Pinus pungens Lamb.

Pinus pungens is a scrub pine and is too small and knotty to be much utilized except for pulpwood and firewood. Its common name refers to a general type of landform, not to a specific, named mountain.

Trees to 12m; trunk to 0.6m diam., straight to crooked, erect to leaning, poorly self-pruning; crown irregularly rounded or flattened. Bark red- to gray-brown, irregularly checked into scaly plates. Branches horizontally spreading; twigs slender, orange- to yellow-brown, aging darker brown, rough. Buds ovoid to cylindric, red-brown, 0.6--0.9cm, resinous. Leaves 2(--3) per fascicle, spreading or ascending, persisting 3 years, 3--6(--8)cm ´ 1--1.5mm, twisted, deep yellow-green, all surfaces with fine stomatal lines, margins harshly serrulate, apex acute to short-acuminate; sheath 0.5--1cm, base persistent. Pollen cones ellipsoid, ca. 15mm, yellow. Seed cones maturing in 2 years, variably serotinous, mostly whorled, downcurved, asymmetric, ovoid before opening, broadly ovoid when open, (4--)6--10cm, gray- to pale red-brown, nearly sessile or on stalks to 1cm; apophyses thickened, diamond-shaped, strongly keeled, elongate, mammillate at cone base abaxially; umbo central, a stout, curved, sharp claw. Seeds deltoid-obovoid, oblique; body ca. 6mm, deep purple-brown to black; wing 10--20(--30)mm. 2 n =24.

Dry, mostly sandy or shaly uplands; Appalachians and associated Piedmont; 500--1350m; Del., Ga., Md., N.J., N.C., Pa., S.C., Tenn., Va., W.Va.

More info for the terms: fire intensity, hardwood, prescribed fire, severity, tree

The effect of fire on Table Mountain pine is related to fire intensity and severity but cannot always be immediately determined. One year after prescribed fire in a mixed hardwood and pine (Table mountain pine and pitch pine) stand, tree mortality was significantly higher in areas of high and medium-high fire intensity, compared to areas with lower fire intensity. However, by the end of the 6th growing season after fire, almost all overstory pines and hardwoods were dead in all study plots burned at all intensities [75,77]. For a more complete summary of this study, see Early postfire response of Table Mountain pine stands burned under prescription at varying intensities.

More info for the terms: competition, cover, density, duff, fire severity, forest, frequency, hardwood, high-severity fire, litter, low-severity fire, moderate-severity fire, prescribed burn, seed, serotinous, severity, wildfire, xeric

While it is commonly perceived that Table Mountain pine communities require
high-severity crown fires for regeneration (e.g., [85,91]), additional evidence
(e.g., [14,52,75,77]) suggests that Table Mountain pine communities can
reproduce and persist under a regime of frequent, low-severity fire, and in some
cases, in the absence of fire [3,6]. High-severity fires certainly result in
reduced shading and competition, the opening of serotinous cones, and removal
of forest floor, thus providing seed and a favorable seed bed for germination and
establishment; and ample data provide evidence that Table Mountain pine reproduces
and persists after high-severity fires [15,33,37,56,57,61,91]. Low-severity
fires may not provide optimum conditions for Table Mountain pine regeneration [32].
For example, about 3 months following a low-severity, spring prescribed burn
on the Grandfather Ranger District, Pisgah National Forest, North Carolina, frequency
and cover of Table Mountain pine seedlings increased; however, seedlings were not
expected to survive due to shading and competition from overstory, midstory, and
understory species [71,79]. For a complete summary of this study, see
Early postfire response of southern Appalachian Table Mountain-pitch pine stands
to prescribed fires.

Several studies have found greater regeneration of Table Mountain pine after
high-severity fire than after low-severity fire. Zobel [91] sampled a recently burned
Table Mountain pine stand in western North Carolina and noted that all Table Mountain
pine cones were open, even though fire had been "light" in portions
of the area. However, 2 years after fire, regeneration persisted only in areas where
the overstory trees were killed, the undergrowth was almost completely killed,
and erosion had exposed mineral soil [91]. High-severity fire provided a
favorable seedbed for germination of Table Mountain pine following a lightning-caused
wildfire on Dovel Mountain, Shenandoah National Park, Virginia. The number of Table
Mountain pine seedlings on burned areas the second year after fire was considerably
higher than on unburned areas with high-severity, low-severity, and unburned patches
averaging 6,997, 1,489, and 386 seedlings/ha, respectively [33]. Regeneration of
Table Mountain pine following an April 1986 fire on Bote Mountain, Great Smoky
Mountains National Park, was most successful where fire severities were high. Study
plots were located on areas that experienced low-severity (surface fire only),
moderate-severity (surface fire with some scorching and torching of individual trees),
and high-severity (crown fire) fires. Overall Table Mountain pine seedling densities
were 1,250/ha, 15,312/ha, 18,304/ha in low-, moderate-, and high-severity plots,
respectively. Of all seedlings counted 75.4% were found growing where the organic
matter depth was less than 0.4 inches (1 cm) [61]. Pine regeneration (Table Mountain
pine and pitch pine combined) was greater after high-intensity fires (mean bark char
height of 25 feet (7.6 m)) than repeated low-intensity fires, when dormant season
prescribed burns were carried out in stands on xeric ridgetops in western North Carolina.
Pine regeneration was 9,440 stems/ha on high-intensity burned plots, 34 stems/ha on
low-intensity burned plots, and 7 stems/ha on control plots. Sparse regeneration on
low-intensity plots could be due to the amount of duff left on these sites, the type of
litter (hardwood litter detrimental to Table Mountain pine seedling survival), or the
amount of shading from remaining overstory trees [56,57].

High-intensity, stand-replacement fires have been recommended to regenerate stands of
Table Mountain pine; however, Mohr and others [52] (see Germination
for details), Brose and Waldrop [14], and Waldrop and Brose [75,77] provide
evidence that successful regeneration of Table Mountain pine can occur after low-
and moderate-severity fires.

Brose and Waldrop [14] provide evidence from 9 uneven-aged Table
Mountain pine-pitch pine stands across Georgia, Tennessee, and South
Carolina, that suggests frequent periodic or continuous pine and
hardwood recruitment under a regime of periodic surface fires
during the past 100 to 150 years. These fires were likely low- to
moderate-severity, given that cores and cross-sections were sampled from
living chestnut oaks which would have been killed by a high-severity
fire [14]. The results of this study suggest that successful seedling
establishment of Table Mountain pine can occur following low- and
moderate-severity fire.

Waldrop and Brose [75,77] report effects of
low, medium-low, medium-high, and high intensity fires on the
establishment of Table Mountain pine seedlings after prescribed burning
in the War Woman Wildlife Management Area of the Tallulah Ranger
District in Georgia. Sixty sample plots were surveyed at the end of the
1st and 6th growing seasons after burning. In the 1st growing season, postburn
pine density ranged from 1,396 seedlings/acre on the high
intensity plots to more

©Thomas Waldrop

than 9,130 seedlings/acre on plots burned at medium-low intensity. First year findings
reveal that fire severity did not have to reach extreme levels for Table
Mountain pine to successfully regenerate. By the end of the 6th growing season, however, seedling density decreased
by nearly half for the low and medium-low intensities, slightly decreased for medium-high
intensity, and increased on sites burned at the highest intensity. Successful Table Mountain
pine regeneration was achieved for all plots burned at all intensities [75,77]. For a more
complete summary of this study, see
Early postfire response of Table Mountain pine stands burned under prescription at varying
intensities.

Recurrent fire and maintenance of genetic diversity:
Evaluation of the morphological features and life-history characteristics of Table Mountain
pine provides strong evidence that the evolution of this species has been uniquely shaped by
fire. Fire is instrumental in opening cones and creating suitable environmental
and edaphic conditions for stand regeneration. Frequent fires are also important
in perpetuating genetic diversity within stands. Routine burning of stands would allow for
regular population turnover, in turn preventing loss of individual genotypes due to stand
senescence and maintaining a majority of the intrapopulational genetic variation in living
individuals [28].

Other information: This fire study provides information on postfire responses of plant species in communities that include Table Mountain pine:

• Research Project Summary: Early postfire response of Georgian Table Mountain pine stands burned under prescription at varying intensities

Table Mountain pine

hickory pine

mountain pine

prickly pine

Information on state-level protected status of plants in the United States is available at Plants Database.

More info for the terms: basal area, cone, forest, litter, presence, seed, serotinous, shrubs, tree, xeric

This description provides characteristics that may be relevant to fire ecology, and is not meant for identification. Keys for identification are available [21,55,66,89].

Morphology: Table Mountain pine is a native, slow-growing conifer. It is often small in stature and exceedingly limby [19]. It rarely grows beyond 66 feet (20 m) tall [21,66,92], though the tallest individual recorded was 95 feet (29 m) [91]. Table Mountain pine is typically around 16 inches (40 cm) DBH [21,66,92]. The maximum recorded DBH was 34 inches (94 cm) [91].

