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Introductory

SPECIES: Quercus prinus | Chestnut Oak
ABBREVIATION : QUEPRI SYNONYMS : Quercus montana Willd. SCS PLANT CODE : QUPR2 COMMON NAMES : chestnut oak rock chestnut oak rock oak tanbark oak TAXONOMY : The currently accepted scientific name of chestnut oak is Quercus prinus L. [38,49]. It has been placed within the subgenus Lepidobalanus, or white oak group [28]. In the past, Quercus prinus was applied to swamp chestnut oak (Q. michauxii) and Q. montana was applied to chestnut oak. Quercus prinus was restored to chestnut oak by Fernald in 1950 [21,38]. Chestnut oak naturally hybridizes with the following species [38]: x Q. alba (white oak): Q. X saulii Schneid. x Q. bicolor (swamp white oak) x Q. robur (English oak): Q. X sargentii Rehd. x Q. stellata (post oak): Q. X bernardiensis W. Wolf LIFE FORM : Tree FEDERAL LEGAL STATUS : No special status OTHER STATUS : Chestnut oak is listed as endangered in Maine's Official List of Endangered and Threatened Plants [15]. COMPILED BY AND DATE : Jennifer H. Carey, December 1992 LAST REVISED BY AND DATE : NO-ENTRY AUTHORSHIP AND CITATION : Carey, Jennifer H. 1992. Quercus prinus. In: Remainder of Citation

DISTRIBUTION AND OCCURRENCE

SPECIES: Quercus prinus | Chestnut Oak
GENERAL DISTRIBUTION : Chestnut oak occurs primarily in the Appalachian Mountains and adjacent hill country. Chestnut oak is distributed from southwestern Maine west through New York to extreme southern Ontario and extreme southeastern Michigan, south through southern Indiana and extreme southern Illinois to extreme northeastern Mississippi, east through northern Alabama to Georgia, and north along the Piedmont to Delaware. Chestnut oak is rare on the Southeastern Coastal Plain, but occurs along the coast in Delaware, New Jersey, New York, and in the New England states [38,49]. ECOSYSTEMS : FRES10 White - red - jack pine FRES13 Loblolly - shortleaf pine FRES14 Oak - pine FRES15 Oak - hickory FRES18 Maple - beech - birch STATES : AL CT DE GA IL IN KY ME MD MA MI MS NJ NH NY NC OH PA RI SC TN VA VT WV ADMINISTRATIVE UNITS : ALPO BISO BLRI CATO CHCH COLO CUGA CUVA DEWA FOCA FODO GWMP GRSM HOBE MACA MANA MORR NATR NERI OBRI PRWI RICH ROCR SHEN BLM PHYSIOGRAPHIC REGIONS : NO-ENTRY KUCHLER PLANT ASSOCIATIONS : K095 Great Lakes pine forest K103 Mixed mesophytic forest K104 Appalachian oak forest K110 Northeastern oak - pine forest K111 Oak - hickory - pine forest SAF COVER TYPES : 21 Eastern white pine 22 White pine - hemlock 43 Bear oak 44 Chestnut oak 45 Pitch pine 51 White pine - chestnut oak 52 White oak - black oak - northern red oak 53 White oak 78 Virginia pine - oak 79 Virginia pine 108 Red maple 110 Black oak SRM (RANGELAND) COVER TYPES : NO-ENTRY HABITAT TYPES AND PLANT COMMUNITIES : Chestnut oak is an important species of eastern upland deciduous and coniferous forests and may occur in pure stands [17]. It constitutes an important component of the subcanopy and canopy layers of Table Mountain pine (Pinus pungens) forests [80]. Chestnut oak codominates with eastern hemlock (Tsuga canadensis) on particularly steep east-facing slopes in the Hudson River Valley in New York [24]. Because of the high mortality of American chestnut (Castanea dentata) caused by the chestnut blight fungus (Endothia parasitica) introduced from Asia in the early 1900's, the former Appalachian oak (Quercus spp.)-American chestnut forest is now dominated by chestnut oak, white oak, and northern red oak (Q. rubra) [29,33,49,79]. Keever [33] recommends that former oak-American chestnut forests be named chestnut oak forests. The following published classifications list chestnut oak as dominant or codominant: Deciduous Forest [26] Vegetation of the Great Smoky Mountains [79] The Natural Communities of South Carolina [54] Eastern Deciduous Forest [74] Forest Vegetation of the Lower Alabama Piedmont [25] The Natural Forests of Maryland: an explanation of the vegetation map of Maryland [8]

