SPECIES: Pinus serotina
AUTHORSHIP AND CITATION:
Archer, Amy J. 2000. Pinus serotina. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ .
The currently accepted scientific name of pond pine is Pinus serotina Michx (Pinaceae) [31,41]. Pond pine hybridizes with pitch pine (P. rigida), longleaf pine (P. palustris), and loblolly pine (P. taeda). Due to pond pine's late pollination season, hybridization with pitch pine and longleaf pine is rare ; however, hybridization with loblolly pine is common throughout southern Delaware .
FEDERAL LEGAL STATUS:
No special status
Pond pine is distributed from Cape May, New Jersey, southward through the coastal plains of Delaware, Maryland, Virginia, North Carolina, South Carolina and Georgia to central Florida and southeastern Alabama [8,36], and is occasionally found in Mississippi and southeastern Texas [25,47]. Pond pine was introduced to England in 1713 where it still exists, although very rare .
BLM PHYSIOGRAPHIC REGIONS:
SAF COVER TYPES :
70 Longleaf pine
73 Southern redcedar
74 Cabbage palmetto
81 Loblolly pine
82 Loblolly pine-hardwood
83 Longleaf pine-slash pine
84 Slash pine
85 Slash pine-hardwood
97 Atlantic white-cedar
98 Pond pine
103 Water tupelo-swamp tupelo
104 Sweetbay-swamp tupelo-redbay
HABITAT TYPES AND PLANT COMMUNITIES:
Pond pine dominates the overstory of many swamps and other poorly drained areas that usually have striking changes in groundwater levels and are frequently subject to fires during dry periods . Pond pine occurs in a variety of coastal plains habitat types. Associates of pond pine in wet pine forests include swamp tupelo (Nyssa sylvatica), water oak (Quercus nigra), baldcypress (Taxodium distichum), pondcypress (T. d. var. nutans), sweetbay (Magnolia virginiana), live oak (Quercus virginiana), loblolly-bay (Gordonia lasianthus), and redbay (Persea borbonia) [25,35,47,57]. Tree species associated with pond pine in savannas of the coastal plain include longleaf pine, loblolly pine, and slash pine (P. elliottii) [3,7,13,56,57,86]. Although pond pine usually dominates pocosin overstory, red maple (Acer rubrum), sweetbay, and redbay may codominate [13,86]. The wettest flatwoods of the longleaf pine communities are generally dominated by pond pine with loblolly-bay and a dense shrub layer .
GENERAL BOTANICAL CHARACTERISTICS:
Pond pine is a relatively small tree that grows to 82 feet (25 m) tall by 2.6 feet (0.8 m) dbh with open and irregular crowns. Its needles are 4 to 8 inches (10-20 cm) long, 0.06 to 0.8 inches (1.5-2.0 mm) wide, with long tapering tips, in bundles of 3 or sometimes 4. Seed cones are persistent and frequently remain closed for several years. Seed cones are stalked but later may become sessile or imbedded as branches enlarge. Branches are moderately stout and may have clusters of needle bundles on the trunk. Winter buds are 0.2 inches (5 mm) in diameter and usually coated with hardened resin. The seed, including the wing, is 1.0 to 1.2 inches (2.5-3 cm) long and the wing is 0.3 inches (7 mm) wide. Pond pine has medium-thick basal bark, a moderate to high and open crown, and moderately open stands. The root systems of pond pine seedlings are superficial , and mature trees have a "medium" rooting habit [10,19]. Pond pine sprouts from the base and roots after fire or other damage [19,20,24,82] and produces epicormic branches after fire damage .
The specific epithet of pond pine, serotina, means "late" and refers to the delayed opening of cones. Cones often persist for 2 to 8 years, giving the tree the appearance of being a prolific seed producer [8,14]. All pond pine cones are probably closed when they first mature . Viability does not decrease in cones that remain closed for as long as 3 years . Cones older than 5 years may open individually following weathering and insect depredations; sound seed may be sparse in cones more than 5 years old . Seed is released from older, open cones of pond pine during 2 seasonal periods: April through September and October through January [8,16]. Regeneration of pond pine is hampered by 2 problems under average conditions. First, serotinous pond pine cones often require heat to open the scaled cones, and 2nd, continual moistness of the organic soil provides an excellent growth medium for intense competitors such as other trees, evergreen brush, and switchcane (Arundinaria gigantea ssp. tecta) .
Pond pine cones open and release seeds soon after exposure to heat from fire. Intensity of heat does not adversely reduce viability of seeds. Even badly charred cones release seeds that are capable of germination. Mature cones can be opened by exposure to 333 to 336 oF (167-169 oC) dry heat for 20 seconds or by immersion in boiling water for a similar period . Optimum stratification of pond pine seeds is 30 days at 33-41oF (0.5-5 oC) in a moist medium [5,46]. Germination occurs in 24 days . Pond pine seeds germinated in a nursery were unable to survive full saturation conditions .
Among pine species, pond pine is somewhat unique in that it usually sprouts after being damaged by disturbance [8,19,20,24,82]. Seedlings or saplings usually sprout prolifically when cut, and older trees will sprout vigorously along the stem and branches even after intense scorching and defoliation by fire. These sprouts come from dormant buds in the axils of primary needles of young seedlings. Buds are dormant and are protected by bark. When the tree is injured, these buds resume growth and may give rise to clusters of buds and numerous short shoots . In pocosins, seedlings of pond pine sprout prolifically, and stands up to sapling size are primarily reestablished by sprouting . However, 1 study showed that most seedlings on pond pine plots, which were completely burned, did not sprout . Evidence suggests that pond pine may not sprout indefinitely, and that frequent burning of any age pond pine stand may kill it .
Pond pine is found on a variety of sites in the southeastern United States. Pond pine occupies the wettest sites within the wet pine type. It is found in swamps, pond borders, and pocosins, characterized by undrained peat soils and/or wet, sandy flats. Pond pine grows extensively in the broad, poorly drained interstream areas of peaty soils in the lower coastal plains of North Carolina. Further south, it is found on wet pine flatwoods (pine forests on poorly drained soils) and savannas of the lower coastal marine terraces and in bays and ponds throughout the coastal plain .
In eastern North Carolina, pond pine usually occurs on wet flats and extensive, poorly drained evergreen shrub bogs called pocosins. These pocosins have organic soils varying from 1 to 25 feet (0.3-7.6 m) deep [16,61]. Pocosins are highmoor, ombrotrophic bogs dominated by a continuous shrub cover 3.3 to 13 feet (1-4 m) high with scattered emergent trees, usually pond pine [61,87]. Pond pine is the only "important" pocosin tree, probably because of recurrent fire. Pocosins apparently represent the lower limit of site quality tolerated by pond pine .