The trunks of Table Mountain pine are often crooked and have irregularly shaped cross-sections [23,91,92]. Older trees tend to be flat-topped, while young trees can vary in form from that of a large bush when open-grown, to slender with relatively small limbs when grown in a dense stand [91]. Table Mountain pine typically has long, low-lying, thick limbs on much of the trunk [23,91,92]. The limbs remain alive on over half of the length of the bole. Even in closed stands, where branches are smaller and limited to the upper trunk, Table Mountain pine retains branch stubs for long periods of time [91,92]. The bark of Table Mountain pine is broken by fissures into irregular plates [66] and is nearly smooth to flaky [55]. Zobel [92] states that bark thickness increases linearly with the increase in diameter.

Throughout its range, the needles of Table Mountain pine average 2.7 inches (6.8 cm) in length, ranging from 1.3 inches (3.3 cm) to 4.1 inches (10 cm). They are borne in a 2-needle fascicle. Three-needle fascicles may occur on the same tree but are rare [21,92]. Needles and fascicles are retained for 2 to 3 years [22,60].

Male cones are 0.6 inches (1.5 cm) long. Female cones are sessile, range from 1.7 to 4.1 inches (4.2-10 cm) long, and occur in whorls of 2 to 7. Cone scales are tough and armed with broad, stout, upwardly curving spines [21,23,55,66,92]. Female cones are typically serotinous, although serotiny varies among individuals [3] and populations [91,92]. See Seed dispersal for more information on the serotiny of this species. Seeds are triangular and approximately 0.25 inches (0.63 cm) long, with wings about 1 inch (2.5 cm) long [49]. The size of cones and seeds decreases with increasing elevation [92].

Table Mountain pine seedlings generally anchor their taproot into a rock crevice. Secondary or lateral roots then spread through soil and litter, taking up both moisture and nutrients. Other sinker roots descend into additional crevices, utilizing accumulated soil and the thin skins of finely weathered, nutrient-rich, moist soil-like rock which coat the crevice surface [19].

Age class structure: The age class structure of many Table Mountain pine stands suggests that fire is an important influence on stand structure and regeneration [91]. Many Table Mountain pine stands observed on the Chattahoochee National Forest, Georgia are represented by large DBH size classes, suggesting older age classes, and show no evidence of pine regeneration [70]. Williams and Johnson [86] report that the age distribution of Table Mountain pine in pine-oak forests near Blacksburg, Virginia, was bimodal with peaks in the seedling (10 year old) and large tree (45 to 80 year old) age classes. Estimated stem ages for the study area ranged from 1 to 124 years. There were few individuals in the 20- to 35-year age classes. This gap in age distributions suggests that recruitment may be episodic [86] (e.g., following periodic fire) in these populations.

The presence of many size classes suggests that some Table Mountain pine stands can persist for long periods in the absence of fire. Table Mountain pine stands that appear to be "permanent" are generally associated with a sparse understory, shallower litter, more rock outcrop, and less basal area, than stands that are moving towards a late-seral, oak-dominated forest, with 1 size class of pines, no pine reproduction, and a dense understory of shrubs [91]. Three Table Mountain pine stands located on a xeric, southwest shoulder of Looking Glass Rock, North Carolina, were reported to have continuous recruitment in the absence of fire for 87 years. The age structure of these stands suggests relatively continuous recruitment during each of 3, 20-year periods with a mortality rate of 50% per period, except for one period in which 2 severe droughts were thought to decrease recruitment and increase mortality of saplings and seedlings. Sites supporting continuous recruitment had shallow, rapidly draining soils and little or no mountain-laurel present [2].

A maximum age of 250 years is reported for Table Mountain pine. The oldest recorded individuals in the Appalachians were 227 and 205 years, and were found in the southwest corner of North Carolina [91].

Table Mountain pine is endemic to the Appalachian Mountains of the eastern United States. Its distribution is primarily in the Blue Ridge and Valley-and-Ridge provinces of the Appalachian Highlands although its range extends from central Pennsylvania, southwest to eastern West Virginia and southward into North Carolina, Tennessee, and the extreme northeast corner of Georgia. There are many outlying populations of Table Mountain pine to the east of the Appalachians and a few to the west [16,19,91]. Table Mountain pine is present in 1 county in the northeast corner of Illinois [53]. Flora of North America provides a distributional map of Table Mountain pine.

More info for the terms: crown fire, fire exclusion, fire interval, fire regime, fire-free interval, fire-return interval, forest, frequency, fuel, fuel loading, fuel moisture, low-severity fire, natural, potential natural vegetation, potential natural vegetation group, seed, serotinous, stand-replacing fire, tree, woodland

Fire adaptations: Table Mountain pine has adaptations to fire that are consistent with both long- and short-return-interval fire regimes. Medium-thick to thick bark, a deep rooting habit, self-pruning limbs, and pitch production to seal wounds are characteristics of Table Mountain pine that suggest it is adapted to survive frequent, low-severity fire [3,39,47,52,67,73,76,85]. Delayed seed release from serotinous cones and trees that produce viable seed at a young age also allow Table Mountain pine populations to persist after infrequent, high-severity fire [47,73].

Table Mountain pine can sprout following injury [39,46,56,65], such as that from fire or animal damage [91], but no data in the literature describe resprouting after fire in Table Mountain pine. Table Mountain pine seedlings of "natural origin" usually have a crook just above or just below ground level which may serve to protect basal buds against fire [19,91].

FIRE REGIMES: Table Mountain pine was historically subject to a full range of fire types: frequent low-severity surface fires, mixed-severity fires, and stand-replacement fires [27,36,74]. Table Mountain pine is also adapted to a range of fire frequencies [3]. Fire occurs infrequently on contemporary Appalachian landscapes [41] where Table Mountain pine is common.

Fire histories developed for 2 Table Mountain pine communities in southwestern Virginia revealed that between 1758 and 1944, fires burned approximately every 5 to 10 years during the dormant season. Recent regeneration failure of Table Mountain pine and increasing dominance of oak species appear to coincide with fire exclusion practices initiated after 1950 [67,68]. Based on USDA Forest Service records, Wade and others [74] state that between 1800 and 1944 the fire interval in sampled Table Mountain pine stands averaged 10 to 12 years, and that it is currently 7 to 70 years with an average of 40 years [74]. The mean interval between fires, based on fire scars from pine forests in the Great Smoky Mountains National Park, for the period 1856 to 1940 was 12.7 years. Fires were more frequent, however, at lower elevations. Most of these fires were probably anthropogenic and may have been an important influence ever since Native Americans settled in the Little Tennessee River Valley [36].

Frost [27] uses the term polycyclic (communities with 2 or more kinds of fire cycles) to describe presettlement FIRE REGIMES in pitch pine-Table Mountain pine stands on dry, south-facing slopes. Past FIRE REGIMES in these communities consisted of a cycle of high-frequency, understory fires (about 5 to 7 years apart) interrupted periodically by a long fire-free interval (about 75 years) followed by stand-replacing fire. Frost categorizes Table Mountain pine as having a short fire-return interval (25 to 100 years). "Canopy thinning" occurs when fuel loading, fuel moisture, and wind create prolonged or severe fire behavior but fall short of initiating crown fire, and has been observed in pitch pine-Table Mountain pine stands in the southern Appalachians [27].

In the eastern United States, pitch and Virginia pine forests that Table Mountain pine is commonly associated with were subject to mixed FIRE REGIMES. Where burning by Native Americans was common, FIRE REGIMES in pitch pine communities were characterized by understory fires on a 2- to 10-year interval. Fire in Virginia pine was probably less frequent and resulted in higher tree mortality. These forest types are still characterized as mixed fire regime type because the fire return intervals are longer and the majority of wildfires occur during the growing season when damage is greater [74].

Based on species traits and presettlement site descriptions, Landers [42] suggests an "inferred fire regime" for Table Mountain pine stands in the southeastern United States, with "very intense fires" at a frequency of 2 per century [42]. Where Table Mountain pine forms even-aged, pure stands, Wade and others [74] classify the fire regime as typically stand-replacement at intervals of <35 to 200 years.

The following table provides fire return intervals for plant communities and ecosystems in which Table Mountain pine is important. 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".