VALUE AND USE

SPECIES: Quercus prinus | Chestnut Oak
WOOD PRODUCTS VALUE : Chestnut oak wood is cut and utilized as white oak lumber [49]. IMPORTANCE TO LIVESTOCK AND WILDLIFE : Good crops of chestnut oak acorns are infrequent, but when available the acorns are eaten by numerous upland wildlife species, including white-tailed deer, squirrels, chipmunks, mice, and wild turkeys [49]. White-tailed deer occasionally browse young oak sprouts, especially the first year after cutting or burning. The deer only take the top few inches of the sprout unless it is extremely succulent or other food is scarce [41]. Small birds and mammals, as well as insects such as bees, use chestnut oak cavities for nesting. In a survey of 31 oak-hickory (Carya spp.) stands in the Appalachian Mountains, a disproportionate share of cavities were in chestnut oak [9]. PALATABILITY : Chestnut oak acorns are considered sweet [49]. Gray squirrels selected pignut hickory (Carya glabra) nuts and northern red oak acorns over chestnut oak acorns but preferred chestnut oak acorns to those of white oak [36]. White-tailed deer prefer chestnut oak sprouts to seedlings [52]. Chestnut oak sprouts are more palatable than those of bear oak (Q. ilicifolia) [41]. NUTRITIONAL VALUE : Chestnut oak acorns are, on average by dry weight, 5.76 percent crude protein, 10.07 percent crude fat, and 78.9 percent carbohydrates [66]. The acorns are 0.09 percent (dry weight) magnesium and 0.15 percent phosphorus, and contain only a trace of calcium [75]. The average crude energy yield of chestnut oak acorns is 21.8 kJ/kernel, and the average metabolizable energy yield is 15.7 kJ/kernel [36]. COVER VALUE : NO-ENTRY VALUE FOR REHABILITATION OF DISTURBED SITES : In the past, chestnut oak performed well on mine spoils in Ohio [37] and on cast overburden in Illinois and Indiana [73]. However, more recent plantings of chestnut oak on mine spoils have not been as successful. Reclamation practices mandated by federal law are often unfavorable for oak establishment. Top-soiling practices, excess soil compaction caused by grading, and competition from seeded herbaceous covers reduce the growth and survival of planted oak species [73]. Chestnut oak did not show good height growth or survival and is not recommended for planting on graded, top-soiled mine spoils in southern Illinois [2]. OTHER USES AND VALUES : Chestnut oak shows a 7 to 10 day delay in budbreak and leaf flush on sites that have heavy metal (copper, zinc, and lead) enrichment of the soil. This retarded leaf flush may be used in geobotanical remote-sensing techniques for mineral detection [3]. MANAGEMENT CONSIDERATIONS : To regenerate upland oaks successfully, advance regeneration must be 4 to 5 feet (1.2-1.5 m) tall before the overstory is removed. Successful regeneration of a mixed oak forest can only be accomplished after clearcutting if there are adequate numbers of older advance regeneration [62]. Sanders [61] recommended that there be at least 433 well-distributed oak sprouts and saplings per acre (1,070/ha). Otherwise, a shelterwood silviculture system is needed to give oak regeneration time and partial light to grow. For best results, the shelterwood cut should leave a 60 to 70 percent stocking density. All nonoak stems in the understory larger than 4 to 6 feet (1.2-1.8 m) tall should be killed [62]. Forest managers have noticed a decrease in upland oak frequency in newly regenerated stands after clearcutting, especially on good sites. The reason for the decrease is the inability of oak seedlings and sprouts to compete successfully with species that have invaded the oak forest understory in the absence of disturbance [62]. In West Virginia, 59 stands with a history of grazing, thinning, or light fire in the past two decades had more oak regeneration than undisturbed stands [10]. The season of clearcutting appears to have an effect on the regeneration of upland oak stands. On lower quality sites in south-central Ohio, upland oaks (chestnut, scarlet, black, and white) were more favored over mixed hardwoods after summer clearcutting than after winter clearcutting. The season of harvest (dormant season versus growing season) did not affect regeneration on good sites [76]. Site quality affects the ability of upland oaks to regenerate. In the abovementioned study in south-central Ohio, medium-quality sites had higher absolute and relative oak densities 18 to 20 years after clearcutting than did good sites. The oaks showed good early establishment on both medium and good sites but were unable to compete with the faster growing mesic hardwoods on good sites [76]. The seed tree silviculture method was used on fair and good sites in an Appalachian hardwood forest which contained chestnut oak. Twelve years after the seed-tree harvest cut and 9 years after seed trees were cut, chestnut oak regeneration was abundant only on the fair site [68]. Thinning may or may not improve the growth of established chestnut oak stands. Five years after thinning a sawtimber-sized stand, the 75- to 80-year-old chestnut oaks had not responded to the release [67]. Thinning upland oak stands to retain the best acorn producers for wildlife habitat enhancement did not improve acorn yields enough to justify the efforts [16]. Information on thinning, stocking, growth, and yields of upland oaks is detailed [23]. Planting chestnut oak seedlings in old fields in the southern Appalachian Mountains is generally unsuccessful unless the competition is controlled for more than 3 years [18]. Information on storage, seeding, and planting techniques for upland oaks is detailed [60]. In 26 chestnut oak stands in Pennsylvania and Maryland, advance regeneration responded to the gypsy moth (Lymantria dispar) defoliation of the canopy by increasing in height. However, there was a large influx of competing vegetation, and the oak component of future stands will probably be reduced [30]. Chestnut oak is one of the two most preferred host species of the introduced gypsy moth, which defoliates trees [49]. Crow and Hicks [14] developed hazard rating equations from site and stand characteristics associated with chestnut oak mortality caused by insect defoliation. The discriminant function equations correctly classified as dead or alive 59 percent of the chestnut oaks in a study area in West Virginia. The equations use the following variables: d.b.h., height, site index, percent slope, aspect, host preference of insect, shade tolerance, and the number of years of defoliation [14]. Other insects which defoliate chestnut oak include spring and fall cankerworms (Paleacrita vernata and Alsophila pometaria), the forest tent caterpillar (Malacosoma disstria), the half-wing geometer (Phigalia titea) [49], oak leafrollers (Archips spp.) [59], and the linden looper (Erannis tilaria) [49]. Chestnut oak is susceptible to wood-boring beetles, including the Columbian timber beetle (Corthylus columbianus), Platypus spp., and Xyleborus spp. Other wood borers that attack chestnut oak include the oak timberworm (Arrhenodes minutus), the carpenterworm (Prionoxystus robiniae), the little carpenterworm (P. macmurtrei) [49], and the two-lined chestnut borer (Agrilus bilineatus) [51]. Chestnut oak is susceptible to many oak diseases including oak wilt (Ceratocystis fagacearum), twig-blight fungus (Diplodia longispora), and stem cankers caused by Nectria galligena, Strumella coryneoidea, and Botryodiplodia spp. Important decay-causing fungi include Spongipellis pachyodon, Stereum gausapatum, Armillaria mellea, Fistulina hepatica, Wolfiporia cocos, Inonotus dryophilus, Xylobolus frustulatus, and Perenniporia compacta. Decay is common in stump sprouts, although the incidence is lower for those that originate near the ground [49]. Chestnut oak is also susceptible to, but rarely killed by, several gall wasps (Cynipidae), a pit scale (Asterolecanium quercicola), and the golden oak scale (A. variolosum). Acorns are destroyed by nut weevils (Curculio spp. and Conotrachelus spp.), the moth Valentinia glandulella, and cynipid gall wasps [49]. Chestnut oaks that are stressed from drought, gypsy moth defoliation, spring frost defoliation, old age, fire, poor site conditions, or other factors often succumb to secondary agents such as the two-lined chestnut borer. This scenario, in which a primary agent stresses the tree and a secondary agent kills it, is known as "oak decline" and is responsible for considerable chestnut oak mortality [51]. Herbicides have been used to control chestnut oak on sites where pine regeneration is desired. In order to convert a North Carolina Appalachian site to white pine (Pinus strobus), picloram was applied in May as 10 percent acid equivalent pellets at the rate of 4.5 pounds acid equivalent per acre (5.0 kg ae/ha). One year later, 29 percent of the chestnut oaks showed complete crown kill or defoliation; 67 percent showed leaf curling, crown biomass reduction, and/or chlorosis; and 4 percent exhibited no effect from the herbicide treatment [53]. Roundup (glyphosate) was used to control chestnut oak on a white pine plantation in West Virginia. Three subsequent mistblower applications in August and September on small chestnut oak sprouts were 100 percent effective after two growing seasons. In the fall, saplings larger than 1 inch (2.5 cm) in d.b.h. were injected with 0.05 fluid ounce (1.5 ml) of 20 and 50 percent solutions a few inches above the groundline in 1.5-inch (3.8 cm) spacings. Two growing seasons after the injections, 100 percent of the chestnut oak saplings were dead and did not have sprouts [78]. In Georgia, three herbicides were tested on chestnut oak. Each tree received one incision for every 3 inches (7.6 cm) in d.b.h., and each incision was injected with 0.06 ounces (2 ml) of herbicide. One year after injection, chestnut oak injected with Arsenal at two different concentrations (1 and 2 lbs AC 252,925 per gallon) had 100 percent top-kill and no sprouting. Garlon 3A (1.5 pounds triclopyr per gallon) resulted in 40 percent top-kill. Chestnut oak injected with 3,6-dichloropicolinic acid at two concentrations (1.5 and 3 pounds XRM-3972 per gallon) resulted in 0 percent and 20 percent top-kill, respectively [50]. Dead, standing chestnut oak killed by fire had the fastest decomposition rate (11 percent per year) of ten species studied in the Great Smoky Mountains National Park [28].