A pond pine-slash pine association occurs in peaty swamps. Peaty swamps contain surface water only during a part of the growing season. Coniferous trees or shrubs, including many ericaceous species, exist in these swamps. Frutescent and herbaceous plants are also numerous. Peaty swamps are found on poorly drained, flat, interstream areas of lower terraces in the Atlantic and Gulf coastal plains .
Pine flatwoods is one of the most common vegetation types in Florida, and can be dominated by pond pine . Flatwoods with a pH less than 4.5 are usually dominated by pond pine and those with pH above 4.5 by slash pine or longleaf pine [50,51]. Pond pine is a principal tree species characteristic of this wet pine type that is found in the coastal plain physiographic province from Delaware south to Florida and west to eastern Texas, primarily in the flatwoods of the outer marine terraces. Representative sites include boggy, nonriver flatlands; poorly-drained, seasonally-wet coastal flatlands; and lowlands adjacent to ponds, streams, and other wet areas. Soils range widely in texture from clay to sand. Moisture conditions are influenced by an impermeable clay soil layer that resists downward movement of water. Poor drainage limits aeration of the soils on many sites and peat mats commonly develop over mineral surfaces. Acidic conditions prevail on most sites .
The bay forest type, occurring exclusively in the Coastal Plain physiographic province ranging from Maryland to southeastern Texas, is restricted to coastal depressions or floodplains where saturated conditions prevail. Soils are usually organic, highly acidic and low in nutrient availability . Bay swamps, dominated by the redbay-sweetbay association, are characterized by a more or less dense growth of evergreen trees, including pond pine .
Other sites that include pond pine are cypress-swamp tupelo (Nyssa biflora) and cane (Arundinaria spp.). On a cypress-swamp tupelo site, pond pine is most abundant where standing water or poor drainage renders the habitat more acidic . Pond pine is often the overstory of cane habitats that generally occur on organic soils having fair internal drainage and 40% to 50% organic matter content in the upper 4 inches (10 cm) of soil, and a species composition of 70% to 100% reeds [6,29,85].
Pond pine has excellent growth on better drained mineral soils, but the faster, earlier growth of slash and loblolly pine usually restricts it to a subordinate position on such sites . Pond pine is relatively tolerant to waterlogging. Mature trees can tolerate prolonged flooding, but growth is slow. Pond pine also endures poor soil aeration and high acidity, and is confined mostly to ponds and bays of interstream areas [27,57].
Another site characteristic important to maintenance of pond pine is the unique climate of the southeastern United States. Weather patterns in the Southeast spawn more lightning storms than in any other region of North America. The lightning fire season begins in early spring, is often most prevalent in May or June, then tapers off in July and August through early fall. The large majority of fires in the Southeast are caused by lightning .
The successional status of pond pine is somewhat controversial. Traditional ecological succession and climax concepts tend to place many of the fire-dependent vegetation types, such as pond pine, as "subclimax" types [2,8,15]. However, others believe the fire-dependent systems are relatively stable "climaxes" under a burning regime, if, historically, fire was a natural part of the environment [54,81]. Walker  considers pond pine climax vegetation in pocosins since the tree is maintained in the absence of fire and regenerated as a consequence of fire. The bay swamps, many of which contain pond pine, are considered climax by some, since the southern white-cedar swamps revert to redbay-sweetbay when protected from fire; however it is stated that the hydric communities "can not be considered climax because none are controlled by climate ."
Fire is an important aspect of succession in many habitat types in which pond pine is found. The long-term perpetuation of flatwoods, sand-pine scrub, pocosins, and savannas are tied to fire. Species diversity is highest immediately following fire and generally decreases thereafter .
Flatwoods ecosystems, along with the rest of the southeastern pine forests, have been classified as fire subclimax within the deciduous forest region . Pine flatwoods are considered to be fire maintained but, without fire, they ultimately give way to hardwoods . Pond pine flatwoods, if unburned, can develop either to bayheads or to mixed hardwood swamps [2,8]. Pond pine communities of North Carolina possess little herbaceous vegetation and are often changed by fire into shrub swamps in which pond pine occurs only as scattered relicts among the fire resistant shrubs . Human alteration of natural fire frequency is thought to be the most common cause of successional change in flatwoods .
Pocosins, often with pond pine overstory, generally can be viewed as intermediate successional communities. They are often maintained in a subclimax stage by fire and hydroperiod, with the mature vegetational stages being suppressed for long periods on the wettest sites but developing relatively quickly on drier sites .
The bay forest type, occurring exclusively in the Coastal Plain physiographic province, frequently reverts to pond pine or Atlantic white-cedar after severe fires . If the original stand of southern white-cedar is destroyed, together with seed stored in the upper layer of peat, succession is toward a type in which pond pine predominates in Virginia, North Carolina and South Carolina . With recurrent fire, the bay swamp, pond pine-slash pine wetlands, and Atlantic white-cedar bog may revert to shrub bogs . It should be noted, however, that the shallow marsh is characterized by a mineral soil, and could not arise in a peaty area without aggradation of inorganic sediments . The patchwork nature of evergreen bay forests is related to their extreme susceptibility to fire, after which they may revert back to any one of several freshwater, hydric vegetative cover types depending on the intensity of the burn and depth to the water table during and following the burn .
Frequency of fire also determines the successional stage of many southeastern swamp forests. Serotiny of species such as pond pine may be selected for in regimes where fires are spaced with a frequency which allows maturation but does not "unreasonably exceed" this. Too frequent firing could eliminate pond pine from an area altogether  and produce a grass-sedge bog or savannah . Wells  observed that fires started by Native Americans, and later even more frequently by European settlers, began to change stream-head swamp forests of the southeastern United States into grass-sedge bogs or savannahs. Under burning at least every decade, the native deciduous forest largely disappeared or became dominated by pond pine. With more frequent burning the pines were replaced by fire-resistant shrubs or shrubby trees with the ability to sprout from stumps or roots. Under annual fires, the shrubs practically disappeared and were replaced by grasses, sedges, and many bulbous herbs. These nearly treeless areas were called sub-tropical savannahs. Under reduced frequency of fires, savannahs revert to shrubby pocosins, and under no fire, savannahs and pocosins rapidly return to swamp forest.
Excluding fire, primary factors affecting plant succession in any hydric area are aggradation of the level of the substratum by inorganic and organic sediments, with consequent lowering of the water table and subsequent increase in oxygen tension above that level . On organic soils subject to prolonged flooding, longleaf and slash pine are replaced by pond pine communities. Pond pine assumes dominance in Virginia and the Carolinas on the wettest sites and on organic soils . On mineral soils, pond pine is in an intermediate stage of plant succession, where it may grow in pure even-aged stands until harvesting or mortality removes the dominant trees .