Fire regime information on vegetation communities in which Table Mountain pine may occur. For each community, fire regime characteristics are taken from the LANDFIRE Rapid Assessment Vegetation Models [44]. These vegetation models were developed by local experts using available literature, local data, and/or expert opinion as documented in the PDF file linked from the name of each Potential Natural Vegetation Group listed below. Cells are blank where information is not available in the Rapid Assessment Vegetation Model. Northeast Southern Appalachians Northeast

• Northeast Woodland
• Northeast Forested

Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics Percent of fires Mean interval
(years) Minimum interval
(years) Maximum interval
(years) Northeast Woodland Eastern woodland mosaic Replacement 2% 200 100 300 Mixed 9% 40 20 60 Surface or low 89% 4 1 7 Rocky outcrop pine (Northeast) Replacement 16% 128     Mixed 32% 65     Surface or low 52% 40     Pine barrens Replacement 10% 78     Mixed 25% 32     Surface or low 65% 12     Oak-pine (eastern dry-xeric) Replacement 4% 185     Mixed 7% 110     Surface or low 90% 8     Northeast Forested Appalachian oak forest (dry-mesic) Replacement 2% 625 500 >1,000 Mixed 6% 250 200 500 Surface or low 92% 15 7 26 Southern Appalachians

• Southern Appalachians Woodland
• Southern Appalachians Forested

Vegetation Community (Potential Natural Vegetation Group) Fire severity* Fire regime characteristics Percent of fires Mean interval
(years) Minimum interval
(years) Maximum interval
(years) Southern Appalachians Woodland Appalachian shortleaf pine Replacement 4% 125     Mixed 4% 155     Surface or low 92% 6     Table Mountain-pitch pine Replacement 5% 100     Mixed 3% 160     Surface or low 92% 5     Southern Appalachians Forested Appalachian oak-hickory-pine Replacement 3% 180 30 500 Mixed 8% 65 15 150 Surface or low 89% 6 3 10 Oak (eastern dry-xeric) Replacement 6% 128 50   Mixed 16% 50 20   Surface or low 78% 10 1 10 Appalachian Virginia pine Replacement 20% 110 25 125 Mixed 15% 145     Surface or low 64% 35 10 40 Appalachian oak forest (dry-mesic) Replacement 6% 220     Mixed 15% 90     Surface or low 79% 17     *Fire Severities:
Replacement=Any fire that causes greater than 75% top removal of a vegetation-fuel type, resulting in general replacement of existing vegetation; may or may not cause a lethal effect on the plants.
Mixed=Any fire burning more than 5% of an area that does not qualify as a replacement, surface, or low-severity fire; includes mosaic and other fires that are intermediate in effects.
Surface or low=Any fire that causes less than 25% upper layer replacement and/or removal in a vegetation-fuel class but burns 5% or more of the area [35,43].

More info for the terms: competition, duff, fire exclusion, fire regime, forest, hardwood, litter, natural, seed, succession, tree

Table Mountain pine is typically fire-dependent, requiring fire to ensure optimal regeneration and maintenance [33,85]. Fire plays an important role in creating the environmental and biological conditions necessary for natural regeneration of Table Mountain pine forests [34]. Beginning in the 1930's, changes in land use and effective fire exclusion programs slowed or reversed pine forest expansion [85]. Fire exclusion in the southern Appalachians is thought to decrease regeneration in Table Mountain pine [17] and allow for accelerated succession to hardwood-dominated forests that ultimately threaten the perpetuation of Table Mountain pine populations [41,67].

Findings by Welch and others [79] on the first-year response of southern Appalachian Table Mountain pine and pitch pine stands to prescribed burning, suggest that prescribed fires in which objective is to restore these communities must open the canopy, reduce accumulated litter and duff layers, and expose regenerative buds of hardwoods to lethal temperatures in order to lessen postburn sprouting. Prescribed burns that do not accomplish these goals may further encourage succession towards hardwood-dominated stands [79]. For complete details of this study, see Early postfire response of southern Appalachian Table Mountain-pitch pine stands to prescribed fires.

Declines in Table Mountain pine populations during the era of fire exclusion have greatly reduced the seed source for this species. Oaks have spread into these communities and, in many cases, dominate the canopy. In addition, there is tremendous litter build-up on these areas. A review by Buckner and Turrill [15] recommends intense, small scale prescribed burning to clear the forest floor down to mineral soil, eliminate hardwood competition, and allow Table Mountain pine regeneration. If the previous Table Mountain pine community was much depleted, this process might have to be repeated on a 5- to 7-year cycle, similar to the "pulsed" prescribed fire regime used to manage other yellow or hard pines of eastern North America [84]. Such burning practices could restore Table Mountain pine populations on a local scale [15].

If management of declining populations of Table Mountain pine is to be effective, the development of a prescribed burning plan should consider tree age and burn season to ensure that an adequate and viable seed source is present. This conclusion was drawn from a study on the patterns of seed production in Table Mountain pine, which revealed that the number of seeds and viability of seeds was greatest in cones collected in the winter, even though cones are mature in the fall of the second year [31].

More info on this topic.

More info for the term: phanerophyte

RAUNKIAER [58] LIFE FORM:
Phanerophyte

More info for the terms: series, tree

Table Mountain pine is endemic to the Appalachian Mountains and commonly dominates steep, rocky ridges and exposed escarpments too marginal for many tree species [23,28,41,91]. Throughout its range, Table Mountain pine is most commonly found on south- and west-facing slopes at elevations ranging from 1,000 to 4,000 feet (305-1,220 m) [7,12,91]. Elevation extremes are noted from about 150 feet (46 m) near Delaware to over 4,700 feet (1,430 m) in Tennessee. In the Great Smoky Mountains of Tennessee and North Carolina, it has been reported in elevations up to 5,800 feet (1,767 m). Outlying populations are associated with monadnocks (isolated mountain or rocky mass in an otherwise level area), and other eastern range extensions are along major rivers. Table Mountain pine has been reported around bogs on Mt. Pisgah, North Carolina, and at Big Meadows, Shenandoah National Park, Virginia [91].

Soils: Stands dominated by Table Mountain pine are usually associated with exposed rocky sites with shallow soil over bedrock or a very high rock content [23,42]. The soil series where Table Mountain pine dominates are generally stony; shallow, sometimes without profile development; strongly acidic (pH 4.5-5.5); infertile and of low productivity; and well to excessively drained [12,42,91].

On Looking Glass Rock, western North Carolina, stunted (<33 feet (<10 m) tall) Table Mountain pine survive where the soil depth in cracks and depressions in the granite is 4 to 16 inches (10-40 cm) [5].

Climate: The climate throughout Table Mountain pine's narrow range is warm, humid, and continental. Average monthly high temperatures range from 20 °F (-7 °C) in January to 85 °F (29 °C) in July. Mean annual precipitation ranges from 30 to more than 80 inches (760-2,030 mm) distributed evenly throughout the year [13,19]. The average number of frost-free days in Pennsylvania varies from 150 to 170, and from 170 to 180 in the mountains of Tennessee, North Carolina, and northern Georgia [19]. Table Mountain pine most often occurs on the warmest and driest microsites within these areas [91].

More info for the terms: seed, serotinous

Table Mountain pine often survives fire, but trees can be top-killed or killed [71,75,77,79].

After passage of fire, the resin of the serotinous cones melts and the scales reflex for seed release and dispersal. Barden [3] reveals that 78% of cones opened after 2 minutes but before 4 minutes when heated in an oven at 210 °F (100 °C).

More info for the terms: cover, seed, serotinous

Table Mountain pine's serotinous cones are of special importance as a year-round food source for the American red squirrel [23,91]. Red squirrels may limit the growth and competitive ability of Table Mountain pine, as well as limiting the reserve seed crop, by destroying past growth and photosynthetic tissue when they "cut" branches and then remove cones on the ground [91].

Palatability/nutritional value: No information is available on this topic.

Cover value: No information is available on this topic.

More info for the terms: cover, shrub, shrubs, tree, xeric

Table Mountain pine is common in upland yellow pine communities that are dominated by
Table Mountain pine, pitch pine, shortleaf pine, and Virginia pine (Pinus virginiana)
with an understory of chestnut oak (Quercus prinus) [7,13]. It is
also often found in pure stands or co-dominant with pitch pine [30]. Other
understory tree species that may be present in Table Mountain pine stands are
red maple (Acer rubrum), black tupelo (Nyssa sylvatica), sourwood (Oxydendrum arboreum),
scarlet oak (Quercus coccinea), bear oak (Q. ilicifolia), white oak (Q. alba),
post oak (Q. stellata), southern red oak (Q. falcata), blackjack oak (Q. marilandica),
black oak (Q. velutina), American chestnut (Castanea dentata), black locust
(Robinia pseudoacacia), eastern hemlock (Tsuga canadensis), and sweet birch (Betula
lenta) [4,7,12,19,69]. Table Mountain pine populations in northeastern
Georgia are also associated with hickory (Carya spp.) and eastern white pine (Pinus
strobus) [12]. On Looking Glass Rock, western North Carolina, Table Mountain pine grows with
eastern redcedar (Juniperus virginiana) [5]. Mountain-laurel (Kalmia
latifolia) is the most common shrub species associated with Table Mountain pine stands. Other
ericaceous shrub species such as mountain fetterbush (Pieris floribunda), hillside blueberry
(Vaccinium pallidum), black huckleberry (Gaylussacia baccata), bear huckleberry
(G. ursina), and several species of rhododendron (Rhododendron spp.) are common
associates [12,23,85].

Table Mountain pine is associated with the Virginia pine-oak [10], Virginia pine
[11], pitch pine [45], shortleaf pine [80], and chestnut oak [18] cover
types, as recognized by the Society of American Foresters.

