Marsh Pine

Pinus serotina is fire successional and sprouts adventitiously after crown fires. It is part of a distinct forest type including Taxodium distichum (Linnaeus) Richard, Nyssa biflora Walter, Magnolia virginiana Linnaeus, Persea sp., and Ilex sp. Of good form when protected from fire, P . serotina then much resembles P . taeda , with which it hybridizes naturally. It is of increasing importance as pulpwood.

Trees to 21 m tall; trunk to 0.6 m d.b.h. in native range, usually with adventitious sprouts; bark red-brown, irregularly furrowed into oblong, flat, scaly plates; crown often rounded or flat; branchlets orange- or yellow-brown, often glaucous, stout; winter buds red-brown, ovoid or narrowly ovoid, 1-1.5 (-2) cm. Needles tufted at branchlet tips, 3 (or 5 in adventitious or disturbed growth) per bundle, slightly twisted, (12-)15-21 cm × 0.3-1.5(-2) mm, stomatal lines present on all surfaces, fine, base with persistent sheath 1-2 cm, margin serrulate, Seed cones whorled, sessile or pedunculate (when peduncles to 1 cm), pale red-brown or creamy brown, broadly ovoid or globose when open, 5-8 cm, maturing in 2 years, late dehiscent. Seed scales with dark red-brown border adaxially distally; apophyses rhombic, low cross keeled; umbo with a short, weak prickle, sometimes unarmed. Seeds pale brown, mottled darker or nearly black, somewhat compressed, ellipsoid, 5-6 mm, apex oblique; wing to 2 cm.

Trees to 21m; trunk to 0.6m diam., straight or more often crooked, commonly with adventitious sprouts; crown becoming ragged, thin, often broadly rounded or flat. Bark red-brown, irregularly furrowed and cross-checked into rectangular, flat, scaly plates. Branches spreading to ascending; twigs stout, orange- to yellow-orange, frequently glaucous, aging darker. Buds ovoid to narrowly ovoid, red-brown, 1--1.5(--2)cm, resinous. Leaves 3 per fascicle (to 5 in adventitious or disturbed growth), spreading to ascending, persisting 2--3 years, (12--)15--20(--21)cm ´ 1.3--1.5(--2)mm, slightly twisted, tufted at twig tips, straight, yellow-green, all surfaces with fine stomatal lines, margins serrulate, apex acuminate; sheath 1--2cm, base persistent. Pollen cones cylindric, to 30mm, yellow-brown. Seed cones maturing in 2 years, in some populations beginning to shed seeds then but more often variably serotinous, long-persistent, often whorled, symmetric, ovoid to lanceoloid before opening, broadly ovoid to nearly globose when open, 5--8cm, pale red-brown to creamy brown, sessile or on stalks to 1cm, scales with dark red-brown border on adaxial surface distally; apophyses slightly thickened, low, rhombic, low cross-keeled; umbo central, low-conic, with short, weak prickle, sometimes unarmed. Seeds ellipsoid, oblique at tip, somewhat compressed; body 5--6mm, pale brown, mottled darker or nearly black; wing to 20mm. 2 n =24.

Cultivated. Jiangsu, Jiangxi, Zhejiang [native to SE United States]

Flatwoods, flatwoods bogs, savannas, and barrens; 0--200m; Ala., Del., Fla., Ga., Md., N.J., N.C., S.C., Va.

Pinus rigida Miller var. serotina (Michaux) Loudon ex Hoopes.

Pinus rigida Miller subsp. serotina (Michaux) R.T. Clausen; P. rigida var. serotina (Michaux) Hoopes

More info for the term: bog

The data of Wilbur and Christensen [87], 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 [87].

More info for the terms: competition, fuel, ground fire, seed, seed tree, serotinous, shrub, tree

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 [42]. However, one study suggests that the ability to sprout is generally restricted to seedling and saplings [45]. 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 [18].

Heat generated by a ground fire is usually of
sufficient intensity to open the serotinous cones of pond pine [6]. Pond pine cones do not
characteristically remain closed for longer than 4 to 5 years [79], 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 [83]. 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 [6].

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
[16]. After a fire on Cumberland Island, Georgia, pond pine, loblolly pine, and slash pine reseeded where the pine canopy was killed
[9]. 