BOTANICAL AND ECOLOGICAL CHARACTERISTICS

SPECIES: Quercus prinus | Chestnut Oak
GENERAL BOTANICAL CHARACTERISTICS : Chestnut oak is a medium-sized, native, deciduous, monoecious tree. It is long-lived and slow-growing. At maturity, chestnut oak is usually 65 to 80 feet (20-24 m) tall and 20 to 30 inches (51-76 cm) in d.b.h., but on good sites it can reach a maximum size of 100 feet (30 m) in height and 72 inches (183 cm) in d.b.h. Seedlings initially develop a deep taproot, but saplings and larger trees have six to ten main lateral roots extending 10 to 33 feet (3-10 m) from the root crown. These roots occur from near the soil surface to a depth of 36 inches (91 cm) [49]. The acorns are large [6]. RAUNKIAER LIFE FORM : Phanerophyte REGENERATION PROCESSES : Sexual: Seed production begins when the tree is about 20 years old. Acorn crop sizes vary considerably from year to year with heavy crops occurring only once every 4 to 5 years [49]. Good crops are dependent on spring temperature patterns. Above normal temperatures in early April followed by subnormal temperatures in early May result in the best acorn crops. The early warm temperatures induce the early development of staminate flowers and increase the development of viable pollen. The cool weather delays the pollen dispersal to coincide with pistillate flower development, and the delay may also enhance ovary development. A gradual increase in the temperature from early spring to summer results in poor crops. Occasionally, a chestnut oak will produce 100 to 300 pounds (45-136 kg) of acorns, but this is rare. Often a tree will produce less than 10 pounds (4.5 kg) [64]. Chestnut oaks generally produce fewer acorns than other upland oak species [49]. Dissemination is by gravity and squirrels [49], although white oak group acorns are not dispersed by squirrels to the extent that red oak group acorns are [66]. Very few (0.5 percent) available chestnut oak acorns were buried by gray squirrels in a study of acorn preference [36]. Most chestnut oak acorns germinate at day/night temperatures of 65/50 degrees Fahrenheit (18/10 deg C). Chestnut oak germination is enhanced by 1 inch (2-3 cm) of leaf litter, but litter deeper than 2 inches (5 cm) is unfavorable. The germination capacity of sound acorns is 90 percent. A thick paenchyma layer in the chestnut oak acorn pericup allows it to absorb and retain more moisture than acorns of other oak species. Consequently, they can germinate in dry soil [49]. The germination of chestnut oak acorns was not greatly affected by treating the soil with copper, lead, and zinc solutions, although acorns from sites naturally high in metals had slightly higher survival than acorns from sites with low background levels of metals [4]. In an oak-pine forest in the New Jersey Pine Barrens, 1-year-old chestnut oak occurred in areas with deeper litter (an average of 1.4 inches [3.5 cm]) and less light (22 percent of available) than its upland associate, scarlet oak (Q. coccinea). Chestnut oak seedlings, however, are not highly site specific because of the large energy reserves in the acorns [11]. The roots of chestnut oak seedlings penetrate 5 to 6 inches (12.7-15.2 cm) before the unfolding of primary leaves, which are borne on stems 2 to 3 inches (5-7.6 cm) tall [40]. Seedling growth is slow. Ten years after establishment, seedling were 6 inches (15 cm) in a unthinned forest, 9 inches (24 cm) in a thinned forest, and 58 inches (146 cm) in a clearcut [49]. Vegetative: If top-killed, chestnut oaks sprout vigorously from dormant buds at the root crown. Sprouts grow faster than seedlings. Ten years after clearcutting, some stump sprouts were larger than 21 feet (6.4 m) tall. Probably 75 percent of chestnut oak reproduction in the southern Appalachian Mountains is of sprout origin [49]. Chestnut oak sprouting frequency is high compared to other upland oaks. In one study in the Virginia Piedmont, the sprouting frequency of chestnut oak was over 90 percent, regardless of season of harvest or stump diameter [32]. Although chestnut oak initially produces large numbers of sprouts, sprout clumps tend towards the survival of one to three stems. In one study, 5, 10, 15, 25, and 35 years after cutting, the average number of sprouts per stump was 7.3, 3.8, 2.8, 2.3, and 1.9, respectively [48]. Between the ages of 4 and 8, competition is dominated by interaction between sprout clumps, not stem-to-stem competition within a sprout clump [12]. SITE CHARACTERISTICS : Chestnut oak, an upland xerophytic species, commonly occurs on ridgetops and upper slopes. It occurs from sea level on the coastal plain of New Jersey and Long Island, New York, to about 4,600 feet (1,400 m) in the southern Appalachians [49]. It can occur on all aspects; however, it is usually on south- and west-facing upper slopes and on north and easterly aspects in the southern Piedmont [25,49,54]. In the Ridge and Valley Province of central Pennsylvania, chestnut oak dominated the steep inclines and xeric ridgetop communities. It decreased in importance on mesic sites, although on some coarse-textured valley and cove sites, chestnut oak was codominant with white oak [56]. Chestnut oak is usually found on dry, rocky, infertile soil with a low moisture-holding capacity, although it grows best in rich, well-drained soils along