The minimum seed-bearing age of pond pine is between 4 and 10 years; however, cone production usually begins when pond pine seedlings are approximately 10 years old [8,36]. The tree flowers between March and April [22,36], and pollen flight and female flower receptivity occur in late March in Florida and about 1 month later in North Carolina . Cones ripen in September or October [8,36], and seed dispersal is often in the spring, but as cones are serotinous they may remain closed for several months or years. Interval between large seed crops is 1 year .
FIRE ECOLOGY OR ADAPTATIONS:
Pond pine has adaptations that not only allow it to exist, but to thrive in frequent fire regimes [34,76]. Requirements for fire adapted species such as pond pine are (1) bare mineral or exposed substrate for seed germination, (2) removal of toxic or allelopathic chemicals or disease-causing conditions, (3) reproductive structures requiring fire to disperse propagules, and (4) ability to vegetatively reproduce by root suckering, stump sprouts or epicormic stems . Pond pine readily sprouts after fire, both from root collar and stem, and is one of the conifers most able to endure fire defoliation [10,32,70] because it also produces epicormic branches . Other characteristics of pond pine that contribute to its fire resistance include: medium-thick basal bark; moderate-to-high and open crown; moderately open stands; and medium rooting habit [10,19].
Many southeastern habitats have fire regimes with frequent fire. Fires recurring at intervals of 20 to 40 years have long been recognized as an integral part of the ecology of pocosins dominated by pond pine . The high incidence of fires in longleaf pine communities, which often contain pond pine, is related to their high flammability, a consequence of the volatile oils and resins in longleaf pine needles and wiregrass that is common in the understory. However, human fire suppression has reduced the high natural occurrence of fire in this habitat. Noss  stated that an immediate priority for longleaf sites is to reestablish the natural regime of summer fires. Wet pine forests, which include pond pine, depend on fire for their continued existence . Pond pine can be found in wet savannahs in North Carolina that are dependent on fire to eliminate encroaching forests and to remove grass and sedge foliage that casts a heavy shade and can lead to a loss of smaller grasses and forbs which grow between pine clumps .
Pond pine communities are wet and nutrient-poor and are most susceptible to fires during droughts that allow the organic soils to dry. The large amount of fuel produced by the understory makes fires severe. However, all dominant species sprout readily and the shrub layer grows to its former height in just a few years. In the wetter, shrub-dominated slash pine and pond pine wetlands, presettlement fire frequency was probably 10 to 30 years. Frequency in these types depends on occurrence of drought conditions sufficient to increase the flammability of the understory to where it will burn readily. Such fires are intense and usually burn all aboveground vegetation, especially in pond pine woodlands . Burning may be severe in both swamps and marshes during prolonged periods of low rainfall. In fresh-water marshes, in which pond pine occurs, fires cause little damage when surface water is present, but may destroy all the marsh dominants as well as the component animal life, such as beaver and muskrat, during drought years when the water table is well below the marsh surface .The following table is a summary of the fire frequency of nonalluvial wetland communities of the southeastern United States in which pond pine is a canopy dominant .
|Community||Soil||Hydroperiod/water source||fire frequency|
|slash pine flatwoods||mineral||<3 mos./groundwater||3-10 years|
|wet longleaf pine- slash pine flatwoods||mineral||<3 mos./groundwater||3-10 years|
|pond pine woodland||shallow organics and peat||6-9 mos./rainfall||10-20 years|
|low pocosin||deep peat > 1.64 ft (0.5 m)||6-9 mos./rainfall||15-30 years|
|small depression pocosin||shallow peat <1.64 ft (0.5 m)||6-9 mos./rainfall||15-30 years|
|high pocosin||shallow peat < 1.64 ft (0.5 m)||6-9 mos./rainfall||15-30 years|
To learn more about fire regimes and fire ecology of communities where pond pine occurs with other dominants, refer to the FEIS review of the dominant species listed below.
|Community or Ecosystem||Dominant Species||Fire Return Interval Range (years)|
|Atlantic white-cedar||Chamaecyparis thyoides||35 to > 200|
|slash pine||Pinus elliottii||3-8|
|slash pine-hardwood||P. e.-variable||< 35|
|longleaf-slash pine||P. palustris-P. e.||1-4 |
|pitch pine||P. rigida||6-25 |
|loblolly pine||P. taeda||3-8|
|cabbage palmetto-slash pine||Sabal palmetto-P. e.||< 10|
|baldcypress||Taxodium distichum var. distichum||100 to > 300|
|pondcypress||T. d. var. nutans||< 35 |
IMMEDIATE FIRE EFFECT ON PLANT:
Pond pine is well adapted to fire [6,13,39,45] and is one of the conifers most able to endure fire defoliation [10,32]. Because of pond pine's ability to resprout, mature trees are only killed by the most severe fires [6,13]. An immediate effect of fire on pond pine is the opening of serotinous cones [6,8,14,16,44,83]. Burning also may result in a considerable enrichment of most available nutrients  in pond pine dominated habitats.
DISCUSSION AND QUALIFICATION OF FIRE EFFECT:
The data of Wilbur and Christensen , suggest that pocosin peats are deficient in nutrients, particularly phosphorus. Burning resulted in considerable enrichment of most available nutrients, and some amelioration of the prefire phosphorus deficiency. They believe these nutrient changes are responsible in part for increased postfire production. However, increased fertility following fire was short-lived, and by the 2nd growing season concentrations of most nutrients were back to prefire levels.
Not only were mean concentrations of particular nutrients elevated following fire, but spatial variation in those concentrations was also increased. This was particularly true for nitrogen and phosphorus. Thus, high and low nutrient patches were created in an area characterized by uniformly low prefire nutrient concentrations. This redistribution of resources might lead to the creation of patches of high and low production within a bog .
PLANT RESPONSE TO FIRE:
Wildfire has been responsible for the rehabilitation of pond pine stands in the past  because of pond pine's successful regeneration after the disturbance [6,13,39,45]. Seedlings sprout from dormant buds near the root collar after being top-killed by a low intensity fire [13,45,75], and mature trees sprout from buds along the bole, main branches and root crowns [10,13,32,39,60]. Pond pine also reproduces after fire by the opening of serotinous cones and subsequent seed release [6,8,14,16,44,83].
Pond pine may not sprout after severe fire. For example, it did not sprout after a severe wildfire on the Air Force Bomb Range in Dare County, North Carolina. See Fire Case Studies for further information.
DISCUSSION AND QUALIFICATION OF PLANT RESPONSE:
Mature trees sprout vigorously from protected buds along the bole, main branches, and root crowns [10,13,32,39,60], and are only killed by severe fires [6,13]. Repeated fire would favor pond pine over loblolly pine because of pond pine's ability to sprout vigorously . However, one study suggests that the ability to sprout is generally restricted to seedling and saplings . After a fire burned through a pine scrub on Cumberland Island in Georgia, tree mortality was very high (95-100%) in plots where pond pine was a strong dominant. Pond pine seedlings found in the first 2 postfire seasonal samples did not survive .