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 [14]. Low-severity winter fires are valuable in preparing favorable seedbeds for establishment of pine seedlings and in partially controlling
associated hardwoods [43]. Most wildfires in the pocosins occur during the early spring and before the evergreen vegetation has increased its moisture content through spring growth
[44].

More info for the term: marsh

pond pine

marsh pine

pocosin pine

More info for the term: cover

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 [30]. Trees must be infected with heart rot disease that softens the wood and allows red-cockaded woodpeckers to excavate roost and nest cavities [52].

More info for the terms: seed, tree

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 [6], 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 [60]. 

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 [49].

More info for the terms: fire frequency, fire regime, fire suppression, forbs, frequency, fuel, marsh, natural, organic soils, peat, root collar, seed, shrub, woodland

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 [54]. 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 [60]. 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].

FIRE REGIMES:
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 [87]. 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 [55] 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 [25]. 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 [80].

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 [25]. 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 [57].

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 [72].

Community Soil Hydroperiod/water source fire frequency slash pine flatwoods mineral 3-10 years wet longleaf pine- slash pine flatwoods mineral 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 Species Review of the dominant species listed below. Find further fire regime information for the plant communities in which this species may occur by entering the species name in the FEIS home page under "Find FIRE REGIMES".

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 longleaf-slash pine P. palustris-P. e. 1-4 [11] pitch pine P. rigida 6-25 [12] loblolly pine P. taeda 3-8  cabbage palmetto-slash pine Sabal palmetto-P. e. baldcypress Taxodium distichum var. distichum 100 to > 300  pondcypress T. d. var. nutans 11]

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More info for the term: phanerophyte

RAUNKIAER [58] LIFE FORM:
Phanerophyte

More info for the terms: association, cover, forest, herbaceous, organic soils, peat, shrub, shrubs, tree

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 [25].

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 [66].  

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 [57]. 

Pine flatwoods is one of the most common vegetation types in Florida, and can be dominated by pond pine [50]. 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 [25].

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 [47]. Bay swamps, dominated by the redbay-sweetbay association, are characterized by a more or less dense growth of evergreen trees, including pond pine [57].  

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 [4]. 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 [25]. 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 [38].

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This species is known to occur in association with the following cover types (as classified by the Society of American Foresters):

More info for the term: cover

SAF COVER TYPES [21]:




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

100 Pondcypress

102 Baldcypress-tupelo

103 Water tupelo-swamp tupelo

104 Sweetbay-swamp tupelo-redbay

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This species is known to occur in the following ecosystem types (as named by the U.S. Forest Service in their Forest and Range Ecosystem [FRES] Type classification):

ECOSYSTEMS [23]:




FRES12 Longleaf-slash pine

FRES13 Loblolly-shortleaf pine

FRES16 Oak-gum-cypress

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This species is known to occur in association with the following plant community types (as classified by Küchler 1964):

More info for the term: forest

KUCHLER [37] PLANT ASSOCIATIONS:




K079 Palmetto prairie

K111 Oak-hickory-pine

K112 Southern mixed forest

K113 Southern floodplain forest

K114 Pocosin

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This species is known to occur in association with the following Rangeland Cover Types (as classified by the Society for Range Management, SRM):

More info for the term: cover

SRM (RANGELAND) COVER TYPES [67]:




812 North Florida flatwoods

814 Cabbage palm flatwoods

More info for the term: serotinous

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 [87] in pond pine dominated habitats.

More info for the terms: marsh, mast, tree

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 [56]. 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 [15].

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 [2].

More info for the term: tree

Tree

AL
DE
FL
GA
LA
MD

MS
NJ
NC
SC
TX
VA

More info for the term: peat

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 [65].

More info for the terms: root collar, seed, serotinous, wildfire

Wildfire has been responsible for the rehabilitation of pond pine stands in the past [16] 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.

More info for the terms: seed, serotinous, stratification, tree

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 [79]. Viability does not decrease in cones that remain closed for as long as 3 years [8]. 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 [79]. 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) [16].

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 [8]. 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 [5]. Pond pine seeds germinated in a nursery were unable to survive full saturation conditions [46].  