Table Mountain pine is a dominant species in the following vegetation types
and plant communities:

North Carolina:

• Table Mountain pine


• Table Mountain pine/sandmyrtle (Leiophyllum buxifolium)


• Table Mountain pine/Tall, evergreen heaths


• Table Mountain pine-Virginia pine


• Table Mountain pine-Mixed oaks and hardwoods/mountain-laurel–rosebay (Rhododendron maximum)


• pitch pine-Table Mountain pine/black huckleberry


• pitch pine-Table Mountain pine/mountain-laurel


• pitch pine-Table Mountain pine/sandmyrtle


• pitch pine-Table Mountain pine/catawba rosebay (Rhododendron catawbiense)


• pitch pine-Table Mountain pine/Carolina azalea (Rhododendron carolinianum)


• Table Mountain pine-Carolina hemlock (Tsuga caroliniana)-bigtooth aspen (Populus
  grandidentata)/mixed shrubs


• Table Mountain pine/mountain-laurel


• Table Mountain pine-pitch pine/mountain-laurel


• Table Mountain pine/black huckleberry


• Table Mountain pine-pitch pine/mixed heaths [62].

Virginia:

• red oak (Quercus rubra) [69].

Black and Craggy Mountains, North Carolina:

• Xeric Pine Forests [50]

Great Smoky Mountains:

• Table Mountain pine heaths [82]

More info for the term: tree

Tree

More info for the terms: basal area, cone, seed, tree

Table Mountain pine is a preferred species for the southern pine beetle and can be selected,
attacked, and killed [1,41,54]. Table Mountain pine is also susceptible to comandra blister rust,
which is a canker disease of hard pines caused by the fungus Cronartium comandrae [51].
Trees of all sizes are attacked by the pine twig gall scale which causes bark to swell and crack,
killing foliage and tree [19].

The needles of Table Mountain pine can be affected by hypoderma needle blight
which causes mortality of infected needles [9]. The European pine sawfly can defoliate trees of their previous year's
needles but seldom kills the trees [19]. The cones of Table Mountain pine are susceptible to Table
Mountain pine coneworm (Diorytria yatesi) [31], and cone boring insects may have a significant
effect on Table Mountain pine. The larva eats the part of the maturing cone between the central core
and the outer layer of the cone, destroying the seeds. Damage varies from year to year, but in some
years it appears to have destroyed the entire seed crop of an area. Heart rot is common in larger Table
Mountain pine trees, usually over 100 years old. It appears to enter most often through fire scars and
from long-persistent branch stubs characteristic of Table Mountain pine [91].

On Brush Mountain, Virginia, seed predators that attack cones of Table Mountain pine include red
squirrels, larvae of the mountain pine coneworm, and the shield-backed pine seed
bug [87].

Ice storms cause extensive damage to Table Mountain pine trees and can decrease basal area
[41,81]. In a study by Whitney and Johnson [81], over 82% of Table Mountain pine were
damaged by heavy glaze. Of these trees, 85% experienced 50% crown loss. Sampling was done two
years after storm. Seedlings did increase after the storm due to the canopy gap
created by fallen trees [81].

More info for the terms: fuel, softwood

Table Mountain pine provides valuable soil and watershed protection [19,23] and wildlife habitat [23].

Wood Products: Table Mountain pine is a softwood species [63] and has limited economic importance for timber products because of its small stature and often poor stem shape. Small amounts of Table Mountain pine are harvested, along with other pines, for fuel and commercially for pulp, paper production, and small roundwood [19,23,49]. Table Mountain pine has some potential for small saw timber when reintroduced in mixture with naturally occurring hardwoods [90].

Table Mountain pine is commonly found on steep slopes with limited access that are likely to be unsuitable for timber production [86].

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In northwestern North Carolina, pollen release at 1,500 feet (457 m) elevation begins the last week of March and ends during the first week in April; at 2,500 feet (762 m), pollen release begins about the second week in April and ends near the end of the third week. Cones ripen in autumn of the second season. The opening of cones depends on the degree of serotiny [19]. Seeds are shed from September to November [55], although on Brush Mountain, Virginia, seedfall of Table Mountain pine occurred throughout the year but was primarily concentrated in the spring and summer months [87].

Table Mountain pine's early spring development and reproductive activity (development of male and female strobili) could exclude it from extensively colonizing areas with later spring frosts. By "flowering" and initiating twig growth earlier, Table Mountain pine is more susceptible to frost damage than other pine species studied [91].

More info for the terms: competition, serotinous

Fire plays an important role in the maintenance of Table Mountain pine in most of the communities where it occurs [48]. Fire opens serotinous cones, prepares a suitable seedbed for germination, and removes competition from the understory and overstory in the short-term [91].

More info for the terms: crown residual colonizer, secondary colonizer, seed

POSTFIRE REGENERATION STRATEGY [64]:
Crown residual colonizer (on-site, initial community)
Secondary colonizer (on-site or off-site seed sources)

More info for the terms: cone, duff, forest, litter, low-severity fire, monoecious, seed, serotinous, severity, shrub, soil moisture regimes, tree, xeric

Table Mountain pine regenerates mainly from seed [23,52,91]. Table Mountain pine has one or more basal buds near the cotyledonary level with additional buds often appearing as a quasi whorl an inch or so above this point, allowing it to sprout after injury [19,39,46,56,65,91].

Fire aids the regeneration of Table Mountain pine in several ways. It opens the serotinous cones, consumes litter, exposes mineral soil, and eliminates competing vegetation, allowing more light and water for pine seedlings and minimizing allelopathic effects [91].

Pollination: Table Mountain pine is pollinated by wind [24].

Breeding system: Table Mountain pine is monoecious [19]. Feret and others [24] found that Table Mountain pine stands in southwest Virginia have large genetic variability within a relatively small area.

Table Mountain pine is predominantly outcrossed [29].

For information on how fire relates to Table Mountain pine genetic diversity see Recurrent fire and maintenance of genetic diversity.

Seed production: Table Mountain pine begins cone and viable seed production when saplings are 5 to 7 years old [15,47,49,76]. Della-Bianca [19] states that, although Table Mountain pine cones shed their seeds irregularly, large numbers of seeds are disseminated annually. On Looking Glass Rock, North Carolina, Table Mountain pine produced one whorl of 2 to 5 cones each year on the new growth of branches. Cones remained closed for 2 to 5 years after maturity, and then about 40% of them opened without fire. Cones remained attached to the stem for a decade or longer, whether or not they opened [3].

Patterns of seed production in Table Mountain pine were studied by Gray and others [31] to determine if Table Mountain pine seed viability and availability varied with tree age, cone age, and season. Cones were collected from stands where Table Mountain pine was the main component and where a wide range of age classes was present. Seed viability was determined in a greenhouse where cones were dried in an oven for a minimum of 12 hours at 140 °F (60 °C) to allow them to open. Following drying, seeds were extracted, wings removed and germination tests conducted. The average number of seeds per cone tended to be higher in the younger age classes (5-25 years); however, viability was lowest in the 5- to 10-year-old age class [31].

Average number of seeds/cone and percent viability of seeds for Table Mountain pine by tree age class [31] Tree age class (years) Average number of seeds/cone* Average percent viability of seed 5 to 10 46.0a, b 8.8b 11 to 25 51.9a 33.3a 26 to 50 43.5a, b 32.7a 51 to 75 41.5b 32.9a 76+ 37.9b 38.6a *Averages followed by the same letter do not differ at alpha=0.05
The number of seeds per cone and percent viability of seeds increased with increasing cone age.

Average number of seeds/cone and percent viability of seeds for Table Mountain pine by cone age [31] Cone age (years) Average number of seeds/cone* Average percent viability of seed 2 38.2b 27.3b 3 46.0a 27.6b 4 47.3a 38.5a 5 47.0a 36.7a *Averages followed by the same letter do not differ at alpha=0.05
Percent viability of seeds was lowest in cones collected in fall and summer.

Average number of seeds/cone and percent viability of seeds for Table Mountain pine by season collected [31] Season collected Average number of seeds/cone* Average percent viability of seed Fall 45.9a 28.1b Winter 49.8a 40.0a Spring 39.2b 35.4a Summer 37.5b 21.2c *Averages followed by the same letter do not differ at alpha=0.05
Seed dispersal: Cones of Table Mountain pine are typically serotinous, requiring high temperatures from solar radiation or fire to loosen the resin and release the seeds [3,12,49,91]; however, serotiny varies among populations [92] and individuals [3]. Zobel [92] observes that some populations of Table Mountain pine show no serotiny while in others, cones generally remain closed for over 5 years and may remain closed for 25 to 30 years. Serotiny was positively correlated with elevation and negatively correlated with latitude based on range-wide observations of Table Mountain pine in the Appalachians [91]. McIntyre [49] suggests that serotiny is more common in closed stands on southern and eastern exposures, although the area in which these observations were made is not specified. On a single xeric site in North Carolina, Barden [3] found that Table Mountain pine cones remained closed for 2 to 5 years after maturity, and then about 40% of cones >2 years old opened without fire. Cones remained attached to stems for a decade or longer, whether or not they opened [3].