Heat generated by a ground fire is usually of sufficient intensity to open the serotinous cones of pond pine . Pond pine cones do not characteristically remain closed for longer than 4 to 5 years , and observations suggest that cones open rapidly on felled trees during midday on clear summer days [6,83] when the outside temperature may be sufficient to open the cones. Seeds up to 14 years old have been germinated . Pond pine cones germinate best after a burn on soil that is moist. This may be because the root system of pond pine is superficial and the seedlings many not be able to obtain enough water to survive in dry conditions .
Fire removes competition, exposes the soil and causes the seed of pond pine to fall. Volume of seed caught in traps averaged about 3,000 sound seed per acre per year for undisturbed and unburned seed tree stands. From 2 uncut but burned stands the average was 53,000 sound seed/acre over a 6-week period following the burn . After a fire on Cumberland Island, Georgia, pond pine, loblolly pine, and slash pine reseeded where the pine canopy was killed .
Season of burning is largely determined by availability of ignition sources and seasonal variation in fuel flammability. A major factor affecting flammability is seasonal variation in moisture. Seasonal variation in the dead-to-live ratio is also important in shrub bogs . Low-severity winter fires are valuable in preparing favorable seedbeds for establishment of pine seedlings and in partially controlling associated hardwoods . Most wildfires in the pocosins occur during the early spring and before the evergreen vegetation has increased its moisture content through spring growth .
FIRE MANAGEMENT CONSIDERATIONS:
The range of ecological processes and conditions that recurring fire initiates, terminates, or continues in wet pine forests can not be duplicated by any other disturbance . In the absence of fire, adequate regeneration of pond pine is not obtained . Fire is usually considered to be most damaging to pond pine in the pond pine ecosystems where it is infrequent and intense. Lowering of the water table in pocosins will "undoubtedly" increase the frequency of fire due to increased frequency of dry periods .
Fire exclusion is an unrealistic management objective for sustaining ecological communities, such as those containing pond pine, that are adapted to and dependent upon recurring fires to maintain balance. Prescribed burning is a more reasonable management goal as it approximates natural processes inherent to a fire region. Under natural conditions, frequent fires maintain vegetation, recycle nutrients, and drive vegetation cycles . However, because seedlings are either killed outright or set back by fire during their early years, prescribed burning is not recommended for most pine species until they are at least 9.8 to 13 feet (3- 4 m) tall , and too frequent burning of pond pine at any age may eliminate the species from an area altogether .
Prescribed fire appears to be a potentially valuable tool in the management of pond pine by promoting seed fall from the serotinous cones and in preparing the seedbed. Fires of sufficient intensity to achieve these results, however, are usually difficult to control and tend to burn deeply into the organic soil. The problem has become more acute in the past 10 years because of increased commercial importance of pocosins as pulpwood and timber processing areas . Following harvest of mature pines, flatwoods sites are often prepared for pine regeneration by use of prescribed fires. Objective of this burning is to consume as much surface litter and remaining understory biomass as possible, creating a site easy to plant by machine and removing potential competing vegetation. Burning is also often used in established stands to reduce potential for hazardous fire in young stands and to control disease and insect pests .
Although recommendations for optimum season of burns of southeastern forests are somewhat contradictory, Streng, Glitzenstein, and Platt  concluded that growing season burns do not reduce growth or survival of pines more than do dormant season burns; therefore, growing season prescribed burns can be conducted without undue fear of creating damaging fires.
In North Carolina pocosins, growth and vigor of pond pine seedlings are much better on burned-logged areas than on burned-unlogged. It is speculated that there is less root and top competition for seedlings on the burned-logged areas. Seedlings that germinate during the 1st, 2nd, or 3rd years following logging and burning have the best chance of surviving competition with the brush . Although logging opened up the tree canopy of a Coastal Plains pond pine forest, it did not significantly affect seedling establishment. Rate of seedling establishment was extremely low on unburned sites regardless of logging treatment .
Grazing of pond pine habitats without a fire disturbance may be beneficial to pond pine establishment, although grazing after a fire may reduce the number of pond pine seedlings. The average number of new pond pine seedlings per acre that became established in grazed and ungrazed plots before and after fire disturbance in the southern Coastal Plains follows :
|Ungrazed||before fire||1 yr. after||2 yrs. after|
|Grazed||before fire||1 yr. after||2 yrs. after|
Evidence indicates that both grazing and fire substantially influenced seedling establishment, although the effect of fire was of much greater magnitude than that of grazing. On unburned areas grazing increased seedling establishment, but not enough for adequate regeneration of pond pine. Where grazing occurred before an intense burn, it tended to decrease the fire effect, but the interaction of grazing with burning was not apparent on a low-severity burn .
Disturbance of the surface litter by the trampling of cattle apparently increased chances for pond pine seed to reach an environment suitable for germination and growth. Accompanying reduction of competing herbage may also have been a contributing factor. Fire was more effective, however, for providing a favorable seedbed. Grazing contributed to the mortality of very small seedlings, but had little effect after they were 1 foot (0.3 m) tall. On the grazed plots mortality averages 13% higher than on unprotected plots for seedlings in the 1 to 6 inch (2.5-15.2 cm) height class, and 6% higher for the 7 to 12 (18-30.5 cm) inch height class. The higher mortality under grazing is probably due largely to trampling rather than browsing. Browsing damage was seldom encountered, but some trampling damage to small seedlings was observed, especially along cattle trails .
The fires in this study killed practically all seedlings in the plots affected. Although it is generally accepted that pond pine usually sprouts after being burned, small seedlings may not always have this ability. On plots that were completely burned over, all seedlings were killed. This may be due to the severity of the fire. These plots had burned 4 years earlier also, so most of the seedlings were less than 2 feet tall and not more than 4 years old. As a result of cattle trails and limited fuels, some grazed plots did not burn completely, and on those about 33% of the seedlings survived with some damage to the tips .
FIRE CASE STUDY CITATION:
Archer, Amy., compiler. 2000. Effects of a in wildfire pocosin in Dare County, North Carolina, on pond pine. In: Pinus serotina. In: Fire Effects Information System, [Online]. U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory (Producer). Available: http://www.fs.fed.us/database/feis/ .
Wade, Dale D.; Ward, Darold E. 1973. An analysis of the Air Force Bomb Range Fire. Res. Pap. SE-105. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest and Range Experiment Station. 38 p. .