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 [8]. In pocosins, seedlings of pond pine sprout prolifically, and stands up to sapling size are primarily reestablished by sprouting [29]. However, 1 study showed that most seedlings on pond pine plots, which were completely burned, did not sprout [66]. Evidence suggests that pond pine may not sprout indefinitely, and that frequent burning of any age pond pine stand may kill it [26].

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More info for the terms: bog, climax, cover, fire frequency, forest, frequency, hardwood, herbaceous, hydroperiod, marsh, natural, organic soils, peat, seed, shrub, shrubs, succession, swamp, tree

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 [81] 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 [57]."   

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 [13]. 

Flatwoods ecosystems, along with the rest of the southeastern pine forests, have been classified as fire subclimax within the deciduous forest region [2]. Pine flatwoods are considered to be fire maintained but, without fire, they ultimately give way to hardwoods [50]. 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 [57]. Human alteration of natural fire frequency is thought to be the most common cause of successional change in flatwoods [2]. 

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 [15]. 

The bay forest type, occurring exclusively in the Coastal Plain physiographic province, frequently reverts to pond pine or Atlantic white-cedar after severe fires [47]. 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 [35]. With recurrent fire, the bay swamp, pond pine-slash pine wetlands, and Atlantic white-cedar bog may revert to shrub bogs [47]. 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 [57]. 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 [47].  

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 [26] and produce a grass-sedge bog or savannah [8]. Wells [84] 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 [57]. 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 [25]. 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 [8]. 

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 [24]; however, hybridization
with loblolly pine is common throughout southern
Delaware [64].

More info for the term: shrub

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 [68].

More info for the term: tree

The economic value of pond pine is variable. Some consider pond pine a commercially valuable species [40] as a timber source [36]. However, others have determined pond pine has low economic value as a pulpwood source [20]. 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 [8] and may make it more susceptible to fire damage [66]. Pond pine has not been widely cultivated, due to its "low" economic value [59].

Pond pine is the principal commercial species of the pocosins of North Carolina. According to Besse [6], 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 [66].

Pond pine is a major species in the forest cover type Pond Pine (Society of American Foresters Type 98) and is an associate in nine other cover types (5):

  81  Loblolly Pine
  82  Loblolly Pine-Hardwood
  84  Slash Pine
  85  Slash Pine-Hardwood
  97  Atlantic White-Cedar
100  Pondcypress
102  Baldcypress-Tupelo
103  Water Tupelo-Swamp Tupelo
104  Sweetbay-Swamp Tupelo-Redbay

Other associated tree species are sweetgum (Liquidambar styraciflua), red maple (Acer rubrum), and loblolly-bay (Gordonia lasianthus).

In the pocosins and very wet areas, a great variety of evergreen shrubs form a dense understory. Laurel-leaf greenbrier (Smilax laurifolia) is almost always present on pond pine sites. Switchcane (Arundinaria tecta) is locally abundant and, as it sprouts prolifically, it is highly favored by periodic burning. Other frequent shrub species include inkberry (Ilex glabra), large gallberry (I. coriacea), zenobia (Zenobia pulverulenta), swamp cyrilla (Cyrilla racemiflora), southern bayberry (Myrica cerifera), sweet pepperbush (Clethra alnifolia), and saw-palmetto (Serenoa repens).

The climate throughout the pond pine range is mild and humid. Frost-free days range from 210 in the North to 300 in central Florida. The normal January and February daily average temperature is 2° C (35° F) in the northern range, increasing to 16° C (60° F) for the southern range. In July and August, the normal daily average temperature ranges from 29° C (85° F) in the South to 24° C (75° F) in the North. The extremes of temperature within the range have been as low as -23° C (-10° F) and as high as 43° C (110° F). Annual precipitation increases from 1120 to 1370 mm (44 to 54 in) on a transect from north to south and from 1120 to 1420 mm (44 to 56 in) on an inland to coastal transect. July and August have an average rainfall of 100 to 200 mm (4 to 8 in) per month in contrast to 50 to 100 mm (2 to 4 in) for December and January.