streams [49]. In southeastern Pennsylvania, Keever [33] discovered that many of the ridge sites that chestnut oak dominates have good soil moisture. Presumably, these ridges get more precipitation than lower elevations. It is unclear why other species are excluded from these ridgetop sites, although its possible that more mesic species cannot endure occasional drought, which may be more severe on these sites [33]. The infertile rocky soil, steep slopes, and exposed conditions may also select against other forest species [46]. In 51 upland hardwood stands in the Piedmont of Virginia, chestnut oak was important on sites with low soil calcium, magnesium, and pH [19]. Chestnut oak is commonly found on acidic soils derived from sandstone, quartzite, and coarse-grained schists [8,54]. In the Hudson River Valley in New York, chestnut oak forests differed significantly (P<0.05) from white oak-black oak-pignut hickory (Carya glabra) forests and red maple (Acer rubrum) forests in several site characteristics. Chestnut oak forests were more likely to have exposed bedrock (67 percent of the stands), have a higher percent cover of bare ground by rocks (5.28 percent), and have deeper litter (1.3 inches [3.4 cm]) [24]. Overstory associates not mentioned in Distribution and Occurrence include scarlet oak, post oak (Q. stellata), hickories, sweet birch (Betula lenta), yellow-poplar (Liriodendron tulipifera), blackgum (Nyssa sylvatica), sweetgum (Liquidambar styraciflua), black cherry (Prunus serotina), black walnut (Juglans nigra), red maple, sugar maple (Acer saccharum), sourwood (Oxydendron arboreum), and black locust (Robinia pseudoacacia). Shrub associates include blueberry (Vaccinium spp.), dwarf chinkapin oak (Q. prinoides), mountain-laurel (Kalmia latifolia), Rhododendron spp., sumac (Rhus spp.), greenbrier (Smilax spp.), grape (Vitis spp.), and Ceanothus spp. [25,49]. Pure and almost pure stands of chestnut oak have sparse ground vegetation [6]. SUCCESSIONAL STATUS : Facultative Seral Species Chestnut oak is intermediate in shade tolerance. Chestnut oak reproduction dies after a few years under a closed canopy, but if some light penetrates to the forest floor, seedling sprouts may persist for years. The sprouts will respond to release. Chestnut oak is excluded from mesic sites by more rapidly growing species including yellow-poplar, sugar maple, red maple, black cherry, northern red oak, black oak, and white oak. Post oak, scarlet oak, and pitch pine (Pinus rigida) are better adapted than chestnut oak to some extremely xeric sites [49]. In the absence of disturbance, red maple and other shade-tolerant species will succeed old-growth chestnut oak on good sites [43]. On some poor sites in the Appalachian Mountains, chestnut oak stands are considered a physiographic climax [49]. Little [39] suggests a mixed oak forest of black, white, chestnut, and scarlet oaks may represent a physiographic climax association on upland sites in the New Jersey Pine Barrens. In a study of forest composition in North Carolina, chestnut oak showed good regeneration over a 30-year period on low density rhododendron (Rhododendron maximum) sites, suggesting chestnut oak will continue to dominate these forests. However, it may diminish with time in areas where high rhododendron density inhibits regeneration [57]. Advance regeneration is released by gypsy moth defoliation of the overstory canopy. However, stands defoliated by gypsy moth in Pennsylvania and Maryland will probabaly have a smaller oak component in the future because of competing vegetation and insufficient numbers of advance regeneration [30]. In forests in the Hudson River Valley in New York, the percent occurrence of chestnut oak has increased from 2.1 percent in the period before 1800 to 13.7 percent in 1984. However, early land surveys may have underrepresented chestnut oak because it occurs on poor sites and inaccessible areas. If it were not underrepresented, frequent logging may have increased its importance in stands because of the superior ability of chestnut oak to sprout from stumps [24]. A forest stand growth model was developed and used to compare the pre- and post-chestnut-blight forest. After 500 years without American chestnut, the model showed chestnut oak increased in frequency in the forests [65]. SEASONAL DEVELOPMENT : Flowers develop in the spring at the same time as leaf development [49]. In a 3-year study of chestnut oak phenology in Pennsylvania, staminate flowers, borne on ephemeral catkins, usually emerged during the first week in May, and leaves unfolded several day later. Pistillate flowers appeared in the axils of leaves on the current year's shoots, usually 5 to 10 days after the staminate flowers emerged [63,64]. Pollen dispersal, largely controlled by weather, usually occurs 10 to 20 days after the staminate flowers emerge [49]. Cool weather delays pollen dispersal [64]. Acorns mature in one growing season and drop from early September to early October, usually 2 to 5 weeks before the acorns of other upland oaks drop. Acorns exhibit no dormancy and germinate in the fall. If the temperature is below 61 degrees Fahrenheit (16 deg C), shoot development is inhibited by an induced epicotyl dormancy, but root development continues. Normal shoot development resumes in the spring [49].