Table Mountain pine cones open in response to heat. When cones were heated in an oven at 210 °F (100 °C), 9% opened within 2 minutes, 78% opened between 2 and 4 minutes, and 100% opened after 7.5 minutes [3]. McIntyre [49] observed that Table Mountain pine cones placed by a furnace "opened readily" at approximately 95 °F (35 °C), and that the temperature at the end of a Table Mountain pine branch in full sunlight in the field averaged 95 °F (35 °C). This suggests that cones in the shade may not be exposed to temperatures sufficient to open cones, and may therefore remain closed for several years until exposed to adequate temperature regimes [49].

Table Mountain pine seeds are dispersed with the aid of large wings [23].

Seed banking: Table Mountain pine maintains an aerial seed bank of viable seeds in serotinous cones an average of 9 to 11 years [3,6,49] and up to 30 years [91]. Table Mountain pine establishes from seed released from canopy seed banks after fire [59].

Germination: Germination of Table Mountain pine occurs after cones open and seeds are released, as long as there is a suitable seedbed [91]. Groeschl and others [33] state that overstory canopy reduction, a decrease in forest floor litter, and the elimination of dense shrub layers are necessary to ensure adequate germination and establishment of Table Mountain pine. In the past, infrequent, high-severity crown fires provided these conditions [3,33,85,91]. However, a field and greenhouse study by Waldrop [78] reported that germination was higher under shade than in full sun, and was highest under 63% shade. Germination on 2 and 4 inches (5 and 10 cm) of duff was comparable to germination on no duff. Results suggest that lower severity fires can produce a suitable seedbed for germination of Table Mountain pine [78]. Barden [2,3,6] describes stands of Table Mountain pine on an extremely xeric site in western North Carolina that exhibited continuous recruitment despite the absence of fire since 1889.

A greenhouse study by Mohr and others [52] compared germination rates of Table Mountain pine seeds on different duff depth and shade level combinations to determine the best microhabitat for germination and survival of Table Mountain pine seedlings. Seeds were placed on duff 0, 2, and 4 inches (0, 5, and 10 cm) deep, and shade levels were 0% (full sun), 38%, 52%, and 98%. Samples were watered twice a week during June, once a week in July, and every 10 days in August. Germination was not statistically different on different duff depths or under different shade levels (see table below). Seedling height varied little among duff depths, while shade had a significant effect on seedling height (see table below). Seedling survival was greatest with 2 or 4 inches (5 or 10 cm) of duff. Moderate shade (52%) with either 2 or 4 inches (5 or 10 cm) of duff was the best treatment combination in this study for Table Mountain pine seedling survival. These results suggest that germination and establishment of Table Mountain pine is possible after low-severity fire or possibly without fire.

Mean percent germination and seedling height by duff depth and shade level [52] Treatment level Mean percent germination Seedling height* (inches) Duff depth (inches) 0 70.7 3.5a 2 68.5 3.6a 4 63.3 3.8b Shade level (percent) 0 62.6 3.5a 38 67.3 3.9b 52 69.7 4.1b 98 70.6 3.0c *Means followed by the same letter within a treatment group are not significantly different at the 0.05 level

Seedling establishment/growth: The requirements for successful Table Mountain pine seedling establishment and growth are unclear, and may vary among sites and plant communities. Zobel [91] states that very few Table Mountain pine seedlings occur on sites without exposed mineral soil or on sites with a dense canopy or shrub layer. These microsite conditions often occur after fire [75]. Whether or not high-severity, stand-replacing fires are needed to create these conditions is unclear. A greenhouse study [52] found that seedling establishment of Table Mountain pine was most successful on 2 or 4 inches (5 or 10 cm) of duff and under moderate shade (52%) (see table above). These results suggest that establishment of Table Mountain pine may be possible after low-severity fire or possibly without fire. For complete details on how seedling establishment and regeneration is effected by fire, see Discussion and Qualification of Plant Response.

In Virginia, the low availability of suitable seedling habitat strongly limits recruitment of Table Mountain pine in pine-oak forests. Williams [84] found that Table Mountain pine seedlings occurred almost exclusively in open microsites with shallow pine litter, which are relatively rare in mature pine-oak forest. Summer drought also contributed to low seedling survivorship, particularly in very young seedlings. He concludes that optimal regeneration and maintenance of Table Mountain pine are unlikely in the absence of fire when occurring in pine-oak forests suitable for growth of hardwoods, particularly oaks. Increased oak dominance leads to site modifications that are detrimental to Table Mountain pine recruitment such as litter accumulation and shading [84,86,87].

Oak litter accumulation and available soil moisture impact Table Mountain pine seedling establishment. A greenhouse study was conducted by Williams and others [88] to distinguish these effects. Soil and litter were taken from a pine-oak forest on Brush Mountain, Virginia. Oak litter was comprised of chestnut oak and scarlet oak. Pine litter was comprised of Table Mountain pine and pitch pine needles. Litter-free flats lost more water than flats with litter, and there was little difference in soil moisture between the pine and oak litter flats. Leaf litter, water regime, and the interaction of these factors had significant (P=0.0001) effects upon seedling mortality and survival. Mean seedling emergence and mortality for Table Mountain pine are provided in the table below [88].

Mean seedling emergence and mortality (percent) for Table Mountain pine under different soil moisture regimes and leaf litter accumulations [88]   Litter-free Pine litter Oak litter Watering interval Seedling emergence Mortality Seedling emergence Mortality Seedling emergence Mortality Daily 83.3 0.01 80 1.7 56 3.7 Eight-day 54.7 100 65.3 14.0 37.3 3.6 Fifteen-day 0.02 100 60.7 98.9 30 82.2

Oak litter had a negative effect on Table Mountain pine seedlings when compared to that of pine litter and litter-free treatments, although it did not completely hinder recruitment. Several germinated but desiccated Table Mountain pine seeds were recovered from oak litter treatments during harvest of seedlings. Allelopathic effects are not believed to be the reason for lowered seedling emergence in oak litter, since the seedling mortality was similar between the oak and pine litter treatments. Further studies are needed to understand possible allelopathic effects of oak litter on establishment of Table Mountain pine [88].

Vegetative regeneration: Table Mountain pine can sprout from the base after the stem is injured [19,39,46,56,65,91]. It has one or more basal buds near the cotyledonary level with additional buds often appearing an inch or so above this point that elongate readily only after injury [19,65,91]. Seedlings of "natural origin" usually have a crook just above or just below ground level which may serve to protect basal buds against fire [19,91]. Some buds appear to be short-lived, but the sprouting of cut stumps of large Table Mountain pine trees suggest that some persist [65].

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More info for the terms: hardwood, succession

Table Mountain pine is a shade-intolerant and drought tolerant pioneer species that typically establishes stands following disturbance, such as fire [12,15,25,40,41,70]. In later seral stages, on sites suitable for growth of hardwood species, it is replaced by more shade-tolerant hardwoods, specifically oaks, with mountain-laurel understories [12,15,56,68]. Stands with dense mountain-laurel understories and established oaks lack regeneration niches needed by Table Mountain pine [70]. Outbreaks of southern pine bark beetle and ice storms hasten succession toward oak dominance because they both select against Table Mountain pine [12]. Regeneration of Table Mountain pine on sites favorable for the growth of oaks and other hardwoods, like the pine-oak forests of Brush Mountain, Virginia, occurs only after fire. On sites favorable for the growth of oaks, incorporating fire at frequent intervals would facilitate the maintenance of Table Mountain pine [87].

Table Mountain pine may produce self-maintaining or nonsuccessional populations [83]. Populations of Table Mountain pine occur and reproduce in the absence of fire on dry exposed ridges and bedrock outcrops [6].

More info for the term: natural

The scientific name of Table Mountain pine is Pinus pungens Lamb. (Pinaceae) [26,38].

Hybrids: Dorman [20] suggests that natural hybridization occurs,
to a small degree, between Table Mountain pine and shortleaf pine (Pinus echinata),
and between Table Mountain pine and pitch pine (P. rigida). This conclusion is based on
occurrence of trees with intermediate morphological characteristics.

In Table Mountain pine stands of the Great Smoky Mountains associated tree species are red maple (Acer rubrum), blackgum (Nyssa sylvatica), sourwood (Oxydendrum arboreum), pitch pine (Pinus rigida), and chestnut oak Quercus prinus). In Table Mountain pine-pitch pine stands, additional associated species include scarlet oak (Q. coccinea), American chestnut (Castanea dentata), and black locust (Robinia pseudoacacia) (16).

Table Mountain pine is also a minor associate in five forest cover types: Pitch Pine (Society of American Foresters Type 45), Shortleaf Pine (Type 75), Chestnut Oak (Type 44), White Pine-Chestnut Oak (Type 51), and Virginia Pine-Oak (Type 78) (13). In the Shortleaf Pine type, Table Mountain pine is included only in the mountains; other associates are pitch pine, eastern hemlock (Tsuga canadensis), black, white, southern red, scarlet, chestnut, post, and blackjack oaks Quercus velutina, Q. alba, Q. falcata, Q. coccinea, Q. prinus, Q. stellata, and Q. marilandica, respectively), hickories (Carya spp.), and blackgum. In the Virginia Pine-Oak type, associated species in the foothills include shortleaf pine (Pinus echinata) and pitch pine, black, white, scarlet, chestnut, post, and blackjack oaks, hickories, and blackgum (13,30,31).