PREFIRE VEGETATIVE COMMUNITY:
Pond pine (Pinus serotina) was the dominant overstory tree on the burn, averaging 1,250 stems/acre (3,100/ha). About 64% of the forested area was well-stocked with 13-year-old pond pine that regenerated after a burn 14 years earlier. Another 13% contained poorly or medium-stocked, 13-year-old pond pine. The remainder was comprised of reproduced pond pine from a burn 5 years earlier and older trees that survived the burn 14 years previous. In addition, 5,000 acres (2,025 ha) of marsh, 5,000 acres (2,025 ha) of low, open pocosin, and 1,200 acres (490 ha) of species other than pond pine also burned. The 13-year-old pond pine averaged 16 feet (4.9 m) in height and 2.9 inches (7.4 cm) dbh. Its principal associates were redbay (Persea borbonia) and sweetbay (Magnolia virginiana), which together averaged 13 feet (4 m) in height, 1.6 inches (4.1 cm) dbh, and 220 stems/acre (550/ha). The weight, height, and blowup potential for 5 of the more common fuel types burned in this fire follow .
|Fuel type||Fuel weight before burn- lb/acre (kg/ha.)||Fuel weight after burn-lb/acre (kg/ha)||Blowup potential||Avg. height of understory fuel in feet (m)|
|high pocosin||25,600 (28,700)||2,000 (2,250)||3.4 (med. high)||14 (4.3)|
|very high reeds||30,900 (34,650)||2,100 (2,350)||5.8 (med. high)||10 (3.0)|
|medium reeds-brush||21,900 (24,550)||2,100 (2,350)||2.8 (med)||5 (1.5)|
|grass-low brush||12,900 (14,450)||na||1.1 (med. low)||3 (0.9)|
|low pocosin-open||12,800 (14,350)||2,000 (2,250)||1.1 (med. low)||2-3 (0.6-0.9)|
TARGET SPECIES PHENOLOGICAL STATE:
Analysis of this fire does not describe the phenological state of pond pine at time of disturbance. According to other studies of pond pine seasonal development in North Carolina, it is likely that it was actively growing and had just begun to flower [8,22,36].
Dare County, where this fire occurred, is a sparsely populated peninsula comprised primarily of pocosins and marshland. Elevation of this site is 10 feet (3 m) above mean sea level. Most of the area is managed for forest production. A network of drainage canals and woods roads is the only interruption in an otherwise unbroken expanse of fuel. At the time of the fire, the conditions were low humidity (below 60 %), winds 15 to 20 mph (7 to 9 m/sec.), and full sunshine. The temperature reached 70 oF (21 oC) on the day the fire began .
The 29,300-acre (11,860-ha) Air Force Bomb Range fire was a severe, blowup fire. The spring fire exhibited several blowup features including high-density, short-distance spotting, a well-developed convection column, and rates of spread exceeding 2 mph (2.2 m/sec.) during a 4 hour period with a maximum of almost 5 mph (2.2 m/sec.) during one 4 mile (6.4 km) run. The fire traveled 14 miles (22.5 km) prior to the passage of a cold front, then changed direction, and ran an additional 6 miles (9.7 km) as a postcold-frontal fire. The fire crowned through more than 15,000 acres (6,070 ha) of pond pine during a 20 hour period and eventually burned 29,300 acres (11,860 ha) of timber and nonforested watershed .
FIRE EFFECTS ON TARGET SPECIES:
Pond pine typically sprouts readily from both the root crown and bole, and such sprouting was expected to occur after the Air Force Bomb Range fire. Defoliation by wet-season fires generally has little effect on the survival of pond pine and causes only a temporary reduction in height and diameter growth. However, observations 1 year after the fire did not follow expectations. Although no numbers were recorded, it was noted that a much lower percentage of trees sprouted from the bole than had been expected. The lack of bole sprouts is 1 more indicator of the high intensities and severities associated with this fire. However, a high proportion of trees sprouted profusely from the root collar. The degree of basal sprouting suggests that a well-stocked stand of pond pine will again occupy the burned area, but the loss of 13 years' growth must be considered more than a temporary setback in a 50-year rotation .
FIRE MANAGEMENT IMPLICATIONS:
When this study was completed in 1973, researchers believed that further work was needed to adapt the National Fire Danger Rating System to this area . Most federal land managers in the area now burn pocosins periodically to perpetuate pocosin communities and reduce fuel loads. Prescriptions are designed to meet objectives without igniting the organic soil .
WOOD PRODUCTS VALUE:
The economic value of pond pine is variable. Some consider pond pine a commercially valuable species  as a timber source . However, others have determined pond pine has low economic value as a pulpwood source . The wood is course-grained, resinous, and of fair quality. Pond pine is apt to have more defects than other southern pines on the market, largely due to its susceptibility to red heart disease which substantially reduces its lumber value  and may make it more susceptible to fire damage . Pond pine has not been widely cultivated, due to its "low" economic value .
Pond pine is the principal commercial species of the pocosins of North Carolina. According to Besse , in 1952 the tree occurred on approximately 2 million acres of the coastal plain of North Carolina and the total board foot volume was 1.75 billion board feet. On wet sites characteristic of present day pond pine forests, yields are comparatively low, and logging costs are high. Observations indicate that growth rates of pond pine may be 25% to 33% lower than loblolly pine where the species grow together .
IMPORTANCE TO LIVESTOCK AND WILDLIFE:
Bottomland forests, in which pond pine is found, tend to be among the most productive habitats for wildlife, with surface water that moderates temperature extremes and serves as escape habitat. There is a predominance of broad-leaved evergreen plants in the understory and mast-producing trees in the overstory that provide an abundance of branch cavities and mast for over-wintering migrant birds . Animals found in pond pine habitat in North Carolina include the marsh rabbit, silver-haired bat, eastern cottontail, grasshopper mouse, meadow vole, black bear, long-tailed weasel, and bobcat .
Pocosins, in which pond pine dominates the overstory, serve as habitat for the specialized swallowtail, Hessel's hairstreak butterfly, the rare pine barrens tree frog, the eastern diamondback rattlesnake (endangered in North Carolina), and the federally endangered American alligator. Pocosins are also refuges for native species such as black bear, white-tailed deer, and smaller mammals such as the bobcat, marsh rabbit, and gray squirrel [61,62]. The endangered red-cockaded woodpecker inhabits mature pond pine in pocosins [17,28,61,62,73].
Pond pine often dominates the flatwoods habitat that supports various populations of birds and small mammals such as the cotton rat, cotton mouse, and short-tailed shrew. Avian densities are low throughout the year with some increase in winter due to the influx of migratory winter residents. No mammal is exclusive to the flatwoods, although the fox squirrel is highly characteristic of flatwoods with open understory. Three large mammals native to Florida use flatwoods: white-tailed deer, black bear, and the endangered Florida panther .
Pond pine is an important cover species for the endangered red-cockaded woodpecker [30,52,61,62,63,73,88]. The red-cockaded woodpecker usually inhabits pond pine trees between the ages of 30 to 170 years old; in South Carolina and Mississippi, however, they primarily use 54 to 119 year old pond pine trees . Trees must be infected with heart rot disease that softens the wood and allows red-cockaded woodpeckers to excavate roost and nest cavities .