The most serious disease of pond pine is red heart (Phellinus pini), which is common in most of the older pocosin stands. On the Hofmann Forest in eastern North Carolina, 90 to 95 percent of the older pond pines are infected. P. pini is a white rot fungus which occurs relatively high in the bole of the tree. It primarily dissolves the lignin of the cell walls, while cellulose is relatively unaffected. Red heart reduces the value of pond pine lumber to such an extent that many stands of sawtimber-size trees may be sold for pulpwood only; and even pulpwood yields are reduced in the severely infected trees (14).

Pond pine is subject to both fusiform rust (Cronartium quercuum f. sp. fusiforme) and eastern gall rust (C. quercuum), which cause stem and branch cankers on pines and have alternate stages on oaks. In a study of three half-sib families of pond pine, significant differences were observed in rust susceptibility. Pond pine was only half as susceptible to fusiform rust as loblolly pine (13). Comandra blister rust, caused by the fungus Cronartium comandrae, has been found on pond pine planted in Tennessee (3). The needle cast fungus (Hypoderma lethale) and brown-spot fungus (Scirrhia acicola) sometimes cause browning and foliar damage to needles but severe damage is uncommon. Coleosporium spp.may attack the foliage but have little effect on tree growth. Pond pine is also attacked by the southern pine beetle (Dendroctonus frontalis), the black turpentine beetle (D. terebrans), and engraver beetles (Ips spp.).

In natural stands, numerous pests attack cones and seeds and reduce the final seed crop. Specific insects known to damage pond pine cones and seeds are Dioryctria spp.and Leptoglossus corculus.

During drought periods, the fire hazard of pond pine sites is extremely high because of the heavy fuel accumulation in the dense understory. Dry fuel weights of the understory and litter layer are frequently more than 22.4 t/ha (10 tons/acre). When fires occur in pocosin areas the excessive fuel and large areas of unbroken forest make fire control extremely difficult. Very intense fires consume not only the trees, shrubs, and litter, but the peat soils as well.

Pond pine is monoecious. Pollen flight and female flower receptivity occur in late March in Florida and about 1 month later in North Carolina. At any given latitude, pond pine flowering is considerably later than that of loblolly (Pinus taeda) or slash (P. elliottii) pine. It is possible that in some years the flowering period of pond pine may overlap with other southern pines, but hybridization occurs infrequently. Cones ripen in September and October.

Population Differences No definite pattern of variation was found in pond pine wood specific gravity or tracheid length associated with geographic area or depth of organic material in the coastal area of North Carolina (12). Differences in both wood traits were noted among individual plots and among individual trees, however.

The average wood specific gravity of unextracted increment cores of 554 pond pine trees was 0.492 with a standard deviation of 0.049, about the same as the variation among trees in other southern pines. The oleoresin of pond pine is unique for the high proportion of limonene and low proportion of alpha- and beta-pinene.

From a study of 30 traits of pond pine and loblolly pine, slight differences were reported in some Coastal Plain trees compared with those from drier, inland areas (10). Both species, however, had a general uniformity in all locations. Pond pine is included in the North Carolina State Tree Improvement Cooperative. First-generation select trees are located in four seed orchards with a total of 12 ha (29 acres) (9).

Races and Hybrids Distinct races of pond pine have not been identified. Pond pine hybridizes with several southern pines. Although pitch and pond pine are usually separated geographically, they readily hybridize and intergrade when found together in New Jersey and Maryland. When loblolly pine occurs in association with natural hybrids of pitch and pond pines, hybrids of the complex are frequently found (17). Pond pine also hybridizes with loblolly pine, but in most localities the flowering dates are distinct enough to restrict hybridization. Evidence of introgressive hybridization, however, has been found at several locations in the North Carolina Piedmont and Coastal Plain provinces (15).

Pond pine grows surprisingly well on the better sites with soils that have good internal drainage. In fact, pond pine may be difficult to distinguish from loblolly pine in the sapling stage when in natural or planted stands on comparable upland sites. More typically, however, pond pine is found in the pocosins where it grows very slowly.

Average sizes of pond pine in a North Carolina pocosin were as follows:

Age Height D.b.h. yr m ft cm in 20 8.5 28 14.7 5.8 50 14.9 49 26.4 10.4 100 20.1 66 35.1 13.8 Stands of pond pine normally have low volumes with seldom over 70 m³/ha (5,000 fbm/acre). The trees in these stands usually have been repeatedly burned and have poor form and low quality (19).