FIRE ECOLOGY

SPECIES: Quercus prinus | Chestnut Oak
FIRE ECOLOGY OR ADAPTATIONS : Chestnut oak is moderately resistant to fire [7]. In three separate rankings, chestnut oak was listed as the most fire resistant of four oak species: scarlet, chestnut, black, and white [69]. Large chestnut oaks have fairly thick bark and, while more susceptible to basal wounding than pines, they survive most ground fires [7]. Top-killed chestnut oaks sprout vigorously from the root crown after fire [49]. Because bark thickens with age, the larger the fire interval is, the greater is the chance of survival. Based on a semilogarithmic model of bark thickness and mortality, chestnut oak requires a fire interval of 14 years for 50 percent survival of a low-severity surface fire. The model was developed from data collected after low-severity surface fires on south-facing slopes in the Great Smoky Mountain National Park [29]. A litter covering deeper than 2 inches (5 cm) is unfavorable for chestnut oak acorn germination [49]. Fire removes excess litter and may facilitate chestnut oak regeneration. In an oak-pine forest in the New Jersey pine barrens, chestnut oak seedling density was lowest in the forest fragment that had not been recently burned and that had an average litter depth of 2.4 inches (6.1 cm) [11]. However, the primary mode of regeneration after fire appears to be sprouting. The prevalent presettlement upland oak forests in the eastern and central United States were associated with recurring fire. These forests probably burned at an intermediate frequency (50 to 100 year intervals), which promoted the dominance and stability of oak [1]. Oak-hickory forests are usually not conducive to high-severity fires, but surface fires are easily ignited and spread rapidly under favorable conditions [13]. Periodic fire opens the canopy and sets back competition, providing an opportunity for upland oaks to regenerate and maintain dominance [1]. Fifty-five years after a late summer fire in south-central Connecticut, absolute and relative amounts of oak (chestnut, scarlet, black, white, and northern red) were higher on burned areas than adjacent unburned areas [77]. However, a fire in a 5 to 6-year old mixed hardwood stand did not affect relative species dominance, it merely retarded stand development [47]. The exact timing and conditions of fire that favor oak dominance have not been determined. POSTFIRE REGENERATION STRATEGY : Tree with adventitious-bud root crown/root sucker