VALUE FOR REHABILITATION OF DISTURBED SITES:
Seedlings of pond pine were planted on cleared shrub oak areas in South Carolina. The average survival was 72% after 2 years. Although pond pine is characteristically a wet site species, it showed good early response on this relatively dry site .
OTHER USES AND VALUES:
Subsequent to regional timber removal in early development of the Southeast, pocosin ecosystems have generally been considered of low economic value for agriculture, although they have recently been recognized as a source of peat .
OTHER MANAGEMENT CONSIDERATIONS:
The draining of pond pine habitats has created much controversy. The southern forested wetlands are home to a wide variety of endangered, threatened, and candidate plant and animal species. It is thought that these species, along with pond pine, may be jeopardized if the wetlands are drained [1,61]. In general, forested wetlands comprise about 60% of all the U.S. wetland areas (exclusive of deepwater habitats, such as submerged grasslands and lake bottoms), and loss of forested wetlands acreage is particularly high. From 1940 to 1980, forested wetlands areas were decreasing at an annual rate of 0.3% nationally .
1. Abernethy, Y.; Turner, R. E. 1987. US forested wetlands: 1940-1980: Field-data surveys document changes and can guide national resource management. BioScience. 37(10): 721-727. 
2. Abrahamson, Warren G.; Hartnett, David C. 1990. Pine flatwoods and dry prairies. In: Myers, Ronald L.; Ewel, John J., eds. Ecosystems of Florida. Orlando, FL: University of Central Florida Press: 103-149. 
3. Baker, James B.; Langdon, O. Gordon. 1990. Pinus taeda L. loblolly pine. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 497-512. 
4. Beaven, George Francis; Oosting, Henry J. 1939. Pocomoke Swamp: a study of a cypress swamp on the eastern shore of Maryland. Bulletin of the Torrey Botanical Club. 66: 376-389. 
5. Belcher, Earl W., Jr.; Hitt, Robert G. 1965. Eastern Tree Seed Laboratory: 12th annual report--fiscal year 1965. Macon, GA: Eastern Tree Seed Laboratory. 66 p. On file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT. 
6. Besse, John David. 1952. Initial influence of fire on the regeneration of pond pine. Raleigh, NC: North Carolina State College. 53 p. Thesis. 
7. Binkley, Dan. 1986. Soil acidity in loblolly pine stands with interval burning. Soil Science Society of America Journal. 50(6): 1590-1594. 
8. Bramlett, David L. 1990. Pinus serotina Michx. pond pine. In: Burns, Russell M.; Honkala, Barbara H., technical coordinators. Silvics of North America. Volume 1. Conifers. Agric. Handb. 654. Washington, DC: U.S. Department of Agriculture, Forest Service: 470-475. 
9. Bratton, Susan P. 1993. Survivorship of evergreen hardwoods after wildfire in maritime forest, Cumberland Island National Seashore, Georgia. Castanea. 58(1): 34-44. 
10. Brown, Arthur A.; Davis, Kenneth P. 1973. Forest fire control and use. 2nd ed. New York: McGraw-Hill. 686 p. 
11. Brown, James K.; Smith, Jane Kapler, eds. 2000. Wildland fire in ecosystems: Effects of fire on flora. Gen. Tech Rep. RMRS-GTR-42-vol. 2. Ogden, UT: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station. 257 p. 
12. Buchholz, Kenneth; Good, Ralph E. 1982. Density, age structure, biomass and net annual aboveground productivity of dwarfed Pinus rigida Moll. from the New Jersey Pine Barren Plains. Bulletin of the Torrey Botanical Club. 109(1): 24-34. 
13. Christensen, Norman L. 1981. Fire regimes in southeastern ecosystems. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; Lotan, J. E.; Reiners, W. A., technical coordinators. Fire regimes and ecosystem properties: Proceedings of the conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 112-136. 
14. Christensen, Norman L. 1985. Shrubland fire regimes and their evolutionary consequences. In: Pickett, S. T. A.; White, P. S., eds. The ecology of natural disturbance and patch dynamics. Orlando, FL: Academic Press, Inc.: 85-100. 
15. Clark, Mary K.; Lee, David S.; Funderburg, John B., Jr. 1985. The mammal fauna of Carolina bays, pocosins, and associated communities in North Carolina: an overview. Brimleyana. Raleigh, NC: North Carolina State Museum of Natural History. 11: 1-38. 
16. Crutchfield, D. M.; Trew, I. F. 1961. Investigation of natural regeneration of pond pine. Journal of Forestry. 59(4): 264-266. 
17. Czuhai, Eugene. 1971. Synoptic review of forest resource and use within the range of the red-cockaded woodpecker. In: Thompson, Richard L, ed. The ecology and management of the red-cockaded woodpecker: Proccedings of a symposium; 1971 May 26-27; Folkston, GA. Tallahassee, FL: Tall Timbers Research Station: 108-124. 
18. Davison, Kathryn L.; Bratton, Susan P. 1988. Vegetation response and regrowth after fire on Cumberland Island National Seashore, Georgia. Castanea. 53(1): 47-65. 
19. Dean, George W. 1969. Forests and forestry in the Dismal Swamp. Virginia Journal of Science. 20: 166-173. 
20. Duncan, Wilbur H.; Duncan, Marion B. 1988. Trees of the southeastern United States. Athens, GA: The University of Georgia Press. 322 p. 
21. Eyre, F. H., ed. 1980. Forest cover types of the United States and Canada. Washington, DC: Society of American Foresters. 148 p. 
22. Gaddy, L. L. 1982. The floristics of three South Carolina pine savannahs. Castanea. 47: 393-402. 
23. Garrison, George A.; Bjugstad, Ardell J.; Duncan, Don A.; Lewis, Mont E.; Smith, Dixie R. 1977. Vegetation and environmental features of forest and range ecosystems. Agric. Handb. 475. Washington, DC: U.S. Department of Agriculture, Forest Service. 68 p. 
24. 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. 
25. Harms, William R. 1996. An old-growth definition for wet pine forests, woodlands, and savannas. Gen. Tech. Rep. SRS-2. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 7 p. 
26. Hendrickson, William H. 1972. Perspective on fire and ecosystems in the United States. In: Fire in the environment: Symposium proceedings; 1972 May 1-5; Denver, CO. FS-276. [Washington, DC]: U.S. Department of Agriculture, Forest Service: 29-33. In cooperation with: Fire Services of Canada, Mexico, and the United States; Members of the Fire Management Study Group; North American Forestry Commission; FAO. 
27. Hook, D. D. 1984. Waterlogging tolerance of lowland tree species of the South. Southern Journal of Applied Forestry. 8: 136-149. 
28. Hooper, Robert G.; Robinson, Andrew F., Jr.; Jackson, Jerome A. 1980. The red-cockaded woodpecker: notes on life history and management. General Report SA-GR-9. Atlanta, GA: U.S. Department of Agriculture, Forest Service, Southeastern Area, State and Private Forestry. 8 p. 