The lower canopy vegetation in Table Mountain pine stands (8,16,31) includes rosebay rhododendron (Rhododendron maximum), Catawba rhododendron (R. catawbiense), Piedmont rhododendron (R. minus), mountain-laurel (Kalmia latifolia), mountain winterberry (Ilex montana), hobblebush (Viburnum alnifolium), blueberries (Vaccinium spp.), sawbrier (Smilax glauca), greenbrier (S. rotundifolia), fetterbush (Pieris floribunda), white-alder (Clethra acuminata), black huckleberry (Gaylussacia baccata), bear huckleberry (G. ursina), wild grape (Vitis spp.), and male blueberry (Lyonia ligustrina). Mean shrub cover in the Great Smoky Mountains amounted to 65 percent in Table Mountain pine stands and 84 percent in Table Mountain pine-pitch pine stands.

Bear oak (Quercus ilicifolia), mapleleaf viburnum (Viburnum acerifolium), and low sweet blueberry (Vaccinium angustifolium) are most important stand components only in the northern part of the range of Table Mountain pine. Among tree species, black oak is important only in the north; blackjack oak and sourwood, in the south. Black birch (Betula lenta) is an associate only on rocky, nonshale areas, but Virginia pine is an associate at lower elevations, especially on shale. The evergreens, trailing arbutus (Epigaea repens) and galax (Galax aphylla), are important only south of Virginia. Understory species in Table Mountain pine stands stabilize the shallow soil surrounding the pines.

Precipitation within the range of Table Mountain pine varies with latitude and elevation. Warm season rainfall from April through September varies annually from 460 to 610 mm (18 to 24 in) in Pennsylvania to 690 to 990 mm (27 to 39 in) in the Southern Appalachians (27). Mean annual precipitation ranges from 760 mm (30 in) in Pennsylvania to more than 2030 mm (80 in) in the Great Smoky Mountains. At the northern end of the range, average July temperature varies from 21° to 27° C (70° to 80° F); in the Great Smoky Mountains, the range is 15° to 21° C (60° to 70° F). Average January temperatures in Pennsylvania range from -7° to -1° C (20° to 30° F); in the Great Smoky Mountains, 2° to 5° C (35° to 40° F). In Pennsylvania, the average number of frost-free days varies from 150 to 170, and in the mountains of Tennessee, North Carolina, and north Georgia, from 170 to 180.

Weather-related factors such as high gusty winds, glaze, heavy wet snowfall, cloudbursts, tornadoes, and lightning have been known to damage stands and isolated trees of Table Mountain pine (7,15,17,31). The most serious diseases of Table Mountain pine are Phaeolus schweinitzii, which causes butt and root rot, and Phellinus pini, which causes heartrot in older or damaged trees (18). Dioryctria yatesi, a cone-boring insect, can in some years destroy entire local seed crops. Periodically, the southern pine beetle, Dendroctonus frontalis, decimates entire stands (3). The European pine sawfly, Neodiprion sertifer, at times defoliates trees of their previous year's needles, but seldom kills the trees. Trees of all sizes, from 3-year-old seedlings to mature specimens, are attacked by the pine twig gall scale, Matsucoccus gallicola, which causes bark to swell and crack, killing foliage and tree.

Table Mountain pine is monoecious. Cones are commonly seen on trees of sapling size and minimum seed-bearing age is 5 years. In northwestern North Carolina, pollen release at 457 m (1,500 ft) elevation begins the last week of March and ends during the first week of April; at 762 m (2,500 ft), pollen release begins about the second week in April and ceases near the end of the third week. Growth and reproductive activities of Table Mountain pine generally occur as early as, or earlier than, those of associated species (31). Table Mountain pine is reproductively isolated from other pine associates by early pollen release, so hybridization is restricted.

The staminate cones of Table Mountain pine are a reddish purple (31). The pollen is very large for eastern pines, being 50.2 ± 4.6 µ in inside diameter. The cone is heavy and egg shaped; the scales are much thicker at the ends and are armed with stout, hooked spines. Young ovulate strobili have a peduncle about 1 cm (0.4 in) long which is visible at maturity; as branch diameter increases, cones appear sessile (32). From two to seven cones are often arranged in whorls on branches, around the stems of saplings, or on leaders (21,25). Cones average 72 mm (2.8 in) in length, ranging from 42 to 103 mm (1.7 to 4.1 in); 54 mm (2.1 in) in width, ranging from 33 to 75 mm (1.3 to 3.0 in); and 64 cm³ (3.9 in³) in volume, ranging from 27 to 134 cm³ (1.6 to 8.2 in³). Cone dimensions and degree of serotiny decrease with increased elevation. Cones are largest at northern latitudes. In general, cones at higher elevations are well developed. Immature cones are deep green to brown, ripe cones are lustrous light brown, and old cones retained on branches are gray. Cones ripen in autumn of the second season; cone opening depends upon the degree of serotiny.

Population Differences The length-width ratio of Table Mountain pine needles is higher south of latitude 37° N. than north (31). Stands in which most cones open the first and second year after ripening are in the northern end of its range. Seed weight is negatively correlated with elevation, but seedcoat characteristics and cotyledon number are not. There is no difference in specific gravity (0.470 unextracted cores) of Table Mountain pine over its natural geographic range (12). In the Appalachian Mountains of southwest Virginia, in three stands of Table Mountain pine, as far as 42 km (26 mi) apart, there were significant differences in needle length, number of stomatal rows, and needle margin serrulations among trees and half-sib progeny within each stand (14). In seedlings grown from seed from trees in the three stands, no significant differences among stands were observed for any of the characteristics measured.

Races and Hybrids Racial variation, if any, among Table Mountain pines is not known.

Two hybrids of Table Mountain pine are known Pinus pungens x P. echinata and P. pungens x P. rigida. In the former, the needles are long and there are two or three as in shortleaf pine, rather than the two, mostly intermediate in width, and twisted as in Table Mountain pine (12). Needles of Table Mountain pine are stiff and prickly, while those of shortleaf pine are flexible; the intermediate needles of the hybrid are stiff and short pointed, but not prickly. The cones are small as in shortleaf pine, but intermediate, and more like Table Mountain pine in keel of apophysis and spine length. A P. pungens x P. rigida natural hybrid from Pisgah Ridge in Henderson County, NC, had external needle characteristics of Table Mountain pine, a cone intermediate in size and development of apophysis and umbo, and a dense group of short, small-diameter branches about 4.6 m (15 ft) up the bole, suggesting the sprouting characteristics of pitch pine (31).

Stand and stocking values for Table Mountain pine are shown in table 1 by diameter classes. The table shows that trees in the largest d.b.h. classes are in Virginia and North Carolina; this peculiarity of distribution of large trees may be due to differences in the time of stand establishment caused by fire or insect kills.

Table 1- Inventory of Table Mountain pine growing stock combined for Virginia, North Carolina, Tennessee, and Georgia (20)
D.b.h. class Number of trees Merchantable stem volume in thousands¹ Sawlog volume in thousands¹ thousand m³ ft³ m³ fbm 15 cm or 6 in 5,633 606 21,399 - - 20 cm or 8 in 4,315 913 32,256 - - 25 cm or 10 in 2,201 994 35,140 550   96,509 30 cm or 12 in 1,379 858 30,320 588 103,110 36 cm or 14 in    882 660 23,308 557   97,648 41 cm or 16 in    257 242   8,557 226   39,623 46 cm or 18 in      77 104   3,687 113   19,829 51 cm or 20 in      23   36   1,268   45     7,870   (Virginia only) 53 to 71 cm or   21 to 28 in      22   29   1,016   36     6,400   (N. Carolina only) Total 14,789 ³4442 ³156,951 2115 370,989 ¹Includes West Virginia.
²International 0.25-inch log rule.
³An additional 176.8 thousand m³ (6,243 thousand ft³) is found in Pennsylvania. In the southeastern mountains, Virginia has the largest area occupied by Table Mountain pine; the species accounts for 3.4 percent of all trees on the Jefferson National Forest in Virginia (15). Georgia has the least acreage, but average volumes per unit area increase southward from Virginia, with 165.8 m³/ha (2,368 ft³/acre), to Georgia with 282.0 m³/ha (4,028 ft³/acre), probably because of more growing season precipitation, higher temperatures, and longer frost-free periods in stands growing farther south. Because of the large area and volume of Table Mountain pine in Virginia, this State grows and cuts more than other States; the species is virtually nonexistent as a viable timber type in South Carolina mountains and is at a very low volume in Georgia (20).

