey barrels [43]. White oak wood has also been used as a source of clapboard shingles and woven baskets, although demands for these products are decreasing [43]. Its high fuel value makes white oak an attractive firewood [95]. IMPORTANCE TO LIVESTOCK AND WILDLIFE : Browse: The young shoots of many eastern oak species are readily eaten by deer [57]. Dried oak leaves are also occasionally eaten by white-tailed deer in the fall or winter [120]. Rabbits often browse twigs and can girdle stems [57]. The porcupine feeds on the bark, and beavers eat twigs of white oaks [135]. Acorns: Acorns of white oak are considered choice food for many wildlife species [118], including the white-footed mouse, fox squirrel, black bear, pine mouse, red squirrel, and cottontail rabbits [22,27,135]. The gray squirrel consumes white oak acorns but prefers the acorns of other oak species [80]. Many birds, including the bluejay, northern bobwhite, mallard, ring-necked pheasant, greater prairie chicken, ruffed grouse, and wild turkey, eat white oak acorns [60,66,135]. In some areas, the abundance of fall mast crops, such as acorns, can affect black bear reproductive success during the following year [44]. Sprouted acorns are often eaten by deer, mice, and bobwhite [135]. PALATABILITY : The palatability of oak browse is relatively high for domestic livestock and for many wildlife species [135]. Eastern oaks are preferred by white-tailed deer in some locations [135]. New growth is particularly palatable to deer and rabbits [57]. The acorns of most oaks are highly palatable to many species of birds and mammals [57,90]. Palatability of white oak acorns to fox squirrels, and presumably to some other species, declines after the acorns have sprouted [123]. NUTRITIONAL VALUE : Browse: The nutritional value of white oak browse varies geographically, and with site history and phenological development. Annual variation has also been observed [40]. Foliar nitrogen content was measured at 1.40 percent in Tennessee but averaged only 0.7 percent in New York [114]. The calcium content of leaves tends to increase slowly as the growing season progresses [15]. Calcium levels of twigs, and protein and phosphorus content of the foliage, are generally higher on recently burned sites [12,15,40]. Total solids, ash, ether extract, crude fiber, and N-free extract appear to be unaffected by fire [40]. [see Fire Management Considerations]. Winter nutrient content of white oak browse in Texas has been documented as follows [78]: protein fat fiber N-free ash phosphoric Ca extract acid percent at 15 percent moisture 3.89 1.46 34.22 42.43 3.00 0.13 1.67 Acorns: Acorns are nutritious [57] and high in carbohydrates [59]. White oak acorns are relatively low in protein, crude fiber, and potassium [16,123,142] but high in digestible cell contents such as fats, starches, sugars, and pectins [123]. White oak acorns tend to be lower in fats than the acorns of many other oak species [124]. Primary stored energy reserves are in the form of carbohydrates [16]. Specific nutritional values are reported below [123,124]: crude crude crude Si Ca P ash N-free protein fat fiber extract percent dry weight - 5.9 4.3 18.7 0.01 0.15 0.09 --- --- 4.6 5.8 18.6 0.06 0.18 0.09 2.7 68.3 Tannin levels of white oak acorns are relatively low, generally ranging from 0.5 to 2.5 percent [126]. Lipid concentrations are also low, averaging 5 to 10 percent [126]. However, Lewis [80] reported tannin and lipid levels of 2.94 and 4.6 percent, respectively. Metabolizable energy content has been estimated at 72 percent [80]. Taproot: The taproot of white oak is high in fibers, lignin, and cellulose [126]. COVER VALUE : White oak provides good cover for a wide variety of birds and mammals. Oak leaves often persist longer than many other plant associates and in some areas, young oaks may represent the only brushy winter cover in dense pole stands [120]. Oaks frequently serve as perching or nesting sites for various songbirds [29]. The well-developed crowns provide shelter and hiding cover for small mammals such as tree squirrels. Many birds and mammals use twigs and leaves as nesting materials [90]. Large oaks provide denning sites for a variety of mammals [29]. VALUE FOR REHABILITATION OF DISTURBED SITES : White oak is potentially valuable for use in reforestation projects [79] and appears to have potential for use on other types of disturbed sites. It has been planted on strip-mined lands in Ohio, Indiana, and Illinois [6,23,81] and has exhibited good growth and survival on cast overburden and graded topsoil overlying mine spoils [6,139]. It is well adapted to loamy and clayey spoils with a pH of 5.5 to 8.0 [81]. White oak is difficult to transplant and grows slowly [148]. It can be readily propagated through seed which is generally planted in the fall [99]. Seed collection, storage, and planting techniques have been documented [16,99]. OTHER USES AND VALUES : Acorns were traditionally an important food source for many Native American peoples [134]. White oak acorns have been described variously as sweet and edible [131] and as slightly bitter [43]. The acorns were often boiled to remove bitter tannins [35]. Oils obtained from pressed acorns were used to alleviate pain in the joints [63]. White oak is commonly used in landscaping [125] and is often planted as a shade tree or ornamental [43,148. Its colorful purplish-red to violet-purple foliage enhances its ornamental value in autumn [125,148]. White oak was first cultivated in 1724 [99]. OTHER MANAGEMENT CONSIDERATIONS : Chemical control: Oaks often produce basal sprouts in response to herbicide treatments [50]. Damage: White oak can be damaged by frost or drought. It is also sensitive to periodic flooding [64]. Environmental considerations: White oak is sensitive to excessive ozone [64]. Grazing: Intensive grazing can reduce the number of trees present and aid in the regeneration of white oak through seed [8]. Wildlife considerations: Acorns are a particularly important food source for black bears in many areas. Acorn crop failures have been correlated with increases in damage to crops, livestock, and beehives by bears [112].