29. Hughes, Ralph H. 1966. Fire ecology of canebrakes. In: Proceedings, 5th annual Tall Timbers fire ecology conference; 1966 March 24-25; Tallahassee, FL. No. 5. Tallahassee, FL: Tall Timbers Research Station: 148-158. 
30. Jackson, Jerome A.; Lennartz, Michael R.; Hooper, Robert G. 1979. Tree age and cavity initiation by red-cockaded woodpeckers. Journal of Forestry. 77(2): 102-103. 
31. Kartesz, John T. 1994. A synonymized checklist of the vascular flora of the United States, Canada, and Greenland. Volume I--checklist. 2nd ed. Portland, OR: Timber Press. 622 p. 
32. Keeley, Jon E. 1981. Reproductive cycles and fire regimes. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; Lotan, J. E.; Reiners, W. A., tech. coords. Fire regimes and ecosystem properties: Proceedings of the conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 231-277. 
33. Kologiski, Russell L. 1977. The phytosociology of the Green Swamp, North Carolina. Tech. Bull. No. 250. Raleigh, NC: North Carolina State University, Agricultural Experiment Station. 101 p. 
34. Komarek, E. V. 1983. Fire as an anthropogenic factor in vegetation ecology. In: Holzner, W.; Werger, M. J. A.; Ikusima, I., eds. Man's impact on vegetation. Boston, MA: Dr W. Junk Publishers: 77-82. 
35. Korstian, C. F. 1924. Natural regeneration of southern white cedar. Ecology. 5: 188-191. 
36. Krugman, Stanley L.; Jenkinson, James L. 1974. Pinus L. pine. In: Schopmeyer, C. S., tech. cood. Seeds of woody plants in the United States. Agric. Handb. 450. Washington, D.C.: U.S. Department of Agriculture, Forest Service: 598-638. 
37. Kuchler, A. W. 1964. United States [Potential natural vegetation of the conterminous United States]. Special Publication No. 36. New York: American Geographical Society. 1:3,168,000; colored. 
38. Landers, J. Larry. 1991. Disturbance influences on pine traits in the southeastern United States. In: Proceedings, 17th Tall Timbers fire ecology conference; 1989 May 18-21; Tallahassee, FL. No. 17. Tallahassee, FL: Tall Timbers Research Station: 61-95. 
39. Langdon, O. Gordon. 1981. Some effects of prescribed fire on understory vegetation in loblolly pine stands. In: Wood, Gene W., ed. Prescribed fire and wildlife in southern forests: Proceedings of a symposium; 1981 April 6-8; Myrtle Beach, SC. Georgetown, SC: Clemson University, Belle W. Baruch Forest Science Institute: 143-153. 
40. Layman, Harry. 1990. Fire--a paradox. In: Are forests the answer?: Proceedings of the 1990 Society of American Foresters national convention; 1990 July 29 - August 1; Washington, DC. Bethesda, MD: Society of American Foresters: 127-130. 
41. 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. 
42. Little, Silas, Jr. 1953. Prescribed burning as a tool of forest management in the northeastern states. Journal of Forestry. 51: 496-500. 
43. Little, Silas. 1974. Effects of fire on temperate forests: northeastern United States. In: Kozlowski, T. T.; Ahlgren, C. E., eds. Fire and ecosystems. New York: Academic Press: 225-250. 
44. Martin, Robert E.; Anderson, Hal E.; Boyer, William D.; [and others]. 1979. Effects of fire on fuels: A state-of-knowledge review. Gen. Tech. Rep. WO-13. Washington, DC: U.S. Department of Agriculture, Forest Service. 64 p. [Prepared for: National fire effects workshop; 1978 April 10-14; Denver, CO]. 
45. McCune, Bruce. 1988. Ecological diversity in North American pines. American Journal of Botany. 75(3): 353-368. 
46. McIninch, Suzanne; Garbisch, Edgar. 1994. Response of plants with severed taproots to different hydrologic regimes. Wetland Journal. 6(1): 22-23. 
47. McKevlin, Martha R. 1996. An old-growth definition for evergreen bay forests and related seral communities. Gen. Tech. Rep. SRS-3. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southern Research Station. 14 p. 
48. Mengel, Dennis L.; Tew, D. Thompson, eds. 1991. Ecological land classification: applications to identify the productive potential of southern forest: Proceedings of a symposium; 1991 January 7-9; Charlotte, NC. Gen. Tech. Rep. SE-68. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 149 p. 
49. Mitchell, Alan F. 1972. Conifers in the British Isles: A descriptive handbook. Forestry Commission Booklet No. 33. London: Her Majesty's Stationery Office. 322 p. 
50. Monk, Carl D. 1965. Southern mixed hardwood forest of north-central Florida. Ecological Monographs. 35: 335-354. 
51. Monk, Carl D.; Brown, Timothy W. 1965. Ecological consideration of cypress heads in north-central Florida. The American Midland Naturalist. 74: 126-140. 
52. Mosher, James A.; Taylor, Gary J.; Devlin, William J. 1979. Endangered species program: red-cockaded woodpecker study. Project No: MD.E-001-R1/Study 011/FIN. Baltimore, MD: Maryland Department of Natural Resources. 78 p. 
53. Nelson, John B. 1986. The natural communities of South Carolina. Columbia, SC: South Carolina Wildlife & Marine Resources Department. 54 p. 
54. Niering, William A. 1981. The role of fire management in altering ecosystems. In: Mooney, H. A.; Bonnicksen, T. M.; Christensen, N. L.; Lotan, J. E.; Reiners, W. A., technical coordinators. Fire regimes and ecosystem properties: Proceedings of the conference; 1978 December 11-15; Honolulu, HI. Gen. Tech. Rep. WO-26. Washington, DC: U.S. Department of Agriculture, Forest Service: 489-510. 
55. Noss, Reed F. 1988. The longleaf pine landscape of the Southeast: almost gone and almost forgotten. Endangered Species UPDATE. 5(5): 1-5. 
56. Noss, Reed F.; Harris, Larry D. 1990. Habitat connectivity and the conservation of biological diversity: Florida as a case history. In: Forestry on the frontier: Proceedings of the 1989 Society of American Foresters national convention; 1989 September 24-27; Spokane, WA. Bethesea, MD: Society of American Foresters: 131-135. 
57. Penfound, William T. 1952. Southern swamps and marshes. The Botanical Review. 18: 413-446. 
58. Raunkiaer, C. 1934. The life forms of plants and statistical plant geography. Oxford: Clarendon Press. 632 p. 
59. Rejmanek, Marcel; Richardson, David M. 1996. What attributes make some plant species more invasive? Ecology. 77(6): 1655-1661. 