A well-stocked stand of pond pine at 50 years of age with a site index of 21 m (70 ft) may be expected to have 25.9 m²/ha (113 ft²/acre) of basal area and 860 trees per hectare (348/acre), with an average d.b.h. of 19.6 cm (7.7 in). The entire stand might have 179 m³/ha (2,560 ft³/acre) of wood inside the bark or 195 m³/ha (31 cords/acre) of merchantable wood. The total wood volume per hectare could also be allocated as 51.8 m³ of sawtimber (3,700 fbm/acre) and an additional 146 m³/ha (23.2 cords/acre) of pulpwood (16).

Pond pine stands with a site index of more than 27 m (90 ft) at age 50 have been measured. These stands could yield 129 m³/ha (9,200 fbm/acre) of sawtimber and 151 m³/ha (23.9 cords/acre) of pulpwood. Subsoil texture is directly correlated with site index. Height growth increases with increasing amounts of fine particles (silt and clay) in the subsoil. Other soil properties that are inversely related to site index include depth to mottling, percent of organic matter in the surface soil, and total depth of organic matter (1).

Pond pine is classed as a species intolerant of shade. It is almost as intolerant as loblolly pine, but less tolerant than slash pine and more tolerant than longleaf pine (Pinus palustris). Pond pine responds favorably to drainage. In an eastern North Carolina site, basal area and height growth of pond pine were nearly doubled by drainage (6).

On mineral soils, pond pine is an intermediate stage in plant succession. As it is established primarily after fire, it may grow in pure even-aged stands until harvesting or mortality removes the dominant trees. Pond pine stands yield eventually to wetland hardwood species including oaks (Quercus spp.), gums (Nyssa spp.), hickories (Carya spp.), and magnolias (Magnolia spp.). In the pocosins, pond pine follows Atlantic white-cedar (Chamaecyparis thyoides) when the cedar is killed by fire, but cedar may become reestablished.

Fire maintains pond pine as a subclimax type through sprouting or by stimulating seed dissemination from the serotinous cones. Repeated fires during the dry season, however, may completely eliminate pond pine and produce a grass-sedge bog or savannah. In some large pocosins, woody shrubs, including cyrilla, redbay (Persea borbonia), and gallberry may completely dominate the site for a considerable time.

Almost all stands of pond pine originate from natural regeneration and the species has only infrequently been planted. In a study in the South Carolina Coastal Plain, southern pines were planted on a freshly burned site. The planting site was poorly drained, with a seasonably high water table. After 10 years, slash pine trees averaged 3.6 m (11.8 ft), and pond pine 2.3 m (7.5 ft). This study suggested that the growth of pines on the site was limited by poor nutrition and soil aeration (8).

No information is currently available.

Cone production may begin at an early age in pond pine. In eastern North Carolina pocosins, trees less than 10 years old (d.b.h.) produced an average of eight cones per tree. Cone production increased with age and diameter up to 30 years. Seed trees over 30 years (d.b.h.) produced an average of 175 to 200 cones (19).

Mature cones are normally 5 to 10 cm (2 to 4 in) long and remain on the tree for as long as 10 years. Because the cones are serotinous, only a few seeds are released at maturity each year, and the seed crop of several years may be present in an individual cone bearing tree. Viability does not decrease in cones that remain closed for as long as 3 years. Cones open gradually over several years, with two seasonal periods of seed dissemination: April through September and October through January (4).

Like other pine species with serotinous cones, pond pine cones open and release seeds soon after exposure to heat from fire. The intensity of heat does not adversely reduce the viability of seeds; even badly charred cones, following wildfires, release seeds that are capable of germination. In an uncut but burned stand of pond pine, 131,000 seeds per hectare (53,000/acre) were trapped in a 6-week period following the burn.

Mature cones can be opened by exposure to 167° to 169° C (333° to 336° F) dry heat for 30 seconds or by immersion in boiling water for a similar period. An average of 75 to 80 developed seeds are produced per cone and 77 percent of these can usually be extracted. Germination of pond pine seed from North Carolina averaged 44 percent of the extractable seed (including both filled and empty seed) after 60 days (19).