SPECIES: Quercus alba BOTANICAL AND ECOLOGICAL CHARACTERISTICS
GENERAL BOTANICAL CHARACTERISTICS : White oak is a medium to large, spreading, deciduous tree which commonly reaches 60 to 80 feet (18-24 m) in height [31,53,131]. On favorable sites, individuals may grow to more than 100 feet (30 m) in height and exceed 5 feet (1.5 m) in diameter [19,108]. White oak is slow-growing and long-lived (up to 600 years) [35]. White oak is monoecious [131]. Yellowish staminate catkins are borne at the base of new growth, whereas reddish pistillate catkins grow in the axils of new growth [119,131,148]. The short-stalked, glabrous, ovoid acorns are tan to brown [31,53,108]. Acorns are generally borne in pairs [31]. The rough, warty cup covers approximately 33 to 50 percent of the nut [31,55,131]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : White oak reproduces through seed and by vegetative means. Both modes of regeneration appear to be important. Seed: White oak produces good acorn crops at erratic intervals. Good crops have been reported at 4- to 10-year [148] and at 3- to 5-year intervals [118]. Vigorous crowned trees greater than 20 inches d.b.h. (51 cm) generally produce the best seed crops [57]. Pollen, which is produced in abundance, is dispersed by wind, but generally travels less than 656 feet (200 m) [46,58]. Plants generally bear fruit between 50 and 200 years of age, but open-grown trees on good sites may produce seed as early as 20 years of age [99,148]. Reproduction from seed can occur when (1) large seed trees are present within 200 feet (61 m), (2) litter cover is moderate, and (3) the site receives at least 35 percent of full sunlight [148]. Seeds of white oak do not store well [16]. Seed longevity is less than 1 year; white oak is not considered a seed banker [60]. Viability in storage declines from 90 percent for fresh seed to 7.0 percent for seed stored for 6 months [16]. Only 14 to 18 percent of the total seed produced may be sound [148]. Many acorns are damaged or destroyed by insects [144] or bird and mammal seed predators. Several studies have reported that animals consumed 72 to 83 percent of all white oak acorns [135]. In years of poor acorn production, the entire seed crop may be eliminated [148]. Acorn production: Acorn production varies annually with the individual tree or stand [148]. Certain trees tend to produce larger acorn crops on a consistent basis [119]. Weather conditions, and tree size and vigor, also influence acorn production. An individual oak 69 feet (21 m) tall with a d.b.h. of 25 inches (63.5 cm) produced more than 23,000 acorns in a favorable year [148]. However, most forest-grown trees produce less than 10,000 acorns annually. Annual yields may range from 0 to 202,000 acorns per acre (500,000/ha) [148]. Acorn production may be reduced by cool April temperatures [119] and drought [118]. Seed dispersal: In parts of Michigan, the blue jay is the primary dispersal agent of white oak [60]. Blue jays commonly exhibit a preference for burying acorns in bare open areas which are well suited for germination [60]. Gray squirrels are also important dispersal agents in many locations and are the only known long-distance disperser [35,148]. The now-extinct passenger pigeon may have effected long-distance dispersal of many eastern oaks [21]. Wind and gravity also aid in seed dispersal [148]. Germination: White oak acorns do not exhibit dormancy [16]. In storage, seeds germinate readily at temperatures of 33 to 37 degrees F (1-3 deg C) [16]. Under natural conditions, acorns begin germinating soon after they fall [35]. Acorns require a cover of litter for good germination and seedling establishment [86]. Acorns without such protection are often damaged or killed by frost or drought [86]. Germination capacity ranges from 50 to 99 percent [148]. Seedling establishment: Seedling establishment is generally limited to years of abundant acorn production [148]. Light to moderate litter cover and periods of full sunlight are required for establishment. Establishment is best on loose soils [30]. Vegetative regeneration: White oak exhibits a number of modes of vegetative regeneration. Vigorous sprouting from the stump or root crown is commonly observed after fire, mechanical damage, and other types of disturbance. Sprouting generally decreases with increasing stem diameter [64], although trees up to 80 years of age occasionally retain the ability to sprout [42]. Small poles, saplings, and even seedlings sprout readily if cut or burned [51]. Stump-sprouting by diameter class has been reported as follows [99]: d.b.h. (inches) percent of stumps likely to sprout 2 to 5 80 6 to 11 50 12 to 16 15 16 + 0 Repeated sprouting is commonly observed [125]. Seedlings often develop an "s"-shaped curve at ground level, which helps protect dormant buds from fire [100]. Root stools develop under the ground surface after repeated fires or herbivory. These root stools, made up of callus tissue filled with dormant buds, typically sprout vigorously in the absence of further disturbance [100]. Seedling sprouts persist beneath the forest canopy even in the absence of disturbance. Although the top dies back every few years, the root system continues to develop and plants may persist for up to 90 years or more [99]. As the forest canopy is opened, the seedling sprouts grow rapidly [99]. Epicormic branches or water sprouts often develop from dormant buds located on the boles [16,23]. Buds are stimulated to sprout by sudden shifts in light intensity, partial removal of the crown, and a loss of plant vigor [16]. Bud dormancy in oaks is largely controlled by auxins rather than by levels of carbohydrate reserves [125]. Apical dominance can restrict the development of belowground buds when buds survive on aboveground portions of the plant. Sprouting is reduced by low light levels [125] and decreases as the stand ages [82]. McIntyre [82] reported that the number of sprout groups decreases from poor to good sites. Silviculture: Oaks often regenerate poorly after timber harvest. Hannah [51] reported that the use of natural seedbeds and standard silvicultural practices are often ineffectual in promoting oak regeneration. The presence of vigorous advanced regeneration is essential for producing good stands of oaks after timber harvest [29,88,102]. For adequate regeneration of oaks, advanced regeneration of at least 4.5 feet (1.4 m) in height should number at least 435 per acre (1,074/ha) prior to harvest [99,102]. A series of selection cuts can produce stands with several age classes and can generate sufficient advanced regeneration for well-stocked postharvest stands. Initial cuts should reduce overstory densities to no less than 60 percent stocking [102]. Reduction of competing understory species may also be necessary in some instances [102]. Mechanical treatment: Sprouts tend to be larger and taller when white oaks are cut during the dormant season [64]. Sprout growth by season has been reported in detail [64]. SITE CHARACTERISTICS : White oak grows in rich uplands, moist bottomlands, along streams, on hammocks, sinks, sandy plains, and on dry, gravelly slopes [17,28,30,99,116]. It occurs on all upland aspects, and slope positions [99], but in the southern Appalachians, it exhibits best growth on northern lower slopes and in coves [32]. White oak is absent on ridgetops with shallow soil, on poorly drained flats, and on very wet bottomlands [99]. Latitude, aspect, and topography are important factors influencing the distribution of white oak within its range [99]. White oak grows in a variety of dry to mesic woodland communities [131] including pine-oak-hickory woods, beech-maple, and mixed hardwood forests [30,131]. It also occurs in relatively open post oak savanna [110] and oak savanna codominated by bur oak [8]. Plant associates: White oak grows in pure or mixed stands in the Southeast [38] but towards the northern portion of its range it rarely occurs in pure stands [57]. Important tree associates are numerous and include beech (Fagus grandifolia), sugar maple, black cherry (Prunus serotina), white ash (Fraxinus americana), yellow poplar, shortleaf pine (Pinus echinata), loblolly pine (P. taeda), eastern white pine (P. strobus), jack pine (P. banksiana), eastern hemlock, sweet gum, black gum (Nyssa sylvatica), American basswood (Tilia americana), shagbark hickory (Carya ovata), and other hickories (Carya spp.) [28,57,83,110]. Scarlet oak, post oak, bur oak, black oak, and northern red oak are also important associates [99], Upland oaks and hickories are the most common associates [99]. Many herbaceous species grow in association with white oak. Climate: White oak is often associated with a cool, temperate, continental climate [12] but can grow under a variety of climatic regimes [99]. Mean average temperatures range from 45 degrees F (7 deg C) in the North to 70 degrees F (21 deg C) in eastern Texas and northern Florida [32]. Annual precipitation averages 80 inches (203 cm) in the southern Appalachians but is less than 30 inches (77 cm) in southern Minnesota [99]. Growing season length ranges from 5 months in the North to 9 months in the South [99]. Soils: White oak grows on a wide variety of soils [28] derived from many types of parent materials [42]. It grows on silty loam, clay loam, silty clay loam, fine sand, and loamy clay [12,43,110] but grows best on deep, well-drained loamy soils. Low soil-nutrient levels limit growth of white oak only on sandy soils [99]. White oak is common on rocky soils [116]. Elevation: White oak grows from sea level to 5,900 feet (0-1,800 m) [38]. In the North, it generally grows under 500 feet (152 m) in elevation, but in the southern Appalachians, it grows as a "scrub tree" at 4,500 feet (1,372 m) [99]. It is absent from higher elevations in the northern Appalachians. In the Smoky Mountains, two populations are separated by an elevational gap of 1,000 feet (305 m) [130]. White oak grows below 2,000 feet (610 m) in the Cumberland Mountains [130]. SUCCESSIONAL STATUS : White oak readily regenerates after disturbances such as fire or logging and often assumes prominence in mid to late seral stages [2,61]. In the North, white oak is commonly seral to sugar maple and other species characteristic of mixed mesophytic stands [42]. In much of its range, it is succeeded by beech and other shade-tolerant species on well-drained second bottoms and in protected coves [99]. White oak is a pioneer on frequently burned sites in southern Wisconsin [8], and in Michigan, readily colonizes agricultural land 15 years after abandonment [54]. In much of the eastern deciduous woodlands, forests formerly dominated by white oak, beech, red maple, yellow poplar, and northern red oak are now being replaced by more shade-tolerant species such as sugar maple and American basswood [8,91]. White oak cannot regenerate successfully beneath a dense canopy and in many areas, grows in forests transitional to climax sugar maple or mixed mesophytic forests [2,34]. Because of the longevity of white oak, climax development proceeds very slowly [2]. White oaks may persist on exposed sites within climax stands [8]. White oak is considered a climax tree in oak-hickory stands in the central and southern hardwood forest zone [99]. It grows as a climax dominant or codominant on certain lower elevation sites in the Smoky Mountains [130] and occurs in climax pine-oak forests of New Jersey [77]. It also assumes importance in climax floodplain oak-hickory forests of Tennessee [107]. White oak is represented in mixed hardwood old growth stands of northwestern Ohio [12]. Old-growth oak-hickory forests of southern Michigan [50], and in old-growth oak communities of eastern Tennessee [81]. Pine-oak forests cyclically replace beech-magnolia forests after disturbance in parts of southeastern Texas [47] and Louisiana. SEASONAL DEVELOPMENT : Leaves begin to develop and new shoots are initiated in mid-March to late May, depending on geographic location [99]. The timing of bud break is largely dependent on latitude [99] but also depends on soil nutrient levels [18] and weather. Bell and others [18] observed delayed budbreak on copper, lead, and zinc-mineralized sites. Most vegetative growth takes place during the spring, with up to 50 percent of seedling height growth attained in April [99]. Fowells [42] reported that seedling height growth was 90 percent complete by July 1. Plants may become dormant in late fall, although leaves commonly persist into winter [28]. Flowering generally occurs in spring when the new leaves are elongating [32] but varies according to latitude, weather conditions, and with the genetic composition of individual trees [99,104,105]. Flowering can occur from late March to May [99] or June [103]. In Pennsylvania, pistillate catkins emerge in late April or May, approximately 5 to 10 days after the emergence of staminate flowers [105]. Sharp and Chisman [104] observed trees within the same population flowering early (May 5 to May 11) and late (May 13 to May 19). Three distinct waves of flowering (early, middle, and late) have been reported. Warm weather speeds up floral development, which begins after exposure to minimum temperatures of 50 degrees F (10 deg C) for at least 10 days [104]. Pollen is generally shed within 3 days, but light winds can accelerate shedding [104]. Pollen shedding is often delayed by prolonged rainy weather [104]. Acorns typically ripen approximately 120 days after pollination [99]. In Pennsylvania, embryos generally begin development after July 24, grow rapidly by August 4, and reach full size by August 25 [105]. Acorns fall from the trees by September or October [99,105]. Generalized flowering and fruit ripening dates by geographic location are as follows: Location Flowering Fruit ripe Authority PA April-May ---- Sharp and Sprague 1967 NC, SC April Sept.-Nov. Radford and others 1968 New England May 21-June 3 ---- Seymour 1985 Blue Ridge Mtns. April-May ---- Wofford 1989 Adirondack Mtns. May Sept. Chapman & Besette 1990 nc Great Plains May Oct. Stephens 1973 WV ---- Oct. 3 Park 1942
SPECIES: Quercus alba FIRE ECOLOGY
FIRE ECOLOGY OR ADAPTATIONS : White oak is unable to regenerate beneath the shade of parent trees and relies on periodic fires for its perpetuation. The exclusion of fire has inhibited white oak regeneration through much of its range [121]. Following fire, white oak typically sprouts from the root crown or stump. Some postfire seedling establishment may also occur on favorable sites during favorable years. Northeast and central states: Fire has played an important role in deciduous forests of the eastern United States [100,128]. Evidence suggests that most oaks (Quercus spp.) are favored by a regime of relatively frequent fire. Many present-day oak forests may have developed in response to recurrent fire. Declines of oak forests have been noted throughout much of the East and are often attributed to reduced fire frequency [2,7,100]. The Southeast: Fire was also a major influence in presettlement forests of the Southeast [121,123]. In the southern Appalachians, many present-day oak stands may have developed 60 to 100 years ago with widespread burning associated with agricultural activities or timber harvest. Increased fire suppression has evidently favored more shade-tolerant hardwoods and resulted in a decrease in oaks [123]. Oak savannas: White oak formerly assumed importance in open oak savannas of Wisconsin and Iowa, but with increased fire suppression, fire-tolerant species such as white oak are being replaced by sugar maple and other more shade-tolerant species. Many open savannas are being converted to dense, forested stands [19,37]. FIRE REGIMES : Find fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find Fire Regimes". POSTFIRE REGENERATION STRATEGY : survivor species; on-site surviving root crown or caudex survivor species; on-site surviving roots off-site colonizer; seeds carried by animals or water; postfire yr1&2