An overview of the natural range of Table Mountain pine indicates that it is not reproducing itself well and that regeneration is extremely scattered in patches in openings. Seedlings grow well, forming a dense sapling stand that attains its maximum growth rate at about age 30 when the trees are about 18 cm (7 in) d.b.h., but thereafter growth is exceedingly slow. Most of the present stands range from 60 to 100 years old; some originated after the severe beetle kills of the late 1800's and the fires that followed (4,7,15,16).

The keys to successful natural regeneration of Table Mountain pine are heat and light (21). It is classed as intolerant of shade. Even-age management precipitated by clearcutting, or in some cases, a seed-tree cut followed by a hot, fast-moving, prescribed fire to open cones without damaging enclosed seed, should suffice to regenerate it on harsh sites. A severe fire appears necessary for regeneration on sites with a heavy shrub layer (31). It has least competition from associated species on shallow soils, erosion pavements, and rocky areas, and it is on such sites that its management can be most successful. On deeper soils, pitch and Virginia pine should be favored; where considerable microclimatic and soil variation occurs, a mixture of all three species would be best adapted to fully utilizing the site and offering protection against fire and other damaging agents. In remote areas where its perpetuation is desirable, small block or strip cuts on rocky southwesterly exposures with natural seedfall should suffice to regenerate it (7,15,21,31).

Table Mountain pine seedlings are generally anchored into a rock crevice by their taproot (7,15). Secondary or lateral roots then spread through the available soil and litter cover, taking up both moisture and nutrients. Other sinker roots descend into additional crevices, utilizing accumulated soil and the thin skins of finely weathered, nutrient-rich, moist soil-like rock which coat the crevice surfaces. In this way the trees can survive on the most xeric sites.

Another adaptation to xeric sites shown by isolated or sparse trees is the production of extremely long branches--even longer than the height of the tree, where the lowermost ones sweep downward, often touching the ground (31). These branches shield the underlying soil, rock, and roots from direct radiation and conserve soil moisture. Under such trees there is more protection against soil creep and the litter cover is relatively stable, with slow decomposition that contributes to available nutrient supplies and permits survival and growth under harshest conditions. Table Mountain and pitch pines are comparable in windfirmness; both surpass Virginia pine in this characteristic (7).

The seeds of Table Mountain pine are more or less triangular (32). They average 5.3 mm (0.2 in) long, 3.1 mm (0. 12 in) wide, and 13.8 mg (894 grains) in weight. Wing length varies from 19 to 25 mm (0.7 to 1.0 in) and wings are about 7 mm (0.3 in) wide; they range from a transparent light tan to dark brown. Seed coat roughness varies from smooth to a wart-like surface; the seed coat may be ridged. Cones averaging 58 mm (2.3 in) long and weighing 45 g (1.6 oz) produce about 50 seeds per cone, with an average viability of 81 percent. A bushel of cones contains about 0. 18 kg (0. 4 lb) of seeds. Cleaned seeds average 75,240/kg (34,200/lb) with a range of 67,540 to 83,600/kg (30,700 to 38,000/lb). A temperature range of 16° to 32° C (60° to 90° F) for 30 days is required to extract seed from cones (26); kiln drying expedites the process. Seed weight without coat is about 7.6 mg (0.0003 oz); these large seeds may have adaptive value in drier regions, giving Table Mountain pine an advantage in establishing seedlings (31). Frosts, drought, and heavy rains exert a greater influence on flower initiation, cone growth, seed development, and viability than tree age (21). Although cones shed their seeds very irregularly, large numbers of seeds are disseminated annually.

Table Mountain pine seed kept at 24° C (75° F) for 8 hours daylight and at 5° C (40° F) for 16 hours darkness in sand and peat has shown 65 percent germinative capacity with no pretreatment stratification (26). Germination is epigeal. There can be four to nine cotyledons per seedling (32). At 0.5 and 1.5 years, northern sources have a higher rate of water loss at high soil moisture contents than southern sources. Rate of water loss from seedlings with primary needles is greater than from seedlings with secondary needles (31). Growth of seedlings can be increased by adding calcium and magnesium as soil amendments; other fertilizers have not been tried.

Of the three dry-site pines, Table Mountain pine roots are most affected by temperature; it grows a longer root than Virginia pine only between 16° to 28° C (61° to 82° F); pitch pine seedlings have the shortest roots at all temperatures (31). Small top size to root length during seedling establishment would probably be the optimal condition for dry-site survival. Table Mountain pine grows more slowly without mycorrhizae, the root-shoot ratio of non-mycorrhizal seedlings is larger than that of mycorrhizal seedlings. Number of mycorrhizal types was higher in limed than unlimed soil, but liming caused no change in numbers of mycorrhizae (31).

Competition from other vascular plants is probably an important factor in determining the distribution of the species, especially during establishment. Table Mountain pine occurs on areas least favorable for rapid plant growth; these harsher sites seemingly limit its early growth less than that of other species, thus allowing it to develop fast enough to compete successfully for canopy space, minerals, and soil moisture. The roots of very young seedlings are longer than that of pitch pine, and 0.5-year-old seedlings have lower transpiration rates (31). Seedlings of Table Mountain pine develop mature needle fascicles earlier than Virginia pine seedlings.

Another advantage shown by this species is that seedling height growth and twig and needle elongation occur earlier than in pitch and Virginia pine during the period of least drought stress. The ultimate size of Table Mountain pine on xeric sites is constrained less than that of other tree species, allowing the pine to continue to compete, which it could not do on more fertile sites where hardwoods dominate. Growing conditions associated with poor oak growth characterize Table Mountain pine habitat.

Table Mountain pine is regarded as intolerant of shade, even more so than pitch and Virginia pine. The shrub stratum presents greatest competition for seedlings (16), and generally, no seedlings of Table Mountain pine occur on sites without exposed mineral soil. Without fire and anthropic disturbances, Table Mountain pine would probably be found only on extremely dry and sterile rock outcrops and steep shale slopes, where the overstory canopy would be quite open and litter cover would never remain complete (31).

In Pennsylvania, Table Mountain pine grows on substrates of upper Silurian and lower Devonian rocks. Elsewhere it is known to grow on substrates of the Unicoi, Alligator Back, Loudon, Weverton, Erwin, Pottsville, Chemung, Pocono, Portage, and Hampshire or Catskill Formations (17,31). Table Mountain pine is not found over limestone substrates. Although surface rock varies in Table Mountain pine stands in the Great Smoky Mountains, it is often less than 15 percent of total surface cover; however, on narrow ridges, surface rock can range from 35 to 55 percent or more (16). In about half the stands bedrock occurs less than 50 cm (20 in) below the soil surface. Large amounts of gravel and rocks up to 30 cm (12 in) or more commonly occur in the solum; stone content has been found to range from 24 to 59 percent of air-dry weight (31).

Some eleven soil series and rough stony land are associated with Table Mountain pine stands (16,28,31). The most common soil series are Ashe, DeKalb, Ramsey, and Porters. In the Great Smoky Mountains, 57 percent of associated soils are Lithic Dystrochrepts, and 43 percent Typic Dystrochrepts of the order Inceptisols. Generally, A-horizon soils contain an average of 61, 28, and 11 percent sand, silt, and clay, respectively. Most of the B-horizon textures are loams and silt loams; silty clay loam occurs infrequently. Litter depth is about 5 cm (2 in). Organic matter content of the A horizon averages 5.5 percent, and in the B horizon 2.5 percent. Soil pH varies from 4.0 to 4.4 for the A and B horizons, respectively. Water-holding capacity in the solum ranges from 5 to 16 percent. Average nutrient content of a 15-cm (6-in) thick A horizon is as follows: calcium 120 kg/ha (107 lb/acre), magnesium 30 (27), phosphorus 5.8 (5.2), potassium 102 (91), and manganese 40 (36). In the B horizon, nutrient content averages are as follows: calcium 50 kg/ha (45 lb/acre), magnesium 27 (24), phosphorus 7.6 (6.8), potassium 76 (68), and manganese 24 (21). Nitrate and ammonium of the A horizon amount to 2.0 and 19 p/m, respectively; B horizon values are slightly less. Generally, Table Mountain pine stands grow on soils that have minimum profile development and are shallow, stony, strongly acid, excessively drained, infertile, and of low productivity.

Table Mountain pine can grow over a wide range in elevation. Known extremes are 46 m (150 ft) in Delaware and 1762 m (5,780 ft) in the Great Smoky Mountains (16,31); most often it occurs between 305 and 1220 m (1,000 to 4,000 ft) above sea level. In the Great Smoky Mountains, elevation affects the distribution of Table Mountain pine and Table Mountain pine-pitch pine stands (8,16,29). Above 1060 m (3,480 ft) Table Mountain pine-pitch pine stands were mostly restricted to south-facing ridges; with increasing elevation, pitch pine decreased in abundance. Dominance of Table Mountain pine stands occurred at elevations above 1300 m (4,270 ft). On typical sites, the southwest-facing aspect is of critical importance to the presence of Table Mountain pine although the species can and does grow on other aspects (16,17,31). Slopes on which it occurs in the Great Smoky Mountains average 42 percent (16).