FIRE EFFECTS

SPECIES: Quercus prinus | Chestnut Oak
IMMEDIATE FIRE EFFECT ON PLANT : Small chestnut oaks are top-killed by low-severity fire. Surviving trees may have basal fire wounds [49]. Acorns cannot withstand the amount of heat usually generated in leaf litter fires [34]. DISCUSSION AND QUALIFICATION OF FIRE EFFECT : NO-ENTRY PLANT RESPONSE TO FIRE : Chestnut oak sprouts vigorously after being top-killed. After a fire in south-central New York, 100 percent of the top-killed chestnut oak saplings (less than 4 inches [10 cm] in d.b.h.) sprouted, with an average of 5.9 sprouts per top-killed stem. Eleven percent of the sampled saplings were not top-killed [71]. The mortality of oak trees from fire is often delayed. Six months after two surface fires of different severity in southern New York, living butt-scorched trees (larger than 1 inch [2.5 cm] in d.b.h.) were tagged for future study. In the less severely burned area, 22 percent of the tagged chestnut oak were dead 1.5 years after the fire. Small diameter trees, especially those less than 5 inches (12.7 cm) in d.b.h., had the highest mortality. In the other area that burned more severely because of a dense understory of mountain-laurel, 41 percent of the tagged chestnut oak were dead 1.5 years after the fire. The authors concluded that at least one postfire growing season must elapse before fire damage to oaks can be accurately determined [70]. Fire may increase the growth rate of chestnut oak. Three chestnut oaks, that had suffered no crown damage from a winter fire, averaged 38 percent higher diameter growth rate in the first postfire year than the 4 years prior to the fire. Unburned chestnut oaks did not show increased growth rates. Foliar phosphorus and calcium concentrations in burned chestnut oak trees were higher than control trees through most of the growing season. In addition, the proportional phosphorus resorption and both proportional and absolute calcium deposition in leaves was higher in burned trees. However, it is uncertain what caused the increase in growth rate because other factors, such as a decrease in competition from understory shrubs, also may have contributed to the increased growth [5]. DISCUSSION AND QUALIFICATION OF PLANT RESPONSE : NO-ENTRY FIRE MANAGEMENT CONSIDERATIONS : Prescribed fire is often used to control hardwoods and promote the establishment of pine. In a study on the South Carolina Piedmont, spring felling of leafed-out residual oaks (chestnut, scarlet, and black) followed by summer burning produced greater reductions of dominant sprout height and sprout clump crown diameters at the end of the first postfire growing season than did winter felling followed by summer broadcast burning. Spring felling was probably more effective because carbohydrate root reserves are low after leaves emerge [22]. Equations have been developed to predict lumber value losses due to fire wounding of chestnut oak [42]. An equation has also been developed to predict the size of a fire wound on a chestnut oak from the area of the exterior discolored bark and the diameter of the damaged tree [55]. While fire has been suggested as a tool for improving upland oak regeneration, it has been used with only mixed success [44]. Five- to six-year-old naturally regenerating mixed hardwood stands were prescribed burned in order to increase the relative dominance of oak. The former harvested stands were 60 to 90 percent oak, but the regenerating stand had a large number of yellow-poplar, black cherry, and white ash (Fraxinus americana). The fire retarded the development of the young stand but did not increase the relative dominance of oak, which was estimated to be not more than 30 to 40 percent of the future stand. The season of fire (spring versus fall) did not change the outcome [47]. In such a stand, there may not be sufficient differences in fire resistance between oak stems and those of other species for fire to give oaks a distinct advantage [44].