SPECIES: Quercus alba FIRE EFFECTS
IMMEDIATE FIRE EFFECT ON PLANT : White oak is moderately resistant to fire [44,59]. Aerial portions may be killed by fire [11], but underground regenerative structures protected by overlying soil usually survive [10,76]. The rough, scaly bark of white oak is more fire-resistant than the solid bark of many other oaks [114]. Oaks typically become more fire resistant as the bark thickens with age [51]. Most oaks will survive periodic fires. In parts of the New Jersey Pine Region, most white oaks 25 years and older possessed fire scars; four fire scars were observed on a 65-year old tree [76]. However, frequent fires can damage or kill white oaks, and recurrent fires at less than 8-year intervals could eliminate white oak [77]. Approximately 76 percent of white oaks were killed following fire in a loblolly pine stand in Virginia [4], and an estimated 56.5 percent were killed after a fire in a New Jersey pine-oak community [106]. Many of the observed differences in susceptibility of oaks to fire can be attributed to variation in fire severity and intensity, site characteristics, plant age or size, form, vigor, season of burn, and stocking levels [100]. Most acorns are characterized by a relatively high moisture content. As the moisture within the acorns is heated, the seeds swell and often rupture [100]. Therefore, "average" fires kill all white oak acorns present on-site [44]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : Oaks tend to be less susceptible to fire during the dormant season [100]. Mean white oak mortality after fires in the dormant season was 23 percent, as compared with 69 percent after fires occurring in the growing season. Individuals of poor vigor are less likely to heal than healthy vigorous specimens. Oaks growing in overstocked stands typically are less vigorous and thus more susceptible to fire damage. Crooked or leaning trees are particularly susceptible to damage because the flames are more likely to be directly below the stem, thereby increasing the amount of heat received at the bark's surface. Higher fire intensity and severity increase mortality and serious injury. Topographic factors such as aspect and slope can also influence mortality [78]. Fire mortality also varies with the size of the tree; fire is more likely to kill smaller white oaks than large ones [76]. A fire in an oak-pine stand in New Jersey killed 44 percent of trees 1 inch in d.b.h., 5 percent of tree 2 to 4 inches in d.b.h., but no trees greater than 5 inches in d.b.h. were killed [76]. Toole [117] reported that approximately 20 percent of white oaks examined were uninjured by fire despite discolored bark. Bark sloughed off wounded white oaks within 5 years [117]. Following an early-season fire in the Pine Barrens of New Jersey, some white oaks exhibited partial crown mortality later in the summer, while others showed no evidence of significant crown damage [14]. White oak is reportedly susceptible to fire scars [93] which can permit the entry of insects or decay that may ultimately kill the tree [100]. However, Kaufert [63] reported that 50 percent of all fire scars on white oak had healed within 15 years in a southern bottomland forest [63]. Studies suggest that basal wounding does not affect growth rates [59]. Large white oaks can survive bark scorch up to two-thirds of their circumferences [100]. Mortality equations based on d.b.h., and the width and height of bark blackening have been developed for white oak [55,78,86]. These equations can be useful in predicting if a fire-damaged oak will survive [78]. PLANT RESPONSE TO FIRE : White oak commonly sprouts vigorously from the stump or root crown after aboveground portions of the plant are damaged or killed [11,99]. Sprouting depends on such factors as plant vigor [84], genetic composition, size, and fire severity and intensity. White oak probably stump-sprouts after moderate fires [51], and when completely top-killed, underground portions often regenerate [100]. Hannah [51] reported that the "best" sprouts often originate from buds located at or below ground level. These sprouts may be more vigorous and less susceptible to rot or other damage. White oak seedlings generally sprout after fire, and in many instances, numbers remain essentially unchanged [97]. Damaged seedlings can often resprout several times and may ultimately grow beyond the fire-susceptible stage [51]. Seedlings often develop an enlarged root crown after frequent fires [11,42]. Sprouting ability typically decreases with increasing d.b.h. [64]. Pole-sized trees sprout readily from stumps [32], but older, faster-growing, or taller trees often fail to sprout [64]. Multiple sprouts, which resemble seedlings, commonly develop after fire [75] and plant density is often increased. In the southern Appalachians, Keetch [65] reported an average of six to seven sprouts per clump 4 years after fire and 10 to 15 per clump 2 years after several consecutive fires. White oak generally responds quickly to release [99]. Previously suppressed individuals often grow rapidly into the understory soon after fire [90]. Initial postfire sprout growth is also rapid, and prolific seed production occurs at an early age [10,14]. Sprouts are commonly present within one growing season after fire [13]. Rouse [100] reported that most surviving oaks are "capable of minimizing fire-caused losses due to damaged cambium by rerouting the functions of fire-killed portions within weeks after a fire." Large oaks that survive fire frequently serve as seed sources for burned areas [51]; dying trees often produce a massive seed crop [100]. Birds and mammals may transport seeds from adjacent unburned areas, and seedling establishment may occur. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : Postfire increases in white oak have been documented as follows after fire in a mixed hardwood community of Rhode Island [22]: size class burned unburned density % BA % density % BA % overstory 42.7 23.40 23.6 27.60 1-10 ft. tall 42.4 ----- 17.0 ---- < 1 ft. tall 42.3 ----- 28.7 ----
The following Research Project Summaries and a Research Paper by
Bowles and others 2007
provide information on managment using prescribed fire and postfire response of
several plant species, including white oak, that was not available when this species
review was written:
FIRE MANAGEMENT CONSIDERATIONS : Prescribed fire can be an important tool for regenerating oak stands. Fire may favor seedling establishment by creating suitable seedbeds and reducing competing vegetation [100]. A series of low-intensity prescribed fires prior to timber harvest can promote advanced regeneration [123]. In the southern Appalachians, biennial summer burns are often most effective in promoting advance regeneration [123]. Single preharvest or postharvest burns generally have little effect [123]. Protein content of white oak browse was higher during the year following low- and high-intensity burns [36]. Calcium levels also tend to increase in twigs on recently burned sites [11]. Changes in nutritive value after fire have been documented [11,12,36].