Table Mountain pine has been found growing on an island in the Susquehanna River in southern Pennsylvania where the roots were continuously washed by water. It has also been seen around bogs on Mount Pisgah, NC, and at Big Meadows, Shenandoah National Park, VA (31).

Table Mountain pine is used commercially for pulpwood, low-grade sawtimber, and firewood (7,15). The serotinous cones on many trees make seed available for wildlife on a year-round basis. Many of the short stubby limbs seen on Table Mountain pine are caused by squirrels that prune off the cone-bearing section of the limb to get at seed in the heavily armed cone cluster (22). Also, the heavy heath layer in Table Mountain pine stands provides plentiful wildlife food and cover (16,31). The often gnarly Table Mountain pines are a welcome sight on rocky, cliff-like areas of the Appalachians, but perhaps the most important use of the species is as protection forest; it stabilizes soil, minimizing erosion and runoff from the vast shale barrens and other rugged topographic features within its natural range (17).

The prominent epicormic sprouts protruding from the bark of pitch pine are absent in Table Mountain pine; also absent are the dormant buds that occur along the bole and branches of pitch pine-an adaptation which allows recovery from defoliation. Table Mountain pine also has fewer basal buds than pitch pine; basal buds sprout after injury to the stem, allowing recovery of saplings after fire injury and animal damage (31). Table Mountain pine seedlings of natural origin usually have a crook just above or just below ground level which may serve to protect the basal buds against fire.

Table Mountain pine, an Appalachian endemic, grows almost entirely within the range of pitch pine (Pinus rigida) and Virginia pine (P. virginiana), but is less frequent. In general, Table Mountain pine occupies xeric sites of Appalachian rocky and shaly mountainous areas from Georgia into Pennsylvania (8,16,17,29,31). It is frequently found on ridges of the precipitous gorges that dissect the Blue Ridge Mountains.

Table Mountain pine is unevenly distributed; its range extends from the Northern Appalachians in central Pennsylvania southwest to eastern West Virginia and southward into the Southern Appalachians, ending on the steep western edge of the mountains of North Carolina and east Tennessee (1,2,6,9,10,11,19,25,29). To the east and south, its range includes the crest and eastern escarpment of the prominent Blue Ridge Front with its numerous rocky gorges and torrential mountain streams. Toward the southern end of its range, Table Mountain pine reaches its highest elevation of 1760 m (5,780 ft) in the Great Smoky Mountains (8,16,29). There are numerous outlying populations of Table Mountain pine to the east and a few to the west of the Appalachians; many outliers are associated with monadnocks which rise considerably higher than the surrounding Piedmont (5,6,23,24,30).


- The native range of Table Mountain pine.

Pinaceae -- Pine family

Lino Della-Bianca

Table Mountain pine (Pinus pungens) is also called hickory pine, mountain pine, or prickly pine. It is most often small in stature, poor in form, and exceedingly limby. One large tree near Covington, VA, measures 70 cm (28 in) in d.b.h. and 29.6 m (97 ft) tall and has a crown spread of 10.4 m (34 ft). These pines are used locally for fuel and commercially for pulpwood, and they provide valuable watershed protection.

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 > 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 2, 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 obvious prickle, Bracts of seed cone included, Seeds black, Seeds purple, Seeds winged, Seeds unequally winged, Seed wings prominent, Seed wings equal to or broader than body.

Pinus pungens (lat. Pinus pungens) - şamkimilər fəsiləsinin şam ağacı cinsinə aid bitki növü.

Pinus pungens ist ein immergrüner Nadelbaum aus der Gattung der Kiefern (Pinus) mit meist 5 bis 7 Zentimeter langen, in Gruppen von zwei oder seltener drei wachsenden Nadeln und 5 bis 10 Zentimeter langen Samenzapfen. Das natürliche Verbreitungsgebiet liegt in den Appalachen im Südosten der Vereinigten Staaten. Sie wird in der Roten Liste der IUCN als nicht gefährdet eingestuft. Die Art ist wirtschaftlich nur wenig bedeutend und wird als Feuerholz und zur Herstellung von Zellstoff genutzt.

Table Mountain pine, Pinus pungens, also called hickory pine, prickly pine, or mountain pine, is a small pine native to the Appalachian Mountains in the United States.

Cônes de pollen et feuillage

Pinus pungens est une espèce de conifères de la famille des Pinaceae.

Skíðafura (fræðiheiti: Pinus pungens) er smávaxin furutegund sem er ættuð frá Appalasíufjöllum í Bandaríkjunum.

Sosna kłująca (Pinus pungens Lamb.) – gatunek drzewa z rodziny sosnowatych. Sosna kłująca występuje w stanie dzikim na półkuli północnej, we wschodnich regionach Ameryki Północnej - USA, Appalachy od Georgii do Pensylwanii. W Polsce rzadko spotykana, przeważnie w arboretach lub ogrodach botanicznych.

Pinus pungens é uma espécie de pinheiro originária do Novo Mundo. Faz parte do grupo de espécies de pinheiros com área de distribuição no Canadá e Estados Unidos da América (com excepção das àreas adjacentes à fronteira com o México).

Поширення

Поширення: Сполучені Штати Америки (Джорджія, Меріленд, Північна Кароліна, Пенсильванія, Південна Кароліна, Теннессі, Вірджинія, Західна Вірджинія). Ця сосна віддає перевагу сухим умови і зустрічається в основному на кам'янистих схилах на висотах 300-1760 м над рівнем моря. Зазвичай росте у вигляді окремих, розкиданих дерев або невеликих гаїв.

Опис

Щільність деревини: 0,49 гр/см³. Колір квітки жовтий; колір листя зелений; насіння коричневе. Це дерева 6-12 м м висотою. Стовбур до 0,6 м діаметром. Кора від червоно- до сіро-коричневого кольору, в лускатих пластинах. Гілки горизонтальні, оранжево-жовто-коричневі, при старінні темно-коричневі, грубі. Голки зібрані в пучки по 2-3, вони 4-7 см довжиною. Пилкові шишечки яйцеподібні, щоб циліндричні, червоно-коричневі, 0,6-0,9 см, смолисті. Шишки яйцеподібні, від блідо-рожевого до жовтувато-бурого кольору 4-9 см довжиною.

Найбільше дерево діаметром 78 см, висота 29 м, крона діаметром 14 м. Найбільший вік 232 років, Pederson (2006).

Використання

Ці сосни використовують локально як паливо і на комерційній основі для деревини і це цінний захист вододілу.

Загрози та охорона

Періодично південний сосновий жук, Dendroctonus frontalis проріджує цілі популяції цієї сосни. Цей вид зустрічається в кількох охоронних територіях в своєму діапазоні поширення.

Посилання

• The Gymnosperm Database

Це незавершена стаття про родину Соснові.
Ви можете допомогти проекту, виправивши або дописавши її.

Pinus pungens là một loài thực vật hạt trần trong họ Thông. Loài này được Lamb. miêu tả khoa học đầu tiên năm 1806.

Международное научное название

Pinus pungens Lamb., 1805

Охранный статус
Систематика
на Викивидах
Изображения
на Викискладе ITIS 183369NCBI 164241EOL 1033068GRIN t:400599IPNI 927546-1TPL kew-2562461

Сосна колючая^[1] (лат. Pinus pungens) — североамериканский вид растений рода Сосна (Pinus) семейства Сосновые (Pinaceae).

Ботаническое описание

Сосна колючая — сравнительно небольшое дерево до 12 м в высоту, ствол которого достигает 0,6 м в диаметре, с округлой или неправильной кроной. Кора красно- или серо-коричневая, чешуйчато-бороздчатая. Молодые ветки рыже- или жёлто-коричневые, затем темнеющие до красно-коричневых.

Почки красно-коричневые, покрытые смолой, цилиндрические или яйцевидные, менее 1 см.

Хвоя сохраняющаяся на протяжении 3 лет. Хвоинки собранные в пучки по 2 или 3, до 8 см длиной, прямые, различных оттенков жёлто-зелёного цвета; края хвоинок зазубренные.

Мужские стробилы цилиндрической формы, около 1,5 см длиной, жёлтого цвета. Женские стробилы двулетние, изогнутые, асимметричные, незрелые — узко-яйцевидные, затем раскрывающиеся и становящиеся широко-яйцевидными, серо- или красно-коричневого цвета, 4—10 см длиной. Чешуйки жёсткие, с заострённой верхушкой.

Семена обратнояйцевидной формы, около 6 мм, фиолетово-коричневые, с крылом до 3 см.

Число хромосом — 2n = 24.

Ареал

Сосна колючая встречается только в Аппалачах, на высоте от 500 до 1350 м над уровнем моря. Северная граница ареала — центральная Пенсильвания, южная — Северная Каролина и север Джорджии.

Примечания

Сосна колючая (лат. Pinus pungens) — североамериканский вид растений рода Сосна (Pinus) семейства Сосновые (Pinaceae).