Pond pine seeds are relatively small, ranging from 104,000 to 139,000/kg (47,000 to 63,000/lb) and averaging 119,000/kg (54,000/lb). Seeds are winged and can be dispersed a horizontal distance several times the height of parent trees.

Adequate moisture is normally available for pond pine germination in the peat and organic soils of the Coastal Plain. Seeds also germinate well on exposed mineral soil. Establishment of seedlings, however, is much better if the competition from weeds, grass, and woody vegetation is reduced by mechanical site preparation or by the use of fire. Germination is epigeal.

Without question, fire has been a major influence on the perpetuation of pond pine in coastal regions of the Southeastern United States. The majority of present-day stands date to a previous wildfire. For natural regeneration, prescribed fires reduce the dense understory of shrubs, prepare a seedbed, and open the serotinous cones. Research in eastern North Carolina indicated that seedling establishment was most favorable in plots that were burned before clearcutting. Clearcutting before burning was also favorable for establishment of an adequate number of free-to-grow seedlings (4). In another study, a headfire was more effective than a flanking fire in the establishment of seedlings.

Techniques for the use of successful prescribed fire for the regeneration of pond pine have been outlined (18). Fires must generally be of high intensity to consume understory shrubs, yet not so intense that serious damage or mortality occurs to the overstory pines. There is usually only a narrow range of weather and fuel conditions where silvicultural objectives and controlled fire can be attained in a given pocosin. Key elements in a prescribed fire plan include light to moderate winds with relative humidities between 35 and 50 percent. The water table also should be close to the surface to prevent spotting and burning of the organic soils.

Seed for regeneration may be provided by seed trees. If seed trees are 23 to 25 cm (9 to 10 in) in d.b.h. and 30 years old, each tree can be expected to produce 5,000 sound seeds. On this basis, 15 to 20 seed trees per hectare (6 to 8/acre) are required to provide adequate seed for establishment. Scarification with fire plows, disks, or logging equipment is sometimes effective in securing seedling establishment but is not as reliable as burning because without heat from a fire seedfall is limited by the serotinous cones of pond pine.

Pond pine seedling growth frequently is limited by excessive soil moisture, low nutrition, and competition from herbaceous and woody vegetation. Under the most severe conditions, seedlings grow only 30 cm (12 in) or less per year. In a pond pine study in a pocosin, seedlings required an average of 18 years to reach a height of 1.5 m (5 ft). The fastest growing seedlings reached this height in less than 10 years (19).

The lower terraces of the Coastal Plain in the Southeastern United States are characterized by sandy and organic soils. Typical soils are in the Ultisols order, Aquults suborder, Albaquults and Umbraquults great groups. These soils usually have a dark gray, sandy loam surface soil and a heavy subsoil. The entire area has minor topographic relief and is interspersed with major drainage systems that are frequently very broad. Throughout the region are numerous streams, swamps, pocosins, marshes, and bays that are characterized by poorly developed drainage patterns. The pocosins, in particular, have unique topographic features in the Carolinas. These areas are upland bogs with streams draining from them on all sides. The pocosins have a considerable peat accumulation, frequently as deep as 2 m (6 ft). Also, in the southeastern Coastal Plain there are shallow, poorly drained depressions called bays or ponds. Pond pine is frequently found as the major overstory species in these pocosins and bays in association with a heavy understory of shrubby vegetation.

Although pond pine is most frequently found on poorly drained lands, the species can make excellent growth on mineral soils or on land that is not continuously waterlogged. The slow growth of pond pine is primarily a function of prolonged water saturation and reduced soil aeration. Poor aeration retards decay of organic material and results in the accumulation of muck and peat with high acidity. Although availability of mineral nutrients is usually adequate, nitrogen fixation and nitrification proceed very slowly. Consequently, the amount of available nitrogen is small even though total nitrogen content is high in the undecayed organic matter. Soil saturation also deprives roots of the oxygen required for respiration and growth and tends to keep the soil temperature low (19).

Although pond pine has poor form and slow growth, it produces forest stands of pulpwood and sawtimber where other species will not grow. The pocosins and pond pine stands are considered a major wildlife sanctuary for many wetland species.