1. Abell, Margaret S. 1932. Much heart rot enters white oaks through fire wounds. Forest Worker. 8(6): 10. [6593]
2. Abrams, Marc D.; Downs, Julie A. 1990. Successional replacement of old-growth white oak by mixed mesophytic hardwoods in southwestern Pennsylvania. Canadian Journal of Forest Research. 20: 1864-1870. [13328]
3. Allard, H. A. 1949. An analysis of seedling progeny of an individual of Quercus saulii compared with seedlings of a typical individual of the white oak... Castanea. 14: 109-117. [10795]
4. Allen, Peter H. 1960. Scorch and mortality after a summer burn in loblolly pine. Res. Note No. 144. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 2 p. [12256]
5. Archambault, Louis; Barnes, Burton V.; Witter, John A. 1990. Landscape ecosystems of disturbed oak forests of southeastern Michigan, U.S.A. Canadian Journal of Forest Research. 20: 1570-1582. [13448]
6. Van Sambeek, J. W.; Rink, George; Johnson, Paul S., eds. 1988. Proceedings, 3rd workshop on seedling physiology and growth problems in oak plantings; 1986 February 12-13; Carbondale, IL. (Abstracts). Gen. Tech. Rep. NC-121. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 19 p. [10878]
7. Auclair, Allan Nelson Douglas. 1968. Dynamics of Prunus serotina in southern Wisconsin. Madison, WI: University of Wisconsin. 125 p. Thesis. [12759]
8. Auclair, Allan N.; Cottam, Grant. 1971. Dynamics of black cherry (Prunus serotina Erhr.) in southern Wisconsin oak forests. Ecological Monographs. 41(2): 153-177. [8102]
9. Bernard, Stephen R.; Brown, Kenneth F. 1977. Distribution of mammals, reptiles, and amphibians by BLM physiographic regions and A.W. Kuchler's associations for the eleven western states. Tech. Note 301. Denver, CO: U.S. Department of the Interior, Bureau of Land Management. 169 p. [434]
10. Boerner, Ralph E. J. 1981. Forest structure dynamics following wildfire and prescribed burning in the New Jersey Pine Barrens. The American Midland Naturalist. 105(2): 321-333. [8649]
11. Boerner, Ralph E. J. 1983. Nutrient dynamics of vegetation and detritus following two intensities of fire in the New Jersey pine barrens. Oecologia. 59: 129-134. [8648]
12. Boerner, Ralph E. J.; Cho, Do-Soon. 1987. Structure and composition of Goll Woods, an old-growth forest remnant in northwestern Ohio. Bulletin of the Torrey Botanical Club. 114(2): 173-179. [8711]
13. Boerner, Ralph E. J.; Forman, R. T. T. 1982. Hydrologic and mineral budgets of New Jersey Pine Barrens upland forests following two intensities of fire. Canadian Journal of Forest Research. 12: 503-510. [8647]
14. Boerner, Ralph E. J.; Lord, Thomas R.; Peterson, John C. 1988. Prescribed burning in the oak-pine forest of the New Jersey Pine Barrens : effects on growth and nutrient dynamics of two Quercus species. The American Midland Naturalist. 120(1): 108-119. [8646]
15. Bonner, F. T.; Vozzo, J. A. 1987. Seed biology and technology of Quercus. Gen. Tech. Rep. SO-66. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 21 p. [3248]
16. Bowersox, T. W.; Ward, W. W. 1968. Auxin inhibition of epicormic shoots in white oak. Forest Science. 14: 192-196. [9961]
17. Braun, E. Lucy. 1961. The woody plants of Ohio. Columbus, OH: Ohio State University Press. 362 p. [12914]
18. Bell, R.; Labovitz, M. L.; Sullivan, D. P. 1985. Delay in leaf flush associated with a heavy metal-enriched soil. Economic Geology. 80: 1407-1414. [11014]
19. Brewer, Richard; Kitler, Steven. 1989. Tree distribution in southwestern Michigan bur oak openings. Michigan Botanist. 28(2): 73-79. [13005]
20. Briggs, John M.; Smith, Kimberly G. 1989. Influence of habitat on acorn selection by Peromyscus leucopus. Journal of Mammalogy. 70(1): 35-43. [10387]
21. Brothers, Timothy S. 1988. Indiana surface-mine forests: historical development and composition of a human-created vegetation complex. Southeastern Geographer. 28(1): 19-33. [8787]
22. Brown, James H., Jr. 1960. The role of fire in altering the species composition of forests in Rhode Island. Ecology. 41(2): 310-316. [5935]
23. Burns, Paul Y.; Nichols, J. Milford. 1952. Oak pruning in the Missouri Ozarks. University of Missouri Agricultural Experiment Station Bulletin. 581(Apr): 1-8. [10156]
24. Cahalane, Victor H. 1942. Caching and recovery of food by the western fox squirrel. Journal of Wildlife Management. 6: 338-352. [12141]
25. Callahan, John C. 1979. Hardwood resource perspectives: how much, where, who, why, and what. In: North America's forests: gateway to opportunity: Proceedings of the 1978 joint convention of the Society of American Foresters and the Canadian Institute of Forestry. Washington, DC: Society of American Foresters: 174-181. [10018]
26. Carey, Andrew B.; Gill, John D. 1980. Firewood and wildlife. Res. Note 299. Broomall, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 5 p. [9925]
27. Carvell, K. L.; Tryon, E. H. 1961. The effect of environmental factors on the abundance of oak regeneration beneath mature oak stands. Forestry Science. 7: 98-105. [10115]
28. Chapman, William K.; Bessette, Alan E. 1990. Trees and shrubs of the Adirondacks. Utica, NY: North Country Books, Inc. 131 p. [12766]
29. Clark, F. Bryan; Watt, Richard F. 1971. Silvicultural methods for regenerating oaks. In: Oak symposium: Proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 37-43. [9080]
30. Clewell, Andre F. 1985. Guide to the vascular plants of the Florida Panhandle. Tallahassee, FL: Florida State University Press. 605 p. [13124]
31. Coffman, Michael S.; Alyanak, Edward; Resovsky, Richard. 1980. Field guide habitat classification system: For Upper Peninsula of Michigan and northeast Wisconsin. Houghton, MI: School of Forestry and Wood Production, Michigan Technical University. 112 p. [8997]
32. Core, Earl L. 1971. Silvical characteristics of the five upland oaks. In: Oak symposium: Proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 19-22. [9077]
33. Crow, Gerald R.; Hicks, Ray R., Jr. 1990. Predicting mortality in mixed oak stands following spring insect defoliation. Forest Science. 36(3): 831-841. [13019]
34. Curtis, J. T.; McIntosh, R. P. 1951. An upland forest continuum in the prairie-forest border region of Wisconsin. Ecology. 32: 476-496. [6927]