REFERENCES

SPECIES: Quercus prinus | Chestnut Oak
REFERENCES : 1. Abrams, Marc D. 1992. Fire and the development of oak forests. BioScience. 42(5): 346-353. [19215] 2. Ashby, W. Clark. 1990. Growth of oaks on topsoiled 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: 20. Abstract. [13147] 3. 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] 4. Bell, R.; Teramura, A. H. 1991. Soil metal effects on the germination and survival of Quercus alba L. and Q. prinus L. Environmental and Experimental Botany. 31(2): 145-152. [15281] 5. 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. American Midland Naturalist. 120(1): 108-119. [8646] 6. Braun, E. Lucy. 1961. The woody plants of Ohio. Columbus, OH: Ohio State University Press. 362 p. [12914] 7. Brown, Arthur A.; Davis, Kenneth P. 1973. Forest fire control and use. 2nd ed. New York: McGraw-Hill. 686 p. [15993] 8. Brush, Grace S.; Lenk, Cecilia; Smith, Joanne. 1980. The natural forests of Maryland: an explanation of the vegetation map of Maryland. Ecological Monographs. 50(1): 77-92. [19035] 9. Carey, Andrew B. 1983. Cavities in trees in hardwood forests. In: Davis, Jerry W.; Goodwin, Gregory A.; Ockenfeis, Richard A., technical coordinators. Snag habitat management: proceedings of the symposium; 1983 June 7-9; Flagstaff, AZ. Gen. Tech. Rep. RM-99. Fort Collins, CO: U.S. Department of Agriculture, Forest Service, Rocky Mountain Forest and Range Experiment Station: 167-184. [17833] 10. 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] 11. Collins, Scott L.; Good, Ralph E. 1987. The seedling regeneration niche: habitat structure of tree seedlings in an oak-pine forest. Oikos. 48: 89-98. [8637] 12. Cook, James E. 1990. Degree of competition and integration in one- to eight-year-old scarlet and chestnut oak sprout clumps. 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: 29. Abstract. [13156] 13. Crosby, John S.; Loomis, Robert M. 1974. Some forest floor fuelbed characteristics of black oak stands in southeast Missouri. NC-162. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 4 p. [8153] 14. 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] 15. Dibble, Alison C.; Campbell, Christopher S.; Tyler, Harry R., Jr.; Vickery, Barbara St. J. 1989. Maine's official list of endangered and threatened plants. Rhodora. 91(867): 244-269. [4258] 16. Drake, William E. 1991. Evaluation of an approach to improve acorn production during thinning. In: McCormick, Larry H.; Gottschalk, Kurt W., eds. Proceedings, 8th central hardwood forest conference; 1991 March 4-6; University Park, PA. Gen. Tech. Rep. NE-148. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 429-441. [15328] 17. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. [905] 18. Farmer, R. E., Jr. 1981. Early growth of black cherry, oaks, and yellow-poplar in southern Appalachian plantings. Tree Planters' Notes. 32(3): 12-14. [12504] 19. Farrell, John D.; Ware, Stewart. 1991. Edaphic factors and forest vegetation in the piedmont of Virgina. Bulletin of the Torrey Botanical Club. 118(2): 161-169. [15694] 20. Fernald, Merritt Lyndon. 1950. Gray's manual of botany. [Corrections supplied by R. C. Rollins]. Portland, OR: Dioscorides Press. 1632 p. (Dudley, Theodore R., gen. ed.; Biosystematics, Floristic & Phylogeny Series; vol. 2). [14935] 21. 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] 22. Geisinger, Donn R.; Waldrop, Thomas A.; Haymond, Jacqueline L.; Van Lear, David H. 1989. Sprout growth following winter and spring felling with and without summer broadcast burning. 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: 91-95. [10262] 23. Gingrich, Samuel F. 1971. Stocking, growth, and yield of oak stands. In: Oak symposium: Proceedings; 1971 August 16-20; Morgantown, WV. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 65-73. [9085] 24. Glitzenstein, Jeff S.; Canham, Charles D.; McDonnell, Mark J.; Streng, Donna R. 1990. Effects of environment and land-use history on upland forests of the Cary Arboretum, Hudson Valley, New York. Bulletin of the Torrey Botanical Club. 117(2): 106-122. [13301] 25. Golden, Michael S. 1979. Forest vegetation of the lower Alabama Piedmont. Ecology. 60(4): 770-782. [9643] 26. Greller, Andrew M. 1988. Deciduous forest. In: Barbour, Michael G.; Billings, William Dwight, eds. North American terrestrial vegetation. Cambridge; New York: Cambridge University Press: 288-316. [19544] 27. Harlow, William M.; Harrar, Ellwood S., White, F. M. 1979. Textbook of dendrology. 6th ed. New York: McGraw-Hill, Inc. 510 p. [18070] 28. Harmon, Mark E. 1982. Decomposition of standing dead trees in the southern Appalachian Mountains. Oecologia. 52: 214-215. [13735] 29. Harmon, Mark E. 1984. Survival of trees after low-intensity surface fires in Great Smoky Mountains National Park. Ecology. 65(3): 796-802. [10997] 30. Hix, David M.; Fosbroke, David E.; Hicks, Ray R., Jr.; Gottschalk, Kurt W. 1991. Development of regeneration following gypsy moth defoliation of Appalachian Plateau and Ridge & Valley hardwood stands. In: McCormick, Larry H.; Gottschalk, Kurt W., eds. Proceedings, 8th central hardwood forest conference; 1991 March 4-6; University Park, PA. Gen. Tech. Rep. NE-148. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 347-359. [15323] 31. 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] 32. Dieterich, John H. 1983. Fire history of southwestern mixed conifer: a case study. Forest Ecology. 6: 13-31. [5242] 33. Keever, Catherine. 1973. Distribution of major forest species in southeastern Pennsylvania. Ecological Monographs. 43(3): 303-327. [19550] 34. Korstian, C. F. 1927. Factors controlling germination and early survival of oaks. Bull. No. 19. New Haven, CT: Yale University, School of Forestry. 115 p. [19369] 35. 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] 36. Lewis, Allen R. 1982. Selection of nuts by gray squirrels and optimal foraging theory. American Midland Naturalist. 