60. Renfro, James Francis. 1956. Bark and cambium characteristics of pond pine (Pinus serotina Mich.) and loblolly pine (Pinus taeda L.) with special reference to pine hardness. Raleigh, NC: North Carolina State College. 46 p. Thesis. 
61. Richardson, Curtis J. 1983. Pocosins: vanishing wastelands or valuable wetlands? Bioscience. 33(10): 626-633. 
62. Richardson, Curtis J.; Gibbons, J. Whitfield. 1993. Pocosins, Carolina bays, and mountain bogs. In: Martin, William H.; Boyce, Stephen G.; Echternacht, Arthur C., eds. Biodiversity of the southeastern United States: Lowland terrestrial communities. New York: John Wiley & Sons, Inc: 257-311. 
63. Robbins, Louise E.; Myers, Ronald L. 1992. Seasonal effects of prescribed burning in Florida: a review. Misc. Publ. No. 8. Tallahassee, FL: Tall Timbers Research, Inc. 96 p. 
64. Schultz, Robert P. 1997. Genetics and tree improvement. In: Schultz, Robert P. Loblolly pine: The ecology and culture of loblolly pine (Pinus taeda L.). Agricultural Handbook 713. Washington, DC: U.S. Department of Agriculture, Forest Service: 7-3 to 7-50. 
65. Sharitz, Rebecca R.; Gibbons, J. Whitfield. 1982. The ecology of southeastern shrub bogs (pocosins) and Carolina bays: a community profile. FWS/OBS-82/04. Washington, DC: U.S. Department of the Interior, Fish and Wildlife Service, Division of Biological Services. 93 p. 
66. Shepherd, W. O.; Dillard, E. U.; Lucas, H. L. 1951. Grazing and fire influences in pond pine forests. Tech. Bull. No. 97. Raleigh, NC: North Carolina State College, Agricultural Experiment Station. 56 p. In cooperation with: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 
67. Shiflet, Thomas N., ed. 1994. Rangeland cover types of the United States. Denver, CO: Society for Range Management. 152 p. 
68. Shipman, Robert D. 1958. Planting pine in the Carolina sandhills. Station Pap. No. 96. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 43 p. 
69. Stickney, Peter F. 1989. Seral origin of species comprising secondary plant succession in Northern Rocky Mountain forests. FEIS workshop: Postfire regeneration. Unpublished draft on file at: U.S. Department of Agriculture, Forest Service, Intermountain Research Station, Fire Sciences Laboratory, Missoula, MT. 10 p. 
70. Stone, Earl L.; Stone, Margaret H. 1943. Dormant buds in certain species of Pinus. American Journal of Botany. 30(5): 346-351. 
71. Streng, Donna R.; Glitzenstein, Jeff S.; Platt, William J. 1993. Evaluating effects of season of burn in longleaf pine forests: a critical literature review and some results from an ongoing long-term study. In: Hermann, Sharon M., ed. The longleaf pine ecosystem: ecology, restoration and management: Proceedings, 18th Tall Timbers fire ecology conference; 1991 May 30 - June 2; Tallahassee, FL. No. 18. Tallahassee, FL: Tall Timbers Research, Inc: 227-263. 
72. Sutter, Robert D.; Kral, Robert. 1994. The ecology, status, and conservation of two non-alluvial wetland communities in the south Atlantic and eastern Gulf Coastal Plain, USA. Biological Conservation. 68: 235-243. 
73. Thompson, Richard L., ed. 1971. The ecology and management of the red-cockaded woodpecker: Proceedings of a symposium; 1971 May 26-27; Folkston, GA. Tallahassee, FL: Tall Timbers Research Station. 188 p. 
74. U.S. Department of Agriculture, Soil Conservation Service. 1994. Plants of the U.S.--alphabetical listing. Washington, DC: U.S. Department of Agriculture, Soil Conservation Service. 954 p. 
75. Van Lear, David H.; Waldrop, Thomas A. 1991. Prescribed burning for regeneration. In: Duryea, M. L.; Dougherty, P. M., eds. Forest regeneration manual. The Netherlands: Kluwer Academic Publishers: 235-250. 
76. Vogl, Richard J. 1977. Fire: a destructive menace or a natural process? In: Cairns, J., Jr.; Dickson, K. L.; Herricks, E. E., eds. Recovery and restoration of damaged ecosystems: Proceedings of the international symposium; 1975 March 23-25; Blacksburg, VA. Charlottesville, VA: University Press of Virginia: 261-289. 
77. Wade, Dale D. 2001. [Email to Jane Kapler Smith]. April 10. Athens, GA: U.S. Department of Agriculture, Forest Service, Southern Research Station. On file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 
78. Wade, Dale D.; Ward, Darold E. 1973. An analysis of the Air Force Bomb Range Fire. Res. Pap. SE-105. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest and Range Experiment Station. 38 p. 
79. Wade, Dale. 2001. [Email to Janet Howard]. January 17. Athens, GA: U.S. Department of Agriculture, Forest Service, Southern Research Station. On file at: U.S. Department of Agriculture, Forest Service, Rocky Mountain Research Station, Fire Sciences Laboratory, Missoula, MT; RWU 4403 files. 
80. Walker, Joan; Peet, Robert K. 1983. Composition and species diversity of pine-wiregrass savannas of the Green Swamp, North Carolina. Vegetatio. 55: 163-179. 
81. Walker, Laurence C. 1990. Forests: A naturalist's guide to trees and forest ecology. Wiley Nature Editions. New York: John Wiley & Sons, Inc. 288 p. 
82. Ward, Daniel B. 1963. Contributions to the flora of Florida--2, Pinus (Pinaceae). Castanea. 28(1): 1-10. 
83. Warren, Richard; Fordham, Alfred J. 1978. The fire pines. Arnoldia. 38(1): 1-11. 
84. Wells, B. W.; Whitford, L. A. 1976. History of stream-head swamp forests, pocosins, and savannahs in the Southeast. Journal of the Elisha Mitchell Science Society. 92: 148-150. 
85. Wendel, G. W.; Storey, T. G.; Byram, G. M. 1962. Forest fuels on organic and associated soils in the coastal plain of North Carolina. Station Paper No. 144. Asheville, NC: U.S. Department of Agriculture, Forest Service, Southeastern Forest Experiment Station. 46 p. 
86. Whitehead, Donald R. 1972. Developmental and environmental history of the Dismal Swamp. Ecological Monographs. 42(3): 301-315. 
87. Wilbur, Rebecca B.; Christensen, Norman L. 1983. Effects of fire on nutrient availability in a North Carolina coastal plain pocosin. The American Midland Naturalist. 110(1): 54-61. 
88. Zwicker, Susan M.; Walters, Jeffrey R. 1999. Selection of pines for foraging by red-cockaded woodpeckers. Journal of Wildlife Management. 63(3): 843-852.