35. DeWitt, James B.; Derby, James V., Jr. 1955. Changes in nutritive value of browse plants following forest fires. Journal of Wildlife Management. 19(1): 65-70. [7343]
36. Dills, Gary G. 1970. Effects of prescribed burning on deer browse. Journal of Wildlife Management. 34(3): 540-545. [218]
37. Dorney, Cheryl H.; Dorney, John R. 1989. An unusual oak savanna in northeastern Wisconsin: the effect of Indian-caused fire. The American Midland Naturalist. 122: 103-113. [7892]
38. Duncan, Wilbur H.; Duncan, Marion B. 1988. Trees of the southeastern United States. Athens, GA: The University of Georgia Press. 322 p. [12764]
39. Elowe, Kenneth D.; Dodge, Wendell E. 1989. Factors affecting black bear reproductive success and cub survival. Journal of Wildlife Management. 53(4): 962-968. [10339]
40. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905]
41. Farmer, Robert E., Jr.; Pitcher, John A. 1981. Pollen handling for southern hardwoods. In: Agric. Handb. 587. Washington, DC: U.S. Department of Agriculture, Forest Service: 77-83. [12654]
42. Fowells, H. A., compiler. 1965. Silvics of forest trees of the United States. Agric. Handb. 271. Washington, DC: U.S. Department of Agriculture, Forest Service. 762 p. [12442]
43. Garren, Kenneth H. 1943. Effects of fire on vegetation of the southeastern United States. Botanical Review. 9: 617-654. [9517]
44. Garrett, H. E.; Thomas, M. W.; Pallardy, S. G. 1989. Susceptibility of sugar maple and oak to eleven foliar-applied herbicides. In: Rink, George; Budelsky, Carl A., eds. Proceedings, 7th central hardwood conference; 1989 March 5-8; Carbondale, IL. Gen. Tech. Rep. NC-132. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 81-85. [9371]
45. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; [and others]. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. [998]
46. Glitzenstein, Jeff S.; Harcombe, Paul A.; Streng, Donna R. 1986. Disturbance, succession, and maintenance of species diversity in an east Texas forest. Ecological Monographs. 56(3): 243-258. [9670]
47. Godfrey, Robert K. 1988. Trees, shrubs, and woody vines of northern Florida and adjacent Georgia and Alabama. Athens, GA: The University of Georgia Press. 734 p. [10239]
48. Great Plains Flora Association. 1986. Flora of the Great Plains. Lawrence, KS: University Press of Kansas. 1392 p. [1603]
49. Hammitt, William E.; Barnes, Burton V. 1989. Composition and structure of an old-growth oak-hickory forest in southern Michigan over 20 years. In: Rink, George; Budelsky, Carl A., eds. Proceedings, 7th central hardwood conference; 1989 March 5-8; Carbondale, IL. Gen. Tech. Rep. NC-132. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 247-253. [9386]
50. Hannah, Peter R. 1987. Regeneration methods for oaks. Northern Journal of Applied Forestry. 4: 97-101. [3728]
51. Hardin, James W. 1975. Hybridization and introgression in Quercus alba. Journal of the Arnold Arboretum. 56: 336-363. [10553]
52. Harlow, Richard F.; Whelan, James B.; Crawford, Hewlette S.; Skeen, John E. 1975. Deer foods during years of oak mast abundance and scarcity. Journal of Wildlife Management. 39(2): 330-336. [10088]
53. Harrison, Janet S.; Werner, Patricia A. 1984. Colonization by oak seedlings into a heterogeneous successional habitat. Canadian Journal of Botany. 62: 559-563. [11979]
54. Hepting, George H. 1941. Prediction of cull following fire in Appalachian oaks. Journal of Agricultural Research. 62(2): 109-120. [13660]
55. Hepting, George H.; Hedgcock, George G. 1935. Relation between butt rot and fire in some eastern hardwoods. Tech. Note 14. Asheville, NC: U.S. Department of Agriculture, Forest Service, Appalachian Forest Experiment Station. 2 p. [10186]
56. Hosie, R. C. 1969. Native trees of Canada. 7th ed. Ottawa, ON: Canadian Forestry Service, Department of Fisheries and Forestry. 380 p. [3375]
57. Houston, David R. 1971. Noninfectious diseases of oaks. In: Oak symposium: Proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 118-123. [9088]
58. Jemison, George M. 1944. The effect of basal wounding by forest fires on the diameter growth of some southern appalachian hardwoods. Bulletin 9. Durham, NC: Duke University, School of Forestry. 63 p. [8716]
59. Knapp, Eric E.; Rice, Kevin J. 1998. Genetic structure and gene flow in Elymus glaucus (blue rye): implications for native grassland retoration. Restoration Ecology. 4(1): 1-10. [11875]
60. Jones, Steven M. 1989. Application of landscape ecosystem classification in identifying productive potential of pine-hardwood stands. In: Waldrop, Thomas A., ed. Proceedings of pine-hardwood mixtures: a symposium on management and ecology of the type; 1989 April 18-19; Atlanta, GA. Gen. Tech. Rep. SE-58. Asheville, SC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station: 64-69. [10259]
61. Kartesz, John T.; Kartesz, Rosemarie. 1980. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume II: The biota of North America. Chapel Hill, NC: The University of North Carolina Press; in confederation with Anne H. Lindsey and C. Richie Bell, North Carolina Botanical Garden. 500 p. [6954]
62. Kaufert, F. H. 1933. Fire and decay injury in the Southern bottomland hardwoods. Journal of Forestry. 31: 64-67. [2694]
63. Kays, Jonathan S.; Smith, David Wm.; Zedaker, Shepard M.; Kreh, Richard E. 1988. Factors affecting natural regeneration of Piedmont hardwoods. Southern Journal of Applied Forestry. 12(2): 98-102. [5642]
64. Keetch, John J. 1944. Sprout development on once-burned and repeatedly-burned areas in the southern Appalachians. Technical Note No. 59. Asheville,NC: U.S. Department of Agriculture, Forest Service, Appalachian Forest Experiment Station. 3 p. [10995]
65. Kline, Virginia M.; Cottam, Grant. 1979. Vegetation response to climate and fire in the driftless area of Wisconsin. Ecology. 60(5): 861-868. [3420]
66. Kotar, John; Kovach, Joseph A.; Locey, Craig T. 1988. Field guide to forest habitat types of northern Wisconsin. Madison, WI: University of Wisconsin, Department of Forestry; Wisconsin Department of Natural Resources. 217 p. [11510]
67. Kuchler, A. W. 1964. Manual to accompany the map of potential vegetation of the conterminous United States. Special Publication No. 36. New York: American Geographical Society. 77 p. [1384]