Among pine species, pond pine is unique in that it sprouts readily from stumps until quite old. Seedlings or saplings will sprout prolifically when cut, and older trees will sprout vigorously along the stem and branches after intense scorching and defoliation by fire. These sprouts arise from dormant buds in the axils of primary needles of young seedlings. The buds are dormant and are protected by the bark. When the tree is injured, these buds resume growth and may give rise to clusters of buds and numerous short shoots (19).

Not all buds, formed at intermediate and wintergrowth flushes, sprout the following spring. Those that do not sprout remain alive; they sometimes develop into short, weak branches and give rise to many lateral buds. They may also put forth secondary dormant buds directly without growing into branches. Buds may also form in needle fascicles, but these are of little importance in the sprouting of pond pine because they are present for only a short time. Thus, pond pine stems and branches bear many clusters of dormant buds that remain alive and capable of sprouting for many years. When trees are defoliated by fire, these buds sprout and give the stems and branches a feathery appearance. Stem sprouting is one of the primary reasons for the low quality of pond pine in frequently burned areas (19).

Pinaceae -- Pine family

David L. Bramlett

Pond pine (Pinus serotina), also called marsh pine, bay pine, and pocosin pine, is a medium-sized tree that grows on soils with a high water table. The species name, serotina, means "late" and refers to the delayed opening of the cones, up to 2 years before seeds are shed (11). Open cones persist for many years and often become embedded in the growing branches, giving the tree the appearance of being overloaded with cones and a prolific seed producer. One of the largest pond pines grows in North Carolina and measures 94 cm (37 in) in d.b.h., 29 m (94 ft) in height, with a crown spread of 14 m (46 ft) (7). The wood is coarse-grained, resinous, and of fair quality (2).

Pond pine grows 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. Within its native range, pond pine is most frequently found on wet or poorly drained sites.


- The native range of pond pine.

Tree, Evergreen, Monoecious, Habit erect, Trees without or rarely having knees, Tree with bark rough or scaly, Young shoots 3-dimensional, Buds resinous, Leaves needle-like, Leaves alternate, Needle-like leaf margins finely serrulate (use magnification or slide your finger along the leaf), Leaf apex acute, Leaves > 5 cm long, Leaves > 10 cm long, Leaves yellow-green above, Leaves yellow-green below, Leaves not blue-green, Needle-like leaves triangular, Needle-like leaves twisted, Needle-like leaf habit erect, Needle-like leaves per fascicle mostly 3, Needle-like leaf sheath persistent, Twigs glabrous, Twigs viscid, Twigs not viscid, Twigs without peg-like projections or large fascicles after needles fall, Berry-like cones orange, Woody seed cones > 5 cm long, Seed cones bearing a scarlike umbo, Umbo with missing or very weak prickle, Bracts of seed cone included, Seeds brown, Seeds winged, Seeds unequally winged, Seed wings prominent, Seed wings equal to or broader than body.

Pinus serotina, the pond pine, marsh pine or pocosin pine, is a pine tree found along the Southeastern portion of the Atlantic coastal plain of the United States, from southern New Jersey south to Florida and west to southern Alabama. This pine often has a crooked growth pattern and an irregular top and grows up to 21 metres (69 ft) high, rarely to 29 metres (95 ft).

The needles are in bundles of three or four, and 15–20 cm (6–8 in) long. The almost round cones are 5–8 cm (2–3+1⁄4 in) long with small prickles on the scales. Its cones are usually serotinous, requiring fire to open. The pond pine is found in wet habitats near ponds, bays, swamps, and pocosins. Often found among long leaf pines due to their high flammability and the pond pines need for fire to germinate.

The species name serotina is derived from the persistently unopened cones that may remain closed for several years before they release their seeds; the opening is often in response to forest fires.

At the north end of its range, it intergrades and hybridizes with pitch pine (P. rigida); it is distinguished from that species by the longer needles and on average slightly larger cones. Some botanists treat pond pine as a subspecies of pitch pine.

Pond pine cones are smaller and rounder than loblolly pine cones.

Unlike loblolly pines, pond pines have the ability to grow needles directly from the trunk.