107: 250-257. [8391] 37. 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] 38. 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] 39. Martin, S. Clark. 1980. Mesquite. In: Eyre, F. H., ed. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters: 118. [9858] 40. 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] 41. Little, Silas; Moorhead, George R.; Somes, Horace A. 1958. Forestry and deer in the Pine Region of New Jersey. Station Pap. No. 109. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 33 p. [11681] 42. Loomis, Robert M. 1974. Predicting the losses in sawtimber volume and quality from fires in oak-hickory forests. NC-104. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 6 p. [8712] 43. Lorimer, Craig G. 1984. Development of the red maple understory in northeastern oak forests. Forest Science. 30(1): 3-22. [12565] 44. Lorimer, Craig G. 1985. The role of fire in the perpetuation of oak forests. In:, Johnson, J. E., ed. Challenges in oak management and utilization. Madison, WI: University of Wisconsin, Cooperative Extension Service: 8-25. [19543] 45. 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] 46. 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] 47. McGee, Charles E. 1980. The effect of fire on species dominance in young upland hardwood stands. In: Proceedings, mid-south upland hardwood symposium for the practicing forester and land manager; [Date of conference unknown]; [Location of conference unknown]. Atlanta, GA: U.S. Department of Agriculture, Forest Service, Division of State and Private Forestry: 97-104. [12706] 48. McIntyre, A. C. 1936. Sprout groups and their relation to the oak forests of Pennsylvania. Journal of Forestry. 34: 1054-1058. [10086] 49. McQuilkin, Robert A. 1990. Quercus prinus L. chestnut oak. In: Burns, Russell M.; Honkala, Barbara H., tech. coords. Silvics of North America. Vol. 2. Hardwoods. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 726 p. [19551] 50. Michael, J. L. 1985. Hardwood control by injection with two new chemicals. Proceedings of the Southern Weed Science Society. 38: 164-167. [12687] 51. 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] 52. Moore, William H.; Johnson, Frank M. 1967. Nature of deer browsing on hardwood seedlings and sprouts. Journal of Wildlife Management. 31(2): 351-353. [16394] 53. Neary, D. G.; Douglass, J. E.; Ruehle, J. L.; Fox, W. 1984. Converting rhododendron-laurel thickets to white pine with picloram and mycorrhizae-inoculated seedlings. Southern Journal of Applied Forestry. 8(3): 163-168. [10697] 54. Nelson, John B. 1986. The natural communities of South Carolina. Columbia, SC: South Carolina Wildlife & Marine Resources Department. 54 p. [15578] 55. 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] 56. Nowacki, Gregory J.; Abrams, Marc D. 1991. Community and edaphic analysis of mixed oak forests in the Ridge and Valley Province of central Pennsylvania. In: McCormick, Larry H.; Gottschalk, Kurt W., eds. Proceedings, 8th central hardwood forest conference; 1991 March 4-6; University Park, PA. Gen. Tech. Rep. NE-148. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 247-260. [15315] 57. Phillips, Donald L.; Murdy, William H. 1985. Effects of Rhododendron (Rhododendron maximum L.) on regeneration of southern Appalachian hardwoods. Forest Science. 31(1): 226-233. [19372] 58. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. [2843] 59. Rexrode, Charles O. 1971. Insect damage to 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: 129-134. [9089] 60. Russell, T. E. 1971. Seeding and planting 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: 49-54. [9082] 61. Sander, Ivan L. 1977. Manager's handbook for oaks in the North Central States. Gen. Tech. Rep NC-37. St. Paul, MN: U.S. Department of Agriculture, Forest Service, North Central Forest Experiment Station. 35 p. [11002] 62. Sander, Ivan L. 1988. Guidelines for regenerating Appalachian oak stands. 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: 189-198. [13945] 63. 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] 64. 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] 65. Shugart, H.H., Jr.; West, D.C. 1977. Development of an Appalachian deciduous forest succession model and its application to assessment of the impact of the chestnut blight. Journal of Environmental Management. 5: 161-179. [67] 66. Smallwood, Peter D.; Peters, W. David. 1986. Grey squirrel food preferences: the effects of tannin and fat concentration. Ecology. 67(1): 168-175. [10519] 67. Smith, H. Clay; Miller, Gary W. 1991. Releasing 75- to 80-year-old Appalachian hardwood sawtimber trees: 5-year d.b.h. response. In: McCormick, Larry H.; Gottschalk, Kurt W., eds. Proceedings, 8th central hardwood forest conference; 1991 March 4-6; University Park, PA. Gen. Tech. Rep. NE-148. Radnor, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station: 402-413. [15327] 68. Smith, H. Clay; Rosier, Robert L.; Hammack, K. P.. 1976. Reproduction 12 years after seed-tree harvest cutting in Appalachian hardwoods. Res. Pap. NE-350. Upper Darby, PA: U.S. Department of Agriculture, Forest Service, Northeastern Forest Experiment Station. 11 p. [10887] 69. 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] 70. Stickel, Paul W. 1935. Forest fire damage studies in the Northeast. II. First-year mortality in burned-over oak stands. Journal of Forestry. 33: 595-598. [18764] 71. Swan, Frederick R., Jr. 1970. Post-fire response of four plant communities in south-central New York state. Ecology. 51(6): 1074-1082. [3446] 72. 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] 73. Vogel, Willis G. 1990. Results of planting oaks on coal surface-mined lands. 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Index

Related categories for Species: Quercus prinus | Chestnut Oak

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