68. Lay, Daniel W. 1957. Browse quality and the effects of prescribed burning in southern pine forests. Journal of Forestry. 55: 342-347. [7633]
69. Ledig, F. Thomas; Wilson, Robert W.; Duffield, John W.; Maxwell, Gerald. 1969. A discriminant analysis of introgression between Quercus prinus L. and Quercus alba L. Bulletin of the Torrey Botanical Club. 96(2): 156-163. [10665]
70. Lewis, Allen R. 1982. Selection of nuts by gray squirrels and optimal foraging theory. The American Midland Naturalist. 107: 250-257. [8391]
71. Limstrom, G. A.; Merz, R. W. 1949. Rehabilitation of lands stripped for coal in Ohio. Tech. Pap. No. 113. Columbus, OH: The Ohio Reclamation Association. 41 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Central States Forest Experiment Station. [4427]
72. Little, Elbert L., Jr. 1971. Atlas of the United States trees. Volume 1. Conifers and important hardwoods. Misc. Publ. 1146. Washington, DC: U.S. Department of Agriculture, Forest Service. 320 p. [1462]
73. Little, Elbert L., Jr. 1979. Checklist of United States trees (native and naturalized). Agric. Handb. 541. Washington, DC: U.S. Department of Agriculture, Forest Service. 375 p. [2952]
74. Little, Silas, Jr. 1938. Relationships between vigor of resprouting and intensity of cutting in coppice stands. Journal of Forestry. 36: 1216-1223. [9964]
75. Little, S. 1946. The effects of forest fires on the stand history of New Jersey's Pine Region. Forest Management Paper No. 2. Upper Darby, PA: U.S. Department of Agriculture,Forest Service, Northeastern Forest Experiment Station. 43 p. [11619]
76. Little, S.; Moore, E. B. 1949. The ecological role of prescribed burns in the pine-oak forests of southern New Jersey. Ecology. 30(2): 223-233. [11107]
77. Loomis, Robert M. 1973. Estimating fire-caused mortality and injury in oak-hickory forests. Res. Pap. NC-94. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 6 p. [8740]
78. Lyon, L. Jack; Stickney, Peter F. 1976. Early vegetal succession following large northern Rocky Mountain wildfires. In: Proceedings, Tall Timbers fire ecology conference and Intermountain Fire Research Council fire and land management symposium; 1974 October 8-10; Missoula, MT. No. 14. Tallahassee, FL: Tall Timbers Research Station: 355-373. [1496]
79. Martin, Alexander C.; Zim, Herbert S.; Nelson, Arnold L. 1951. American wildlife and plants. New York: McGraw-Hill Book Company, Inc. 500 p. [4021]
80. Martin, William H.; DeSelm, Hal R. 1976. Forest communities of dissected uplands in the Great Valley of east Tennessee. In: Fralish, James S.; Weaver, George T.; Schlesinger, Richard C., eds. Central hardwood forest conference: Proceedings of a meeting; 1976 October 17-19; Carbondale, IL. Carbondale, IL: Southern Illinois University: 11-29. [3810]
81. McIntyre, A. C. 1936. Sprout groups and their relation to the oak forests of Pennsylvania. Journal of Forestry. 34: 1054-1058. [10086]
82. Millers, Imants; Shriner, David S.; Rizzo, David. 1989. History of hardwood decline in the eastern United States. Gen. Tech. Rep. NE-126. Bromall, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 75 p. [10925]
83. Monk, Carl D.; Imm, Donald W.; Potter, Robert L.; Parker, Geoffrey G. 1989. A classification of the deciduous forest of eastern North America. Vegetatio. 80: 167-181. [9297]
84. Moser, Harold C. 1971. Manufacture of oak furniture, cabinets, and panels. In: White, D. E.; Roach, B. A., co-chairmen. Oak symposium proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 100-102. [13732]
85. Nelson, Ralph M.; Sims, Ivan H.; Abell, Margaret S. 1933. Basal fire wounds on some southern Appalachian hardwoods. Journal of Forestry. 31: 829-837. [160]
86. Olson, David F., Jr. 1974. Quercus L. oak. In: Schopmeyer, C. S., ed. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, DC: U.S. Department of Agriculture, Forest Service: 692-703. [7737]
87. Olson, David F., Jr.; Boyce, Stephen G. 1971. Factors affecting acorn production and germination and early growth of seedlings and seedling sprouts. In: Oak symposium: Proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 44-48. [9081]
88. Ontario Department of Lands and Forests. 1953. Forest tree planting. 2d ed. Bull. No. R 1. Toronto, Canada: Ontario Department of Lands and Forests, Division of Reforestation. 68 p. [12130]
89. Oosting, Henry J. 1944. The comparative effect of surface and crown fire on the composition of a loblolly pine community. Ecology. 25(1): 61-69. [9919]
90. Pallardy, S. G.; Nigh, T. A.; Garrett, H. E. 1988. Changes in forest composition in central Missouri: 1968-1982. The American Midland Naturalist. 120(2): 380-390. [9043]
91. Park, Barry C. 1942. The yield and persistence of wildlife food plants. Journal of Wildlife Management. 6(2): 118-121. [7446]
92. Paulsell, Lee K. 1957. Effects of burning on Ozark hardwood timberlands. Res. Bull. 640. Columbia, MO: University of Missouri, College of Agriculture, Agricultural Experiment Station. 24 p. [11885]
93. Radford, Albert E.; Ahles, Harry E.; Bell, C. Ritchie. 1968. Manual of the vascular flora of the Carolinas. Chapel Hill, NC: The University of North Carolina Press. 1183 p. [7606]
94. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843]
95. Reid, Vincent H.; Goodrum, Phil D. 1957. The effect of hardwood removal on wildlife. In: Proceedings of the Society of American Foresters meeting; 1957 November 10-13; Syracuse, NY. Washington, DC: Society of American Foresters: 141-147. [10477]
96. Reich, Peter B.; Abrams, Marc D.; Ellsworth, David S.; [and others]. 1990. Fire affects ecophysiology and community dynamics of central Wisconsin oak forest regeneration. Ecology. 71(6): 2179-2190. [13326]
97. Rogers, Lynn. 1976. Effects of mast and berry crop failures on survival, growth, and reproductive success of black bears. Transactions, North American Wildlife Conference. 41: 431-438. [8951]
98. Rogers, Robert. 1990. Quercus alba L. white oak. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 2. Hardwoods. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 605-613. [13973]
99. Rouse, Cary. 1986. Fire effects in northeastern forests: oak. Gen. Tech. Rep. NC-105. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 7 p. [3884]
100. Rundel, Philip W. 1980. Adaptations of Mediterranean-climate oaks to environmental stress. In: Plumb, Timothy R., technical coordinator. Proceedings of the symposium on the ecology, management and utilization of California oaks; 1979 June 26-28; Claremont, CA. Gen. Tech. Rep. PSW-44. Berkeley, CA: U.S. Department of Agriculture, Forest Service, Pacific Southwest Forest and Range Experiment Station: 43-54. [7014]
101. Sander, Ivan L. 1979. Regenerating oaks. In: Proceedings of the National siviculture workshop. Theme: The shelterwood regeneration method; 1979 September 17-21; Charleston, SC. Washington, D. C.: U.S. Department of Agriculture, Forest Service, Division of Timber Management: 212-22. [11670]
102. Seymour, Frank Conkling. 1982. The flora of New England. 2d ed. Phytologia Memoirs 5. Plainfield, NJ: Harold N. Moldenke and Alma L. Moldenke. 611 p. [7604]
103. Sharp, Ward M.; Chisman, Henry H. 1961. Flowering and fruiting in the white oaks. I. Staminate flowering through pollen dispersal. Ecology. 42: 365-372. [3910]
104. Sharp, Ward M.; Sprague, Vance G. 1967. Flowering and fruiting in the white oaks, pistillate flowering, acorn development, weather, and yields. Ecology. 48: 243-251. [3909]
105. Shaw, Samuel P. 1971. Wildlife and oak management. In: Oak symposium: Proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 84-89. [9087]
106. Shelford, V. E. 1954. Some lower Mississippi valley flood plain biotic communities; their age and elevation. Ecology. 35(2): 126-142. [4329]
107. Short, Henry L. 1976. Composition and squirrel use of acorns of black and white oak groups. Journal of Wildlife Management. 40(3): 479-483. [10590]
108. Short, Henry L.; Epps, E. A., Jr. 1976. Nutrient quality and digestibility of seeds and fruits from southern forests. Journal of Wildlife Management. 40(2): 283-289. [10510]
109. Simpson, Benny J. 1988. A field guide to Texas trees. Austin, TX: Texas Monthly Press. 372 p. [11708]
110. Smallwood, Peter D.; Peters, W. David. 1986. Grey squirrel food preferences: the effects of tannin and fat concentration. Ecology. 67(1): 168-175. [10519]
111. Smith, Christopher C.; Follmer, David. 1972. Food preferences of squirrels. Ecology. 53: 82-91. [2942]
112. Sork, Victoria L.; Stacey, Peter; Averett, John E. 1983. Utilization of red oak acorns in non-bumper crop year. Oecologia. 59: 49-53. [4593]
113. Spalt, Karl W.; Reifsnyder, William E. 1962. Bark characteristics and fire resistance: a literature survey. Occas. Paper 193. New Orleans, LA: U.S. Department of Agriculture, Forest Service, Southern Forest Experiment Station. 19 p. In cooperation with: Yale University, School of Forestry. [266]
114. Starker, T. J. 1932. Fire resistance of trees of northeast United States. Forest Worker. 8(3): 8-9. [81]
115. Stephens, H. A. 1973. Woody plants of the North Central Plains. Lawrence, KS: The University Press of Kansas. 530 p. [3804]
116. Toole, E. Richard. 1965. Fire damage to commercial hardwoods in southern bottom lands. In: Proceedings, 4th annual Tall Timbers fire ecology conference; 1965 March 18-19; Tallahassee, FL. Tallahassee, FL: Tall Timbers Research Station: 144-151. [8715]
117. U.S. Department of Agriculture, Soil Conservation Service. 1982. National list of scientific plant names. Vol. 1. List of plant names. SCS-TP-159. Washington, DC. 416 p. [11573]
118. Van Dersal, William R. 1938. Native woody plants of the United States, their erosion-control and wildlife values. Washington, DC: U.S. Department of Agriculture. 362 p. [4240]
119. Van Dersal, William R. 1940. Utilization of oaks by birds and mammals. Journal of Wildlife Management. 4(4): 404-428. [11983]
120. Van Lear, David H.; Johnson, Von J. 1983. Effects of prescribed burning in the southern Appalachian and upper Piedmont forests: a review. Forestry Bull. No. 36. Clemson, SC: Clemson University, Collage of Forest and Recreation Resources, Department of Forestry. 8 p. [11755]
121. Van Lear, David H.; Waldrop, Thomas A. 1988. Effects of fire on natural regeneration in the Appalachian Mountains. In: Smith, H. Clay; Perkey, Arlyn W.; Kidd, William E., Jr., eds. Guidelines for regenerating Appalachian hardwood stands: Workshop proceedings; 1988 May 24-26; Morgantown, WV. SAF Publ. 88-03. Morgantown, WV: West Virginia University Books: 56-70. [13934]
122. Van Lear, David H.; Waldrop, Thomas A. 1989. History, uses, and effects of fire in the Appalachians. Gen. Tech. Rep. SE-54. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 20 p. [10126]
123. Vogel, Willis G. 1990. Results of planting oaks on coal surface-mined lands. In: Van Sambeek, J. W.; Larson, M. M., eds. Proceedings, 4th workshop on seedling physiology and growth problems in oak plantings; 1989 March 1-2; Columbus, OH. (Abstracts). Gen. Tech. Rep. NC-139. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 19. Abstract. [13146]
124. Vogt, Albert R.; Cox, Gene S. 1970. Evidence for the hormonal control of stump sprouting by oak. Forest Science. 16(2): 165-171. [9872]
125. Voss, Edward G. 1985. Michigan flora. Part II. Dicots (Saururaceae--Cornaceae). Bull. 59. Bloomfield Hills, MI: Cranbrook Institute of Science; Ann Arbor, MI: University of Michigan Herbarium. 724 p. [11472]
126. Wainio, Walter W.; Forbes, E. B. 1941. The chemical composition of forest fruits and nuts from Pennsylvania. Journal of Agricultural Research. 62(10): 627-635. [5401]
127. Ward, Jeffrey S.; Stephens, George R. 1989. Long-term effects of a 1932 surface fire on stand structure in a Connecticut mixed hardwood forest. In: Rink, George; Budelsky, Carl A., eds. Proceedings, 7th central hardwood conference; 1989 March 5-8; Carbondale, IL. Gen. Tech. Rep. NC-132. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station: 267-273. [9389]
128. Weckerly, Floyd W.; Sugg, Derrick W.; Semlitsch, Raymond D. 1989. Germination success of acorns (Quercus): insect predation and tannins. Canadian Journal of Forest Research. 19: 811-815. [10150]
129. Whittaker, R. H. 1956. Vegetation of the Great Smoky Mountains. Ecological Monographs. 26(1): 1-79. [11108]
130. Wofford, B. Eugene. 1989. Guide to the vascular plants of the Blue Ridge. Athens, GA: The University of Georgia Press. 384 p. [12908]